FL'IAL REPORT (4 December 1997)

PROJECT REPORT FOR CONSTJLTING CONTRACT

Title: The Risk ofNonindigenous Species Invasion in Prince William Sound Associated with Oil Tanker Traffic and Ballast Water Management: Pilot Study.

RFP Number: 632.97.1

PRESENTED TO:

Regional Citizens' Advisory Council of Prince William Sound P.O. Box 3089 (154 Fairbanks Drive) Valdez, Alaska 99686 USA

Telephone: 907-835-5957 Fax: 907-835-5926 Email: [email protected]

PRESENTED BY:

Principal Investigators: Dr. Gregory M. Ruiz, Ph.D., Marine Ecologist Dr. Anson H. Hines, Ph. D., Marine Ecologist

Mail Address: Smithsonian Environmental Research Center Smithsonian Institution P.O. Box 28 (647 Contees Wharf Road) Edgewater, iv!D 21037-0028 USA

Telephone: 301 261-4190 ext. 227 (Ruiz), 208 (Hines) FAX: 301 261-7954 Email: [email protected], [email protected]

The opinions expressed in this RCAC commissioned report are not necessarily those of RCA C. Synopsis

Although nonindigenous species are common in marine environments, and some cause significant environmental and economic impacts throughout the world, there is very little information available for the frequency or impact of invasions by nonindigenous species at high latitudes. This Pilot Study was therefore conducted over a one-year period as an initial step in defining the problem and potential risks in Port Valdez and Prince William Sound. The study briefly summarized the current state of knowledge about nonindigenous species and risk of invasions that are relevant to Prince William Sound. Although limited in scope, the study also examined the transfer of organisms into Prince William Sound that arrive in ballast water of oil tankers. This initial analysis indicates that risk of invasion exists for Prince William Sound, in that large quantities of diverse plankton is transported during spring in ballast water of tankers transiting from west coast ports. Although tankers arriving from foreign ports exchange their ballast water in open ocean, some residual foreign plankton remains in this diluted ballast water; but numbers of tankers arriving from overseas remain low at present. The available information is currently inadequate to assess the magnitude of risk for ballast-mediated invasions, because seasonal and annual variability of plankton in the ballast water has not been measured for tankers. Moreover, the potential for survival and establishment of these organisms has not been studied. Over the next two years, we will collect the necessary information for a detailed analysis of the risks, mechanisms, and patterns of species introductions for Prince William Sound. The Pilot Project and on-going research represent a cooperative and successful partnership of industry, citizen, agency, and scientific groups.

ii Executive Summary

Project Overview

This Pilot Study begins to assess the risk of nonindigenous species ~iS) invasion associated with oil tanker traffic and ballast water management for Port Valdez I Prince William Sound, Alaska. This study included four major components: • Review and analysis of existing knowledge of invasions and ship-mediated transfer of species relevant to Prince William Sound. • Analysis of plankton communities associated with segregated ballast water on tankers that arrived to Prince William Sound. • Experimental measurements of the effect of ballast water exchange and voyage duration on plankton communities arriving on tankers to Prince William Sound. • Characterization of plankton communities in non-segregated ballast water passing from oil tankers through the Alyeska shore-side ballast water treatment facility in Port Valdez.

Although this is a preliminary analysis of risk, the Pilot Stody advances our understanding of invasion processes in many significant ways. • Our study provides the most detailed and thorough analysis to date of plankton communities in segregated ballast water of tankers. • This is the "first-ever" analysis of plankton in non-segregated ballast water, and especially at different stages of the Alyeska (or any other) ballast water treatment process. • We report the first experimental and quantitative measurements of (a) the effectiveness of ballast water exchange to reduce unwanted organisms and (b) the effect of voyage duration on plankton survivorship in ballast tanks of oil tankers. • We provide a summary ofl'HS known from Alaska to California that begins to examine latitudinal patterns of invasions, including invasions to 60' , for the first time.

The scope of this initial stody "ill be expanded over the next two years to provide a more complete and detailed analysis of the risks, mechanisms, and patterns of invasion in Prince William Sound. We have now developed a strong cooperative program to address critical gaps in our understanding of invasion risks as well as facilitate information exchange and participation among a broad spectrum of industry, citizen, agency, and scientific groups. • From a science perspective, this program "ill result in a comprehensive analysis of invasion processes and risks for Prince William Sound, representing the first such study in the world for a high-latitude I cold-water marine ecosystem. • From an industry and management perspective. this program will assess the effectiveness and trade­ offs involved for various management strategies that are now required in Prince William Sound, and are being promoted on a national and international scale. • From a public perspective, this program will disseminate findings and serve as a key source of information, especially through groups like Alaska Sea Grant and the Regional Citizens' Advisory Council of Prince William Sound.

iii Results

Biological invasions of coastal bays and estuaries are common throughout the world and are having significant ecological and economic impacts. • Within the lower 48 states of the U.S., 70-212 NIS per estuary are known for sites that have been explicitly surveyed for invasions, and more than 400 marine NIS are documented for these contiguous states. • Reports of marine NIS for other global regions range from tens-to-hundreds, indicating the widespread nature of this phenomenon. • These reports actually underestimate the full extent of invasions, because many NIS are not documented. • Some NIS can and do significantly impact community structure and function, productivity, commercial fisheries, and human health.

Transport of coastal planktonic organisms in ballast water of commercial ships appears to be the major source of new invasions worldwide in recent years. • Large bulk carriers and tankers can carry individually 30,000 to 60,000 metric tons of segregated (i.e., non-oily) ballast water from one port to another in voyages lasting days to weeks. • This water often contains a rich diversity of planktonic organisms that inoculate recipient ports upon ballast water discharge. • Single ports can receive> I 0,000.000 metric tons of segregated ballast water annually.

High-latitude I cold-water regions are also subject to biological invasions by many species with potential ecological and economic consequences. • Over 100 marine NIS are known from high-latitude regions (40'-60' latitude) around the world, despite the limited effort devoted to documentation. • These invasions have a range of impacts similar to those reported for more temperate latitudes.

A preliminary literature review and field survey for Alaska and Prince William Sound, respectively, identified 10 known :"<1S and many cryptogenic species (i.e., possible NIS), indicating that invasions have occurred and very limited information currently exists to evaluate l'<1S in Alaska.

SLx major risk factors exist in Prince William Sound that may favor successful invasions by NIS. • 600 tankers currently arrive per year to Prince William Sound and release an estimated 20,000.000 metric tons of segregated ballast water. • The voyage duration of these mnkers is usually short (3-7 days), favoring high survivorship of transported plankton. • A pattern of repeated delivery of ballast water from the same, limited donor locations provides repeated inoculations of the same NIS. • Environmental conditions of source pons match those in Prince William Sound for some portions of the year. • A lack of mid-ocean exchange by most tankers of domestic origin improves transfer rate ofNIS • A large number ofNIS are kno\\11 from both domestic and foreign source ports of ballast water arriving in Prince William Sound.

iv A large quantity of taxonomically diverse plankton is released in Prince William Sound with segregated ballast water from oil tankers from domestic ports. • During a period of high plankton productivity in May and June, segregated ballast water from domestic ports contained an average density of 7,000 large (>80 micron) planktonic organisms/m3 across 13 different ships. • This is equivalent to an average of 244,000,000 planktonic organisms per ship. • We have identified a minimum of 69 different taxonomic groups in this ballast water, and a minimum average of 19 species/ship. • Most of the 600 tankers that arrive annually to Prince William Sound come from domestic ports. l'oiS are present in segregated ballast water released in Prince William Sound. • We have identified 4 NIS arriving in ballast water from domestic ports, and many others are certainly arriving from these locations. • The 4 NIS identified to date are all copepods from Asian waters which are now abundant in San Francisco Bay. • Some coastal organisms are also arriving from foreign ports, despite ballast water exchange; these may include additional species that are nonindigenous to Alaska.

Ballast water exchange is effective at reducing the abundance of coastal organisms that arrive to Prince William Sound in oil tankers. • Fewer coastal organisms appear to be present on tankers arriving from overseas which have undergone ballast water exchange. • Over 90% of coastal plankton was removed in ballast water exchange experiments on oil tan.l<:ers.

The abundance of some coastal organisms in ballast water declined with voyage duration for short­ term voyages, but this relationship was not as strong as that observed in other studies. • The densities of annelids and molluscs declined with voyage duration (3-7 days) among tankers arriving to Port Valdez from domestic ports. • However, the total number of organisms showed no relationship with voyage duration for these same ships, differing from the negative relationship observed for longer (14-20 day), trans-Atlantic voyages of bulk grain and coal ships. • Also, densities did not decline for most taxonomic groups over three consecutive days of sampling the same ballast water within a single ship.

A large quantity of non-segregated (or oily) ballast water arrives to Port Valdez in oil tankers. • An average of 29.909 metric tons of non-segregated ballast water is delivered to Port Valdez per tanker. compnsing 49% of the total ballast water per ship.

The density and diversity of organisms present in non-segregated ballast water as it arrives and travels through the Alyeska Ballast Water Treatment Facility is extremely low. • Nearly all organisms present in the first 3 stages of the ballast water treatment fac:lity (Chicks an Arms, 90s Tanks, and DAF Tanks) are dead. • Only at the last stage (BT Tanks) were live organisms regularly detected, and these consisted primarily of nematodes and diatoms. • It is unclear whether these nematodes and diatoms derived from ballast water or wind-borne dispersal from the local environment.

v Conclusions

Prince William Sound is at some risk of invasion by NIS that arrive in segregated ballast water of oil tankers from domestic ports. • Large, frequent. and dense inoculations of plankton in ballast water from oil tankers occur in Prince William Sound. • This ballast water, and its plankton. derive from ports that include tens-to-hundreds of species nonindigenous to the eastern Pacific ocean. • Some of the known NIS in Alaska also occur in San Francisco Bay, demonstranng that emironmental conditions are not always a barrier to invasion by some species in current domestic source ports of ballast water. • Surveys of Prince William Sound are not now available to assess the rate of invasions in many key habitats that are likely to be invaded by these NIS.

Ballast water exchange appears effective at reducing resident plankton on tankers from overseas, although a risk of invasion still exists. • Preliminary data indicate >90% reduction of coastal plankton. • Yet, 300,000 or more organisms/ship can remain from the original source port following ballast water exchange.

Despite the large volume of non-segregated ballast water delivered to Port Valdez, we surmise the risk of NIS invasion associated with oily ballast water is extremely small. • Few to no live organisms are present in water entering the ballast water treatment facility from each oil tanker. • The nematodes and diatoms present at the last stage may well be oflocal (Port Valdez) origin. • Even if these organisms derive from ballast, the cumulative abundance and diversity of this community is effectively zero in comparison to that arriving in segregated ballast water of tankers.

Further investigation is needed to assess adequately the risk of invasion for Prince William Sound, and this effort should concentrate on segregated ballast water and existing evidence for invasion. • Analysis of segregated ballast water from more ships is needed to sufficiently characterize associated plankton density and diversity by port, season, and voyage duration. • Laboratory experiments should measure the viability of these organisms upon arrival and their tolerance of local environmental conditions. • Shipboard experiments should measure the survivorship and transfer of organisms in segregated ballast water during operation under various environmental conditions and ballast water management practices. • Intensive sur:eys of local biota should be focused on key taxonomic groups and habnats to measure the occurrence ofNIS invasions in Port Valdez and Prince William Sound.

vi Acknowledgments

This project was funded by the Regional Citizens Advisory Council (RCAC) of Prince William Sound, and included contributed support from the U.S. Coast Guard and U.S. Fish & Wildlife Service. The project was stimulated and encouraged by the members of the Non-Indigenous Species Working Group ofRCAC. We thank Mr. Joel Kopp ofRCAC for his able oversight and friendly assistance at all stages of this project. Lt. Larry Greene of the U.S. Coast Guard and Mr. Gary Sonnevil of U.S. Fish & Wildlife Service also helped in many ways. We are grateful to Mr. Rex Brown and the managers and staff of the Alyeska Pipeline Service Company, the Valdez Marine Terminal, and especially the staff of the Ballast Water Treatment Facility for access and help in sampling. Quanterra Corporation generously allowed our use of their Water Chemistry Laboratory at the Marine Terminal; Richard Nenahlo, Kelly Hawkin, and Satch Tapanco of Quanterra were especially helpful. The Masters, Officers, and crew members of the tankers gave willingly of their rime and help; and we especially thank the following agents for their logistical assistance, advice and suggestions. and support: Kurt Hallier and Wayne Brandenburger of ARCO Marine, Phil Eichenberger and Bill Deppe of SeaRiver, Steve Herring and Tom Colby of British Petroleum, and Carla Hilgendorf of Alaska Maritime. The full cooperation of the oil companies operating these ships is gratefully acknowledged. Scott Godwin of the Smithsonian Environmental Research Center (SERC) coordinated all of the field work and sample sorting. Assistance in sample sorting was given by Kimberly Philips and Matt Nicklin at SERC. Field assistance was provided by John Chapman of Hatfield Marine Science Center, Oregon State University. Joe Bridgman ofRCAC provided assistance and use ofhis boat for field collections. John Chapman and Gayle Hansen of Oregon State University provided the information for the section on invertebrates and algae, respectively, which were collected in Port Valdez. John Chapman and Gayle Hansen also pro,ided information on introduced species in Oregon and Alaska. We obtained additional assistance with identifications of invertebrates and algae collected from Port Valdez: Jeff Cordell (University of Washington), Leslie Hanis (Los Angeles County Museum), Faith Cole (U.S. EPA), and James Carlton (Williams College). Identification of plankton species in ballast water samples was provided by: Frank Ferrari and Chad Walter for copepods: Oithona davisae, Limnoithona senensis, Pseudodiaptomous forbesi, and P. marimlS. Report preparation, data analysis, and preparation of figures and tables were assisted by Lynn Kurzava, Linda McCann, and Laura Rodriguez at SERC. Helpful comments on an early draft of the report were provided by: Joel Kopp, Joe Bridgman, Peter Armato, John .Armstrong, Howard Feder, Tom Colby, Kurt Hallier, and Bill Deppe.

Sections of this report on "Known NIS in Prince William Sound and Alaskan Waters", "Field Surveys ofNIS in Port Valdez", and "Non-Indigenous Marine/Estuarine Species \vith Potential for Introduction to Prince William Sound" were based upon a subcontracted report provided by John W. Chapman (Department of Fisheries and Wildlife, Hatfield Marine Science Center. Oregon State University, Newport, OR 97365-5296, USA, Ph. 541 867-0235/FA.,'X 541 867- 0105, CHAPMA..'J.JOHN@ EPA.t\1AIL.EPA.GOV) and Gayle I. Hansen (Department of Botany and Plant Pathology, Hatfield Marine Science Center. Oregon State University, Newport, OR 973565-5296. USA, Ph. 541 867-0200/FA.. X 541 867-0105, [email protected]).

The cooperative support by all of these individuals made this project possible. SERC Invasions Biology Program Contribution No. 11.

vii TABLE OF CO~TE:'."TS

Synopsis ...... ii Executive Summary ...... iii Acknowledgments ...... vii

Table of Contents ...... viii

A. Introduction ......

Overview Objectives of the Pilot Study Approach to Project

B. Existing knowledge ofNIS relevant to Prince William Sound ...... 3

Purpose Methods Results Information Exchange & Consultation ...... 5 State of Knowledge ofNIS in Marine/Estuarine Ecosystems Outside of Prince William Sound ...... 5 i'HS in Marine Ecosystems at High Latitude ...... 8 NIS in Marine/Estuarine Ecosystems in Prince William Sound and Alaska Hypotheses About Why NIS Are Not Evident at High Latitude Factors Adding Risk for NIS Invasions in Prince William Sound Kno"TI N1S in Prince William Sound and Alaskan Waters...... 11 Purpose Methods Results Submerged Aquatic Angiosperm Plants Macro-Algae Invertebrates Fish Unsuccessful Invertebrate Introductions into Alaska Unsuccessful Macro-Algae Introductions into Alaska Field Surveys of N1S in Port Valdez ...... 16 Purpose Methods Results Invertebrates Macro-Algae Discussion Non-Indigenous Marine/Estuarine Species with Potential for Introduction to Prince William Sound ...... 19 Purpose Methods Results High Latitude Invertebrate Introductions Outside of Alaska Invertebrate Introductions from Alaska

viii High Latitude Macro-Algae Introductions Macro-Algae Introductions from Alaska High Latitude Micro-Algae Introductions Discussion

C. Plankton in Segregated Ballast Water ...... 21

Purpose Methods Results Discussion

D. Experimental Analysis of Effects of Ballast Water Management Practices ...... 27

Ballast Water Exchange Experiments

Purpose Methods Results Discussion

Duration of Time in Ballast Tanks

Purpose Methods Results Discussion

E. Plankton in Non-Segregated Ballast Water Passing Through the Alyeska Ballast VVater Treatment Facility...... 30

Purpose Methods Results Discussion

F. References ...... 34

ix A. Introduction

Overview

Aquatic nuisance species have invaded many, perhaps most, freshwater and marine ports around the world; and ballast water from commercial shipping is increasingly recognized as the most significant vector currently for those invasions occurring (Carlton and Geller, 1993). Ballast water consists of water pumped into dedicated tanks or cargo holds/tanks for trim and stability during oceanic voyages, especially when the vessel is empty or only partially full of cargo. Ballast water is usually taken from coastal water containing a rich diversity of planktonic organisms. Ballast water is often discharged into a receiving port prior to loading cargo, inoculating the ecosystem with exotic species. If the plankton is viable and becomes established, these non-indigenous species (NIS) can cause maJor ecological and economic disruption in the coastal ecosystem, with numerous examples in San Francisco Bay (Cohen and Carlton, 1996), the Great Lakes (Mills et al., 1993), Chesapeake Bay (Ruiz et al., unpubl. database), and elsewhere (Ruiz et al., 1997a). In San Francisco Bay, the rate of invasion has increased to about one new NIS invasion every 12 weeks, probably as a result of increased ballast water discharge (Cohen and Carlton, 1996). \Vnether the invasion is Eurasian zebra mussels in the Great Lakes, Asian clams in San Francisco Bay, or ctenophores in the Black Sea, impacts of ballast introductions have been devastating and irreversible. Despite the profound impact of ballast-mediated invasions, the biological characteristics of ballast water and the factors which regulate invasion success are little studied and poorly understood. In the USA, biological characteristics of ballast water have only been studied in two port systems: Coos Bay (Carlton and Geller, 1993) and Chesapeake Bay (Smith et al., 1996; Ruiz et al., unpubl. data); and in other countries the biology of ballast water has similarly received little quantitative analysis (Carlton, 1989; however see Williams et al., 1988; Hallegraeff and Bolsch, 1992).

While the significance ofballast-mediated invasions has recently focused on temperate zone ports, little consideration has been given to NIS invasions at high latitude. despite :he volume of shipping and crittcal importance of certain cold-water ports to the world economy and especially to U.S. energy interests. Port Valdez in Prince William Sound, Alaska, is an especially crucial region at high latitude that ships approximately 20% of U.S. domestic oil production. The terminus of the 800-mile Trans-Alaska Pipeline lies on the south shore of Port Valdez (Fig. A-1). The pipeline and terminal for loading crude oil are operated by Alyeska Pipeline Service Company and receive oil from Alaska's North Slope fields at the rate of approximately 1.35 million barrels per day. This oil is loaded into tankers at the terminal's-+ ber:hs and transported mostly to refineries on the U.S. West Coast. but with occasional deliveries to orher cold-water ports in Korea and Japan and some warm-water pons (Figs. A-2 and A-3). now that Congress has authorized sale of North Slope crude on foreign markets. Tankers have made more than !5,000 ,·oyages through Prince William Sound to Port Valdez since the startup of the terminal in 1977. In recent years, 1987-1994, tanker arrivals to Valdez have averaged 799 per year bur have declined to less than 600 per year currently (Fig. A-4). Tankers arriving to Prince \\iilliam Sound discharge two types of ballast water: (1) Segregated ballast water from tanks dedicated solely to ballast water and (2) non-segregated ballast water from tanks which are used to carry petroleum products. Approximately 20 million metric tons of segregated ballast water are discharged annually by tankers into the port and sound. which is a quantity of domestic ballast water that greatly exceeds the volumes of foreign ballast water released in other U.S. West Coast ports and approaches the volumes of foreign ballast water released into New Orleans/Baton Rouge and Chesapeake Bay (Fig. A-5). All non-segregated. oily water (about 50% of total) discharged by tankers in Port Valdez must pass through the Ballast Water Treatment Facility located on shore at the Valdez Marine Terminal. Effects of the treatment plant on ).lS were unknown prior to this Pilot Study. Segregated ballast water is discharged directly into the sound/port without treatment.

1 In light of the problem of aquatic nuisance species invading various shipping ports around the world via ballast water, the Regional Citizens Advisory Council (RCAC) of Prince William Sound, shipping industry, U.S. Fish & Wildlife Service, U.S. Coast Guard, other state and federal governmental agencies, and other citizens are concerned that Port Valdez and Prince William Sound may also be at risk of such invasions. At the root of this concern is the millions of tons of segregated ballast water discharged by these tankers into the Port and Sound each year, as well as the effluent from Alyeska's Ballast Water Treatment Facility.

Objectives of the Pilot Studv

As stated in the Request For Proposals, this project was designed with the following objectives over a short (< one year) period: • To participate in a Nonindigenous Species Working Group in the Prince William Sound; • To review NIS literature and studies relevant to Prince William Sound, and conduct initial field survey for potential NIS in Port Valdez; • To conduct a biological analysis of tanker ballast water and Ballast Water Treatment Facility effluent; • To assist in the development and conduct of a one-day workshop to discuss future evaluation ofNIS invasion in Prince William Sound; • To inform and educate the RCAC, shipping industry, scientists, state and federal governmental agencies, and other citizens about this issue; • To make recommendations based on findings, and identify specific information needs, as appropriate.

Approach to Project

There is little organized knowledge of marine/estuarine NIS at high latitudes generally or for Alaska specifically. This Pilot Study was therefore conducted over a one year period as an initial step in defining the problem and the risks ofNIS in Prince William Sound. The Pilot Study briefly surveyed major elements of environmental risk for NIS in Prince William Sound, Alaska. and provided qualitative and preliminary quantitative information on the potential for ballast-mediated invasions. The Pilot Study involved 4 major components: 1. Analysis of, and communication about, existing knowledge ofNIS relevant to Prince William Sound, supplemented by an initial field survey of potential NISin Port Valdez; 2. Characterization of plankton communities in segregated ballast water coming into Port Valdez aboard oil tankers during late spring, when is abundant and presumed to be of maximal risk for introductions; 3. Experimental assessment of the effect of ballast water management practices of oil tankers (mid-ocean exchange and duration oftime in ballast tanks) upon plankton communities in segregated ballast water; and 4. Characterization of plankton communities in non-segregated ballast water passing from oil tankers through the Alyeska shore-side ballast water treatment facility. Each of these components is discussed in separate sections below.

Although this Pilot Study was limited in scope, it begins to provide important insights into the issue ofNIS in a high latitude/cold water ecosystem and the role of oil tankers in transporting plankton in ballast water. The high yield of information from this initial effort derived from a special blend of cooperation and professional commitment of the partnership of pri,·ate citizens. the oil and shipping industry, government managers, and scientists working closely together. Individually, several elements of this study provided the first such analyses in the world. This is the first detailed, replicated sampling

2 of plankton communities in oil tankers. It provides the first experimental analysis of the potential for ballast management practices to reduce risk ofNIS transfer in ballast water of tankers. The analysis of the ballast water treannent plant is the first of its kind. And, combined with studies in the eastern L.S., this begins to assess the importance ofNIS transfer among domestic ports of the U.S. Cumulatively. these elements point to the value of cooperation for problem analysis.

Tne Pilot Srudy will be expanded in an extended analysis over a 2-year period. The proposal for the expanded study was written during the Pilot Study and submitted to the National Sea Grant Program with matching contributions from RCAC, U.S. Fish & Wildlife Service, Smithsonian Environmental Research Center, Williams College, and Alyeska Pipeline Service Company. Full funding was granted for the period from Falll997 to 1999. Thus, this Pilot Study forms a foundation upon which we will build a more comple:e and detailed analysis of the risk of invasions in Prince William Sound and the coastal zone of Alaska.

B. Existing knowledge of NIS relevant to Prince William Sound

Purpose

A major goal of the Pilot Study was to review existing information about i'i1S in marine ecosystems and to evaluate its relevance and quality for assessing biological invasions of high latitude/cold water regions, especially in Alaska and Prince William Sound. This review and evaluation was communicated to the constiruencies of the RCAC with a variety of methods ranging from expert advice and informal conversation, telephone conferences, a public Workshop, and elements of this written report. Communication of the review and evaluation served to inform the Nonindigenous Species Working Group of the RCAC, the constituents of the RCAC as a whole, and the general public about the risk of marine invasions and the state of knowledge about ballast water transport as a vector for NIS introductions. Clearly, the focus of the analysis was risk of invasion of Prince William Sound by organisms transported in ballast water of tankers arriving at the Valdez Marine Terminal. Initially, the state of knowledge was judged to be so incomplete and poorly resolved for Prince William Sound that the analysis needed to be placed in a global perspective that conveyed information for regions which have received more intensive analyses .

.\Iethods

To review and evaluate the existing knowledge ofNIS in Prince William Sound, we used a hterarchical approach. (I) We provided a synopsis of the NIS problem in marine/estuarine ecosystems world-wide. which is based primarily on temperate ecosystems. (2) We developed an overview of reported information for NIS in marine/estuarine ecosystems at several high latirude locations around the world, which allowed us to assess whether high latitude/cold water ports generally may be subject to NIS invasions. (3) Vie examined known patterns of invasions by NISin west coast source ports for tankers traveling to Port Valdez. (4) To determine whether the known i'i1S in Alaska provide an accurate indicator of the risk of biological invasions, we considered the factors that contribute to risk of invasion for Prince \Vil!iam Sound. The risk factors include the pattern of ballast water delivery to Port Valdez, and the latirudinal pattern of biological invasions for source ports along the west coast where tankers take on most of the ballast water transported to Port Valdez. (5) We assembled information about known marine/esruarine invasions of Prince William Sound and south-central to southeastern Alaska. (6) To assess whether previous workers may have overlooked NIS established in Prince William Sound, we initiated a preliminary field survey of species in habitats of Port Valdez. (7) To identifY species with a risk of invasion to Port Valdez, we assembled a list ofi'i1S which have invaded high latitude ecosystems and have the potential for ballast water transport. 3 In this initial analysis of the literature, we utilized state-of-the-art computer searches of bibliographic cross-references, and we reviewed our existing extensive reference library on marine/estuarine NIS world-wide. We focused on high latitude and cold water species and regions (including areas of strong upwelling). From our review, we created a taxonomic list of species for high latirude/cold water ecosystems by region. For the known introduced species along the west coast of North America that comprises the main source ports for ballast water in tankers to Port Valdez, we determined the following variables: site of introduction (southern California, San Francisco Bay, northern California, Oregon, Puget Sound area of Washington and British Columbia); native geographic range; mechanism of introduction (highlighting species introduced in ballast water); date of mtroduction; and reference documenting the primary source of information. This information allowed us to assess the abundance and diversity of invasions along the latirudinal gradient. It also allowed us to consider the probability of species invading multiple ports by comparing the similariry of invasive species in locations relative to San Francisco Bay, the ecosystem with the most complete analysis ofi'•HS. By considering the similarity ofNIS at locations both south and north of San Fnancisco Bay, we attempted to gain insight about the probability ofNIS moving along this broad latirudinal gradient to Alaska.

