Biological Invasions of Cold-Water Coastal Ecosystems - Aquatic ...

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BIOLOGICAL INVASIONS OF COLD-WATER COASTAL ECOSYSTEMS: 

BALLAST-MEDIATED INTRODUCTIONS IN 

PORT VALDEZ / PRINCE WILLIAM SOUND, ALASKA 

PRESENTED TO: 

FINAL PROJECT REPORT 

15 March 2000 

Regional Citizens’ Advisory Council of Prince William Sound 

P.O. Box 3089, 154 Fairbanks Drive 

Valdez, AK 99686 

USA 

Telephone: 907-835-5957 Fax: 907-835-5926 

Email: www.pwsrcac.org 

U.S. Fish and Wildlife Service 

PO Box 1670, 43655 Kalifornski Beach Road 

Soldatna, AK 99669 

National Sea Grant Program 

Washington, DC 

Alaska Sea Grant Program, University of Alaska Fairbanks 

Oregon Sea Grant Program, Oregon State University 

SeaRiver Maritime, Inc. 

ARCO Marine, Inc. 

British Petroleum, Inc. 

American Petroleum Institute 

Alyeska Pipeline Company 

PRESENTED BY: 

Lead Principal Investigators: 

• Dr. Anson H. Hines, Ph.D., Marine Ecologist 

• Dr. Gregory M. Ruiz, Ph.D., Marine Ecologist 

Smithsonian Environmental Research Center 

P.O. Box 28, 647 Contees Wharf Road 

Edgewater, MD 21037-0028 

USA 

Telephone: 301-261-4190 ext. 208 (Hines), 227 (Ruiz) Fax: 301-261-7954 

Email: hines@serc.si.edu, ruiz@serc.si.edu

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Co-Investigators: 

• Dr. John Chapman, Ph.D., Marine Biologist 

• Dr. Gayle I. Hansen, Ph.D., Marine Phycologist 

Hatfield Marine Science Center 

Oregon State University 

Newport, OR 97365-5296 

USA 

• Dr. James T. Carlton, Ph.D., Director 

Maritime Studies Program 

Mystic Seaport 

Williams College 

Mystic, CT 06355-0990 

USA 

• Ms. Nora Foster, M.S., Curator 

University of Alaska Museum 

University of Alaska Fairbanks 

Fairbanks, AK 99775 

USA 

• Dr. Howard M. Feder, Ph.D., Professor 

Institute of Marine Sciences 

University of Alaska Fairbanks 

Fairbanks, AK 99775 

USA 

Contributing Taxonomic Experts: 

• Dr. Jeffrey Cordell. School of Fisheries, University of Washington 

• Dr. Jeffrey Goddard, University of California, Santa Barbara 

• Dr. Paul W. Fofonoff, Smithsonian Environmental Research Center 

• Dr. Jerry Kudenov, University of Alaska, Anchorage 

• Dr. Gretchen Lambert, Friday Harbor Laboratories, University of Washington 

• Dr. Claudia Mills, Friday Harbor Laboratories, University of Washington 

• Ms. Lise Schickel, University of California, Santa Barbara 

• Dr. Judith Winston, Virginia Museum of Natural History 

• Ms. Lea Ann Henry, University of Toronto 

• Ms. Olga Kalata, School of Fisheries, University of Washington 

Biological Technicians: 

• Melissa Frey, Smithsonian Environmental Research Center 

• George Smith, Smithsonian Environmental Research Center 

• Max Hoberg, University of Alaska Fairbanks 

Field Assistants: 

• James Wade, Valdez

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• Stephan Benda, Valdez 

• Todd Miller, Hatfield Marine Science Center, Oregon State University 

Ballast Water Exchange Experiments and Plankton Analysis: 

• Safra Altman, Smithsonian Environmental Research Center 

• Sara Chaves, Smithsonian Environmental Research Center 

• Tami Huber, Smithsonian Environmental Research Center 

• Dani Lipski, Smithsonian Environmental Research Center 

• Kate Murphy, Smithsonian Environmental Research Center 

• Kimberly Philips, Smithsonian Environmental Research Center 

• Brian Steves, Smithsonian Environmental Research Center

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EXECUTIVE SUMMARY 

PROJECT OVERVIEW 

This study assesses the risk of biological invasion by nonindigenous species (NIS) associated 

with oil tanker traffic and ballast water management for Port Valdez / Prince William 

Sound (PWS), Alaska. This study included 8 major components: 

• Review of risk factors for NIS invasions and ship-mediated transfer of species relevant to 

PWS, a high latitude, cold-water marine ecosystem. 

• Analysis of ballast water delivery patterns and plankton communities associated with ballast 

water on tankers that arrived to PWS. 

• Experimental analysis of initial survivorship of ballast water organisms in temperaturesalinity 

combinations typical of receiving waters of Port Valdez. 

• Experimental measurements of the effect of ballast water exchange and voyage duration on 

plankton communities arriving on tankers to PWS. 

• Characterization of organisms fouling hulls and in sea chests of crude oil tankers. 

• Characterization of organisms in sediments of tanker ballast tanks. 

• Determination of NIS established within Alaska, as detected by field surveys and reviews of 

existing collections and literature conducted by experienced naturalists and taxonomic 

experts. 

• Analysis of the biodiversity of PWS. 

This study advances our understanding of invasion processes in many significant ways. 

• Our study provides the most comprehensive analysis worldwide of the abundance and 

taxonomic composition of plankton communities in the segregated ballast water of tankers as 

well as domestic ballast transfer by any vessel type. 

• We have undertaken an ambitious set of experimental and quantitative measures to (a) 

compare directly, for the first time, the relative efficiency of exchange methods (Empty– 

Refill and Flow-Through) for any vessel type or taxon, and (b) the effect of voyage duration 

on plankton survivorship in the ballast water of oil tankers. 

• We provide the first synthesis of NIS known in Alaska, resulting from