The graded criteria (derived from J.T. Carlton, e.g., Carlton, 1979) used to determine whether each species in our database is introduced, native, or cryptogenic are described below "Cryptogenic species" cannot be identified clearly as native or introduced, and thus have Utiknown origin (Carlton, 1996a). Further discussion of criteria for identifying species as introductions are given in Chapman (1988), Chapman and Carlton (1991) and Eno (1996). Often a single criterion is not sufficient to designate a species as being introduced, but combinations of several factors increase the probability of an accurate reconstruction of introductions and invasions. • Paleontological - NIS are absent from fossil record even though they are presen.t in other locations; native species are found locally as recent fossils; cryptogentic species are not in the local fossil record, but they are not reliably fossilized generally. • Archeological - NIS are absent from shell middens and other archeological deposits; native species are in local deposits; cryptogenic species would not be expected to be found in archeological deposits. • Historical - NIS are not recorded by direct observation at early periods, especially by trained naturalists, but suddenly appear where trained observers did not find them previously; native species are recorded in the earliest observations of trained observers; cryptogenic species are species that were not srudied by early trained observers. • Biogeographic- NIS exhibit grossly disjunct patterns of distribution (we took care to evaluate artifacts of the distribution ofbiologists/taxonomists); nattve species have continuous geographic ranges which include Alaska/Prince William Sound or other high latitudes; cryptogenic species have poorly known distributions or "cosmopolitan" distributions. • Ecological - NIS have habitats in close association with other NIS (co-evolved species; specialized predator-prey, commensal or host-parasite relations); native species are closely associated with other native species; cryptogenic species are more generalized. lacking close. specialized association with other species. • Dispersal Mechanisms- NIS presence cannot be plausibly explained by narural dispersal mechanisms and have documented human-mediated mechanisms which could effect the:r distributions; native species have natural dispersal mechanisms and lack known human-mediated mechanisms of introduction; cryptogenic species have both natural and human-mediated mechanisms of dispersal that could account for their distnoution. • Evolutionarv/Genetic - NIS have isozyme or DNA frequencies which match distant proposed source populations and are significantly different from adjacent natural populations: native species have

4 population genetics which blend with adjacent natural populations; cryptogenic species have not been srudied with molecular techniques.

Results

Information Exchange & Consultation The Pilot Srudy included participation in the RCAC NIS Work Group, providing information, analysis and advice about NISin coastal marine systems worldwide, focusing on the risk of biological invasions of cold water/high latirude coastal ecosystems. Advice and information were conveyed to the Working Group through monthly telephone conference calls from December 1996 through November 1997. Additional consultation was provided to Mr. Joel Kopp, project manager for RCAC, throughout the course of the project.

In February 1997, we also \\Tote a major proposal to extend and expand the scope of the Pilot Srudy in response to the Request For Proposals on NISin U.S. coastal waters from the national Sea Grant Program; subsequently the proposal has been funded for a 2 year period (1997-1999). The extended proposal provides a recommended action plan to follow up the Pilot Study. including: • Detailed analysis of abundance and diversity of plankton organisms in segregated ballast water in tankers (N= approximately 60 ships) arriving to Port Valdez, to estimate variation by season, year, source port and duration of voyage; • Experimental laboratory analysis of effects of temperature-salinity combinations on survivorship for certain plankton arriving to Port Valdez, to test for risk of short-term survival of plankton released with segregated ballast water imo Port Valdez; • Experimental analysis of the efficacy of mid-ocean exchange of segregated ballast water in tankers in route to Port Valdez from west coast ports; • Experimental analysis of plankton survivorship during voyages as determined by comparing samples of segregated ballast water collected upon departure from west coast source ports and arrival to Port Valdez; • Characterization of the fouling communities on the bottom and in the sea chests of tankers, which will be sampled in dry docks during routine maintenance; • Review of potential NIS in existing extensive samples of organisms held in the University of Alaska Museum collections and other voucher collections for the Prince William Sound region; • Field studies of possible C\lS in communities of Prince William Sound. including fouling communities of the Marine Terminal and soft-bottom intertidal areas, which are hypothesized to have elevated risk of invasions and which have not received adequate study in the past; and • Further literature analysis ofNIS in Alaska, with assessment ofbiolog~cal invasions in high latirude/cold water ports. Another major element of the process to inform and educate the Working Group was consultants' lead participation in a public Workshop organized in Anchorage by RCAC during March 1997. Presentations at the March Workshop reviewed the scope of the NIS problem and articulated the potential elements of risk for NlS in Prince William Sound. Video tape (full-length and edited, condensed versions) are available form RCAC in Valdez. A summary of the rationale and in forman on about the risk ofNIS in Prince William Sound is provided below.

State ofKnowledge ofNIS in Jlarine/Estuarine Ecosystems Outside ofPrince William Sound The significance ofNIS in marine environments (including bays, esruaries, and open coasts) has received relatively little attention compared to )ITS in terrestrial and freshwater habitats (Carlton, 1989). The few areas with some historical assessments of invasive species in manne/esruarine ecosystems demonstrate the potential importance off,'IS invasions (Table B-1). The number of known NIS for each of five different U.S. estuaries (all those intensively srudied to date) vary between 70 and 212 spec1es.

5 Combined with various additional published records, we can now identifY approximately 400 individual NIS from coastal estuaries of the continental U.S. (Ruiz et al., unpubl. database). Analyses for other global regions (Table B-1) indicate that NIS invasions are common throughout the world and that NIS often form a significant component of marine communities, where they can have major ecological effects and serious economic impacts. The NIS identified in these analyses, within and outside of the U.S., include a broad range of taxonomic and trophic groups that occupy diverse habitats (e.g., soft-sediment, rocky substrata, marsh surface, and water column). Estuaries and embayments appear to be particularly subject to invasions by N1S, in part because these are the sites of ports and associated shipping activities.

Despite the large number ofNIS identified from some sites and regions, all available studies underestimate the actual extent of invasions for two major reasons: (a) many species simply are not examined; and (b) of those examined, the native versus non-native status often is not clear (Carlton, 1996a). Estimates for the Great Lakes and Hudson River estuary resulted from a !-year study that focused on "accessible" ta.xonomic groups, or those for which historical distribution records and taxonomic identities were well established. In that limited time, however, it was possible to examine status of only a portion of the species present. By contrast, the estimate for San Francisco Bay is very comprehensive, deriving from >20 years of research (e.g., Carlton !979a,b, 1985, 1987, 1992b; Chapman, 1988; Carlton eta!., 1990; Chapman and Carlton, 1991; Cohen and Carlton, 1996). Even though that study includes "less accessible taxa" as well as conspicuous ones, the presence of "cryptogenic taxa" indicates that many NIS go undetected.

Within these and other coastal ecosystems, NIS are often numerically dominant organisms, and they can dramatically alter community function (see Ruiz eta!. 1997 for review). For example, the Asian clam Potamocorbula amurensis achieves densities of>lO,OOO clams/m2 in San Francisco Bay, where it is replacing other benthic organisms and clearing plankton communities from overlying waters (Nichols et al., 1990; Alpine and Cloern, 1992; Kimmerer, 1994; Cloern, 1996). Recently (1989), the European green crab (Carcinus maenas) invaded San Francisco Bay, along with Australia and South Africa, resulting possibly from ballast water (Cohen and Carlton. 1996). On the west coast, this crab has spread rapidly to the south to Monterey and north to at least Coos Bay, Oregon (Grosholz and Ruiz. 1995a,b; N. Richmond, pers. comm.). Our detailed studies of this invasion in Bodega Bay, California show that this voracious predator is drastically reducing populations of native clam and infaunal invertebrates, which also are key food items for native shorebirds, fish and Dungeness crabs (Grosho!z and Ruiz. in review); this crab appears to be having similar effects in Tasmania, Australia (Thresher, !997).

Although poorly understood, many NIS are microbial, parasitic or disease producing; and these can have major adverse impacts on marine ecosystem function and economics. One of the major diseases (Perkinsus marinus) decimating oyster populations in Chesapeake Bay appears to have been introduced via ballast water in the late 1950s in nearby Delaware Bay (Andrews, 1979). A parasitic barnacle (Loxothylacus panopaei) was introduced into Chesapeake by in the 1960s and produced epidemic infections that castrated xanthid crabs (Hines eta!., 1997). Cholera (Vibrio cholerae) was recently transported in ballast water to Mobile Bay, Alabama, temporarily closing the local oyster fishery (McCarthy and Khambaty, 1994), and we have documented this pathogen in ballast water arriving to Chesapeake Bay (Ruiz et al., in prep.). Furthermore, such direct effects are thought to have many indirect effects on ecosystem characteristics, from food web structure to nutrient dynamics and sedimentation rates (e.g., Vitousek. 1986).

Human-mediated invasions have increased dramatically over the past century (Carlton. 1989; Mills et al., 1993; Cohen and Carlton, 1996). Human activities have allowed dispersal to occur between donor · and recipient regions where natural barriers existed historically, increasing both the potential pool of species that can invade a region and the number of donor regions from which invasions occur. This 6 increased rate of transfer and introduction ofNIS among coastal regions has resulted largely from (I) movement of fouling communities on the bottom of ships; (2) movement and/or intentional release of aquaculture and fishery species along with their rich assemblages of associated (free-living and parasitic) organisms; (3) release of species associated with pet industries or management; and (4) release of organisms in ballast materials of ships (Elton, !958; Carlton, 1979a,b, !987, 1989, 1992a),

The relative importance of particular transfer mechanisms and source regions ofNIS exhibits both temporal and spatial variation. For example, in San Francisco Bay, transplantation of oysters from the Atlantic in the late 1800s and early 1900s, and then from Japan from 1930-1960, brought many NIS associated as fouling species and with trapped sediments; whereas, ballast-mediated invasions now appear to have become the single largest source of invasions in the past 30 years (Carlton, 1979; Cohen and Carlton, !996). In contrast, construction of canals (such as the Suez and Panama Canals, but also many smaller canals in the midwestern and northeastern states of the U.S.) has played an important role in facilitating transfers between bodies of water and drainage basins at some sites, creating bursts of invasions during certain historical periods.

It is e;ident that invasions continue to occur, even at heavily invaded sites, despite the centuries-long history of marine invasions associated with global ship transport. There are many reasons why invasions continue to occur, and why all potential invaders have not already arrived (Carlton, 1996b). Invasions result largely from stochastic (probabilistic) processes, rather than strictly deterministic processes that produce immediate cause and effect consequences. Moreover, changes in environmental conditions (climate) and water quality (both pollution increase and abatement) in donor and/or source regions can promote or inhibit invasion. And transport mechanisms or patterns of delivery may change as global and regional technology, commerce, and politics shift.

Despite considerable spatial variation for the importance of different NIS transfer mechanisms, the worldwide movement of ballast water appears to be the single largest source of coastal invasions today (Carlton and Geller, 1993; Carlton eta!., 1995; National Research Council, 1996). Most known marine and estuarine invasions are now occurring in or near ports with international shipping traffic and are linked to ballast water as a plausible source. The U.S. and Australia each receive over 79 million metric tons of ballast water from foreign ports per year (the equivalent of 2.4 million gallonsibour; Carlton, 1995c; Hutchings, 1992, respectively). Because ballast is usually taken from bays and estuaries, with waters rich in plankton and nekton, most ships carry a diverse assemblage of organisms in their ballast water (e.g., Medcof, 1975; Carlton, 1985; \Villiams eta!., 1988; Carlton and Geller, 1993; Smith et al., 1996), which inoculate receiving ports with large doses of larval invertebrates, fish, phyto- and zooplankton, and microorganisms.

Although coastal invasions have been "idespread and continue to occur, it is not clear whether susceptibility to invasion differs among regions. Some have suggested that particular geographic sites or habitats are more invasible than others, forming "hotspots" as donor or invasion sites (Vermeij, !991; Carlton, !995c). Sites may be more or less susceptible to invasion due to their particular species composition. disturbance, and/or environmental conditions (Erlich, 1986; Roughgarden, 1986; Simberloff. !986). It would be useful to compare the inoculation and invasion rates among various coastal sites. to test for correlation and possible variation in susceptibility to invasion. Unfortunately, most data available on ballast water and NIS frequencies for most coastal sites are too limited, even for many sites which are well studied. For example. San Francisco Bay has the best studied history of1'i1S. but has received no analysis of the characteristics of ballast water delivered there. In marine systems, only Coos Bay, Oregon (by J.T. Carlton) and Chesapeake Bay (by our research group) have received comparable comprehensive studies ofNIS invasion patterns, ballast water delivery patterns, and ballast water biota. We are now comparing these data between sites.

7 Our pre:Ominary findings indicate that delivery oflarge quantities of ballast water alone is not a good predictor o:· :he rate of in,·asion of a site by NIS, nor of the type ofNIS which are likely to be successful. While Chesapeake Bay receives 10 times more plankton rich ballast water annually than any other port on the U.S. east coast or than San Francisco Bay, many more NIS associated with ballast water transport appear to have invaded San Francisco Bay than Chesapeake Bay over the past 20-40 years. Rather, the delivery pa::em of ballast water inoculations and the match of environmental conditions in donor and recipient regions may be critical. San Francisco Bay receives ballast water repeatedly from a limited number of :.'1e same sites (especially Japan) which have similar moderately fluctuating seasonal patterns of closely catching conditions; whereas, Chesapeake Bay receives ballast from a large number of sites spread avec Europe and the 'v!editerranean regions, which often do not have matching environmental condinons. [mponantly. we plan to extend this comparison with comparable data obtained from Prince William Soc:nd over the next rwo years.

NIS in :VIarine/Estnarine Ecosvstems at High Latitude

Although there has been no significant analysis ofNIS in polar marine ecosystems, there have been a limited number of':'·fiS surveys in high temperate latitudes between 40'- 60' and a study of the Baltic Sea, which i.."!cludes a major bay that extends substantially above 60'(Tables B-2, B-3, B-4). These studies derr.onstrate that invasions are not limited to lower latitudes. For two regions in the northern hemisphere (Baltic Sea and Cnited Kingdom)(Tables B-2, B-4) and one region in the southern hemisphere (Tasmania)(Table B-3), the number of known NIS at each location ranges between 32 and 54 species. For each region, the species include a broad range of taxonomic groups, and some of the invasions have generated serious concerns about their ecological and economic impacts. However, the actual impacts of these species, as well as most invasions, remain unmeasured (e.g., Ruiz eta!., in review). 'Konetheless, based upon reported abundances and known ecology, species such as the green crab Carcin:ts maenas (on the North American east and west coasts, Tasmania), the seastar Asterias amurensis (in Tasmania). and the laminarian kelp Undaria sp. appear likely to cause significant and irreversible changes. Furthermore, the cumulative effects of the entire NIS assemblage may cause many changes in ecosystem function that are not easily identified with any one invasion event (Cohen and Carlton. 1996).

NISin Jfarine!Estuarine Ecosystems ofPrince William Sound and Alaska The nu:::bers oiNIS at high latitudes may be lower than those for temperate regions, although it is not clear wi:ether low numbers of documented NIS reflect lack of invasion in high latitude ecosystems or lack of rese2rch focused on the invasion biology of these areas. At the outset of this Pilot Study, the number of',lS documented in Alaskan waters appeared to be lower than other high latitude/cold water ecosystems with more extensive analysis, despite the extensive environmental studies associated with the Exxon Va!Cez oil spill in Prince William Sound and other ecological research throughout the region.

Hypotheses About W7ty .VIS Are Not Evident at High Latitude \'ie ca."l advance hypotheses why NIS have not been as evident at high latitude as at mid-latitudes: (I) ]'.lS are :ruly rare at high latitude. • High la::rude communities may be resistant to invasion (e.g., severe seasonal stress requires specialized eYolutionary adaptations not possessed by non-native species). • Transport patterns may not have been conducive to inoculation. Shipping/ballast water is major source o:' rap1dly escalatmg invasions in temperate latitudes, but perhaps neither the relatively recent (20 yrs1 surge m tanker traffic to Alaska with very large ballast capacity, nor the current shift in tanker traffic to foreign ports has had time to produce invasions. Note. however, that NIS invasions mediated by ballast water have been common over the past 20 years in some cold temperate ports such as San Francisco Bay (Cohen and Carlton, 1996).

8 (2) NIS are actually common at high latitude, but have not yet received concentrated study by experts of invasion biology. For example, Carlton (1979) identified some 160 NISin San Francisco Bay and Pacific northwest coast. but as of 1995 the number ofNIS documented in San Francisco Bay is 212 species (Cohen and Carlton, 1995) and is now more than 220 species (J.T. Carlton, personal communication). Three years ago. the number of:\lS in Chesapeake Bay was considered to be only about 25 species; yet pursuant to our on-going literature search of the historical records, we have documented> 130 NIS, and the list is still growing with on-going research. Despite extensive biological/ecological assessments of coastal ecosystems associated with oil spills in Alaska, NIS probably remain inadequately studied. A review of the existing reports from past and on-going surveys from oil spill work in Prince William Sound is a good start to assessing prevalence of:\1S, but such a review is probably not adequate because those surveys were designed for purposes other than detecting introduced species, and they focused on rocky shores rather than on soft-bottom and fouling communities that are most invaded in other regions (e.g., Cohen and Carlton, 1996). Most introduced species have been discovered by taxonomic experts systematically examining specimens previously identified by non-specialists or conducting field surveys of their own.

Factors Adding Risk for NIS Invasions in Prince William Sound. To determine whether the known l'i1S in Alaska provide an accurate indicator of the probability for biological invasions, we considered 6 factors which contribute elements of risk for invasions of Prince William Sound: 1. Hu2:e volume of ballast water. The greatest quantities of ballast water are transported by bulk cargo carriers and tankers (Carlton et al., 1995: Smith et al., 1996). Chesapeake Bay receives 10-fold more ballast water than other ports on the east and west coasts of the U.S. because of the high volume of bulkers arriving to the ports of Baltimore and Norfolk. Obviously, the tanker traffic to Port Valdez releases similar large volumes of ballast water (Fig. A-4). Other things being equal, larger ballast volumes mean larger inoculations of1'•HS. 2. Short vovage time. Our analysis of biological characteristics of ballast water arriving to Chesapeake Bay shows a marked inverse relationship between densities of organisms and length of voyage, such that ballast water after voyages of 14-24 days had more than 10-fold fewer organisms than voyages of 5-13 days (Smith et al., 1996). However, these effects may be confounded by differences in the source of ballast, which co-varies with the length of voyage (Smith et al., 1996). Voyages of tankers delivering ballast water to Prince William Sound average only 3-6 days, quite short compared to most trans-oceanic voyages that average 12-22 days. Short voyages mean that many larvae and other organisms are likely to be in good health when they are discharged. This element of risk is assessed further below in section Con "Characteristics of Plankton in Segregated Ballast Water". 3. Pattern of repeated deliverv from same donor locations. Although Chesapeake Bay receives about 10-fold more ballast water than does San Francisco Bay, San Francisco Bay appears to be invaded by many more ballast-mediated NIS. This greater risk could be due to San Francisco Bay receiving repeated ballast inoculations delivered from relatively fewer ports than does Chesapeake Bay. Similarly, Pnnce William Sound could be at increased risk not only by the large volume of ballast water, but also by the repeated inoculation of ballast from a small set of west coast ports (Figs. A-2. A-3). 4. The match of environmental conditions of source and receiving ports. Environmental conditions in Alaska are often perceived as being harsh and inhospitable to most potential invaders from temperate latitudes where moderate conditions prevail. Obviously, temperature,light and other conditions during winter are indeed more extreme than those in temperate regions of North America and Asia. However, temperature-salinity conditions in Prince William Sound during spring and summer often approximate conditions in source ports of northwest North America, especially during productive periods of cold water up-welling. In fact. many of the native marine/estuarine species in Alaska have geographic ranges which extend to British Columbia, Washington, Oregon, and Northern California. Below in Section Con "Characteristics of Plankton in Segregated Ballast Water", 9 temperature-salinity conditions in segregated ballast water of tankers arriving from several west coast source pons is shown to be similar to the waters of Port Valdez during late May-early June. 5. Lack of mid-ocean exchange of ballast water delivered to Prince William Sound. Mid-ocean exchange of ballast water reduces concentrations of larvae and plankton by 50-90% (Smith et aL !996). Exchange presumably limits the risk of invasion, as mid-ocean species are generally thought to be incapable of invading nearshore habitats. Delivery of ballast from coastal pons of the U.S. West Coast without oceanic exchange before release into Prince William Sound poses an elevated risk. Funher experimental assessment of this role of mid-ocean exchange in reducing plankton abundance and diversity in ballast water is presented in Section D below on "Experimental ."ulalysis of Effects of Ballast Water Management Practices". While ballast water exchange is required for tankers from foreign pons, the National Invasive Species Act of 1996 considers tankers from U.S. west coast pons to be domestic, coast-wise traffic that does not require exchange. 6. High freguencv of known NIS - esneciallv those transported by ballast water- in source regions of ballast coming to Prince William Sound. Some workers consider that there may be "hotspots" of invasion or donation ofl'I1S. If such hotspots exist, certainly San Francisco Bay and other ports of the U.S. west coast qualify as having among the highest prevalences of documented ballast-mediated invasions, and these in turn form the sources donating much of the ballast water delivered to Prince William Sound. The 309 known NIS of the west coast of North America vary considerably in abundance among 6 latirudinally separate regions (southern California, San Francisco Bay, northern California, Coos Bay Oregon, northwest region from the Columbia River esruary to British Columbia, and Alaska) (Table B-5). The number of known NIS varies from 75 species in southern California to 38 species in the northwest region of Washington and British Columbia, with the largest number of 218 species occurring in San Francisco Bay. At each location along the west coast, NIS are common in a diverse array of taxonomic groups, with arthropods, molluscs, and annelids comprising major fractions ofNIS at most locations (Fig. B-1). In several locations (San Francisco Bay, Northern California, Oregon), vascular plants and chordates also comprise major portions of the NIS. Much of the variation in number ofl'I1S probably reflects the level of srudy and state of knowledge for each location, especially since the highest numbers occur at two locations (San Francisco Bay and Coos Bay) where J.T. Carlton has focused his past research. Many NIS occur at several locations along the west coast, indicating that invasions by the same species have occurred widely across latirudinally separate sites. In fact. all of the NIS now recognized in Alaska also occur in San Francisco Bay and other west coast regions (Table B-5).

10 Known NISin Prince William Sound and Alaskan Waters Based on a Report Prepared by John Chapman Deparrment ofFisheries and Wildlife Hatfield Marine Science Center Oregon State University Newport Oregon 97365-5296 and Gayle Hansen Deparrment ofBotany and Plant Pathology Hatfield Marine Science Center Oregon State University Newport Oregon 97365-5296

Purpose

To identify species which already may have been introduced into Prince William Sound, we surveyed available published and unpublished in formation which may document known invasions. At this initial stage of analysis, we have attempted to document any known reports ofi'-HS in the region, mcluding failed or unsuccessful introductions; but we also provide comments about the potential status and qualiry of the information for each species.

:\Iethods

Recognition of species as being introduced into Alaskan waters and Prince William Sound is based on the same criteria applied to other west coast locations, as presented abo,·e (Carlton, 1979; and discussed by Chapman, 1988; Chapman and Carlton, 1991; Eno, 1996). Methods of literature review and survey followed the standard approaches of electronic bibliographic searches and surveys of grey literature as described above for reviews ofNIS in other regions. Personal, unpublished accounts of observations ofNIS were also documented.

Results In an initial review and literature search, we identified 12 P.1S that have been reported to occur in Pnnce William Sound and Alaskan waters (Table B-6), plus 4 species of macro-algae which have been introduced but not established successfully (Table B-7). Although this preliminary list for NIS in .-\laskan waters is smalL it is of significant concern from several perspect;ves. Frst. collectively there are more species than previously recognized by other assessments (e.g .. \\'iegers eta!., 1997). Only. the soft-shelled clam ,t~va arenaria was commonly acknowledged as an introduced species in Prince \\'illiam Sound (e.g., Foster, 1989). Second. these known N1S represent a diverse array of taxonomic groups. including invertebrates, fish, and plants. Several of these are spec;es wh;ch have histories of invasions in many places around the world, and which are associated with ecolog:cal and economic impacts. Third, the species also occupy a wide range of ecological niches: howeYer, the abundances and realized niches of these species is not well understood for Prince William Sound and Alaska. Fourth, while the mechanisms of transport and introduction for most these species are not known for Alaska, most of them have larval or fragmented stages which may be taken up and transported by ballast water. In fact. larvae of taxonomically affiliated species are abundant in the plankton of segregated ballast water m tankers arriving to Port Valdez (see section Con "Characteristics of Plankton m Segregated Ballast Water"). Fifth, the documentation of some of these introductions is not established in the reviewed scientiiic literature but derived from direct observations by reliable sources (Atla;mc salmon landings) 11 and from very recent reports (Eurasian milfoil in the Juneau region). Thus, these and other invasions may be occurring without formal scientific detection. Finally, the geographic range of some of these species extends well down the west coast of North America to several source ports of ballast water for tankers, including the heavily invaded port system of San Francisco Bay. This indicates that some "N1S in the temperate source ports are capable of invading Prince William Sound. We also identified cases of unsuccessful or faiied introductions ofNIS, primarily associated with aquaculture, indicating that multiple mechanisms of introduction are active in Alaskan waters.

Submerged Aquatic Angiosperm Plants Myriophyllum spicatum. Eurasian waterrnilfoil is a freshwater species that has widely invaded tidal freshwater of estuaries as a result of release from the ornamental aquarium trade. A 1997 report by the Alaska Deparnnent of Environmental Conservation communicated by Susan Walker ofU.S. Fish & Wildlife Service indicated Eurasian milfoil occurs in the Juneau area of southeast Alaska. However, this report needs further documentation. Native to Europe and Asia, M. spicatum can form dense unsightly beds of vegetation that may clog waterways, but it also creates refuge habitat for juvenile fish.

Macro-Algae Codiumfagile subsp. tomentosoides (Suringar, 1867). Hariot, 1889. In addition to the species listed (Table B-6). an algal specimen which appears to be Codium fagile subsp. tomentosoides is currently under study by G. Hansen. The specimen comes from Green Island in Prince William Sound; and if this tentative identification is validated as distinct from C f subsp.fragi/e, it will represent a new location for this "idely distributed invasive species. This weedy subspecies of Coduim fragile is native to Japan (Cohen and Carlton, 1995). C f. tomentosiodes was introduced to Europe in the 19th century from Asia and to New York, probably as ship fouling, around 1956. C f. tomemosoides spread from Kew York, north to Maine and south to North Carolina (Carlton and Scanlon 1985, Cohen and Carlton 1995). It was first collected from San Francisco Bay in 1977 and suggested to be an introduction with ship fouling (Carlton et, a! 1990, Carlton and Cohen 1995) even though nearly all other introductions to the bay from the 1970s to 1995 have been attributed to ballast water (Carlton and Cohen 1995). Sputnik weed, a common name for the subspecies, is attributed to the fragmentation of the thalli that commonly occurs during cold periods (Carlton and Scanlon 1985). Many workers have noted the striking capacity for natural dispersal of Codium via I) motile reproductive cells with can be produced apomictically, 2) vegetatively by thalli fragmentation, particularly in colder months (Fralick and Mathieson 1972, Wassman and Ramus 1973b, Malinowski 1974) and whole plant buoyancy via internal gas entrapment (Bouck and Morgan 1957, Moeller 1969, Ramus 1971, Malinowski 1974). The native north east Pacific species, Codiumfragile Suringar, 1867, ranges from Prince William Sound to Baja California (Pers. Obs., G. Hansen). Other high latitude introductions of the Coduimfragile complex include Codium fragile atlamicum (Cotton) that has spread as far north as the Orkney Islands, U.K. (Trobridge and Todd, In Re,·iew) and C odium fragile scandinavicum which invaded Scandinavian shores (Silva 1957).

Invertebrate Animals We are aware of two hydroids, one clam, two crustacean amp hi pods and one tunic ate that are reported potential or likely introductions of invertebrates to Alaska in addition to .\{va arena ria and Heteromastus filiformis that we report from Port Valdez. Except for Mya arenaria, Carlton ( 1979) doubted the validity of most NIS in Alaska that he considered.

Cnidaria (Hydroids) Sy11cory11e mirabilis (= Sarsia tubulosa) (M. Sars, 1835). Probably introduced to the northeast Pacific with fouling on wooden ships from the North Atlantic, this species is known also as Coryne rosaria. The early north Pacific records of S tubulosa are from Puget Sound in (Agassiz 1859) and San Francisco Bay in 1860 making it, potentially, one of the earliest marine introductions to the North Pacific (Carlton !979) and is likely to have arrived with ship fouling if it is indeed introduced. This species was 12 collected from 90 m off of the Trinity Islands, Alaska in 1914 (Fraser 1914) and Garforth Island, Miur Inlet, Alaska (USNM 34481) in 1899 (Fraser !937). However, we did not find this species in our preliminary survey, and it is not reported from Port Valdez. Syncoryne mirabilis may be misidentified in Alaska. The two northern records and all records south of Monterey are old. requinng confinnation (Carlton 1979). Syncoryne mirabilis occurs in the North Atlantic in Iceland and in northwestern Europe and in Japan and Russia in the west Pacific (Carlton 1979). It could be a ·'circumboreal neritic species" (Russell, 1953:56) extending down the four continental margins of the Arctic Ocean or a complex of species with introductions or multiple, isolated species at lower latitudes and a single, naturally distributed species at high latitudes. Carlton (1979: 237) did not considerS. tubulosa to be a certain introduction pending further taxonomic studies to reveal the relationships between the many populations of this taxon from around the world. It's occurrence in Alaska is uncertain.

Tubularia croacea (Agassiz, 1862). Probably introduced to the northeast Pacific with fouling on wooden ships from the North Atlantic. The Gulf of Alaska record of this species (Fraser 1937) is also doubtful. Fraser's ( 193 7: 51) single record of this distinctive hydroid "Gulf of Alaska" gives no specific locality. Tubularia croacea is a common shallow-water hydroid that occurs on pilings floats and other structures on the Atlantic coast from Newfoundland to Florida and in the northwestern Gulf of Mexico (Fraser 1944: 98, Carlton 1979). It was very likely introduced into the north Pacific with ship fouling and probably again with Atlantic Oysters (Carlton I 979). Since we did not find this species in our preliminary survey and have no additional reports of it in Alaska since Fraser (193 7), it is not reported from Port Valdez. This species also may be misidentified in Alaska.

Annelida Heteromatus filiformis. The occurrence of Heteromastus filiform is in Port Valdez could be associated with ballast water traffic, although it is not clear when it arrived in the area. It is unlikely to have arrived in fouling communities, since it is not found in such habitats and is unlikely to have been actively introduced. H filiformis is native to the Atlantic coast where its range extends from New England to the Gulf of Mexico. This species has been reported from Greenland, Sweden, the Mediterranean, Morocco, South Africa, the Persian Gulf, New Zealand, Japan, and the Bering and Chukchi Seas (Cohen and Carlton !995). This mud-dwelling species may have been introduced to California, Oregon and Washington in the 1930s with introduced Atlantic oysters, Crassostrea virginica as early as the I 930s (Carlton I 979) or it may have been an early ballast water introduction (Cohen and Carlton 1995). The slow growth of oysters in Alaska results in annual oyster importations and creates a significant potential for introductions organisms that are associated with oysters. H.filiformis could have been introduced to Kachemak, Alaska with oysters transplanted from Willapa Bay, \Vashington. H filiform is occurs in Willapa Bay (Carlton 1979) and it has been found in association with oysters (Wells 1961). We are unaware however, of attempts to grow oysters in Port Valdez. H.filiformis occurs in densities of up to 4000 individuals m·' in the western half of Suisun Bay portion of San Francisco Bay (Hopkins 1986, Markmann 1986). This region of San Francisco Bay is a location where some oil tankers traveling to Valdez take on ballast water. Both the larvae and mature adults of this species are likely to tlow into filling ballast water tanks in this area. Thus. ballast water transport appears to be the most likely mechanism for carrying H. filiformis to Alaska.

Mollusca Mya arenaria Linneaeusl758. The occurrence Mya arenaria in Port Valdez is unlikely to be a result of ballast water in tanker traffic. Carlton (1979) reviewed the evidence for the introductions of this species to the northeast Pacific. M. arenaria is a relatively early introduction that appears to have arrived in Port Valdez long before tanker traffic began. M arenaria is native to the northwest Atlantic. It has been introduced into the Pacific an Europe, Asia and the northeast Pacific. Its northeast Pacific range extends from Alaska north of the Aleutian peninsula (Carlton 1979). Despite extensive systematic research on the genus (Foster 1946, MacGinitie 1959, MacNeil 1965, and Laursen 1966), it's name, 13 limits of intraspecific variation, and geographic distribution are subject of considerable disagreement. Tnis poor resolution of the species and the similarities between M. arenaria and M. japonica have caused particular difficulties in resolving the fossil occurrence of M. arenaria in Alaska (MacGinitie 1959, Carlton 1979a). It's endemic status in northern and western most Alaska is therefore questionable but rather clear in southeast Alaska and Port Valdez. M. arenaria is very likely not native in Alaska since there are no fossil records or records of it from aboriginal shell middens (Carlton 1979a). This large. edible clam may have been actively distributed throughout the northeast Pacific in the late 19th century and early into this century (Cohen and Carlton 1995) as a food source. Thus, although lvf. arenaria has a planktonic larva which could be taken up in ballast water, it is not clearly a ballast water introduction.

Venerupis phillippinarum (Deshayes, 1853). The Japanese little neck clam. Reproductive populations of this species may occur in southeast Alaska (Carlton 1979), but these have not yet been confirmed.

Crustacea Corophium acherusicum Costa 1857. This species is probably endemic to the eastern North Atlantic and is perhaps the most widely introduced amphipod of all species in the world. No specific locality was given for a report of this species in Alaska (Crawford 1937: 617, Shoemaker 1947: 76, 1949: 76). We were unable to locate specimens from Alaska reported by Shoemaker ( 1947) in the Smithsonian Museum ofNatural History collections, therefore, Shoemaker's reports may be based on Crawford (1937). The unidentified Corophium in the University of Alaska Museum collections (code 616915020000) should be closely examined to determine whether they are possibly C. ascherusicum. However, this nearly cosmopolitan species is common in southern British Columbia (Bousfield and Hoover 1997, Carlton 1979) and occurs at high latitudes of the North Atlantic, in North America and Europe (Lincoln 1979), New Zealand (Hurley 1954) and Japan (Ishimaru 1984). Hurley (1954) reports that the distribution of C. acherusicum "traces out some of the major shipping routes" of the 19th and 20th centuries. Corophium insidiosum has been introduced to nearly every location in the world where Corophium acherusicum has by the same mechanisms and is morphologically similar. Moreover, C. insidiosum is more commonly reported in the northern end of the confirmed northeastern Pacific ranges (Carlton 1979). Since, if C. insidiosum is clearly established in Prince William Sound (see below), the records of C. acherusicum in Alaska should be regarded with caution. Nevertheless, Crawford's ( 193 7) indication that he had examined the Alaskan specimens of C. acherusicum in the same paper in which C. insidiosum is described confirms that this species was in Alaska despite the absence of subsequent reports.

Corophium insidiosum Crawford, 193 7. This species, like C. ac/zerusicum, is nearly cosmopolitan C. insidiosum commonly occurring in southern British Columbia (Bousfield and Hoover 1997, Carlton 1979) and at high latitudes of the North Atlantic, in North America and Europe (Lincoln 1979), New Zealand (Hurley 1954) and South America (Alonsa de Pina 1997). Bousfield and Hoover (1997: 114) erect the new genus Monocorophium from Corophium and describe a new species, Monocorophiwn carlouensis from 29 lots containing 290 specimens collected from Prince William Sound and S. E. Alaska through the Queen Charlotte Islands. We continue with the old epithet Corophium pending acceptance of this new designation by other workers. Bousfield and Hoover (1997) identify C. carlottensis in collections as far south as British Columbia and Corophiwn insidiosum from British Columbia, Washington and south. Bousfield and Hoover (1997) thus recognize C. carlortensis as northern and "distributionally non-overlapping" that of C. insidiosum. They distinguish C. carlottensis from C. insidiosum Crawford, 1937 by its smaller size, "more tightly setose antenna 2", the relatively greater length of article 6 of pereopod 7 and the lack of setae on the anterior margins of segment 4 of pereopods 3 and 4. Corophium carlottensis appears otherwise identical to C. insidiosum. The variation among specimens of C. insidiosum from a single live cultured population Hawaiian C. insidiosum and from specimens collected from Yaquina Bay, Oregon completely overlaps every 14 combination of dense to light setation on the antennae and anterior margins of segment-+ of pereopods 3 and-+ of C. carlottensis. Additionally, the relative length of article 6 of pereopod 7 illusrrated for C. carlouensis (Bousfield and Hoover 1997: fig. 29) is indistinguishable from that of C. insidiosum (Bousfield and Hoover 1997: fig. 26). We hope to conduct further morphological comparisons and conduct reproductive viability tests between the Yaquina Bay, Hawaiian and Alaskan populations. There is no evidence that C. carlouensis is a good species at present (Chapman eta/. in ?rep.)

l:rocordata (Tunicates) Ciona intestinalis (Linnaeus, 1767). Ritter's (1913) record (USNM 5633, 1903,) of this species in southeastern Alaska does not provide a specific locality (in Carlton 1979). Records of this species in British Columbia are few and relatively old, warranting re-examination; and there are no firm Washington or Oregon records (Carlton 1979). Carlton (1979) therefore concludes that this species is only confirmed in California south.

Vertebrate Animals, Fish Sa/mo salar. Atlantic salmon have been introduced to supplement salmon fisheries in numerous locations around the world, including various locations of the U.S. by the United States Fish Commission. Often these intentional introductions have not become established successfully; however the species' range in Europe extends northward well within the Arctic Circle, indicating that it has potential for establishment in Alaskan waters where it could compete v.ith native species. Atlantic salmon are reared in commercial pen-culture in Puget Sound, British Columbia and apparently Alaska. Some of these fish escape from the pens are caught by commercial and sport fishermen in isolated instances. Atlantic salmon have be landed in recent years at the small boat harbor in Valdez (R. Benda, personal communication). Thus far, this species does not appear to have established a self-sustaining population in Alaska.

Unsuccessful Invertebrate Introductions Into Alaska Crassostrea gigas (Thunberg, 1795). We are aware only of attempts to establish the Japanese Oyster in Ketchican Bay for aquaculture, which have been unsuccessful. Growth and reproduction of this warm water species in cold Alaskan waters appear too limited to maintain local populations. Local growers depend on annual introductions of new spat for each year's harvest (Foster 1991 ).

l:nsuccessful Macro-algae Introductions to Alaska Four species of macro-algae have been introduced into Alaskan waters in association with aquaculture activities, but these have not established successfully (Table B-7). M acrocystis integrifolia Bory. The giant macro-alga Macrocystis integr(iolia was introduced from California by egg fishermen of Prince William Sound to provide sites for he :'ring eggs until its introduction was outlawed in the 1990s. This species has not been successfully es;ablished within the Sound .

.Hacrocystis pyrifera (L.) C. Agardh 1820. Since the introduction of M. inlegrzfolia into Prince William Sound was outlawed in the 1990s, Macrocystis pyrifera from southern Alaska has been used exclusively to provide sites for herring eggs. This species has not been successful m the Sound.

Pachymellia carnosa Introduced cultures of Pachymenia carnosa held in open net-pens in southern Alaska for aquaculture have not resulted in the establishment of self maintaimng populations.

Porphyra ye;oellsis Introduced cultures of Porphyra ye=oensis held in open net-pens in southern Alaska for aquaculture have not resulted in the establishment of self maintaining populations.

15 An additional algal species of Ascophyllum has also been introduced unsuccessfully into waters off British Columbia (Table B-7). These unsuccessful introductions are easier to document than accidental releases, because they have been associated with aquaculture operations which provide recorded activities. They illustrate the point that many introductions may be occurring by various mechanisms, but many such introductions often do not result in populations becoming established successfully.

Field Surveys for NIS in Port Valdez. Based on a Report Prepared by John Chapman Department of Fisheries and Wildlife Hatfield Marine Science Center Oregon Stare University Newport Oregon 97365-5296 and Gayle Hansen Department ofBotany and Plant Pathology Hatfield Marine Science Center Oregon Stare University Ne>vport Oregon 97365-5296

Purpose

To determine whether nonindigenous species have already become established within Port Valdez without being detected by earlier biological surveys, we conducted an initial collecting survey of intertidal invertebrates and algae from several habitats. Our initial collecting effort focused on mudflat and fouling communities, because these habitats have not had as much study as most other habitats of Port Valdez, and because NISin other, better studied locations often invade such habitats (e.g., Cohen and Carlton 1996). However, Port Valdez is characterized by reduced salinities and comparatively low productivity due to low nutrients and high turbidity in glacial melt that flows into the fjord. We now hypothesize that habitats in Prince William Sound, which are characterized by higher biotic diversity, higher salinities and higher nutrients, may be more vulnerable to invasions than low nutrient waters within the Port. Tankers often begin releasing segregated ballast water into Prince William Sound well before they reach Port Valdez, providing inoculations to extensive areas outside the confines of the Port. Our opportunity to sample habitats outside Port Valdez were very limited due to constraints of time and access during the Pilot Study.

Survev Methods

A preliminary survey of native and nonindigenous intertidal flora and fauna of parts of Port Valdez and Valdez Arm was conducted during late May- early June, 1997. The Duck Flat, immediately east of Valdez, was sampled primarily on 22, 26 :v!ay and 3 June of 1997. The rocky intertidal zone of Anderson Cove at the southwest entrance to the Valdez Harbor. the rocky and mudflat tidal areas of Sawmill Bay at the northwest entrance to Valdez Harbor, and the float fauna of the Valdez small boat harbor were sampled on 31 May 1997. Organisms were collected by hand or from sediments washed on an 0.5 mm mesh sieve. Invertebrates were preserved in 10% formalin in the field and later transferred to 70% ethyl alcohol. Half of the specimens of each macro-alga species were immediately preserved in 5% formalin and the remaining half were pressed and dried.

16 Jeff Cordell, Fisheries Unit, University ofWashington, Seattle WA identified harpacticoid copepods and all other Crustacea were identified by John W. Chapman. Polychaetes were identified by Leslie Harris. Los Angeles County Museum of Natural History, Los Angeles, CA, and Faith Cole, US EPA, Newpon, OR Molluscs were identified by James T. Carlton and John W. Chapman. Macro-algae were identified by Gayle L Hansen. We compared our species list with Wiegers et aL (1997) summary of previous taxonomic surveys of the mudflat and rocky intenidal flora and fauna ofPon Valdez to provide an independent estimate of the relative diversities of nonindigenous, cryptogenic and natlve species in Pon Valdez. This comparison was used also to indicate the sufficiency of all surveys combined for indicating the total diversity of the area,

Recognition of species as being introduced into the Port Valdez/Prince William Sound area is based on the same criteria applied to other west coast locations, as presented above (Carlton, 1979; and discussed by Chapman, 1988; Chapman and Carlton, 1991; Eno, 1996),

Results

Invertebrate Survey We collected 49 benthic invertebrate species, not counting insects (Table B-8), Twenty-six of the 49 species (53%) were reported for Port Valdez previously by Wiegers et aL (1997) and 23 (47%) are new. Most of the new records are in taxa such as gammaridean amp hi pods that were not identified in previous surveys, and only Shaerosyllis brandhorsti, Limnoria ligatum and Danielssenia cinctus. of our survey. are similar to other species reported previously (Table B-8). The large proponion of new benthic invenebrate records (Table B-8) appears to result from incomplete collecting in the Pon. Wiegers et aL ( 1997) report the distinctive commensal clam Orobitella rugifera, which arraches to the abdomen of the mud shrimp Upogebia pugettensis. However, U pugettensis was not reponed in their summary list and we did not find the distinctive holes of this long-lived species (Posey eta/. 1991). We also collected several specimens of the conspicuous purple shore crab Hemigrapsus nudus for the first time but did not find Hemigrapsus oregonensis, an equally distinctive shore crab reported previously. Thus, significant changes in benthic community structure may occur in the area over time and despite the preliminary nature of our survey, the diversity of intertidal flora and fauna of the Pon appears to be underestimated.

The geographic range and overlap with southern species by the Alaskan fauna is significant. indicating suitability of Port Valdez for other species from more temperate locations. Of the combined 73 species with known ranges collected from all surveys of the Port, only 6 (8%) appear endemic only to Alaska (Table B-8). Twenty-two (30%) of the species occur as far south as Washington and 29 (40%) occur as far south as California, Of the 89 benthic invertebrates collected, 15 (17%) are cryptogenic. ha,·ing disjunct geographical or cosmopolitan distributions but origins which are not adequately resolved. Two species. the polychaete worm Heteromastusfiliformis (Claparede, 1864) and the clam My a arenaria Linnea us. 17 58, clearly appear to be introduced.

Macro-algae Survey Twenry-one macrophyte species and an unidentified tubular filamentous diatom were collected in the field su1·vey (Table B-9). Three macrophytes, Blidingia chadefaudii, Neorhodomela acu/eata and Neorhodome!a oregona. were newly recorded for the Port. Ail three are similar to other species reported previously from the Pon suggesting even these species may have been collected previously but misidentlfied. These preliminary results suggest that the macro-algae diversity ofPon Valdez is better known than the intenidal invenebrate diversity. Nevertheless, endemism among the macrophytes is even less than among the invertebrates. None of the 101 species included in the analysis are endemic to Alaska (Table B-9). Of the 102 macrophytes, 24 occur in Australia, 60 in Japan, 85 in Washington, 76 in

17 Oregon, 70 in California and 43 occur in Great Britain. Including only species that occur in all 6 geographical regions, 21 (21 %) of the 101 species analyzed are cryptogenic.

Twenty-nine of the 101 macrophyte species of Port Valdez (Table B-8) can survive unattached (in which plant fragments section off and drift free) (Norton and Mathiesson 1983). This process is unlikely to be noticed without systematic observations and, thus, is probably underestimated in general. Thus, if algal fragments are entrained in ship ballast systems, a broad diversity of macrophyte species in Port Valdez have the potential to be transported in ballast water. However, macro-algae are seldom reported from ballast water (Carlton and Geller, 1993), and accidental introductions are generally associated with fouling on ship hulls (Carlton !979a) or transplantation of species for aquaculture purposes or for packing (Cohen and Carlton 1995). Norton and Mathiesson (1983) have discovered numerous algal species that can survive in an unattached state for days to weeks, and these species are likely candidates for ballast-water transport. Even the fragments of many taxa can release spores that colonize the new environments or reattach by entangling in other algae where they can mature and reproduce. Small algae or algal fragments could be easily overlooked in ballast water studies, and many of the cryptogenic algal species in Port Valdez could have been transported there in this way.

Discussion

The small proportion of NIS found in Port Valdez differ strikingly from the large proportions of NIS that have been collected in surveys of other portS of the northeast Pacific such as Puget Sound and San Francisco Bay (Carlton 1979, Cohen and Carlton 1995). However, the high incidence of cryptogenic species suggests that the total number ofnonindigenous species (invertebrates at 17% and macro-algae species at 21% of total species) in Port Valdez may be underestimated. Carlton ( 1996) estimates conservatively that 37% of the species of San Francisco Bay, one of the most intensely invaded ecosystems in the world, are cryptogenic. Many cyptogenic species also occur in Alaska, and some will undoubtedly will be recognized as nonindigenous species as their local, regional, and global distributions are better resolved. In addition, the low species overlap between our survey and previous surveys indicates that many more species are likely to occur in Port Valdez than are presently reported, and/or that the biota is changing. The large proportion of new invertebrate species collected in the Port in this small survey (Table B-8) indicates that many unreported taxa may remain. The low overlap among algal species collected in the three separate surveys (Calvin and Lindstrom 1980, Wiegers eta/. 1996, Chapman and Hansen 1997, and Hansen 1989, respectively) indicates changing diversity over time. These differences are likely due to the seasonality of macro-algae in Alaska. The appearance of all but extremely successful new taxa arriving in the port would be difficult to detect with the background data at hand. A more extensive survey of the biota will correct this problem by providing missing information on the distributions, abundances and diversity of the biota. The number ofNIS discovered may increase as the sampling effort increases.

The taxonomic resolution of the local biota is presently insufficient to estimate whether the large numbers of cryptogenic species are likely to be predominantly introduced or native. Tne tremendous species overlap between the Port Valdez biota and geographically areas remote from Alaskan waters greatly complicates the analysis. Active ballast water introductions of many genetically distinct intraspecific populations from Washington, Oregon and California could occur, for instance, without detection. The presence of genetically distinct, introduced populations of a species, would be a significant effect of ballast traffic. Thus, the genetic composition of some of the more abundant species in the Port, such as lvfytilus trossulus, should be compared to more southern populations for this possibility. Increasing the resolution of the taxonomic and biogeographical information on the Port Valdez biota may be even more valuable. These data are critical for establishing the numbers ofNIS, their ecological significance, their likely origins, timing and mechanisms of introduction. Clear

18 resolution of the local and regional distributions of the biota over time will allow tests of the criteria for introduced species and resolution of many of the presently cryptogenic species.

Non-Indigenous J1arine/Estuarine Species with Potential for Introduction to Prince William Sound Based on a Report Prepared by John Chapman Department ofFishen·es and Wildlife Hatfield Marine Science Center Oregon State University Newport Oregon 97365-5296 and Gayle Hansen Department ofBotany and Plant Pathology Hatfield Marine Science Center Oregon State University Newport Oregon 97365-5296

Purpose

To identizy non-indigenous species with potential for introduction into Prince William Sound, we reviewed the literature for species which have demonstrated introductions into high latitude ecosystems, especially via ballast water transport. We do not consider the absence of previous introductions of a species (which applies to most species) as an indication oflow potential for introduction via ballast water. Furthermore, the introduction and subsequent abundance of a species in a new location provides little information on its expansion in a new geographic area. The green crab expansion on the east coast ofNurth America progressed over many decades but this species is advancing up the west coast of North America in less than a decade. The Asian estuary clam, Potamocorbula amurensis has not expanded its range since arriving in San Francisco Bay, yet its populations there are extremely abundant (Carlton et al. 1990). However, focusing upon high latitude species which are known to invade, and which have introductions apparently by transport in ballast water. provides an indication of the taxonomic and ecological diversity of species with demonstrated potential for invading an ecosystem such as Prince William Sound.

Methods

We surveyed the literature for known introduced species of the world that have native populations at latitudes greater than 40° north or south and that appear to have a significant potential for transport into Alaska via ballast water traffic. The primary criterion for transport potential is the likelihood of entrainme:lt into and short-term survival in ballast tanks. Spionid polychaetes, decapod crustaceans and molluscs. for instance, produce planktonic larvae, zoea and veligers that are common in ballast water samples (Carlton and Geller 1993, pers. obs.). Peracaridan crustaceans are active swimmers at night as adults (Williams and Bynum 1972) and are also common in ballast water (Carlton and Geller 1993, pers. obs.). Micro-algae are common in plankton of coastal waters. These taxa thus have significant potential for ballast water transport to Port Valdez.

Macro-algae are included even though they are rare in ballast water samples (Carlton and Geller 1993), were not collected in this ballast water survey, and are rarely associated with ballast water dispersal. The principal mechanisms usually associated with accidental introductions of macro-algae are fouling on the hulls of ships or aquaculture related transplantations of species such as oysters and as packing material (Carlton 1979, Carlton and Scanlon 1985, Cohen and Carlton 1995, Trowbridge 1995). 19 However, fragmentation can function as a mechanism of dispersal in algae ()iorton and Mathieson 1983). These fragments can become entangled in suitable locations after discharge and marure. These fragments can also release spores or gametes if they have reproductive bodies which can also lead to colonization of new areas. These fragments could be entrained in ballast water and transported to new locations. Thus, ballast water could be an important mechanism for transporting macro-algae since even a small number of such fragments could be sufficient to colonize new areas. These fragments would easily be missed in a ballast water survey that does not coincide with seasons of maximum fragmentation, principally in the fall.

Results

High Latitude Invertebrate Introductions Outside of Alaska We include 321'<1S invertebrates in our list of potential ballast water introductions of Alaska (Table B-1 0). These species represent a wide array of taxonomic and ecological diversity. The potential impact of an invasion by any of these ·species is difficult to predict for Port Valdez/Prince William Sound.

Invertebrate Introductions from Alaska Adult red king crab, Paralithodes camtschatica from the northern Pacific were successfully introduced to the Barents Sea by the Soviet Russians between 1961 and 1969. The Barents Sea populations was estimated to be at least 500,000 by 1995 (Olsvik 1996). It's environmental impact in the Barents Sea is unknown. The seastar Asterias amurensis appears to be native to Alaskan waters and has been introduced into Tasmania, where the Australian government is concerned about its predatory impacts on subtidal communities and food webs. These introductions suggest that important invertebrate species are capable of introduction into high latirude waters equivalent to those in Alaska.

High Latitude Macroalgae Introductions We include 21 potential macrophytes as potential species for introduction to Alaska via ballast water (Table B-11 ). As for invertebrates, the diversity of algal species with potential for invasion is high; and potential impacts are difficult to predict.

Macroalgae Introductions from Alaska Several species of macroalgae from Alaskan waters have been exported and successfully introduced into other cold-water local!ons of the wor!d (Table B-12), indicating that algal species are capable of introduction into high latitude ecosystems companble with those m Alaska.

High Latitude Micro-algae Introductions Harmful phytoplankton blooms in coastal ecosystems are a growing concern world-wide (Taylor, 1987), and phytoplankton are common in ballast water (Carlton and Geller 1993). \Ve include 11 species of micro-algae that may have potential for ballast water introduction into Alaska (Table B-13). All but three of these species already occur in the North Pacific.

Discussion

The pool of foreign species from locations with potential for introduction into Alaskan waters and Prince William Sound is large and diverse. The array of species which already have success tully invaded a cold-temperate and boreal waters around the world represent nearly every major taxonomic group and ecological niche in marine and esruarine waters. :.!any other species do not have recorded introductions but are also at risk for transport in ballast water. While the odds of successful introduction into Alaska are impossible to calculate accurately for any giYen species, this diversity of known introductions clearly indicates that a demonstrable risk of in,·asion occurs in Prince William Sound. 20 C. Plankton in Segregated Ballast Water

Purpose

To begin an evaluation of the magnitude and diversity of biota transferred to Prince William Sound in the segregated ballast water of oil tankers, we sampled and analyzed ballast water on tankers arriving to the Alyeska Marine Terminal during a 2 week time period, 23 May- 6 June 1997. We selected this rime period to correspond to a season of relatively high reproductive activity and plankton abundance in waters of the northern hemisphere. Also, we predicted that conditions in Prince William Sound are most conducive for invasion during the spring/summer, when water temperatures and productivity may most closely match ballast water source regions to the south.

This portion of our study focused entirely on the segregated ballast water of tankers that arrived to Port Valdez. All but one of the vessels that we sampled arrived from domestic ports. This exception, an arrival from Korea, underwent open ocean ballast water exchange, a management practice used by tankers arriving from foreign ports to reduce the abundance ofNIS. Here, we discuss our analysis of abundance and diversity for plankton communities associated with segregated ballast water as it usually arrives from domestic and foreign ports. In the next section we discuss ballast water exchange in greater depth.

Methods

We sampled a total of 16 oil tankers arriving to Valdez between 22 May and 5 June 1997. Although we attempted to collect ballast water from every tanker that arrived during this time period, some of the vessels with double bottom tanks could not be sampled easily without disruption of ship operations and modification of our sampling protocol. We therefore sampled only a portion of ships with double bottom tanks, as well as all other tankers arriving with ballast water. For these vessels, we applied our established methods for qualitative and quantitative analysis of biota transported in ballast water, which evolved from methods pioneered by J.T. Carlton (e.g., Carlton and Geller. !993: Smith et al., 1996). During the sampling period. we adapted our standard methods to ship operat10ns and ballast management pracl!ces for tankers at the Valdez Marine Terminal. Our protocol consisted of collecting the following information and samples: • Shiv and ballast manarrement information: Last port of call, number of tanks by type, capacnv of tanks, amount of segregated and non-segregated ballast water on board, source(s) of ballast water, age of ballast water, date of arrival, ballast management practices; • Phvsical variables of ballast water: Water temperature and salinity were measured (surface and lOrn depth) for each tank sampled (as below), collecting ballast water with a Niskin bottle through the

Butterworth hatches: oxygen (02 ) concentration was not measured because previous extensiYe

analysis of ballast water tanks in other cargo ships indicated that 0 2 concentrations rarely vaned and were not appreciably lower than saturation (Smith et al., 1996). • Biological samoles of ballast water: Plankton samples were collected by towing a standard plankton net (80 micron mesh, 30 em diameter) vertically through the entire height of the water column in each ballast tank; access to ballast tanks was obtained through the Butterworth hatches; at least 2 tanks were sampled for each ship, when ballast water was present and accessible, and two plankton tows were collected for each tank; the height of each plankton tow was measured to the nearest I 0 em. • AdditiOnal observations and oooortunistic samples: Upon initiating sampling of ballast tanks. we routinely examined the surface waters to look for large, mobile biota (e.g., fish) and organisms

21 attached to the sides of tanks; we often take opportUnities to collect any such organisms observed, as well as bottom sediments, since these are usually missed in our plankton tows. • Phvsical variables of port water: Shipside water temperature and salinity were measured (surface and lOrn depth) usually within an hour of sampling ballast water of most vessels; the samples were collected from the berth platform (within 50m of the ship), using a Niskin bottle.

Most plankton samples were returned to the laboratory and examined initially within an hour of collection to assess condition of organisms present. More specifically, we examined each plankton sample with our dissecting microscopes (10-40x), temporarily set-up in the Quanterra chemistry laboratory at the Valdez Marine Terminal, to provide a qualitative assessment of plankton viability, diversity of plankton species, and categorical abundance of major taxonomic groups. Each sample was washed carefully into a finger bowl for examination, and the presence of each morphologically distinct taxonomic group was noted. For each taxon identified, we estimated the categorical abundance as rare (1-10 individuals/sample), common (10-100 individuals), or abundant (>100 individuals). For each taxon, the percent of individuals alive was estimated by evaluating their morphological integrity, movement, and activity; although status of some organisms (e.g., diatoms or eggs) was difficult to discern with confidence during a brief screening. After initial microscopic examination, the plankton samples were preserved in 5% buffered formalin for later identification and enumeration of organisms (as below). Only those plankton samples from last two ships sampled (Prince William Sound and SIR Benecia) received no live analysis. These samples were preserved directly after collection.

Quantitative analysis of the preserved samples was conducted at the Invasions Biology Laboratory of the Smithsonian Environmental Research Center (SERC) in Edgewater, Maryland. Samples were concentrated on an 80 micron sieve and washed into a finger bowl for identification and enumeration. Each whole sample was examined using a stereo microscope, and all morphologically distinct taxa were identified to the lowest taxonomic level. For many groups that included larval invertebrates (e.g., bivalves, gastropods), identification could not progress beyond gross taxonomic groups; further identification can only be accomplished with intensive culture oflarvae to adult stages, upon which taxonomy is based, or the use of molecular probes. For other groups that include adult stages (e.g., copepods), we sought species-level identifications. For those taxa present in abundances< 100 indi,iduals per sample, the total number ofindi,iduals were recorded. For abundant taxa(> 100 individuals/sample), samples were split using a Folsom plankton splitter to achieve counts between 10- 100 indi,·iduals per subsample (usually splits of 1/8 to 1/32). For organisms in split samples, two subsamples were counted.

Throughout this section, we rreat ship as the level of analysis and replication, because the source of ballast water and ballast management practices were similar for all ballast tanks within ships, except for those cases of experimental exchange on two vessels (see next section). Although our replicate plankton samples from each of two tanks per ship provide some important information on variation within ships, these are not statistically independent (since the ballast water originates from the same source and time) and mainly provide greater confidence in estimating plankton communities per ship. Here, we use mean density measures that derive from replicate tanks and tows, and we plan to address explicitly within-ship variance during our 2-year study.

Taxonomic identification of plankton has followed a standard protocol. For those groups of organisms that can be identified using the life stages present in ballast water samples (as discussed above), we made an initial identification based upon our current knowledge and literature that was immediately available to us at SERC. For many copepods, we were able to discern genera without much difficulty. Enumeration proceeded based upon the lowest discernible taxonomic units, and representative specimens were vouchered for taxonomic verification and. wherever possible. species-level identification. These voucher specimens were sent to taxonomists at the Smithsonian Institution· s 22 Natwnal Museum ofNatural History and elsewhere for such verification and idennfication. To date. only a portion of the voucher specimens have completed this process for full verificanon and identification, and we report these results where available.

Results

Of the 16 ships sampled during the Pilot Study, 15 carried and discharged segregated ballast water from domestic source regions (Table C-1); the sixteenth ship (0/S Washington) was delivering diesel fuel to the port. Most of these vessels and their ballast water came directly from the ports of Long Beach (CA; 2 vessels), San Francisco Bay (CA; 4 vessels), and Puget Sound (W' A; 6 vessels). A single vessel and its ballast water came from each Portand (OR) and Cook Inlet (AK). Only two vessels came from outside the west coast ofNorth America: one from Barber's Point, Hawaii and one from Yosu, Korea. Only the vessel from Korea conducted ballast water exchange on all of its tanks, and two of the ships arriving from San Francisco Bay conducted experimental exchange of ballast water in two tanks upon our request.

The 15 ships arriving with segregated ballast water carried an average of30.311 (s.e. = 3,634) m3 of segregated water, or 73% of their total capacity (Table C-1). With the exception of those 3 ships that underwent some ballast water exchange, all of the segregated ballast water derived from the last port of call. The average age of ballast water was 5.3 days old, ranging from I to 10 days. The average temperature and salinity of this ballast water was 13.0 'C and 27.6 ppt, respectively (Table C-2). There was no evidence of stratification within segregated ballast tanks, as salinity and temperature varied less than I unit between measures at the surface and 1Om.

The temperature and salinity of surrounding waters at the time of ballast water discharge was similar to that of the sampled ballast water, although Port Valdez exhibited significant temperature and salinity stratification. The temperature ranged from 10-14 oC at the surface and 5-9 oC at !Om depth, and the salinity varied between 20-29 ppt at the surface and 29-32 ppt at !Om (Table C-2). Mean values for field and ballast water measures varied less 5'C and 9ppt (Fig. C-1). Although this represents an overall significant difference for both salinity and temperature among measures (temperature: F=44.24, df-=2,38, p< 0.001; salinity: F=27.22, df-=2.38, p

To date. we have identified a mimmum of 69 different taxonomic groups amvmg in the segregated ballast water of tankers from domestic pons (Table C-3). This greatly underestimates the species diversity, as many of the categories include multiple species. Some of this will become evident as we obtain further verification and identirication of vouchers. but most larvae cannot be identified to species without culturing them to maturity.

Our quantitative measures of abundance underscore the high level of variation that existed among ships, even over a very short time period. There was as much variation in ships arriving from different domestic ports as those arriving from the same port (Table C-3). For example, the SIR Baton Rouge and Chevron Mississippi each delivered ballast from Anacortes, but exhibited extreme differences 3 in the densities ofOwenid Polychaetes (688 vs. 1 per m ), bivalves (2,897 vs. 109 perm'), barnacle cyprids and nauplii (1,450 vs. 13 perm'), and many other taxa.

Some of the differences in plankton abundance among ships may result from variation in age of the ballast water, since survival may have been lower in the older water of the Che,Ton Mississippi (7 days versus 4 days forthe Baton Rouge; Table C-1). There was a significant decline in the density of annelids and molluscs with increasing age of ballast water from domestic, continental sources (Fig. C-2), when excluding the !-day old water from Cook Inlet (labelled as AK). However, a similar decline in 23 total number of organisms was not apparent for these same sh:ps. Although it is probable that some variation in both density and diversity of plankton among ships also resulted from differences in communities that were entrained in the ballast tanks, the relative contribution of initial conditions and survivorship to observed patterns is unknown at present.

It is evident that species nonindigenous to both Alaska and North America are arriving to Prince William Sound in segregated ballast water from domestic sources. Although we have only verified the species-level identification of a fraction of our vouchers from the vessels, we have identified at least 4 NIS of copepods arriving from domestic ports. In Table C-3, copepods identified originally as Oithona spp. include at least two different copepod species (Limnoithona sp. and Oithona davisae), and Pseudodiaptomus spp. includes at least two nonindigenous species (P.forbesi and P. marinus) (expert identifications by Frank Ferrari and Chad Walter, National Museum of Natural History). These copepods arrived in multiple vessels from San Francisco Bay, which was invaded by these copepods from Asia during the 1980s (Ferrari and Orsi, 1984; Orsi and Walter, 1991). It also appears that some of these NIS may also be arriving to Prince William Sound from other domestic ports. We are now in the process of separating the species complexes for Oithona and Pseudodiaptomus in each sample, to obtain counts of the individual nonindigenous and native species which are very difficult to discern based upon gross morphology.

The large volume of segregated ballast water and high density of organisms that derive from domestic source ports, which are heavily invaded by NIS, indicate a high probability ofreleasing NIS into Prince William Sound. For the 13 tankers sampled and analyzed that originate from domestic, continental ports (Table C-4)., these carried: • An average of 31,755 m' of segregated ballast water; • An average of7,000 organisms m' in the segregated ballast water; • A minimum diversity of 19.13 species per ship (excluding unresolved species complexes or the identification of most larvae). For this time period (late May- early June), we estimate approximately 244,000,000 organisms/ship arrive to Prince William Sound in segregated ballast water, multiplying the total volume of segregated ballast by the density of organisms (Table C-4). Roughly 600 tankers arrive to Prince William Sound per year currently (Tom Colby, pers. comm.). Although the density of plankton likely declines from fall to spring, a large transfer of plankton is occurring on an annual basis from domestic ports with tens to hundreds ofkno\\11 species that are not native to North .-\mer.ca (Tables B-1 and B-5). as well as native species that are not established in Alaska.

For the single tanker we sampled arriving from a fore:gn port, the density of coastal organisms was significantly lower than that for domestic traffic (Table C-5, Fig. C-3). More specifically, the densities measured for polychaetes, bivalves, and barnacles were all at least an order of magnitude lower than the average for the 13 ships from domestic ports analyzed to date, falling outside of the 95% confidence interval for these taxa. Copepods were a conspicuous exception to this pattern, as the total number of organisms were not significantly different between domestic and this foreign arrivals. The differences in relatiYe abundances of these taxonomic groups between domestic versus foreign ships may result from open-ocean ballast water exchange, which can reduce densities of coastal organisms and increase densities of oceanic copepods and phytoplankton (see next section). Although the same genera of copepods (e.g .. Oirhona) appear for both domestic and foreign vessels, these include multiple species that may differ between sources. Alternatively, but not mutually exclusive, the low relative abundance of coastal organisms on the foreign arrival may also reflect (a) low initial density and (b) low survivorship on the 10-day voyage- the longest for any of these ships sampled during this Pilot Study.

24 The only surprise in analysis of the foreign am val was in the abundance of larval barnacles. gastropods, and bryozoans. Although abundances of these coastal organisms were low compared to that for domestic arrivals (Table C-5), they represe:Jt a significant residual population when considering the total volume of segregated ballast water. Assuming a homogenous distribution among ballast tanks, our estimates of 10 organisms perm' for each snails, bryozoans, and barnacles as residual coastal plankton suggests that> 250,000 individuals of each species were delivered with 25,000 m3 of ballast water by this vessel (Table C-1). While these residual organisms may result from very dense initial populations that have undergone severe reductions during the exchange (see next section), some of the larvae may also be generated by resident adult populations within the ballast tanks themselves.

As with the ballast water from the foreign arrival, domestic ballast water from Hawaii on the OM! Columbia was relatively depauperate of plankton compared to ballast water of domestic, continental origin (Table C-3). However, unlike the foreign arrival, this vessel did not undergo ballast water exchange. Thus, the low plankton density reflects either a low initial density or poor survivorship, but we cannot distinguish between these potential causes at present.

Discussion

Quantitative analysis of segregated ballast water from the first 15 tankers demonstrates that these ships deliver relatively large inoculations of planktonic organisms to Prince William Sound on both an individual and cumulative basis. Most tankers arrive to Port Valdez from other domestic ports, and carry segregated ballast water of domestic origin from bays and estuaries. The average density of planktonic 3 organisms (7,000 individuals/m ) that we measured in this ballast water was roughly 10 times higher than previous studies from Coos Bay, Oregon and Chesapeake Bay, Maryland (Carlton and Geller, 1993; Smith et al., 1996). These densities, combined with the volume of ballast water per tanker and the number of tanker visits per year, indicate a large-scale transfer of organisms to Prince William Sound is occurring.

The high densities of planktonic organisms present in our quantitative analyses of domestic ballast water may result from any combination of factors. First, this water was only entrained for a short period of time, 3-7 days. In contrast, the ballast water sampled in other studies was often 7-10 days old, or older, and plankton density has been shown to decline with increasing age of water (Smith et al., 1996; Wonham eta!., 1996). Second, the collection of ballast water samples in Valdez coincided with a probable time of peak plankton abundance in the northern hemisphere. whereas previous studies have measured and averaged plankton abundances year-round. including significantly lower densities in winter. Nonetheless, even controlling for seasonal variation. we surmise the annual transfer of organisms is large relative to many port syster::s within the U.S., based upon magnitude of ballast water involved (e.g., Carlton eta!, 1995; Fig. A-4).

Although we detected a significant relationship between voyage duration (age of water) and plankton density among ships in this study, it was not as strong as reported elsewhere. For example, repeated sampling of the same ballast water \\ithin a ship has shown significant and exponential mortality over only a few days for densities of both individual taxa and total organisms (Gollasch eta!., 1995; Wonham et a!., 1996). However, although Smith et al. ( 1996) found a significant relationship of plankton density and duration of voyage among voyages that included a much larger range of 5-25 days, no relationship was evident for short voyages of 5-l 0 days. It is possible that initial variation in densities at the time of ballasting may swamp a stronger relationship of plankton survival with duration of voyage over this short range of 3-7 days for domestic voyages. The relative importance of initial communities versus survivorship in the observed variation among ships, compounded by voyages of various durations, can only be tested by comparing ballast water communities in the same tank at the start and finish of each voyage (however, see also below for discussion of repeated measures over a 3-day period). 25 The pattern of trans;er for plankton communities on tankers arriving to Port Valdez from foreign ports is very poorly resolved. Although plankton densities from the first and only tanker sampled to date from a foreign port earned few coastal organisms, relative to arrivals from domestic ports, it is premature to draw any conclusions about the magnitude of transfer or effectiveness of exchange. Tnis set of tankers from domestic ports illustrates the variation in plankton communities of segregated ballast water without exchange. Thus, we must increase the sample size to adequately understand the general patterns and variation associated with ballast transport from foreign ports.

For arrivals from both foreign and domestic ports, our focus thus far has been restricted to planktonic organisms that are sampled by an 80 micron mesh net. However, other organisms are certainly entrained in ballast tanks that deserve serious consideration as potential invaders. First, from our research on ballast water arriving to the Chesapeake Bay, we have documented the presence of mobile macrofauna (adult fish, crabs, shrimp) that are often missed by platikton nets (Smith et al., 1996; Ruiz et al., unpubl. data). We did in fact observe fish swimming in ballast tanks of at least 3 of the tankers sampled, although we only captured one in a plankton tow (Table C-3). Second, sediments in ballast tanks of tankers can apparently accumulate quickly (as they must be removed at regular intervals) and are likely colonized by many benthic organisms that include juvenile and adult invertebrates, as well as resting stages of amoebae to dinoflagellates and various invertebrates (Munson eta!., 1996; Smith et a!., 1996; Ruiz eta!., unpubl. data). Third, small zooplankton and phytoplatikton are certainly present that are not included in 80 micron net samples. Fourth, microorganisms such as bacteria and viruses are probably the most abundant organisms in ballast water arriving to Prince William Sound. For example, in the Chesapeake Bay, we routinely measure densities of 106 bacteria and 107 viruses per ml of ballast water, including many genera of potential pathogens. We plan to assess the abundance and diversity of the first two groups during our 2-year study, and will seek additional opportunities to explore the latter two groups.

We have confirmed 4 NIS (all copepods) arriving in domestic ballast water oftatikers to Port Valdez, and many more are surely delivered from both domestic ports and overseas that are not yet established in Alaska, but we cannot yet estimate the potential to invade or potential impacts for most species. The potential to invade is in part a function of physiological or environmental tolerance, which we will measure using laboratory experiments over the next two years with organisms collected from ballast tanks upon arrival to Port Valdez. Invasions are also a function of inoculation density. We will also estimate inoculation density of key taxa on tankers arriving to Port Valdez (from both domestic and foreign ports) over the next 2 years to test for a relationship with invasion success measured through faunal surveys.

26 D. Experimental Analyses of Ballast Water Management Practices

Purpose

To understand the consequences oivarious ballast water management practices for the transfer and potential risk of invasion for l'<1S in Pri.IJce William Sound, and more generally, we i.IJitiated two types of experimental measures i.IJ the Pilot Study. The first type of experiment measured the effect of open-ocean ballast water exchange on the density and diversity of organisms present in segregated ballast tanks, and the second type of experiment measured the effect of time (or voyage duration) on survivorship of organisms in segregated ballast water. As with our other analysis of segregated ballast water, these experiments occurred in May-June !997, corresponding to a season of relatively high reproductive activity and plankton abundance in waters of the northern hemisphere.

Methods

Ballast Water Exchange Experiments We measured the effect of ballast water exchange on the abundance and diversity of organisms in ballast tanks by comparing plankton samnles from tanks that underwent exchange with those on the same ship that were not exchanged. More specifically, two pairs of wing tanks were selected per vessel that initially contained water ballasted from the same source location, and at the same time. One tank (port or starboard) of each pair was selected in advance for ballast water exchange in open ocean, 500- 600 miles offshore, during transit from the source port to Valdez. Upon arrival to Valdez Marine Terminal, each of the four tanks per ship was sampled following the same standard procedure described for segregated ballast water.

The exchange experiments were conducted on two separate vessels: the SIR Long Beach and SIR Benicia. Each vessel arrived to Valdez directly from San Francisco Bay, after a 4-6 day voyage (Table C-1 ). Each vessel carried segregated ballast water from its last port of call, and each vessel exchanged its ballast in the two designated tanks offshore of British Columbia in depths of>2,000m (Table D-1). Although both tankers used the same flow-c.",rough method (instead of empty-refill method) for ballast water exchange, the SIR Benicia replaced 100% of its tanks' volumes and the SIR Long Beach replaced 300% of its tanks' volumes. These differe::ces in the volume of ballast water exchanged were selected to begin measuring the relationship between ,-olume exchanged and replacement of coastal biota.

Quantitative analysis and taxonomcc identification of plankton samples followed the same standard procedure as described above for segregated ballast water, and we compared the plankton communities of exchanged versus non-exc::anged ballast tanks for each vesseL Again, we treated each ship as the level of replication, estimating a mean density of each taxon by similar management condition (based upon the mean of 2 tanks. 'Jsing replicate plankton tows within tank to obtain tank­ specific estimates) and comparing the perce:n dirTerences between these means.

Plankton Survivorship Experimellt We measured the effect of time on plankton density, using repeated sampling over multiple days within the same segregated ballast water tanks of one tanker, to estimate the survivorship of various taxa during voyages of different duration. In th:s initial experiment, we measured changes in two different ballast water tanks aboard the Area Juneau. following its arrival to Port Valdez on 2 June 1997 with 4- day old ballast water from Puget Sound, W a.shington (Table C-1 ). For each of three consecutive days. we took 2 replicate plankton samples from each tank, following the standard sampling protocol described for segregated ballast water.

27 Results

Ballast Water Exchange Experiments For both the SIR Long Beach and SIR Benecia, ballast water exchange was associated with reduced densities of coastal organisms compared to non-exchanged tanks on the same vessels (Table D- 2). The SIR Benicia had noticeably fewer taxa, and often lower densities, in its non-exchanged tanks compared to the S/R Long Beach. However, for those taxa present on the SIR Benecia. the effect of exchange on entrained biota appeared very similar to that on the SIR. Long Beach (Figs. D-1 and D-2), despite a 3-fold difference in the amount of ballast water exchanged on the two vessels. Using the difference in salinity values between exchanged and non-exchanged tanks (Table D-1), we estimated the efficiency of exchange on rhe SIR Benecia as only 60% (= [35ppt -exchanged salinity] I [35ppt- non­ exchanged salinity] x 100), whereas that for the SIR Long Beach was between 70- 100%. In contrast, we estimated that the percent reduction of coastal organisms during exchange varied between 70-100% on each ship for those common taxa (>I 0 individuals I m'; Fig. D-1 ), with the lowest percent reduction appearing for the SIR Long Beach.

Despite the apparent overlap between tankers in estimated percent reduction for the common coastal taxa, our data suggest that the 300% exchange on the SIR Long Beach resulted in somewhat higher reductions of coastal organisms than the 100% exchange on the SIR Benecia. All but one of the coastal taxa exhibited reductions> 95% on the SIR Long Beach. The exception was the copepod genus Oithona, for which measures may be inaccurate due to an oceanic species that could have replaced the coastal congeners, obscuring a higher efficiency. Further taxonomic scrutiny of these samples is now underway. In comparison, three of the five coastal taxa exhibited reductions < 90% for the SIR Bene cia.

In contrast to coastal plankton, the abundance of some common oceanic taxa exhibited even larger differences. but in the opposite direction, between exchanged versus non-exchanged ballast tanks on these two vessels (Table D-2, Fig. D-2). This was most extreme for discoid diatoms. dinoflagellates in the genus Ceratiwn, and foraminefera, which were 200-10,000% more abundant in exchanged tanks.

Plankton Survivorship Experiment Our plankton analysis of segregated ballast for the first sample date indicated that the Area Juneau contained unusually high densities of crab larvae compared to the average across all ships (260 3 versus 25 individuals/m • respectively), while many other taxa appeared to be of roughly average densities (Table C-.\). In total, there were approximately 12 taxonomic groups wnh abundances sufficient to measc:re changes in density over time, given the level of variation among samples. Interestingly, this mcludes groups that are known to have relatively high impact as invaders (crabs, clams, and snails). as well as a nonindigenous species of copepod (Oithona sp.).

There was :10 clear pattern of change in the abundance of organisms in each tank of the Area Juneua among the 3 sample dates (approximately 48 hours; Table D-3). Some taxa. such as gastropods exhibited an order-of-magnitude decline, although it is not clear whether this was due to mortality or metamorphosis (and colonizing the bottom of tanks). Most taxa did not exhibit an appreciable change in abundance, given rhe high level of variation among samples. Also, some ta'la increased in abundance, resulting from metamorphosis of early larval stages into later stages (e.g., the copepods Acarria sp. and Limnoithona sp.) or possibly changes in distribution within the tanks (e.g., crab zoea).

Discussion

Our results from these initial ballast water exchange experiments provide the first measures of exchange efficiency for oil tankers and suggest that this management strategy may be effective at 28 reducing the transfer of nonindigenous species by thts class of vessels. The efficacy of exchange is roughly similar to the few existing measures for other types of commercial and m!litary vessels. With chemical and pamcle tracers. we have estimated a >90% efficiency for exchange on vessels that use an empty-refill method of exchange (Wonham, 1996; Ruiz eta!., 1997b ). In addition, Rigby eta!. (1993) measured exchange efficiencies of95% for water (using dye tracers) and between 75-95% for phytoplankton on a vessel using a flow-through method of exchange. Finally, a collaborative study between Australian and New Zealand researchers has just made similar measures for one voyage across the Tasman Sea; the results are not yet available, but the outcome appears to be similar in the efficiency measures (J. Hall, pers. comm.).

Despite the general similarity of results among these ballast exchange studies, it is important to recognize that these few studies (usually of only one vessel each) comprise all of the direct measures throughout the world on effectiveness ofballast exchange. This handful of vessels represents a fraction of the classes and designs in operation. Even these existing measures have only been made under a limited range of conditions (e.g., temperature, salinity, etc.) and have usually focused on a small subset of the taxa known to be entrained in ballast tanks (e.g., Carlton and Geller, 1993; Smith eta!., !996).

There is almost certainly variation in the effect of exchange among vessel types, "habitats" within tanks (e.g., bottom vs. surface vs. mid-water), and species which include a broad spectrum of sizes, mobilities, and densities. For example. resting stages of dinoflagellates and other organisms can accumulate on the bottom of ballast tanks, along with sediments and many other organisms, that may be relatively difficult to remove via ballast exchange compared to planktonic organisms, due to density and surface cohesion. This may explain the high densities of toxic dinoflagellates, up to 300 million viable cysts per ship, reported from the bottom sediments of cargo vessels entering Australia (Hallegraeff and Baisch, 1991, 1992); these resting stages are also found commonly on the bottom ofballast tanks entering U.S. ports (Kelly, 1993; Smith eta!., 1996).

At present, the relationships between the amount (or percent) of ballast water exchanged within a tank and (a) reduction of resident biota or (b) influence on invasion success remain unresolved. Our preliminary data suggest a slightly greater reduction of organisms with a 3-fold increase in volume exchanged. There are insufficient data for these tankers, or any other ships, to establish the rate function of decreasing biota with increasing exchange or to define an asymptote in this relationship (below which the return per unit effort diminishes. Intuitively, reduction of number of propagules released will diminish invasion success, and the shape of this function likely varies among species, depending upon particular aspects of biology and ecology. Yet, we cannot now evaluate whether a 90% reduction of organisms during ballast exchange results in a substantial decrease in invasion rate, or whether the residual l-1 0 organisms/m3 that we observed (equivalent to 30,000 - 300,000 organisms per ship) represent a significant risk of invasion. ). In any case. the effect of density or inoculation size on invasion success is unknown. Thus, although our two experimental measures of ballast exchange on tankers represent a significant advance in present knowledge. they provide only the initial steps in assessing the general patterns, efficiency, or consequences associated with ballast exchange.

Although limited to only one ship, the repeated measures of plankton densities in ballast tanks of the Arco Juneau suggest that survivorship of many ta'la is relatively high compared to other studies (e.g., Wonham 1986). The variation among samples is relatively high, but it is clear that densities did not decline exponentially. We cannot draw broader conclusions about comparative survivorship among taxa. Despite some apparent differences, this may result from differences in rates of development, metamorphosis, and colonization of bottom sediments. Also, there may be shifts in the distribution within the water column that affects density estimates. It is clear that the community is highly dynamic and that future work should estimate the relative importance of mortality, benthic colonization, and changes in water column distribution to the overall changes observed. 29 We plan an ambitwus senes of experiments over the next two-year study ( 1997 -1999) to extend these initial measures of ballast water exchange efficacy and plankton survivorship in oil tankers.

E. Plankton Characteristics of Non-Segregated Ballast Water Passing Through the Alyeska Ballast Water Treatment Facility

Purpose

Tankers arriving to Port Valdez, Alaska, carry two types of ballast water: segregated ballast water in tanks dedicated to ballast management; and non-segregated ballast water in tanks which are also used to carry oil. The oily ballast water is required to be treated by a shore-side Ballast Water Treatment Facility of the Valdez Marine Terminal to remove and recycle petroleum compounds before the water is discharged into Port Valdez. Plankton communities in non-segregated ballast water passing through the Ballast Water Treatment Facility were sampled to determine if non-indigenous species (NIS) are being released into Port Valdez afTer treatment. To determine whether the Treatment Facility affected plankton communities in oily ballast water, non-segregated ballast water was sampled and compared before and after the treatment process and following each of the two intermediate stages of treatment. Plankton communities in these samples were characterized for abundance and diversity of morphologically distinct taxa in the same marmer as biological sampling of segregated ballast, so that: the plankton communities in segregated and non-segregated ballast water could be compared; effects of each stage of treatment could be assessed for plankton passing through the plant; and the plankton communities entering the Treatment Facility could be compared to the plankton leaving it.

Methods

During May 23-June 4 1997, non-segregated ballast water discharged from 11 tankers arriving to Port Valdez was sampled from the Alyeska Ballast Water Treatment Facility. The First Mate of each ship was interviewed upon arrival to the Valdez Marine Terminal to determine the quantities of segregated and non-segregated ballast water transported during the voyage. Ballast water management practices were recorded on standardized data sheets to characterize ballast quantities by tank, location of water uptake, and date of loading. After recording the ballast water management data. samples of the discharged non-segregated 'Jallast water were obtained from 4 stages of the Treatment Facility to determine abundance. divers tty and viability of planktonic organisms at each stage of the process (Fig. E-1): ( 1) Chicksan Arms, which connect the tankers to the piping system of the Facility; (2) 90s Tanks, which receiYe the ballast water and allow it to settle for a period of ca. 4-12 hours so that oil separated from wate:- by difference in specific gravity, allowing the floating oil fraction to be pumped off and recycled. while the residual oil-water mixture was sent to the second stage of treatment; (3) Dissolved Air Filtration (DAF) Facility, which injects micro-bubbles of air into the water after adding a pol:mer. causmg petroleum chemicals both to adhere to the foam and to volatilize; and ( 4) Biological Treatment (BT) Tanks, which culture petroleum-metabolizing microbes in large outdoor ponds. so that their biological activities may remove remaining oil chemicals before the ballast water is finally discharged into Port Valdez through a diffuser pipe located at about 180 ft depth just offshore of Berth No.3. By sampling from the Chick.san arms, plankton of non-segregated ballast water could be characterized for each ship before it entered the treatment process. Wherever possible. the same parcel of water was sampled through time as it passed subsequent stages of the Treatment Facility; however. in most cases water from more than one ship (sometimes from several ships) was co-mingled in the 90s tanks and 30 subsequent stages as standard practice of the Facility. Therefore. co-mingled ware was often sampled at the subsequent stages of processing. Although this allowed us to test adequately for organisms at the later stages of the treatment process, effects of the process could not be tracked through the plant for specific parcels of water containing plankton from individual ships.

Sampling was timed to collect water during the middle of its transfer to the next stage. Thus, sampling of the Chicksan Arms began after pumping off the ship was well underway. Sampling the 90s Tanks occurred after the settlement period was completed and water was being transferred to the DAF Facility. Water leaving the DAF Facility was sampled after it had reached a midpoint in its transfer to the BT Tanks. And water from the BT Tanks was sampled at the overflow point as it passed into the discharge pipe.

Water at each point was collected as duplicate samples. For each of the first 3 sampling points, water was collected from spigots designed for the purpose of collecting water samples from the piping 3 system of the Treatment Facility. Water from these spigots was collected in 5 gallon (0.006 rn ) plastic buckets. For each sample of the Chicksan arms, the first 5 buckets of water were discarded to clear 3 residual water and/or oil in the pipe. The next I 0 buckets of water (total sample= 50 gallons, 0.06 rn ) were poured through a SO-micron mesh plankton net supponed over a drain to catch the filtered water. The plankton retained by the net was collected from the cod-end jar; care was taken to wash any plankton retained on the net into the sample jar. The duplicate samples of concentrated plankton were returned promptly to the Quanterra water chemistry laboratory on the Terminal for further processing. Water from the 90s Tanks and DAF Facility was sampled similarly to the Chicksan Arms, except that water was collected from the DAF Facility without discarding the first 5 buckets before filtering through the plankton net, because it was evident that the piping system had essentially no residual water in it. The Biological Treatment Tanks were sampled with duplicate plankton tows pulled vertically from near the bottom of the tank up through 4.5 rn of water column with a net 30 ern in diameter (total sample 3 volume= 0.32 rn ). For each sample at each point, the temperature and salinity of the water were measured in the collecting bucket with a calibrated alcohol thermometer and refractometer, respectively.

Within 30 minutes of delivery to the Quanterra laboratory, the water samples were inspected under a microscope (!0-40X zoom magnification) for qualitative assessment of plankton viability, and for initial determination of diversity and "abundance category" of the major taxonomic groups of plankton. Each sample was washed carefully into a finger bowl, and all major rnor,Jhologically distinct categories of taxa were identified. For each taxon identified. we estimated its categorical abundance as rare (present, but <10 individuals), common (10-100 individuals), and abundant (>100 individuals). For each taxon, the percent of individuals alive was estimated by evaluating their morphological integrity, movement and activity. However, viability of discoid diatoms was often difficult to determine with confidence. After initial microscopic examination of the fresh samples. they were either preserved for later detailed quantitative analysis (if they contained numerous organisms) or discarded (if they contained such low diversity and abundance of organisms that they were characterized adequately when fresh). Preserved samples were fixed in 5% buffered formalin solution and transported back to the Invasions Biology laboratory at the Smithsonian Environmental Research Center in Edgewater, Maryland.

Quantitative analysis of the fixed samples in the SERC laboratory used the following procedure. Samples were concentrated on an 80 micron sieve and washed into a finger bowl. Each whole sample was examined carefully under a stereo microscope (I 0-40X zoom magnification) and all morphologically distinct categories of taxa were identified. For taxa present in abundances of less than 100 individuals per sample, the number of individuals in the whole sample were counted. For abundant taxa(> I 00 individuals per sample), the samples were split with a Folsom plankton splitter to achieve

31 counts of 10-100 individuals per subsample (splits of 1/8 :o 1/32). For organisms in split samples, two subsamples were counted.

Results

Non-segregated ballast water comprised a mean of29,692 metric tons (range 0-60,305) per ship or 49% (range 0-75%) of the total ballast water on board oil tankers sampled upon arrival to Port Valdez during the study period (Table E-1). The source locations for the non-segregated ballast water that was sampled included the major port systems and adjacent waters for Valdez tanker traffic: Los Angeles/Long Beach, California; San Francisco Bay, California; Portland, Oregon; Puget Sound, Washington. A few ships also arrived from other locations, including: Barbers Point, Hawaii; and Korea.

A total of70 samples of non-segregated ballast water was collected from the Ballast Water Treatment Facility). Oily ballast water from 11 tankers was sampled with duplicate collections at the Chicksan Arms (N=22 samples total). Due to co-mingling of water, eight pairs of samples were collected as water departed from the 90s Tanks (N=l6 samples total), from water leaving the DAF Facility (N=l6 samples total), and from the BT Tanks (N=l6 samples total).

Temperature of the water samples averaged 11.5'C (range 10.75-15'C) (Table E-2). Salinity averaged 31 ppt (range 13-35 ppt) (Table E-2). Temperature and salinity of duplicate samples were quite similar, varying less than 1'C and 1 ppt.

A total of 23 taxa/morphological stages oflive and dead organisms were identified among the 70 samples (Table E-3). However, the percent occurrence of individual taxa in individual samples ranged from 0-94%. The diversity of taxa changed in the sample from sequential stages of treatment, with highest diversity occurring in Chicksan Arm samples and lowest diversity occurring in the DAF Facility samples. Dominant taxa in the samples were various stages of copepods, discoid diatoms, and nematodes. While the prevalence of discoid diatoms remained relatively constant in occurrence in samples from sequential stages of treatment, the occurrence ofcopepod stages declined and nematodes increased in occurrence in the sequence of treatment. Overall, the diversity of taxa was low compared to that of segregated ballast samples (see above sections).

Quantitative counts of total organisms (live and dead) in the samples showed similar patterns of low diversity (Table E-4). However, mean density of total organisms increased significantly in the 3 sequential stages of the Treatment Plant CANOVA, F3. 19=60.70. P

F3_19=!2.90, P

Discussion

The diversity and abundance of planktonic organisms in non-segregated ballast water discharged from the G.nkers were low as water left the ships and entered the Treannent Facility. Plankton diversity and abundance remained low throughout the treatment process. However, discoid diatoms and nematodes increased markedly in abundance in the Biological Treatment stage, probably reflecting a process of culturing of these organisms in the BT Tanks.

The source of these two taxa is not evident. They could originate at low a':lundances from the non-segregated ballast water, and then increase under conditions of the BT Tanks. Alte:natively, these taxa are readily transported by wind and may simply colonize the BT Tanks, where they find favorable conditions for population growth. Despite the increase of these two taxa in the BT Tanks. their apparent viability was low in the samples collected from water on the way to the discharge ?ipe. The viability of all other planktonic organisms was low at all stages of the treannent process, indicating that conditions in the non-segregated ballast water and/or the Ballast Water Treatment Facility was ::ot fa,·orable for survival of plankton.

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47 Map of Pnncc Will ram Soun d .s·h owrng 1o- { JIJOrl\ fd- ValUe;: and Aiycc.k<.J Tcrm1nal

Valdez

terminal

n

Prince William Sound

L ____Alaska_

The opinio11s expressed in this RCAC commissioned report are 11ot necessarily those of . RCA C. F1gure A-2 MaJor sh1ppmg routes of commercial vessels arriving to Port Valdez I Prince William Sound.

Washmgton l Oregon Korea f)

~-=_Japan c l"f . - vpv ------~\a 1 orrna

Taiwan

Hawaii Additional Ports Panama Netherlands Caribbean Ftgure A-3. l

Columbia River, unknown St. Lucia Japan Oregon 1.1% 0.6% 0.5% Hawaii 3.5% \ Netherlands St. Croix Taiwan 4.3% .... 0.2% 0 2% Nikiski, Alaska 5.7%

Los Angeles/Long Beach, California Puget Sound, 15.9% Washington 38.0%

San Francisco Bay, California 28.7% Figure A-4. Charactensllcs ofOtl Tanker traffic arriving to Port Valdez for 1987-1994. Shown are the number of tankers that VISI! the port each year and the cumulallve number of amvals for these vessels each year. (Source: Wiegers et al. !997.)

1000

900

800

(/) 700 CIJ (/) (/) 600 CIJ > .... !l!l Number of Tankers 0 500 .... • Number of Port Calls for Tankers CIJ .0 400 E :J z 300

200

100

0 1987 1988 1989 1990 1991 1992 1993 1994 Year Figure A-5. btunatcd annual quantity of segregated ballast water discharged into Chesapeake Bay, Prince William Sound, and San FranctSco Bay from commercial vessels. Estimates were based upon information from the following sources: Smith et al., 1996 and Ruiz et al .. unpubl. data (Chesapeake Bay); Wiegers et al. 1997 (Prince William Sound); Carlton et al., 1995 (San Francisco Bay). (Note: San Francisco Bay estumues do not mclude ballast water from domestic sources, whereas estimates for the other two sites include domestic and foreign sources. I

35000000

~ ~ 30000000 0 -0 .::: 25000000 1------·------<1l E ~ "0 20000000 <1l tJ) -co co 15000000 .Q <1l 0 -c: 10000000 :J 0 E soooooo <(

0

0 Chesapeake Bay: Estimated from 1996 Bulker traffic into Baltimore and Norfolk.

[ill Prince William Sound: Estimated from Tanker Traffic.

• San Francisco Bay: Estimated from 1991 Bulker, General Cargo, and Tanker Traffic from Foreign Ports 1nto San Franctsco and Oakland. Table B-1. Number of nomndigenous species reported for marine and estuarine habitats at various sites and global regions.

Region No. of NIS References

San Francisco Bay, California (USA) 212 Carlton, 1979a: Cohen and Carlton, 1996 ' Coos Bay, Oregon (USA) 70 Carlton, unpublished manuscript Great Lakes (USA) 138 Mills et. a!., 1993 Hudson River (USA) 113 Mills et. al., 1996 Chesapeake Bay (USA) 130 Ruiz et. al., 1997 in progress Hawaii (USA) 150 Eldredge ct. al., in progress Australia 80 Pollard and Hutchings, 1990a, b; Thresher and Martin, 1995 Japan 30 Asakura 1992; Ruiz et. al., unpublished database New Zealand 80 B. Hayden, personal communication Mediterranean Sea 240 Por, 1978; Boudouresque, 1994; Ribera, 1994 Northern Europe 75 Jansson, 1994; Leppakoski 1994; Eno, 1997 rablc B 2 Nonlndlgcnou\ '-f1CLIC'- rerortcd Cor mar:nt: ,lnd t:\lUdflllt: h<.Jbii

Slolar<>kl. 1991.Lcrrako<;k 1• 1991 Jann,on. 1994. ()FJVccrcl al 199))

r.T~Ar.X~O~Nr.------~S~Pr.Er.Cr.I~Er.S------~C~O~M~M~O~N~N~A~M~E------~N~A~Tr.I~V~E~------~

Plant a Bacdlanophyceae Odonte!la smens1s lndoPacrflc Coscmod1SCUS waJ!esu PleurosJa tea v1s I polymorpna ThalasSIOSira sp Charaphyceae Chara conmvens Europe 01nophyceae Procentrum minimum cosmopolitan Spermatophyta Elodea canadens1s Canadian water weed N. America Invertebrata Cnideria Hydrozoa Cordyfophora caspia freshwater hydroid Black & Caspian Seas Annelida Polychaeta Poiydora redeki W. Europe Marenzellen'a viridis N. America Mollusca Bivalvia Oreissena polymorpha zebra mussel Black & Caspian Seas Mya arenaria softshell clam N. America Gastropoda Potomopyrgus antepodirum New Zealand mud sna1l New Zealand Arthropoda Decapod a Eriocheir sinensis Chinese mitten crab SE Asia Orconectes limosus N. America Rhithropanapeus harrisi mudcrab N. America Copepoda Acartia tonsa N. America Amphipoda Corophium cutvispinum Black & Caspian Seas Cirripedia Balanus improvisus bay barnacle N. America Mysidacea Hemimysis anoma/a Black & Caspian Seas Mesomysls kowalewski Black & Caspian Seas Cladocera Cercopagus pengo! Caspian Sea Bryozoa Cheilostomata Victore!la pav;da Black & Caspian Seas? Chordata Fish Acipenser guldenstaedtri Europe A baed Central Asia Oncorhynchus mykiss rainbow trout N. Pacific 0. gorbuscha pink salmon N. Pacific 0. nerka sockeye salmon N. Pacific 0 kera chum salmon N. Pacific 0 tshawytscha chmook salmon N. Pacific Safvelinus namaycush lake !rout N. America S. fontinahs brook trout N. America Catostomus catostomus sucker Asia Coregonus pefed CISCO C. Asia lctalurus me/as black bullhead catfrsh N. America Cypnnus carp1o common carp Asra Neogogius melanastomus round go by Slack Sea Branta canadensis Canada goose N Amenca Cygnus olor mute swan C As1a Mammal Ondatra zibeth1ca muskrat N America Musteta vtson m1nk N America L1 blc B- I Non1nU 1gcrHJU' ,rccJC\ n.:rortctl for m

TAXON SPECIES COMMON NAME NATIVE I Pfan!a 01nophyceae Alexandnum tamarense Global Temper ate Gymnod1mum catenatum red !tde E Pacrftc and N Europe I Phaeophyceae Undana pmnatifida Japan. Korea. Cl"llna Invertebrata Annelida Polychaeta Hydro1des elegans Europe Sabella spallanzanii Europe, Med Arthropoda Oecapoda Carcinus maenas green crab N. Europe Cancer novaezelandiae New Zealand Halicarcinus innominatus New Zealand Petrolisthes elongatus New Zealand Echinodermata Asteroidea Asterias amurensis NW Pacific Astrostole scabra New Zealand Pateriella regularis New Zealand Bryozoa Cheilostomata Cryptosula pallasiana N Atlantic, cosmopolitan Membranipora membranacea N Atlantic, cosmopolitan Mollusca Bivalvia Corbu/a gibbs SE Asia Crassostrea gigas Japanese oyster Japan Musculita senhousia NWAtlantic Neilo australis New Zealand Perna canaliculatis New Zealand Soletellina donacioides New Zealand? Teredo navalis common shipwonn Europe Theora lubrica NW Pacific Venerupis largillierti New Zealand Gastropoda Maoricotpus roseus New Zealand Polyp!acophora Chiton glaucus New Zealand Chordata Asc1diacea Ascidiella aspersa N Europe Borryi!us schlossen' NE Atlantic Ctona mtestinalis N AtlantiC Fish Forsterygion varium New Zealand Oncorhynchus mykiss rainbow trout NE Pactfic Satmo salar atlantic salmon N Amenca Saimo trutta brown trout 8nta1n ! dhk B .J '\;rHllfHlrc:·-·" "', ,rc'-''-> :-·~pllrlt:d !rJr rndrrnc .rmi c· ,tiJdflfiC h,rhHJl\ rd ( ift:<~l HrrLrlfr /he

nl.ltrvt:. rcg1r;n .~nJ Ll!fllt: ,,~ n,ttnc r' ·mlH..dtcd lflr ,reLiC' whcr-..: ,~vdtLrhlc Ill (lrrgur

TAXON SPEC:ES COMMON NAME NA Ti'JE Planta Coscrnoalscopnyce;:~? Thaias:;;os,ra puc11gera ThalasstosJra tealata CosonodiSCus waJies11 Odonte!la smens1s Bacr!lanophycoceae Pfeuros,gma pfanctomcum Rhodophyceae Asparagops1s armata Bonnema1S0ma hamtfera Ptkea ciadormca Grateloup1a liiCJna var luxunans Grateloup1a doryphora Agardh1ella subulata Soliena (tenera) filiform1s Solieria chorda/is Antithamnionella splfographidis Anthithamnionella temifolia Polysiphonia harveyi Phaeophyceae Colpomenia peregrina Undaria pinnati!ida Chlorophyceae Codium fragile atlanticum Codium fragile tomentosoides dead man's fingers Magnoliopsida Spartina angtica Invertebrata Cnidaria Hydrozoa Gonionemus vertens lndoPacific? Europe? Rhizogeton nudum Clavopsella navis Anthozoa Haiiptanella llneata striped anemone Japan, Kor~a Annelida Polychaeta Gonidaella gracilis N America (Atlantic) Marenze!leria viridis N America (Atlantic) Clymene/la torquata N America (Atlantic) Hydroides dianthus N America (Atlantic) Hydroides ezoensis Japan, Korea Acopomatus enigmaticus Australia Janua brasiliensis S America Pi/eo/aria berkeleyana Japan. Korea Pycnogonida Ammothea hiigendorfi Japan, Korea Arthropoda Cirripedia Elminius modestus Australia. New Zealand Balanus amphitrite Copepoda Acartia tonsa Ostracoda Eusars1efla zostencola N America (Atlantic) Amph1poda Corophium sextonae New Zealand Decapod a Eriocheir sinems Chtnese mitten crab JndoPaclfic Mollusca Gastropoda Crepidula torn1ca ta shpper shell N Amerrca (AtlantiC) Rapana venosa Urosalpmx cinerea N America (At!anttc) Aufacomya ater S Amenca (PaCific) Btvaivta Crassostrea g1gas Jaoanese oyster Japan Tiostrea lurana New Zealand Ens1s amencanus razor clam N Amenca (AtlantiC) Mercenana mercenana hardshell clam N Amenca (Atlant1c) Oetnco!a phofadllormts N Amenca (AtlantiC) \1ya arenana soflst;ell clam N Amenca {Atlantrc) ASCidJacea Stye/a clava Asta Nematoda Drac>JnCulotdea 4ngwlhcola crassus lndoPac!lrc") Europe? Chordata Mammal Myocastor coypus nutna South Amenca TABLE B-5. Nomndigcuous species repot1cd from marine aud estuarine habitats ffom southern California to Alaska.Siwwn for each species are its native range, its range along the coast of North America, and associated references. This list summarizes the current state of knowledge concerning nonindigenous species in each region based upon existing and often incomplete surveys. (see text fOr discussion)

Geographic ltange Abbreviations and lteferences; SF"" San Francisco Bay (Carlton, 1979:t; Cohen and Carllon, !996) SCA=Southcrn California, below Snn Fmncisco nay (Carlton,1979a; Crooks, 1997; I.amhert

TAXON SPECIES COMMON NAME NATIVE TO SF SCA NCA OR NW AK Chlorophyta Bryopsis sp ? X -- Codiwnfragile tomentasoides dead man's fingers Japan X ' Diatomacea Gonioceros armatum Australasia X X Pseudonitzchia australis X X X Phaeophyta Sar~asswn muticum Japanese weed Japan X X X __x_ ·' ' Rhodophyta ("allithammon byssoides NW Atlantic X X Gelidium vagum .\ - l.omemarw hakodatensis Japan X ' - l'tkeu yoshizakii Japan X Potysiphonta denudara NW Atlantic X Angiospenn Agrostis alba red top grass Europe X Agrostis maritima creeping bent grass Europe X Chenopodium macrospermum gooseMfoot S.America X X Coat! a corvnopifolia brass buttons S.Africa X X ------· Juncus gerardi saltmeadow rush NWAtlantic X X Lepidium latifolium broadleaf peppergrass Eurasia X Limosella subu!ata awiMieaved mudwort Europe, eN.America X Lyihrum salicaria purple loosestrife Europe X Myriophyllum aquaticum parrot's feather S.America X Myriophyllum spicatum Eurasian milfoil Eurasia, N .Africa X ' Po(vgonum patulum smartweed eEurope X Rorippa nasturlium aquaticwn true watercress Europe X Sa/sola soda saltwort s.Europe X ,)'pergularia marina ' S'pergularia media sand-spurrey Europe X

Spergu/aria salina saltmarsh sand-spurrey Europe X

l:'geria densa(m) elodea S.America X

Eichornia crassipes water hyacinth S.America X Ins pseudacorus yellow flag Europe X - ---·· -·-- --·-·~"" ···~"·- Table B- 5, continued ... TAXON SPECIES COMMON NAME NATIVE TO " SF SCA NCA OR NW-- __::___\K Polypogon elongaws S.Ameria X Pvwmvgeton cnspus curly-leaf pondweed Europe X

S'purllna altern~flora smooth cordgrass ow Atlantic X X X .)'part ina anglica English cordgrass England X

Spartmu densijlora dense-flowered cordgrass Chile X Spartina parens saltmeadow cordgrass seUS X Typha angustfo/ia narrow-leaf cattail Eurasia X Zostera j{Iponica Japanese eelgrass Japan X X Protozoa( Foraminifera) Trochammina hadai Japan X Protozoa( Mollusc host) Ancistrocoma pelseneeri Europe X Ancistrwn cyclidoides Europe X Jloveria teredinidi nAtlantic X ------Sphe nophyra dosiniae Europe X Protozoa( Crustacean host) Cothurnia limnoriae ? X Lobochona prorates ? X

Mirofolliculina sp. ? X X Porifera C/iona sp. boring sponge nAtlantic? X ' N llalichondria bowerbanki Bowerbank's halichondria nAtlantic X __x_ __x_ --- /Ia/idona loosanofji Loosanoff's halicolona nAtlantic X X

klicrociona pro/ifera red beard sponge ow Atlantic X X Prosuherites sp nwAtlantic X

Cnidaria(llydrozoa) IJ!ackfordia Vlrginica Black&Caspian Seas X X Cladonema uchidai Japan X Clava mu/Jicornis club hydroid ow Atlantic X ( "only/ophora caspia freshwater hydroid Black&Caspian Seas X X X ' ---- ( 'on'nunplw SfJ nAtlantic? X Garveia /ranciscana nlndian Ocean? X ( ionothyraea clarki nAtlantic X X II ---- A4aeotttl.\" tnexspectata Black Sea X Obe/w spp -----~ nAtlantic X ' II .\'arsia tubulosa ("" .\'yncoryne minabilis) nAtlantic X X f:(:top!eura crocea (..oo Tubularia) _,' ' ow Atlantic X X X ' lane/eo t'O.'italll nwAtlantic X X -"~ .. -·- Cnidaria( Scyphozoa) Aurelia "aurita" moon jelly nwPacific X # Cnidaria(Anthozoa) Diadumene ?cincta orange anemone Europe? X

Diadwnene franciscana San Francisco anemone ? X X

Diadwnene leuco/ena white anemone nwAtlantic X X X Halip/ane/la luciae orange~striped green anemone Japan X X X X X Platyhelm(T urbellaria) Pseudosty/ochus ostreophagus Japan X X X ' Leptoplana limnoria X Table B - S, conluwcd TAXON SI'FCIJ::S•, COMMON NAME NATIVE TO SF SCA NCA OR NW AK Annelida(Oiigochacla) Hranchiura sowerby1 Asia X Limnodri/oides monothecus nwAtlantic X # Parana is Jrici Caspian&Biack Sea X II Potamothrix bavaricus Eurasia X Tubificoides apectinatus nAtlantic X # Tubificoides brownae nAtlantic X X Tuberficoides diazi X Tubificoides wasse/li nwAtlantic X Varichaetadrilus anguslipenis eUS X Annelida(Polychaeta) Boccardiella /igerica nwEuropean coast X Capitella spp. __x_ nAtlantic? wPacific? ------__x_ II \ Crucigera websleri nwAtlantic X X Eteone tchangsii X ·--- --~-- Ficupomatw; enigmaticus Australian tubewonn Australia X Jleleromastus fil iformis ow Atlantic X X __x_ _-2__ \ .\ !lydroides elegans X

Lycastopsis pontica X X

Alanayunkia spec10sa eN.America X Marenzelleria viridis ow Atlantic X ----- ,\larphysa sangumea nAtlantic? X

Nereis acununata ? X X Nereis succmea pile wonn nAtlantic? X X X X Pionosylis uraga X

Polydora cornut a("""' ligm) mudworm nAtlantic X X X X X Potami/la sp. ? X

Pseudopolydora kempi Indian Ocean or nwPacific X X X X X Pseudopo/ydora paucibranchiata Japan? X X X .)'abaco e/ongatus ham boo worm nwAtlantic X Sabella sp. X

Streblospio benedicti Atlantic X X X X X lharyx lessalata X Mollusca( Gastropoda~ prJ /JusycOiypus canaliculatus channeled whelk nwAtlantic X

JJatillaria zona/is (·"'-allramentaria) Japanese false cerith nwPacitic X X X .\

Cecina manchurica nwPacific X ..\

Ceratosroma ino.rnatum Japanese oyster drill nwPacific X ..\ Cipangopahidina chinensis Chinese mystery snail China,Japan X Crepidula convexa convex slipper shell nwAtlantic X

( 'repidulafornicata Atlantic slipper nwAtlantic X .\

('repidula plana e.white slipper shell nwAtlantic X .\

1/yanassa ohsolela e.mudsnail nwAtlantic X ..\ 1-lflormu 1·axatills rough periwinkle nAtlantic -- X -- Table B- 5, conlintJcd

I~~Q!' __~--- ______SI'H '11·;~---~-- ______co~~ON NAMI,c__ _ NATIVE TO SF SCA NCA Oil N\1' C~---~- --- -~ "_,_!,_ --- ..\J.:fanoules tubcrculata red-nun mclania Africa to E.l ndies X N!lntwws /lcllerculus Japanese nassa nwPucitic X --- Oceanebru mornata Japan ' " -- ( Jceanubra mornatu Japan X /juwmupyrgus antipodarwn New Zealand mud snail New Zealan d _,

Urosalpinx cinerea Atlantic oyster drill nwAtlantic X X X X Mollusca( Gastropoda-op) Aeolidie/la takanosimosis Vennillion Japanese aeolis Japan X Babakina fest iva single-stalk aeolis Japan X Boonea bisuturalis two~groove odostome nwAtlantic X Catriona ricketlsi ? X X Cumanotus beaumonti polyp a eo lis nwAtlantic X X # Cuthona perca Lake Merritt cuthona ? X Eubranchus misakiensis Misaki balloon aeolis Japan? X

Okenia plana flat okenia Japan X X Philine auriformis tortetlini snail N.Zealand; Australia? X X X Sakuraelois enosimensis white-tentacled Jap.snail Japan X

Tenel/ia adspersa miniature aeolis Europe X X X Mollusca(Gasropoda-pul) Myosotella myosotis (=Ovate /Ia) Europe? X X X X X Mollusca(Bivalvia} Arcuatula demissa ribbed mussel nwAtlantic X X Anomia chinensis Chinese jingle nwPacific X ------( 'orbicula jluminea Asian clam China,Kore a, Japan X X X ___, ___ ---- ( 'ron·o.\Hea );/gas Japanese oyster Japan _, ~-- X ---- ( ·rassostreu vtrguuca Eastern oyster nwAtlantic _, ( iemma gemma amethyst gem clam nwAtlantic X X ---- Lwerrw/a Jimicola X J.yrodtu pedice/latu.\· blacktip shipwonn ? X X ,\lacouw petaium Ualtic clam nwAtlantic X Mercenaria mercenana northern quahog nwAtlantic X

Musc1t/ista senhousia Japanese mussel Japan,China X X X X Alya arenaria softshell clam nAtlantic X X X X X Mytilus galioprovincia/is Mediterran. mussel Med. Sea X X X Nurlal/ia obscura Japan? Kor ea? __x_ ---- (Jstrea conchophila Olympia oyster ePacific --- X ·------/JetrictJ/a plw/ad~formis flllse angclwing nwAtlantic X X __x_ --- Poramocorbula amurensis Amur River corbula sChina tosS'iberia, Japan X Teredo navalis naval shipwonn ? X X X X 1heorafragi/is Asian semele wPacific X X Trapezium liratum Japanese trapezium nwPacific X X - Venerupis philippinarwn Japanese littleneck clam wPacific X X X X ·' Table B ~ 5, coutiiHJCd ..

TAXON S!'EC!EE~S ______COMMON NAME NATIVFTO" SF SCA NCA OH NW AK~- Arthropoda( Ostracoda) Asp1dochorda Jimnoriae Europe X X Eusarsie/la zosiericola nwAtlantic X .)'pmilebcns quadracu/ata Japan X -- l

Limnothona tetraspina YangtzeR., China X Mytilicola orienta/is wPacific X X X X Oithona davisae Japan X P seudodiap!Omas forbesi YangtzeR., China X

Pseudodiap1omas inopinus Asia X Pseudodiaptornas marinus China,Japan X Sinocalanus doerrii China X Tisbe gracilis Europe X Tor/anus sp. ? X ------Arthropoda(Cirripedia) lJalanw· amphilrile stri ped barnacle Indian ocean X X ' Balanus improvisus bay barnacle nAtlantic X X X Arthropoda(Neballia) E::pinebalia jp. ? X Arthropoda(M ysidacea) Acanthomysis aspera Japan X Acanthomysis sp ? X !Je/IamY.\'iS Jwlmquistae ? X Arthropoda( Cumacca) .Nippo/euum hinumetuis Japan X X Arthropoda( I so pod a) ( 'oenjaeru horvathi X Dynoides denlisinus Japan,Korea X Lurylana arcuata N.Zealand or Chile X -----·------(inorimosphaeroma rayi Japan X /a is californica Australia,N .Zealand X X X X Umnorta quadripunc/ata gri bble ? X X X Umnorw tripuncwta gri bble ? X X X X jp. Paranthura wPacific? X Sphaeroma quoyanum Australia,N.Zealand X X X X .)fJiweroma walkeri Indian Ocean X .':lynidotea /aevidona/is nwPacific X Arthropoda(Tanaidacca) Sinelobus sp (=- Tanais starifordi) ? X X Arthropoda(Amphipoda) Arnpe!isca abdita nwAtlantic X X Ampithoe longimana X A mpit hoe valida nwAtlantic X X x X Capre/la acanthogaster X Capre/la mutica Japan to Vladivostok X X X Chelura terebrans Atlantic X X X Table B · 5, continued ..

TAX Of'! §!'l(f_'!I:~S C'OI\!MON NAME NATIVE TO SF SC!L_i'l~~~ ,Q!! N\1 A h. ( 'orophwm acherusicwn Atlantic X X X X X X Corophium a/ienense seAsia? X ( 'oruphiwn heteroceratum China X

('orophium lm"idio:sum nAtlanlic X X X X X X ( 'orophium uen01 Japan X X X Euobrolgus spinosus (=Paraphoxus ) ow Atlantic X X 1-- Gammarus daiberi nwAtlantic X

G randidierella japonica Japan X X X Ja.na marmurata nwAtlantic X X Leucolhoe sp. ? X

.He/ita nitida nwAtlantic X X Melita sp. ? X

Paradexamine sp. wPacific? X

Parapleustes derzhavini wPacific? X X .)'tenothoe valida ? X X

Transorchestia enigmatica shorehopper Chile? N.Zealand? X

Arthropoda(Decapoda) Carcinus maenas green crab Europe X X X Eriocheir sinensis Chinese mitten crab China,Korea X X

Exopalaemon modestus China,Korea,Russia? X

Oronectes virilis virile crayfish midwU.S. X Pacifastacus leniusculus signal crayfish Oregon to B.C. X

l'alaemon macrodacty/w; oriental shrimp Korca.Japan,nChina X X X Procambrus c/arkii red swamp crayfish seU.S. X ------Uhithi"OJNlllfiJ'CIU harri'>ii llarris nuulcrah __x_ --- nwAIIanlic --- I X Arthropoda( Insecta) Ani:wlabis maritima maritime earwig nAtlantic X Neochetina hruchi Argentina X NeocheJina eichorniae Argentina X ht>-:onotylus uhleri cordgrass bug nwAtlantic X ·--- Karnpti)J.()a Uan.:nlsia hcnedem Europe --- X X X --"- --- l Jrnatdla gracilis e&midwU.S. X

Bryozoa /llcyumdtum sp wPacific? X X ------,-1ngwne!la palmaw ambiguous bryozoan nAtlantic X X -- IJowerlnmkw gracilts creeping bryozoan nwAtlantic? X X X II ·----- Bugula "neritina" ? X X X Bugula stolonifera nwAtlantic X X

Conopeum tenuissimum nwAtlantic X X Ctyplosu/a pal/asiana nAtlantic X X X # No/ella b/akei X .)'chizoporel!a unicornis nwPacific X X X X X hiricella sp wPacific? X X 'tel I orella puv1da Indian Ocean? X TAXON ~l't < IL~!------'(c:'<.:cliV"'l"'M"-ON NAM F. NATIVF TO SF SC \ N( \ OH ' r-'-- Nil \l~ Waterstporu "subwrquata" nwPacific? X X X --- Loobvtryon verllci/lawm subtropical? X X Chordata(T un1cata) Ascidia sp ? X Ascidia interrupta X /Jotryllus aurantius Japan X JJotryllus sch/osseri golden star h micate neAtlantic X X Botry/lus vio/aceus (=Botry/oides ) nwPacific X X Botryllus sp. ? X X Ciona intestina/is nAtlantic X X X .x Ciona savignyi Japan? X X Diplosoma milsakurii nwPacific X X !vficrocosmus squamiger X Molgula manhauensis owAtlantic X X X X Polyandrocarpa zorritensis X Stye/a canopus X Stye/a clava nChina to Okhotsh Sea X X .\ # - S1yela plicata X "')'ymplegma oceania X Chordata(Fi'>h) ------·-----·- Acumhogobius jlavimanus yellow fin go by Japan,sKorea,China X X Alosa sapidissima American sh ad Labrador to Florida X .\ ;fliiL'/1/F/IS CalliS -- white catfish New York to Miss. X Ameturus me/as black bull he ad central N.America X Amewrus nwalts yellow bullh ead central N.America X :lmemrus nehulosus brown bullhead central N.America X ( ·aru.Htus uurotus ------·· -~ .. --·· goldfish China X Cyprinodon variegatus ·--- sheepshead m in now wN.Atlantic X Cyprinus carpio carp Eurasia X Dorosoma petenense threadfin shad midw U.S., Florida·Guat X (iambusia c~/jinis rnosquitofisl1 midw/scU.S., Mexico X ' .\ --·- lctalurus }itrcatus blue catfish midw/seU.S., RioGrand,Mex X Lepomis c.yanellus green sunfis h midw/seU.S., nMex X Lepomis gulosus warmouth midw/seU.S., RioGrande X ---·- --~ .. - l.cpomis lllltcrocl1irus bluegill --- -·-- midw/seU.S.,Mex,RioGR X -- !.epomis microlophus redear sun tis h midw/seU.S. X Lucuma parva rainwater kil lifish Mass.toMex, RioGR X Menidia beryl/ina inland silverside midw/seUS, RioGR X -- Micropferus ch>lomieu srnallrnouth bass central N.Am X Micropferus salmoides ---t-·-- largemouth b ass central N.Am X TAXON SI'FCIFS- -- COMMON NAME NATIVE TO SF SCA NCA Ott NW ..\h ,.,_ A'lorom: chry.sops x saxalili.s white bass (hybrid) wAtlantic X Marone saxatilis striped bass StLawrence - Louisiana X X Notemigonus crysoleucas golden shiner central N.America X Percma macrolepida bigscale logperch Louisiana- N.Mexico X 1 fathead minnow central N.America X l 1mephales promelas -- l1 umoxis amwlaris white crappie midw/se U.S. X J»omoxis nigromaculatus black crappie midw.se U.S. X .\'u/mu sular Atlantic salmon Atlantic .\ ' hulenliger bifacullu.'i shimofuri goby Japan X J rulenllfo:t'r tngonocephalus chnmelcon goby Japan,China,Siberia X ---- Chordata(Amphibian) l

------·---··~---·----· TAXON SPECIES COMMON NAME NATIVE TO MECIIANISM REFERENCE Angiospemt • Myriophyllum spicatum Eurasian mil foil Eurasia, N.Africa Rl Susan Walker, USFWS, pers. COilHlL Rhodophyta Sargassum nwticum Japan Scagel et. al., 1989 Cnidaria(llydrozoa) • Sarsia tubu/osa nAtlantic SF Carlton, 1979a • Tubularia crocea ow Atlantic SF, CO Carlton, 1979a Annelida( Pol ychaeta) * Capitella capilata NeAtlantic Carlton, 1979a

Jleteromastus fil iformis ow Atlantic BW,OA Wiegers et. al., 1997 1, Mollusca(Bivalvia) •• Crassostrea gigas nwPacific co Carlton, 1979a Mya arenaria softshell clam nAtlantic OA,IP Carlton, 1979a A rtl1ropoda( Atnpl1ipoda) • C'orophium acherusicum Crawf()rd, 1937 • Corophium insidiosum nAtlantic OA,SF Crawford, 1937 Chordata(Tunicata) • Ciona inte.stinalis Carlton, 1979a Chor~ata(Pisces) • • Salmo sa/ar Atlantic salmon Atlantic IP Robert _Benda, pers. co!!.l_. --~ .. ---·· r alllo B 1 Alqiil ~;puclo!; w1\h IHI!;uccu~~~~~l u11rodudion In Northwostorn North Amotlcu lrmh Wu!>hlnglon to Alm.>kn. lndlcatud lw uach :>pHclut. are donor reg1on(s). recipient region(s), date of lirst record, probable vector ot introduction and relerence(s). (Source. Table provided by G. Hansen]

Abbreviations: Donor/Recipient Regions: AUS-TAS=Tasmania, Australia, AUS=Austra!ia, BC=Britlsh Columbia, C-SFB=San Francisco Bay, C=Cahlorrua, CAN-NF""Newloundland, CAN-NS=Nova Scotia, CAN-O=Ouebec, CH=China, EUR=Europe, FA-BRIT =France, Brittany, FR-MEO=French Mediterranean. FR""France, GB=Great Britain, HEL=Helgoland-Germany, IT=Italy, J=Japan, MED=Mediterranean, MX=Mexico, N=No!Way, NETH=Netherlands. NE-ATL= Northeast Atlantic. NE-PAC= Northeast Pacific, NW·ATL=Northwest Atlantic, NW-PAC= Northwest pacific, N-ATL= North Atlanllc, N·PAC=North Pactlic. NY=New York, Long Island Sound, NZ=New Zealand, O=Oregon, PAC=Pacific, PWS=Prince William Sound, R·ARC=Auss1an Arcttc, Whtle Sea, A-SIB::::Siberia, S·ATL= South Atlantic, S·AUS=Southern Australia, S·C=Southem California, SCANo:::Scandinavta. SE·AK=Southeast Alaska, SPo:::Spatn, USA·NC=North Carolina, USA-NE=New England, W:::::Washington Vector: A~AquacuHure, B=ballast water. L=lobster or bait packing, M=Marginal dispersal through currents, OA=oystersAIIantic,OJ=oyslersJapan. OBC=oysters Bnttsh Columbia. ROK=roe-on·kelp; S=sCIBntific research, ?::unknown vector or date

~l'f.Qj I'J> QQMQB ~flllii QAIJ; VECTOB ~iff!;~

Ascoptwllum nodosum USA-NE BC 1950's l Scagel, pars. comm. Macrocystis integrilofia SE-AK PWS 1980's ROK AK Fish and Game, pers. comm. Macrocystis pyrifera c PWS 1980's ROK AK Fish and Game, pars. comm. Pachymenia carnosa J W, SE-AK 1970's ? GIH, personal observation Porphym yozoensis J W, BC, SE-AK 1900's A AK Fish and Game, pors. comm fable 3- 8 Checklist of intert1dal invertebrates of Port Valcez. Ales;.:::::: [Source- Tahlc provided by j_ Chapman. Oregon State University i

Surveys W=W:egers et aL. 1997, B=Both Surveys; S=SERC/OSU Sur;ey Local Range A-:::.Alaska: WA=Wash1ngton. BJC=Boja Californrc: C= Ccli'::.;rnlo. SC=SGuthern Colder Status I = Introduced: C =CryptogeniC: N = Native Geographical Range: A=Aystralia; 8S=Biock Sea, CS=Casp1an Sea: l=lrc:o, J=oooon MED=Med1terroneon: NEA=Northeast Atlantic; NEP=Northeost Pac:t:c NWA=Northwest Atlantic: SAM=South America • See text for mechonsims and estimated date of introduction

Soecies Survey Range Status Geograohic Range Cnidaria Anthopleura artemesia w C N Nemertea Paranemertea sp, B C N Aschelminthes Priapulus caudatus 8 C N Echiurida Echiurus echiurus otoskensis 8 WA N Oligochaeta Unidentified spp, 8 Polychaete Abarenicola sp, s Amphitrite cirrata s c c NWA Barantolla americana w c N Brandiomaldane sp, B Capitella capitola sp. complex 8 c c NEA NWA A Eteone Iongo B c c NEA Euchone ana/is w WA c NEA Exogone lourei w c N Fabricinae sp, s Fabricia sabella 8 c c NEA. NWA Glycera capilata w c N Glycinde picta 8 WA N Harmothoe imbricata s c c NEA NWA J. MED Heteromastus filiformis' B c I A NZ. J NEA, NWA. MED, SAF Laonome kroyeri w WA N Leitoscotoplos ponamensis 8 c N Lumbrineris !uti w WA N Microphthamus sczetkowii w A N Nereis vexiifoso s c N Owenta fusiformis B c NEA.. N'vVA Photoe glabra w c Polydora quadnloba w c N"' Potcrn!lla sp, w Pnonospio steenstrupi w c N Pygosoio etegans B c N Sp10 fificornis w c c NEA Sphcerosyllis brandhorsti s A N Svllts so w Tharyx gtandaria w c N Copepoda Oon;eissenio c;nctus spo. s N Oanielssenia typ;co w WA c NEA. NWA MD Horpacticus finmorchicum w Haiecrinosoma gothiceos w Harpacticus superffexus w WA N Horcccticus uniremis B c c NEA, NWA. J • _:::::;;e ::- 3. c~nt1nued s12ecies Surve~ Range Origin Geagra!2hic Rcnge ~·ers·-:;foopnonte sp w r 'v1es.::::.'lro pygrr:oeo w WA "' NE,A,. NV . .:.. A 'v11C.'':; ::thna·ton littorale w WA c NEA NW~. 5S

\Jonrcpus polustns w WA c NEA. NWA. MED. :::s 35. I SAM :Jarc:::cctyiopodio lattpes w .~arc.c:ophonte perplexa w WA c NEA. N'.VA (hiz.C,7lriX sp. w Stenrefia sp. w iisbe Jlffato w Balanomorpha Ba/cf'us glcdula 8 c N Semii::a/anus balanoides B c N Cumacea Curr.e!la vulgaris 8 c N ::udc:ella sp. w !.eptccumo sp. w lsopoda Gnorimosphoeromo oregonensis 8 c N Limncria a/gorum w WA N Umncria ligatum s WA N !dotea acufeato s WA N ldotea wosensenskii 8 c N Amphipoda Al/orcnestes angusta s c N Corinogcmmaurs morkarovi s A N Eogcmmaus confervicolus s c N Parcmoea suchaneki s WA N Callicoius pacifica s WA N Locusiogammarus locustoides s A N Pontcooreia femorata s WA N Oecapoda "ogurus hirsutiuscuius w sc N riemigrapsus oregonensis w BJC N :..;emi{;ropsus nudis s BJC N Arachnida -ialot,sium occidentale 8 c N Gastropoda Agfcjc diomedio B sc N ~gfcjc sp. w :::;ingL:a kotherinae w A N _:,'ton:-a scutula to s BJC N _ 1ori."" J sitka no 8 WA N _::;ttic :::eltc s BJC N -ectL:a persona s c N Bivalvia Axinc :::side serricoto w 8JC N 3ankc setacea s BJC N :::;:inc::;:::rdium nuttfollii B sc N _.:;sec :;dansoni s WA N \!acc."710 bcftico B c c NEA. N\', "- Vaccmo brota w WA N Vvo :;:encrio • 8 c I NEP. NEA \,\VA \/vo ":..,IlCOTO w WA N .V1ytiiLS trossulus 8 c N C'roc··eifa ruglfera w c N ::atric:::io cordito1des s 8JC N Sarrir;es groenlcndicus w WA N Tatlle B · 9 CheckiJSI ol me Mat:rotJlHllnlC Manne Algae ol Po11 Va!Oez. Stlown lor each species are its presence 111 various surveys of Port Valdez anu known geogu:~pluc range Reports ol unauacned surv1val are indica!ed lor some spec1es. (Source: Table as provided ny G. Hansen, Oregon State Univers1tyJ

Geographic Range Abbreviations and References:

AR=- Archc (Loo, 1980, C!uhara, HJ76), AU=Australia (Wornorsloy, 1984, 1987, 1994), BC=Brltlsh Columbia, Canada (Scagel et aL, 1989), C~A=Communder Islands 111 Russ1a (Selivanova & Zhigadlova, 1997), CA:::Calilornia (Hansen, 1997b), EC=Eastern Canada (South, 1984), GB=Great Britain (Burrows, 1991, Fletcher, 1987, Dixon & Irvine, 1977; Irvine, 1983, Irvine & Chamberlain, 1994, Maggs & Hornmersand, 1993), JA=Japan {Yoshida et al., 1995), OR=Oregon (Hansen, 1997a), SA=Southeast Alaska (Scagel et al.. 1989). VA=VALOEZ SURVEYS (CAL=Calvin & Lindstrom 1980~ WEI= Wiegers et al., 1997, CHA= Chapman & Hansen, 5·6/1997; HAN= Hansen, 9/1997), WA"'Washington (Scagel et aL, 1989). Unatt. Surv. (Capable ol unauached survival): Norton & Mathieson, 1983

Symbols. X "'confirmed record~ 0 "'reported but not confirmed ___.swurvev llunattj ------~2WrnQ~9lL-______Approximate Species CAL WEI CHA HAN~ AU JA C·R VA SA BC WA OR CA AR EC NO GB Distributions CHLOROPHYTA Acrostphonia arcta X X X X X X X X X X X X X X N Pac, Arctic, N AU AcrOSI{Jhorna coalita X X X X X X X X NE Pac Acrostphoma saxatlfts X X X X X X X X X N Pac Acrosiphoma sp. 0 0 0 Blidingia chadelaudil X X X X X X X N Pac, ArCtiC, NE AU Blidmgia m1mma X X X X X X X X X X X X X X X widespread Blidingia subsalsa X X X X X X X X X X X N Pac, Arctic, NE All Chaetomorpha cannabina X X X X X X X X NE Pac , NE AU Chaetomorpha recurva X X NE Pac. ChaetomorptJa tortuosa X X X X X X X X X X X X X X X widesprepd Cladophora albtda X X X X X X X X X X widespre~d Cladophora sencea X X X X X X X X X X X X X X X X widespread Enteromorpha c/atlmWl X X X X X X X X X X X X X X widespread Enteromorpha compress a X X X X X X X X X X X X X wKJespread Enterornorpha mtestmalts X X X X X X X X X X X X X X X X X Widespread Enterornorpha Jinza X X X X X X X X X X X X X X X X widespread Enteromorpha sp 0 0 0 Kornmanma leptoclerma X X X X X X X X X X X X X X N All, ArctiC. N Pac Monu~troma yrtJvJJieJ X X X X X X X X X X X X X NAIL ArctiC. N Pac Monostroma sp 0 0 Rhizoclonwm implexurn X X X X X X X X X X X X X X Widespread Rh1zocJonium npanum X X X X X X X X X X X X X X X X widespread Rh1Zoc1onwm sp 0 0 0 Ulothnx llacca X X X X X X X X X X X X X X X Widespread UlottJnx 1mplexa X X X X X X X X X X X X X widespread 'I 31Jlf:l !3 · ':J CUI lhl !Ul!U

.. .S.Y.c"ig'i .... ___ unatt. Approx1mate Spectes CAL WEI CHA HAN Surv. AU JA C-R VA SA BC WA OR CA AA EC NO GB Oistr1but1ons Ulo/hruc sp 0 0 Ulva tenesuata X X X X X X X X X X X X N Pac Ulvcma oCJsctHa X X X X X X X X X X X X X X X N Pac, Arctic. N All PHAEOPHYTA AgarunJ cribroswn X X X X X X X X X X N Pac, Arcuc, NW All A/aria sp. 0 0 0 Cllordaria /Jagelldoarus X X X X X X X X X X X X N Pac, Archc. N All Chordana gracilis X X X X N Pac CoJJodesme btJihgera X X X X X X X X X X NE Pac. Arctic, NW AU Costaria costata X X X X X X X X X N Pac Desmarestla aculeata X X X X X X X X X X X X X X NE Pac, Arctic. NW All Oesmarestia vuidis X X X X X X X X X X X X X N Pac, Au:hc, N All Dosmurus/Ja sp X

Dictyosiphon foenJculaceus X X X X X X X X X X X X X X N Pac. Archc. NW AU Ectocarpus parvus X X X X X X X NE Pac Ectocarpus siliculosus X X X X X X X X X X X X X X widespread Elachtsta lubnca X X X X NE Pac Fucus garanen X X X X X X X X X X X NE Pac. Arctic Lammana ·groenlandJca· X X X X X X X X X NE Pac, Arcllc Lammana saccnanna X X X X X X X X X X X X X X N Pac, ArCtiC. N AU Laml{lana yezoens1s X X X X X X X X N Pac Leames1a dtf/ourus X X X X X X X X X X X X X X Widespread Melanostphon mtest1naiJS X X X X X X X X X X X NE and NW Pac Petalon1a faSCia X X X X X X X X X X X X X X X widespread P11ayella littoralis X X X X X X X X X X X X X X X X X X widespread Ralfsta sp 0 0 Scytos1phon simpJicissimus X X X X X X X X X X X X X X X X X widespread Soranthera ulvotdea X X X X X X X X X X NE Pac Sphace/aria rigidula X X X X X X X X X X X X X widespread Spongonema tomentosum X X X X X X X X X X X X X N Pac, N All RfiOOOPHYT A

Ahnleltia fastigiata X X X X X X X X X X X X N Pac AhnfeWopsis gigartinoides X X X X X X X NE Pac Antithamnionella paCifica X X X X X X X X X NE Pac Auaouinella purpurea X X X X X X X X X X X X X N All, N Pac, Nl W!(lesprt!

" "_S""•'-"-- Unatt" ______l:\<:Q9li!Qhll

~~y___ Unatt. ______g_~291:l!Phll<..BRn9,"------­ 1 Approximate Species CAL WEI CHA HAN Surv. AU JA C-R VA sA BC WA OR CA AR EC NO GB Distributions Palmana necatensts X X X X X X NE Pac Palmana patmata X X X X X X X X X N Pac. N All Phycodrys nggn X X X X X X X N Pac Polystphonta brodtaet X X X X X X X X X X X X Widespread Polystphoma nenary1 v det1quescens X X X X X X X X NE Pac Polysiphoma hendryt v hendryt X X X X X X X X X NE Pac Polystphonia nendryi v luxunans X X X X X X X NE Pac Polystplwma pactl!c:.t v f>

TOTAL "X" 68 81 20 24 29 24 60 46 101 91 90 85 76 70 42 46 43 42 Total Widespread;::; 21 Table B. 10. An1mal spec1es w1th potential lor 1ntroduct1on to Alaska. These Include spec1es which have h1stones of mtroduct1on elsewnere at latitudes above 40° and tor which ballast water IS the possible medium of transfer. Indicated for each species are donor reg1on. date of first record, possible mechamsm ol mtroduct1on, recipient region and references. [Source: Table prov1ded by J. Chapman, Oregon State University]

Mechanism: A~ Aquaculture; B ~Ballast Water; Cu ~Coastal currents; F =Fouling;!~ Intentional; OJ =Oysters Japan Donor/Recipient: AK~ Alaska, BC~ British Columbia, BS~ Black Sea,C=California, CR~Columbia River, EUR~ Europe, GL~ Great Lakes, K= Kachemak, NEA= North East, Atlantic, NWA=North West Atlantic, NEP=North East Pacific, NA=North Atlantic; NWP= North West Pacific, NZ= New Zealand; O=Oregon, PS=Puget Sound, W=Washington, SE AK= south east Alaska, SFB=San Francisco Bay, TAS= Tasmania

Recipient Species Donor Region Date Mechanism Region Reference Protozoa Trochammina hadai Japan 1990s B SFB Cohen and Carlton 1995 Coelenterata Cordylophora caspia BS 1930s F, B BC Carlton 1979a, Cohen and Carlton 1995 Aurelia aurita NWA 1990s B 0 Cohen and Carlton 1995 Polychaeta BoccardJa ligenca NEA 1935 B, F Korea Cohen and Carlton 1995 Euchone limmcola NZ? B SFB Cohen and Carlton 1995 Lyrodus takanosimensiS Japan 1981 B? BC J. Carlton, pers. comm. Neanthes succinea NEA 1850s? F,B? PS Carlton 1979a, Cohen and Carlton 1995, J. Chapman Pers Obs. 1992 Manayunk1a spec10sa NWA 1960s B,F C,O Cohen and Carlton 1995, J. Chapman Pers Obs Marenzellena vtndts NWA 1983 B SF Bay Hopkins 1986, Cohen and Carlton 1995 Polydora ligm BC 1932 B.F C,O,W Cohen and Carlton 1995 Psuedopolydora kempi Japan 1940s B, F,A C,O,W Cohen and Carlton 1995 Crustacea Balanus improvisus NA 1853 B,F C,O,W Carlton 1979a Bythotrephes cederstroemi EUR 1984 B GL Carlton and Geller 1993, Yan et al. 1992 Grandidierella japonica Japan 1930s OJ, B C,O,W,BC Jassa marmorata NA 1800's F,B C,O,W,BC Conlan 1990; Carlton 1979a Cancer novaezelandiae NZ ? B TAS Bennet 1964, Nations 1979, Furlani 199q Carcinus maenus EUR? 1989-97 B,Cu C,O Cohen et al. 1995, Miller 1996,J. Chapman Pers. Obs. Corophium curvispinum BS 1970's B NEA Carlton et al. 1998 Eriocher sinense Asia 1989-97 B C,O Cohen and Carlton 1997, G. Jensen Pers Comm. Exopaleaemon modestus China 1995 B CR Emmett, USFWS, Pers. Com. Table B · 10. contmued ... Recipient Species Donor Region Date Mechanism Region Reference Halicarcinus innominatus NZ 1983? B NZ Furlani 1996 Nippo/eucon hmumensts Japan 1979 B C,O,W Carlton and Geller 1993, Carlton et al. 1990 Petroltsthes elonga/us NZ 1986? B NZ Furlani 1996 Pseudodoptomus tnoptnus Chtna 1990 B O,W Cordell et al. 1992 Mollusca Dresstma polymorpha EUR 1986 B GL Mills et al. 1993 Muscullsta senhous1a NWP 1990s? B, F, A BC Merilees, 1995, Cohen and Carlton 1995 Myti/us gal/oprovmCJallts NEA 1800s? F,B 0 Cohen and Carlton 1995 Mollusca Ens1s dnectus NEA 1983? B EUR Carlton & Geller 1993 Nuttallia obscura/a J 1991-97 B,Cu BC,W,O Merilees,1995, Pars. Obs. 1997. Potamocorbula amurensis NWP 1986 B c Carlton et al.1990 Tritonia plebeia EUR,NWA 1983 B NWA Carlton 1989 Ectoprocta Membranipora membranacea EUR 1987 B NWA Carlton and Geller 1993 Urochordata Botryllus spp. ? 1990s? B? NEP Kozlolf 1996 Botrylloides spp. ? 1990s? B? NEP Kozlolf 1996 Molgula manhallensis NEA 1900s? O,B NEP Kozloff 1996 Stye/a clava Japan 1900s? F,B O,W Carlton 1996, Kozlolf 1996 Vertebrata Gymnocephalus cernuus NWA 1987 B GL Carlton and Geller 1993 Table 8·11 Algal species with potential for introduction to Port Valdez/Prince William Sound. Indicated for each species are donor reg1on(s), rec1p1ent reglon(sJ, date of lirst record, probable vector of introduction and reference(s). fSource: Table prov1ded by G. Hansen!

Abbreviations/Codes: Donor/Recipient Regions: AUS-TAS= Tasmania, Australia, AUS=Australia, BC=Brltish Columbia, C-SFB=San Francisco Bay, C=California, CAN-NF=Newtoundland, CAN-NS=Nova Scotia, CAN·O=Quebec, CH=China, EUR=Europe, FA·BRIT =France, Brittany, FR-MED=French Mediterranean. FA=France, GB=Great Britain, HEL=Helgoland-Germany, IT =Italy, J=Japan, MED=Mediterranean, MX=Mexico, N=Norway, NETH=Netherlands, NE-ATL;;::: Northeast Atlantic, NE-PAC= Northeast Pacific, NW-ATL=Northwesl Atlantic, NW-PAC= Northwest pacific, N-ATL= North Atlantic, N-PAC=North Pacific, NY=New York, Long Island Sound, NZ=New Zealand, 0=0regon, PAC=Pacific, PWS=Prince William Sound, R-ARC=Russian Arctic, White Sea, A-SIB=Siberia, S·ATL= South Atlantic, S-AUS=Southern Australia, S·C=Southern California, SCAN=Scandinavia, SE·AK=Southeast Alaska, SP=Spain. USA-NC=North Carolina, USA·NE=New England, W=Washlngton Vector: A.::oAquacullure, B=ballast water, Ldobsler or bail packing, M=Marginal dispersal through currents, OA=oystersAUantic,OJ=oystersJapan. OBC=oyslers Bntish Columbia, AOK=roe-on-kelp: S=scienlitic research, ?=unknown vector or date

t:lQtlQB B~f,;!f'lliliT QAIE llSs;:[QB BEEi

~f'(;(;lt;~ QQtulata NE·ATL GB, SP 1973 Farnham, 1994 AnuthamnJon den sum S-ATL EUR 1960 ? Ribera and Boudouresque, 1995 Arlllthammon pectmatum - ? FR 1988 A Curiel et al., 1996 ? IT 1996 A Curiel et al., 1996 AntJthammon sp. NY <1986 ? Foertch el al.. 1995 Anllthammonella temifolia S-ATL GB 1906, 1922 F Lyle, 1922; Farnham, 1980; Ribera and Boudouresque, 1995 Asparagops1s armata S-AUS GB 1922, 1939 F Ribera and Boudouresque, 1995; 'Farnham, 1994 Bonnemwsonia tJanutera J GB 1890, 1911 F Guiry and Maggs, 1991, Farnham, 1994, Ribera and Boudouresque, 1995 Chondrus g1ganteus 1/abel/atus J FA-MED 1993·94 OJ ? Cruoria cruonaeformis? FR·BRIT GB 1976 M Blunden et al., 1981 ? Cryptonenua tuberruca '! GB 1971 ? Ribera and Boudouresque, 1995 Furcetana lumbocal!s GB CAN-NS 1800's B Novaczek and Mclachlan, 1989 Geftdtum vagum J BC, C-SFB 1970's OJ Renfrew et al., 1989 Grateloup1a doryphora NE PAC USA·NE 1990's B? Villalard-Bohnsak and Harlin, 1997 J EUR 1969 OJ Ribera and Boudouresque, 1995 Grateloupia lilicina var luxurians PAC GB, SP 1947 ? Farnham, 1994 Lomentaria clavellosa GB, FR USA·NE 1950's ? Wilce and Lee, 1964 Lomentaria llakodatensis J, S-C BC,W early 1900's OJ Soulh, 1968 FR·BRIT, MED 1984 A Cabioch and Magna, 1987, Farnham, 1994 Mastocarpw> stellatus NW-ATL HEL 1970's s Ribera and Boudourosque, 1995 Polystptwma harvey1 CAN-NF GB and FA 1976 F?,B? Maggs and Hommersand, 1990 Porphyra yezoens1s J MED OJ Ribera and Boudouresque, 1995 Solsefla chorda/is FR·BRIT GB 1976 M Farnham and Jephson, 1977 Soliefta tenera NW-ATL GB 1978 Farnham and Irvine, 1979 Seagrasses Zostera Japon1ca J W,BC early 1900's OJ Harrison and Bigley, 1982 Table 8·12. Algal spec1es nahve to Nonhwestern Nonh America that have been introduced to otller global regions. lnd1cated lor each spec1es are donor reg1on(s), rec1p1ent reg1on(s). date ol l~rst record, probable vector of introduction and reference(s). {Source. Table provided by G. Hansen!

Abbreviations: Donor/Recipient Regions: AUS-TAS= Tasman1a, Australia, AUS=Austratia, BC=Britisll Columbia, C-SFB=San Francisco Bay, C=Ca!ifornta, CAN-NF=Newfoundland. CAN·NS=Nova Scotia, CAN·O=Ouebec, CH=China, EUR=Europe, FA-BRIT =France, Brittany, FR·MED=Frencll Mediterranean, FR=France. GB=Great Brita1n. HEL=Helgoland-Germany, IT =Italy, J=Japan, MED=Mediterranean, MX=Mexico, N=Norway, NETH=Netherlands, NE-ATL= Northeast Atlantic, NE-PAC= Northeast Pacilic, NW·ATL=Northwest Atlantic, NW-PAC= Northwest pacific, N-ATL= Nortll Atlantic, N·PAC=North PacifiC, NY=New York, Long Island Sound, NZ=New Zealand, 0=0regon, PAC=Pacific, PWS=Prince William Sound, R·ARC=Aussian Arctic, Wh1te Sea, R-SIB=Siberia, S·ATL= South Atlantic, 8-AUS=-Southern Australia, S-C=Southern California, SCAN=Scandinavia, SE-AK=Southeast Alaska, SP=Spain, USA·NC=North Carolina, USA-NE=New England, W=Washington Vector: A=Aquacullure, B=bal!asl water, L=lobster or bait packing, M=Marginal dispersal through currents, OA=oystersAitantic,OJ=oystersJapan, OBC=oysters Brilish Columbia, ROK=roe-on·kelp; S=scientific research, ?=unknown vector or date

Sf'E<;;I&~ QQt'lQA A~<;;lf'tft'li I:! ATE YfkiQB B£EfBEt'l<;;f Chlorophyta nosenvmnmlla polyftlila NPAC AlJS Hil>era and f3oudourosque, 1995 Chaatornorpha melagonrum N·PAC, N·ATL AUS Ribera and Boudourasque, 1995 Cladophora laetevirens? N·PAC, N·ATL AUS Ribera and Boudouresque, 1995 Chrysophyceae·Phaeophyta CoJpomema peregrma PAC GB 1907 ? Farnham, 1994 ATL MED 1956 PAC FR·BRIT 1906 Ribera and Boudouresque, 1995 PAC USA·NE 1978 Bird and Edelstein, 1978 Leathes1a diflorrrus PAC MED 1979 OJ Ribera, 1994 PiJayella ltttoralis N-PAC, N·ATL NZ F Adams, 1994 Punctana latilo/Ja N-PAC. N·ATL NZ F Adams, 1994 Sargassum mullcum BC FR 1972 OBC Druehl, 1973 ScytosJphon dotyi? NE·PAC MED 1978 Giaccone, 1978, but see Ribera and Boudouresque, 1995 Sphaerotnchm divancara PAC MED 1981 OJ,OBC Ribera, 1994 Macrocystis pynfera c FR·BRIT 1973 A Farnham, 1994 Rhodophyta Antitharnmonelfa splfographidJs Ne and Nw Pac GB 1920's Farnham, 1994 AUS Ribera and Boudouresque, 1995 Bonnema1sonia hamifera PAC GB 1893 Farnham, 1994 CeramiUm wbrum N·PAC, N·ATL NZ Adams, 1994 GonJotnchopsJs sublittoralis NE-PAC FR·BRIT, MED 1992 Magna, 1992 Grateloupia doryphora NE·PAC GB, S 1969 Farnham, 1994 Sarcodiotheca gaudichaudii NE·PAC GB 1973 Farnham and Irvine, 1979 Pikea californica NE·PAC GB 1967 Farnham, 1994 Polysiphonia brodiaei N-PAC, N·ATL AUS, NZ Ribera and Boudouresque, 1995 Polysiphonia harveyi NE-PAC GB 1976 Farnham, 1994 Table B-13 tlarmlul nucroalqal srwc1es w1th potenllal tor mvas1on ol Port Valdez/Prince William Sound. Indicated for each species are donor regJon(s), rec1p1en1 reg1on(s). date ol lust record. probable vector ot introduction and relerence(s). (Source: Table prov1ded by G. Hansen!

Abbreviations: Donor/Recipient Regions. AUS·T AS= Tasmama, Australia. AUS=Australia, BC=British Columbia, C-SFB=San Francisco Bay, C.:Calilornia, CAN-NF=Newtoundland, CAN-NS=Nova Scotia, CAN-O=Ouebec, CH=China, EUA=Europe, FA-BRIT =France, Brittany, FR-MED=french Mediterranean. FR=France. GB-=Great Britain, HEL=Helgoland-Germany, IT ;::Italy, J=Japan, MED=Mediterranean, MX=Mexico, N=Norway, NETH=Netherlands, NE-ATL= Northeast Atlantic, NE-PAC= Northeast Pacific, NW-ATL=Northwest Atlantic, NW·PAC= Northwest pacific, N-ATL= Nor1h Atlantic, N-PAC=North Pacillc. NY=New York, Long Island Sound, NZ=New Zealand, 0=0regon, PAC=Paclfic, PWS=Prince William Sound, A·ARC=Aussian Archc, Wh1te Sea, A-SIB=Siberia, S·ATL= South Atlanlic, S-AUS=Southem Australia, S·C=Southern Calilornia, SCAN=Scandinavia, SE·AK=Southeast Alaska, SP=Spain, USA·NC=North Carolina, USA·NE=New England, W=Washlngton Vector: A=Aquaculture. B=ballast water, L=lobster or bait packing, M=Marginal dispersal through currents, OA=oystersAtlantic,OJ=oystersJapan, OBC=oysters Brihsh Columbia, ROK=roe-on-ke!p; S=scientilic research, ?=unknown vector or date Effect: ASP=Arnnestc Shellfish Poisoning, DSP=Diarrhelic SheUiish Poisoning, invert.=invertebrate, PSP=Paralytic Shellfish Poisoning

BECOBQS EFFECT

1. Cryptogenic species which presently occur between Oregon and Alaska, and which may occur In Port Valdez/Prince William Sound: Pyrrophyta (Dinoflagellates) Alexandnum catenella C to AK-60 N early 1900s PSP Bressner and Middaugh, 1995; Kvitek et at., 1993 OinophySJS acurmnata and 0 fortii FR. CAN·O, long term DSP Cembe!la el al., 1989;Stamman et al., 1987; AC, R·SI£l Taylor ot nl., 1994; Konovalova, 1993 Baclllarlophyta (Dia!Orns) Chaetoceros concavJcomts and BC,AK 1961 fish kills Farrington, 1988; Tester and Mahony, 1995 C convolutus Pseudomtzschia pungens CAN·NS, W, 1987 ASP Wekell et at., 1994; Postel and Horner, 1993 I mu/1/senes. P cwstra/1s O,C Chrysophyta Heteros1gma carterae BC,AK 1976 fish kills Taylor, 1987;Taylor and Haigh, 1993

2. Species with histories of invasions at sites outside of western North America, which could be Introduced to Port Valdez/Prince William Sound: Chrysophyta Aureococcus anophagetferens USA-NE 1985 brown tide Cosper et al., 1990; Shumway, 1990 Chrysochromulina polylepis SCAN 1988 fish and Underdal et al., 1989; invert kills Aune et aL, 1992 Fibrocapsa japontca FR·BRIT Ribera and Boudouresque, 1995 Cyanophyta (Biuegreens) Nodulana spumigena SCAN, AUS 1985 trophic Nehring, 1993; Edler, 1985; effects Jones eta\., 1994 F1gure B-1. Percent distr:buuon of known non indigenous marine spec1es among maJor taxonom1c groups for each of SIX reg1ons 1n western North America. Percentages are based upon Table B-5 and IOta! number of nomnd1genous spec1es is shown for each reg1on (geographiC reg10ns as also from Table B-5) in parentheses; the data shown represent the current state of knowledge concerning nonmdigenous spec1es in each regiOn (see text for discusSion)

40 CA (N:76) 30 SOUTHERN

20

10

0 40

30 SAN FRANCISCO, CA (N=215)

20

10

0 en 40 w NORTHERN CA u 30 w a. 20 en 10 ..J <( 0 1- 40 0 1- 30 OREGON (N=77) u. 0 20 w 0 10 <( 0 1- z w 40 u a: 30 w a. 20 10

0 40

JO ALASKA (N= 11)

20

10

0 0" a:" u 0 0" ">- N w "0 N 0 !: "' a. 0,. 0" ~ ~ >- 0 a: a: ~ z 0 0 " a: a: 0 .. a: a. 0 J:: a. ::; "... u ...a: " " " Table C-1. Charactenstics of segregated ballast water arriving to Port Valdez, Alaska for each of the 16 oil tankers sampled between 23 May ami 6 June 1997. Shown arc dale of arrival, ballast water source, total ballast water capacity, amount of ballast water on board (as volume and percent capac tty) upon arnval, age of ballast water, mean ballast water salinity and temperature. For each ship, the number of tanks that underwent ballast water exchange is also indicated for each ship. Ballast water source corresponds to last port of call in all cases

Open Ocean %Total Total BW Total BWOB Age Temperature Salinity Ship Name Date of Arrival BW Source BW Exchange BW Capacity (m3) (m3) (days) (C') (ppt) (# tanks) Capacity ARCO Spirit 5/23/97 long Beach, CA none 38362.79 26752.33 70% 6 16 25 35 ARCO Anchorage 5125/97 Cherry Pt., WA nona 21873.58 21004.11 96% 4 14 75 29 Baton Rouge 5/26/97 Anacortes, WA none 37676.73 31447.67 83% 3 11.75 28 25 Long Beach 5126/97 San Francisco, CA two 69640.93 60292.88 87% 4 14 32 8 Chevron Mississippi 5/27/97 Anacortes. WA none 22525.93 19282.02 86% 7 11.75 30 75 Potomac Trader 5/27/97 Cook Inlet, AK none 17762.19 10271.06 58% 1 11.5 29 ARCO Fairbanks 5/28/97 Cherry Pt., WA none 36604.02 20979.51 57% 4 14.25 28 5 0/S Washington 5/28/97 Richmond, CA none 15020,92 no sample 0'% 5 no sample no sample BT Alaska 612/97 San Francisco, CA none 56392.66 47719.76 85% 6 11 25

SIR North Slope 6/2197 Portland, OR none 57450.86 47016.85 82% 4 14 5 25

SIR San Francisco 6/2197 Anacorles, WA nona 37684.66 29503.51 78% 6 12 28 25

AACO Juneau 6/2197 Cherry Pt., WA none 24428.46 22028.24 90% 4 12 25 28 5

ARCO lndopendonco 6/3/97 Long Beach, CA no no 43413.06 29103.97 67°/o 6 14 J2

OMI Columbia 6/3/97 Barber's Pt. HI none 22118.90 16293.81 74% 9 13 25 34 75 Prince William Sound 614197 Yosu, Korea au 41972.01 25562.63 61% 10 11 30 Benecla 6/6/97 Banecia, CA two 53360.91 47418.37 89% 6 12.8 175 Tahk C·2 \alnuty .tnd Temperature ul water irl Port Valdez at the site and time of arrival for oil tanker!:~ sampled between 23 May

~-· ·~----"·-~--~--- Shipside Shipside Shipside Shipside Salinity @ Salinity @ Temperature Temperature Ship Name Surface 10 meters @Surface @ 10 meters (ppt) (ppt) (C 0 l _ _(C!._} __ --- - .. --~--~~------· ARCO Spirit 29 30 14 9

ARCO Anchorage 27 32 10 5'I Baton Rouge 23 29 10 6

1 S/R Long Beach 20 32 1 2 Si I ' 27 32 1 4 ei 'Chevron Mississippi ! 9' Potomac Trader 25 31 1 4 ARCO Fairbanks 22 31 1 1 9 0/S Washington no sample no sample no sample no sample

BT Alaska 21 31 1 3 9.

S/R North Slope 21 31 1 3 9 S/R San Francisco 21 31 1 3 9 ARCO Juneau 21 31 1 3 9 'ARCO Independence 21 31 1 3 9

OMI Columbia 20 31 1 2 51 Prince William Sound no sample no sample no sample no sample S/fl Henecia no sample no sample no sample no sample ------·-----·------Tallie C-3 Dens1ty ol rng<~msms \#1m3) tn eacn ta~tonormc gmup mat occurred 111 plankton samples collec1ed lrom segregated ballast water o114 dil!eren1 oil tankers arriving to Port Valdez, Alaska between 23 MJy a no 6 June \997 ShOwn a1e tne means ano stanoaro errors (tor 2tanks/sh1p) obtalfled 11om quantitative analysis ol Pf&Served samples; estimates lor abundance Wl\h!n tanks were based upon two ven1cal net tows tsee teAt 101 t!•Phlna\lon~ A grana mean and stanoal!l BHOt among all stl1ps IS also snown, anr:l was calculated us1ng the estimated density (moan) lor eac/1 ship. All data prasenled here were lrom ballast waltH ol oomesuc .Ill\\ uM~l<~l •'"iJ"' lJ.tll.J~I wahl/ o:H lorl.luJn onyHt anupoHatu analySis. AU organtsms w~:~ru consklured to Uu 1<\!vu at 11ma ol coUuc\lon. uasud upon tnthal obsurv;~.\!ons that ucmoo 1 ;,uat~u /J~", u1

chevron AflCO SIR San ARCO SponC SIR North Slopa ARCO Jui\Miu Ml ..l .. lpf>l Fakbanlu. Franclaco

Mun MaaJ 62 53.65 4.14 4.1( 4•0 3110 Pe~~ IU)H Pann.la 1321> lll7 1119 II ~ 17 34&.53 341U3 $.22 5.22 •PROTOZOA" '"~u n tu F«amlnllwa 21>11 2U US 0.53 1.01 624 I 1>3 I &3 3 61 l )1

unld.M*duu 4l2 35~ 442 442 18104 )338 020 020 4&2 1.81 Ill :l6 0.03 12 07 I 25 4_04 404 87011 6965 0 1& 018 CTUtOPtlORA 041 041 l2ti8 1363 PLAT'l'HEI.Mitfl'HES 003 OOJ 11 53 I OS 4699 &H 423 262 129 1211 130 1.30 1112 708 464 137 0)8 01& MC.U.,.'l/Goi'IU•"o lar. .. 1>4 ~~ \6 78 202 202 "' 1 n 1 n 1 ~o t4{MUHtA '" "' ' l't'l! ~H I !HI un ou ""'1"""' ...... I ~~ ~ij '~ !) i .'mJ f'htlludu<:Wu u ~J '" o&u "" ...... "" 'bJ Sp1<>nld.o.a lfb ~~~ 1~ 61 211 W20S 2501 866 ,., "' ... "' 33 60 18 72 ~~~~ '" "' 21"' 26 '" 1>0 1>4 liaphlhyd•• Mill~ 11 30 ,., '" ... '" 522 Ill "' 2.07 '" "' '" , .. "',,. Syl~4a• 04l 043 '" 0.9~ Po<>Wnlo«n• '"'" MOl.lUI>CA Oaa!lopo2 Oti;> ~3H l~ll 11306 \1646 1Ml-lll8 165SS 20!.209 24978 56733 3S09 14&05 19.71 3U027 1>3.6-4 114204 2&4-41 468&0 104_05 65451 3003 26617 242.66 311~ 16 &599 091 S6906 29S~Ii 3~~31 16i ;., 6~0 0-1 2J9 19 co~pod!Caa 130!4~ S4liJ911242~8 25~71 301339 l!t456 IH36 4908 5327 720 1:1&963 1666 55-4.60 \5-4.71 62289 1~99 704.66 69.11 20-4325 203226 110691 111.23 '" 253 23911CI 519 rol 791 ~~ lll2 9J 1S2t> Uti So-l 18 Hllp.Cik:<>l<:la '"' l,llcll& op 2 !0 i 040 322 322 6l& us 439~ 2025 2413 250 505 500 '" 018 581 SB! I ij\) J ]I (ulloptR• •:> "" "' I "' 81971 16686 2 14 Joe~ J oa 1414 '" '">!<""""'" Hatpa<:lacoul I 10 591 I 29 I 29 ~99 U15 1011 ' " "nl<>wnl.>•"'l>lc H.rp I ~ 46 1 30 \.30 5&3 sa~ Cycropolda I '" 000 I Ollhon. •P9 816&6 4\1~4 12 91 114 1141 "' 1611 8 0 0 ll~ u~ \2') _j\) Uml'l<>llr><>n• op ""' 524 524 .,, 13H 60 Cycloplt>a op 4WH 14627 : ~ ~' bb •ti 1 J3 •o J2 6<1 ..... ' ...... ············ ...... L.· ~ ~ L~.:...• ..:.: :...... ~.. : .... .~...: ... L...... '.. ' .. .1.1 ""' TaOlt! C-3 con!lrlueo

AHCO ' en.., ton A.ACO AACO SjH•U lla!on Am•11• p;::::" SJH North Slop0611 110 62 C•t•mu ap '" "' 4.02 '" 6~1 5< 46~ ~I CamlophOIIU op. "' .., .., l' "' "' "'0" I;5 "' "' 10.29'" 2.57 0.8:1 0" "'0.113 O.lll"' ""' ou ., , l.lfltk1141 •P· "' I '" "' "' '" "' "' . .. "' "' P...-ocolan•u •P· 95 17 '"' "' "'o,; ,.. 2.83 251.11 1113.75 3HI3 18.32 57.00 23 73 1309 \9 78 "' 0 " ... '" 37U ' 6 04 IS !.6 Poou<:•lanu• op. 17081 2621 7649 167 l!t .17.94.. I 27 43 1.15 120.11a 20.511 2!1.33 "' 11.39 20846 21.23 2.02 \.64 Pnu T<>$l<>m0 "' '" '" '"' Co.-yculd 5•2 31 174,5-8 37 94 II 42 41 81 ~.33 1.40 '" 12.26 2.92 11.65 ~.16 224 91 125 57 Homl<:y~l<>pa ... 6J 24 4009 ap '" "' '" '" 'Rulph' ... I o;; 0.83 .,. 19 62 ,., Ampl>tpo7 000 "' o..:.poUo "' "' "' '" "' '"' '" "' '" "' 12 6f> 13 46 10 85 26042 42116 19 57 "' m•9olopo·~· '" "' , .. 2S H) 10 36 '" "" '" "' '" "' '" '" "' ' Cati<:a ..n "' '" '" "' 0 02 "' '" "' 31 29 31 2g "' <>o I ijJ "' "' "' 0 2• "' "' "' "' "' \) '" I "' '" "' " " "' 0 u '" Oou•«> ll 2211 21 54 "~ "' '" ... "00 '•<'"•• I'HOfiONIOA ~1> ., I '"' <11!'1 !4'11 .'U U!AtcJUGNAIHA ',/1 '".,,, "' "" '•1:11 tW "'' 11 I)~ l 311 11112 I 2$ 12 ()() '''-' ' 0 IH , ~ ,.,. '"' ((.;111/'!0UlfiMA I A ''"" ' '" "' "" ... "-••••"''-'•• tUil~l H 6~ '~ I "' .•u fcldu I 1.:n I~ ~~ I '" "' '" "' "' 0 42 ,., ·~ ~.: CHORDA!"- [""" "' '"' "" '" ' > U." "d." l01~0C .. 0 ~f> I 91111 37 6. 60 11 21 34 ,., ,., I 38 21 25 11 len Jl u. ~~ fton "' i "' "' 1: '" "" J'" ••• "' () tltl a '" C~l••l<><:ho<phOI'• 61 SJ 3~ 60 6 24 t0fl06 35 IS 12"' 99 12 99 moo 16 69 19.00 H.OO 11-~6 ~9 51 ~7.52 3763 ]4 6? I! ti4 polt.n "' 0" 114.14 1H.74 H~63 iiOI&: ...... '" "' 0.18 6 16 "' '" '"' Table C-4. Summary of quantitative plankton analysis for segregated water of 13 different oil tankers arriving to Port Valdez, Alaska from continental domestic ports between 23 May and 6 June 1997. Shown are the ship sample number, number of species, cumulative density for all taxonomic gorups, total ballast water on board at the time of arrival, and estimated total number of organisms in all ballast water on board for each vessel; a mean and standard error are also shown for all biological and volume estimates among ships. Taxonomic diversity (minimum # species) and density (organismslm3) measures were obtained from quantitative analysis (see Table C-3 ). Total number of organisms per shop were calculated as the product of total ballast water volume on board and organism density.

I : TOTAL# I MINIMUM# TOTAL TOTAL# I SHIPNAME SHIP# ORGAN IS 3 SPECIES 3 VOLUME (M" ) ORGANISMS ! MS/M" I ARCO SPIRIT ' AK001 ! 26.00 11,215.511 26,752.33 300,041,025 ARCO ANCHORAGE I AK002 3,997.72 I I 17.00 21,004.11 83,968,551 LONG BEACH I AK003 15.25 11,752.03 60,292.88 708,563,735 BATON ROUGE I AK004 29.25 16,493.91 31,447.67 518,695,039 I POTOMAC TRADER I AK005 19.00 1,452.25 10,271.06 14,916,147 I CHEVRON MISSISSIPPI I AK006 I 15.00 4,986.72 19,282.02 96,154,035 ARGO FAIRBANKS AK007 20.00 9,831.76 20,979.51 206,265,507 SIR NORTH SLOPE AK009 11.50 3,128.55 47,016.85 147,094,566 ARGO JUNEAU I AK010 22.00 3,840.57 22,028.24 84,600,998 S/R SAN FRANCISCO I AK011 26.00 4,887.10 29,503.51 144,186,604 i BTALASKA ! AK012 12.75 13,500.41 47,719.761 644,236,325 ARGO INDEPENDENCE! AK013 I 24.00 2,834.68 29,103.97 82,500,442 I BENE CIA AK016 11.00 3,085.96 47,418.37 146,331 '193 - ! I I MEAN 19.13 7,000.55 31,755.41 -~44,~27 ,243 ., . - -·-~------SE I 1.66 1,355.89 4,026.34 63,878,556.94 fatJ!t: C-:. Cornpanson ol quanlltal!ve planKton analysts lor segregated ballast water arnvrng to Port Valdez, Alaska 1n otl lankers of oornes11c ano loretgn ongrn tJetween 23 May and 6 June, 1997. Shown are lhe denstlies (number/rn3; mean and standard errors for two tanks) by taxonomtc group obtatned from a stngle foretgn arrival compared to the mean density measures(number/m3) lor 14 domestic arnvats Ballast water sampled for the single foretgn arrival originated in Korea but all tanks had been exchanged at sea; ballast water mcluded here tor domestic arnvals was of coastal origin (from last port of call) and was not exchanged in transit.

t:f.!HWkhD.O!a.Shi.sll g tfuru:tchange Sblp} (n»14) fxkhllnse..S.Oio (n•t) (n... 14) I (n,J) Mean SE Mean SE Mean SE Mean SE OINOFLAGELLAT A Copepoda, cont.. Ceratium 4.40 3.80 1.60 0.17 Csfanolda Perldinium sp. 59.68 29.55 3.59 3.59 Acartla op. 651.54 469 51 D!ATOMACEA 35 06 18 08 Catanua 1p. 0.98 0.47 45 17 20 30 DISCOid 722.16 255 25 91.64 54.86 Centrophages ap. 1.99 0.74 2 83 2 !13 Centrale 166.67 103.47 29.95 1.65 Metrldla ap. 0.13 0.13 Pennate 27.20 •mao Paracalanus sp. 37.26 18.56 "PROTOZOA" Pseudocalanus sp. 76.58 24.44 34 30 11 66 Foraminifera 3.31 '08 2.67 2.87 Pseudodleptomus spp. 7.25 7.21 Tintinnlda 0.96 040 95.70 26.36 Rhlncelanus sp. 0.03 0 03 CNJDARIA Tortanus sp. 8.38 522 unid. medusa 22.66 13.63 Ponte!lid 6.15 4 04 CTENOPHORA 0.03 003 Labadocera ap. 0.05 0.04 PlATYHElMINTHES Poec/lostomtt Turbellana 5.86 3.34 Corycaeld 63.24 40.09 MiJller's/G6tte's larvae 5.41 4.60 Hemlcyclops sp. 0.65 0.62 NEMERTEA ~RudolphR 1.75 1.41 pillidium larvae 087 0.82 Amphlpode 0.03 0.02 ANNEliDA Gammerldea 0.15 0.10 Polynou1ae 4.58 2.80 Hyperlldee 0.19 0.07 071 071 PhyUouocldao 1 63 '36 lsopoda 043 015 Sp10llld8U 97.71 60 64 0.72 0.72 Oec1-1poda Nephthydae 461 359 0.72 0.72 ::oea 25.10 16 36 071 071 Chaetopteridae megalopa 0.41 0.19 Syllidae 0 :)8 0.23 Carldoa 0.02 0.01 Oorvilleidae 1.87 187 Cladoceran 2.96 2.24 Capitellidae 1.03 0.72 Anomura Owenidae 53.87 48.90 Porcellanld zoea 0.40 0.38 Magelonidae 0.73 0.50 Pegurld zoea 0.52 0.32 unknown larvae 138 68 39.26 Mysldacee 3.66 1.59 MOLlUSCA Cumecea 0.21 009 Gastropoda 206.91 131.40 10.72 3.64 Stomatopode 0.13 013 Bivalvia 490 41 214.71 0.72 0.72 Ostracode 0.24 0.24 Pteropoda 0.03 0.03 BRYOZOA 0.06 0.06 unknown veHger 1.88 187 cyphonautes larvae 10.51 4.38 28 56 7 34 nudibranch 0.07 O.D7 PHORONIOA 5.46 3.37 CRUSTACEA CHAETOGNATHA 966 446 071 071 ClrrtptHJla ECI-IINODERMATA naupli! 627 67 323 55 8.57 2.92 Asleroldea 16.70 15.52 144 144 cyprids 83 44 25 17 6.40 2.09 Echlnoldee 1.02 0 87 Copepoda CHORDATA nauplii 850 04 239.19 2805.51 891.42 ' Larvacea 17.59 8.12 358 217 copepodites 1526.86 504 16 1757.46 274.84 Fish 0.08 0.07 ' Harpact1coida Cephatochordate 0.06 006 Microsetel/a sp. 780 3.31 1.43 0.01 OTtfEA Eutropina sp 5!:196 56 55 Eggs 20.62 8 92 271l2ti !J5 :w unknown Harpactacold 5.75 3.34 unknown larvae 3.50 3.29 unknown benthic Harp. 3.07 2.02 lrocophore 34.67 11.64 Cyclopoida 3 92 0 39 pollen 8.84 8.16 Oithona spp. 215.05 123.30 769.29 150.27 I Umnoilnona sp. 72.91 66.46 Cyctopina sp 33 40 32 88 ~_o_ee_po__?~ :_o~t- ______------Frgure C-1. Compansons of salinity and temperature measured for segregated ballast water arriving to Port Valdez, Alaska rn oil tankers and for surroundrng port waters at the time of deballasting. Shown are mean (and standard errors) for ballast water and port water. For the port, data are shown for measures at both the surface and 10m, as significant stratification existed; such stratiftcation was not present within the ballast tanks. Data are shown in Tables C-1 and C-2. For each salinity and temperature, letters which differ above histograms indicate significant drfferences rn pa1rw1se compansons.

b a • Tank 30 U Port: Surface OJ] Port: 10 M

-l -a. 20 (1) -__.a. 3 "0 p ....(1) c a a P.)..... c ro .... en (1) 10 b -0 10 __.(')

0+-- Salinity Temperature Figure C-2. Relationship between density of organisms in segregated ballast water arriving to Port Valdez, Alaska in oil tankers from domestic ports and age of ballast water. Density measures were those derived from quantitative analysis of plankton samples, and age indicates the number of days Since the ballast water was gravitated or pumped aboard.

('")- 100000 E • Annelida ~ 6 Mollusca 0 0 g g 0 Total ._...z 10000 AK 6 0 >- e g 0 6 ·--(J) r::: 1000 • Q.) 0 ~ 6 ~ •6 E 100 (J) • • r::: •6 ctl ...Ol 10 0 Ol 0 .. L..l _J 1 0 2 4 6 8 Ballast Water Age (days) Figure C-3. Companson of dens1ttes for planktonic organ1sms measured 1n non-exchanged ballast water of sh1ps from domestiC ports versus exchanged ballast water of a smgle sh1p from a fore1gn port Shown are means and 95% confidence Intervals of all non-exd1anged ballast samples for the respective taxa.

500- Polychaetes 400- 300- 200- 100- l

0 I I

1250- Molluscs 1000- (") E 750- :... 500- Q) 0. 250- :... Q) .c 0 I I E ::s Barnacles z 1250- 1000- 750- 500- 250-

01 I Total 7500

5000

2500

0 Nonexchange Exchange Table 0-l. Characten.\!lcs of ballast water sampled on two od tankers that conducted (.;X pen mental ballast water exchange w!lh segregated ballast water en route to Port Valdez from San Francisco Bay Shown are the sources of ballast warer. !ocatJOn of ballast water exchange, amount n~r}; of ha!!ast water exchanged, method of exchange, tanks (exchanged and non-exchanged) that were sampled. and the salinJtJes of sampled tanks for each sh1p

Ship Long Beach Benecia Original BW Source San Francisco, CA Benecia, CA Exchange BW Source 500-600 mi offshore British Columbia 500-600 mi offshore British Columbia %Exchanged 300% 100% Method of Exchange Flow Through Flow Through Original BW Tanks 2 Starboard. 4 Port 2 Starboard. 4 Port Exchange BW Tanks 4 Starboard. 2 Port 4 Starboard. 2 Port Original BW Salinity 32 ppt 10 ppt Exchange BW Salinity 35ppt 25 ppt Tabie 0~2. Comparison of plankton dcns1tics m exchanged vcrsu.<.; non·exchtmgcd <.;egrcgatcd balla:.t wa~cr followmg ballast water exchange experiment aboard two tanker:. en n'utc to Port Valdez from San

FrJ.ncJsco Bay tn May-June 1997. Dcnstues (#/m l) and standard error an.: <..hf>Wn for each taxonom 1c group by treatment and ship: values shown represent means for 2 tanks (c\tlmatccJ by 2 net tow-.;), ba-.;cd upon quantitative analysis.

Long Beach I Benecia

I ~ E};cbangf!Q: 6Yi w~mu;~bangeg 6W I E~C:tlQ:09"tl 6Y:l tlQD~};~baog~ Mean SE Mean SE I Mean SE Mean SE I Olnon.agellata 182 48 162 63 120 120 I 246 24 201 95 Olatom.acea I 01SCOtd 1686.09 590.31 499 ss 473.29 I 2670 42 2422 57 9013 341)4 Centrale 781.93 332.70 I 18 11 \6 33 ~Protozoa • I Foraf'nlnilera 35.44 30.47 3.61 3.61 I 541 I 72 I TIOIIOOlda 3.07 1.21 I Platyt\4tlmlnlhes I Turbellana 1.20 1.20 I Nem&f"lu 0.16 0.16 I Annelid.a I POlydtaele Chaetopteridae 0.62 0.62 Magelon!dae 0.67 0.67 4.78 4.78 Nephthydae 4.78 4.78 Phyilododdae L\9 1.19 Potynotdae 0.27 0.27 35.89 19.03 Spionidae 9.58 -4.62 852.55 252.96 0.62 0.62 0.62 0.62 unknOwn larvae 9.92 0.01 64.69 11.72 0.59 0.59 MoiiU$C-1 BNaMa 3.12 2.50 437.39 49.70 0.62 0.62 Gastropoda 8.90 0.21 156.21 82.19 3.05 1.87 1.19 1.19 Pleropoda 0.16 0.16 - 0.15 0.15 - Crume< a Clmpe468 28 14 4.27 I Harpactac01d )4 42 33 24 I An~"upoda 0.15 I 0.15 Cumacea 0 IS 0 15 I '20 J 28 0 31 031 ')e<:apoda I 'oea 015 0 15 I I 0 75 044 Mystdacea I 2'0 ! I 7 20 68 2.60 '"""""' I Bryotoa 5054 )1 84 I 20 I Phoronld11 ' 20 I Chae!.:~gnatha 016 0 16 I Chordata I '..aNacea )9.59 16 94 3823 31 01 I ! 19 I 19 "'sl'l I 015 015 Oth

TANK 1 TANK 2 Day1 Day2 Day3 Oay1 Day2 Day3 •EAN SE MEAN SE -'•lEAN SE MEAN SE MEAN SE MEAN SE O!NCFLAGELLA TA C11r;a!lum 8.28 Perldlnlum sp. 4.311 4.311 2:.87 2.81 "' 8.28 5.28 1 ..... • OIA TOMACE.A DiscOid 3711.U- TT.57 3611.111 122.18 2715.63 31.62 81.88 1 T&.H 4118.87 4;.811 75.J.l l C•ntrate 155.12 37.35 t47.n 1311.56 11.411 11.411 22.8] 22.$7 322.H 256.78 5.75 11 P•nn;ate 1.-4-4 ,_ ....

~PROTOZOA- Fon;mlnlfen. 5.53 5.52 Tintlnnldo. l I CNIOARIA 6.73 11.43 12.17 unld. me<:IU$0. 11.311 ... 31.15 11.67 8.-43 57.-46 111.32 " PLATYHELMINTHES ,_,, 5.53 2.77 2.77 l TurtaiLJ!'bi - 1.38 MOUer'SJGl!tte'st.arvn 1.38 1.38 NEMERTEA pillldlum larvu - 1.38 1.]8 ANNEUOA : Polyno«sn - 1.]8 ,_,.1.]8 -- Phyllodocldu 1.38 1.38 1.38 ,_.... 19..23 ,., 42.711 ua 6<1.87 4.33 12...37 SploniC..• 2-4.-42 .. ... "-" " NephthydU 11.66 IUS 4.15 4.14 22.83 22.87 SyttldU 0.12 0.72 0.38 1 Ow.nklae - - 15.87 1.35 Mo.gelonldo.e 2.77 3 unknown l.atvilll 185..vt 55.34 25.68 17.48 68.1M 68.116 187.18 "-" 1411.62 111.17 1.36 2l MOLLUSCA -C.1-C 11 ~stropod.J 211.511 l.St «5.78 1211.11 33.J.l 66.78 338.86 21.40 -4118..25 ill.-48 ,.,_ 1t.17 e88.58 373.5-C 126.-Cil 656.118 88.35 8-43.36 1.17 157.85 Slvalvl.l .. •u.s. " CRUSTACEA C!rrlpedl.a rntuplil l7Ut ,_,. 2111.36 1.2< 215.-4-4 -C$11.68 3<51 27-C.67 51.75 25.11 17 811.-48 '"8.82 28..211 3.-CS 55.28 2.77 23.30 c:yprlds "-" 13.85 n.13 16.112 ' Cop1podo. n:.uplll ,..__,, ,.,. 51.37 11.12 11!3.47 821.66 572.711 21111.55 182.81 356.15 17.35 .,,_,, _, 111.611 tl-47..27 215.48 817.n 326.41 12-C6.33 112.-CII -«1.55 t7 c~podltes 221.22 ,.. " H~rp.aetkoirU Mlcro·uteltl sp. 2.3711 2.15 33.7-4 2.14 2!.73 17..2-C 6<1.18 2.2.7! 56.511 15.18 2-4.80 23 Eutropln.a sp. - .... 4.85 - unknown Ho.rp.act..c:old - unknown benthic: Hillrp.. U2 8.62 5.75 5.75 -C.15 -C.1-C '-" ' ~l;moJd.;, Aeartla sp. 0.12 5.70 1.113 2115.87 238.-C5 -C.72 !.iT 1-C2.18 51.75 245.18 317 Cal.anus sp. '-'-'' l~3 2.51 us 2.12 1.:17 Centrop;~ges sp. 2.87 '-" 2.18 Po.r.aeaLJnus sp. 1U2 ... 87.57 55.118 $.32 2.87 53.14 -C.8l 128.36 75.112 15.13 ' Pseudocat.anus sp. 17.tS ,_ .. 51..2& 211.18 71.81 -C3.t-4 -C.12 O.Sil 85.57 19.33 8.82 3 Puudodt.apromus spp. U8 1.38 5.32 2.87 1.38 4.15 .. ,. 12.37 Tort.anus sp. 1.« 1.« 22.112 2.17 "' ' Cyclopold.ii Olthornt spp. 111.311 0.34 1.38 1.38 4.26 37.27 4.15 1.38 UmnoothOIU sp. 11.66 11.66 22.!111 11.2-C 26.22 12.42 '-" " PO 111 Tal be D-2. M1ssmg bars tndicate that the respective taxa were not of sufficient abundance to estimate changes for that vessel (see text)

• Benecia

~ Long Beach

20 40 60 80 100 Percentage Reduction Figure D-2. Percent mcreasc during ballast water exchange experiment of taxa that derived from oceanic walcrs and were relattvely abundant tn exchanged tanks. Shown is the percent increase for each taxonomic group by ship, as calculated from mean densities in Table D-2 MrsStng bars indrcate that the respective taxa were not of sufficient abundance to estimate changes for that vessel (see text) .

• Benecia benthic Harpacticoid ~ Long Beach

Foramtnifera

Discoid Diatom

Ceratium

10 100 1000 10000 100000 Percentage Increase (Log) Table E-1. Charactensttcs of all ballast watecarnvtng to Port Valdez, Alaska for each of 16 oil tankers. Shown are the sources and volumes of clean (segregated) and oily (nonsegregated) ballast water, percent of total ballast water that IS nonsegregated. and date of arnval for each vessel. Mean and standard errors are also gtven for the three volume characteriStiCS (EX tndtcates shtps that conducted some open ocean ballast water e'change; BWOB denotes Ballast Water on Board tn mctnc tons (MTJ)

Segregated Segregated BW Nonsegregated Nonsegregated BW Ship Name Date of Arrival "!.Nonsegregated BW BWOB (Mn Source I BWOB (MT) Source ARGO Spirit 23.10511997 27421 '14 long Beach. CA 55415.16 long Beach, CA I 67% 29174.45 ARGO Anchorage 25/0511997 21529.21 Charry Pt. WA I Cherry Pt. WA 58% 14414.80 Baton Rouge 2610511997 32233.86 Anacortes, WA I Anacortes. WA 31''/., 37-47N 122·34 W, 38· long Beach (El<.) 2610511997 28306.00 San Francisco. CA 36786.44 I 01N 124·19W 57''!.. Potomac Trader 27/0511997 10527.83 Cook Inlet, AK I 0.00 none QOfo Chevron Mississippi 27/0511997 19764.07 Anacortes. WA I 2594.36 Anacortes. WA 12% Cherry pt., WA ARCO FairbankS 2810511997 21504.00 Cherry Pt .. WA 28635.50 57% 25 % -Richmond, CA 01$ Washington 2810511997 508.10 Richmond, CA 43680.19 75% ·lOOmioHshore 99% SIR North Slope 02106/1997 48192.27 Portland, OR 50362.50 Portland, OA 51% AACOJuneau 02106/1997 22578.95 Cherry Pt., WA 34292.68 Cherry Pt., WA 60% SIR San Francisco 02105/1997 30241.10 Anacortes, WA 11383.47 Anacortes, WA 27% ST Alaska 02'06'1997 57497.61 San Francisco, CA 14668.72 San Francisco, CA 20"/., ARCO Independence 03106/1997 29831.57 Long Beadl, CA 60304.74 Long Beach, CA 67% OMI Columbfa 03106/1997 16701.16 Barber's Pt .• HI 50168.32 Barber's Pt., HI 75% 45-40N 156-.SOE, Prince William Sound 0410611997 26201.70 40-l 16963.43 Yosu. Korea 27N 134·39E 39% Senecia(E><.) 06106/1997 22706.44 Beneda. CA I 29692.35 Beneda, CA 57% Mean :25984.06 MT Mean =29908.58 MT Mean=49%

SE = 3317.15 MT SE = 4692.81 MT SE=0-06 Table E-2 Characteristics of nonsegregated ballast water sampled at 4 stages of the Ballast Water

Treatment Facdtty 1n Valdez, Alaska. Shown are the date. source. temperature. and saitntty of water -;amp led at each stage. Temperature and salinity measures represent means of two consecutive samples (sec text) Multiple sources indicated water was commmg!cd 1n the Treatment Facdily.

TREATMENT DATE SHIP SOURCE(SJ TEMPERATURE SALINITY (°C ( pt) A CHICK ARivlS 05/24/97 ARCO SPIRJT 12 32 05/25/97 ARCO ANCHORAGE 10.75 29 05/27/97 SIR LONG BEACH 12 32 05/27/97 SIR BATON ROUGE 10 28 05/27/97 CHEVRON MISSISSIPPI II 31 05/28/97 ARCO FAIRBANKS II 29.5 06/02/97 SIR NORTH SLOPE 14 13 06/02/97 SIR SAN FRANCISCO II 31 06/02/97 BT ALASKA IU 31 06/03/97 ARCO rNDEPENDENCE l3 32 06/03/97 OM! COLUMBIA 15 35

B. 90's TANK 05/24/97 ARCOSPIRIT 12 32 05/25/97 ARCO ANCHORAGE 12 30 05/27/97 SIR LONG BEACHIBA TON ROUGE 12 32 05/28/97 CHEVRON MISSISSIPPI 14.5 30 06/02/97 SIR NORTH SLOPE/ BT ALASKA 10 l3 06/03/97 SIR SAN FRANCISCO/ BT ALASKA 12 32 06103197 ARCO rNDEPENDENCE I2 32 06104191 OM! COLUMBWARCO rNDEPENDENCE 13 34

C DAFT 05/24/97 ARCO SPIRJT 12 32 05115197 ARCO ANCHORAGE 13 30 05/28197 SIR LONG BEACHIBA TON ROUGE 14 31 05/28/97 CHEVRON MISSISSIPPI/OS WASHINGTON 13 30 06/02/97 SIR NORTH SLOPE/ BT ALASKA 12 12.5 06/03/97 SIR SAN FRANCISCO/BT ALASKA 13 32 06/03/97 ARCOINDEPENDENCE 13 30 06/04/97 OM! COLUMBINARCO INDEPENDENCE 14 34

D BIT 05/25/97 ARCO SPIRJT 12 32 05/26/97 ARCO ANCHORAGE 12.75 33 05/28/97 SIR LONG BEACHIBA TON ROUGE IS 3 I 0610 l/97 CHEVRON MISSISSIPPI 14 30 06103197 SIR NORTH SLOPEIBT ALASKA 13 23 5 06/03/97 SIR SAN FRANCISCO/BT ALASKA 13 32 06/03/97 ARCOINDEPENDENCE ll 29 06/04/97 OM! COLUMBINARCO INDEPENDENCE 14 32 Table E-3. Percent occurrence of organisms m each taxonomic group that occurred in samples collected from the Ballast Water Treatment Facility 1n Valdez. Alaksa. Shown are the mean number of organisms counted from preserved samples from each of the four locations (with indicated samples stzes) lmponamly. these counts do not distinguish between live and dead organisms (see text for discussion). All organisms encountered were included in the classification scheme below.

CHICK ARMS (n=22) 90's TANK (n•16) DAFT (n•16) en (n=l6) ARTHROPODS

AMPHIPOOS 4.55 0.00 0.00 0.00 BARNACLE CYPRIOS 1364 6.25 0.00 000 BARNACLE NAUPUI 45,45 25.00 6.25 2500 COPE PODS

CALANOIDS $4.55 50.00 6.25 625 COPEPODITE 50.00 43.75 12.50 6.25 CYCLOPOIDS 27 27 31.25 18.75 12.50 HARPAC TlCOIOS 31 82 6.25 0.00 12 50 NAUPUI 4545 37.5 18.75 12.50 POE.CtUSTOME. 13.64 0.00 0.00 000 CUMACEANS 4.55 0.00 0.00 000 MITE 909 0.00 0.00 000 MOLLUSCS BIVALVES 18.18 6.25 0.00 000 GASTROPODS 13.64 0.00 0.00 000 NEMATODES 36.36 37.50 93.75 100.00 PLA TYHELMINTHE.S

TURBELLARIANS 9.09 0.00 0.00 000 DINOFLAGELLATES CERA TIUM 18.18 43.75 25.00 2500 PERIOIN!UM 36.36 - 18.75 12.5 18 75 DIATOMS

CHAIN F ORMlNG l8.18 18.75 31.25 625 DISCOID 81.82 93.75 81.25 93 75 ROTIFERS 0.00 6.25 0.00 12 50 PROTOZOANS

TINTINNIDS 31.82 25.00 18.75 56.25 EGGS 50.00 37.50 12.50 25.00 1 E 4 D · f · ms (#im') in each taxonomic group that occurred in samples collected from the Ballast Water Treatemem Factloty tn Valdez, Tab e - ensoty o organts f f · ( · h · d d ·· le · Alaska S.hown are the means, standard errors, and maximum counts obtained from preserved samples for each o the our 1ocauons wtt tn tcate s.1 mp s sizes). ·Importantly, these counts do not distinguish between live and dead organisms (see text for dtscusston). All orgamsms encountered were tncluded tn the classification scheme below.

CHICK ARMS (n=16) 90's TANKS (n-9) DAFT (n•6) BIT (n•t2) Mean SE Maximum Mean S.E. Maximum Mean S.E. ARTHROPODS Maximum Mean S E Max1mum AMPHIPOOS 000 000 0.02 0.00 0.00 0.00 0.00 0.00 0.00 BARNACLE CYPRIOS 0.00 000 000 0 02 0 02 032 0.00 0,00 0.02 0.00 0.00 0.00 0.00 000 BARNACLE NAUPLH 0 14 006 0.76 000 0.01 0.00 0.04 0.00 0.00 COPE PODS 0.02 0.13 006 064 CALANOIDS 0.14 o.oa 1.12 0.04 0.03 0.30 0.00 0.00 0.00 0.11 011 127 COPEPODITE 0 07 002 0.23 0.06 0.04 0.34 0.02 0.02 0.11 CYCLOPOIDS 003 0 OJ 032 HARPACTICOIDS 002 001 0.10 0.01 0,01 0.06 0.00 0.00 0.00 0.05 NAUPLII 0.04 004 0 32 0.01 0.19 0.03 0.02 0.17 0.02 0.01 o.oa 0.05 004 032 POECIUSTOME 000 000 0.04 0.00 0.00 0.00 0.00 0.00 0.00 0.00 CUMACEANS 000 000 0.00 002 0.00 0.00 0.00 0.00 0.00 0.00 0.00 000 MITE 000 0.00 0.04 0.00 d: 0.00 0.00 0,00 0.00 0.00 MOLLUSCS 0.00 0.00 000 BIVALVES 002 001 0.19 0.00 0.00 0.04 0.00 0.00 GASTROPODS 0.00 0.00 000 000 000 0.00 0.02 0.00 0.00 0.00 0.00 0.00 0.00 0.00 000 NEMATODES 002 001 000 o.oa 0.53 0.44 4.03 0.28 0.11 PLATYHELMINTHES 0.76 100 36 1 l 35 159 95 TURBELLARIANS 000 000 004 0.00 0.00 0,00 0.00 0.00 DINOFLAGELLATES 0.00 000 000 000 CERA TIUM 0 02 001 0.15 0.27 0.24 2.15 0.19 0.12 072 011 005 032 PERIDINIUM 0 31 o'Xf 3.06' 0.01 0.00 0.04 0.01 0.00 002 DIATOMS 019 011 127 CHAIN FOf~MitK~ 0 01 001 011 0 01 0.00 0.04 0.01 0.00 0 02 DISCOID 000 000 DOl 5 37 205 29.15 43 32 25.26 230.28 20.16 10.63 55 06 98 02 3J25 ).35 l i ROTIFERS 000 000 000 0.00 0.00 0.02 0.00 0.00 0.00 CILIATED PROTOZOANS 268 265 31 80 TINTINNIDS 0 07 0.04 0.65 0.06 0.05 0.49 0.08 0.05 0.29 EGGS 0.56 0.26 318 0.13 0.04 0.51 0.07 0.03 0.23 0.00 0.00 0.00 064 050 TOTAL 7 32 604 2.62 36.73 45.36 26.27 236.94 21.33 11.13 58.31 285.35 53 71 605 85 Port ValdcL

Chicksan Arm Outfall Diffuser

Biological * Samples Taken Treatment (BT) Tanks

* Dissolved Air Filtration (DAF) Oil Recovery Fadlity (80's) Tanks Primary Separation (90's) Tanks '

Figure E-l. Diagram of the Ballast Water Treatment f'acdny in Port Valdez. Alaska. Asterisks (•) tndicate the four locations of sample collecl!ons tn the tre,ument process: Chick san Arm. 90s Tanks. DAF Tanks. and BT Tanks. 150 1()() Ocns1ty Nematodes C'l 0 '% Al1ve E 75 100 ;t' ~ -J ;>. 0 ... 50 ):> <11 c < Q) ro 0 25

0 150 100 ..- C'l Diatoms E .... 75 100 # "-' 0~ ~ 50 ):> 0 c ID 50 (I)< 0 25

0 0 1 100 ,.... Other (I) E ..... 75 ._,# 100 ~ ;>. 0 ... 50 ):> 0 c < ID 50 ro 0 25

---f--0 Chick 90's DAFT BTT

Figure E-2. Total density of organisms and percent alive in samples collected from the Ballast Water Treatment Facility in Valdez, Alaska. Shown are means ( +s.e.) for density and% alive for each of three taxonomtc groups at the four different stages of treatment. Taxonomic groups include nematodes. discoid diatoms, and all other taxa combined. Question marks denote the rough but conservative estimate on percent alive for diatoms, since this was impossible to determine for some individuals (see text). [Treatment stages: Chick= Chicksan Arms; 90s= 90s Tanks: DAFT'= Dissolved Air Filtration Tanks: BTT =Biological Treatment Tanks; Sample sizes as shown in Table 4. )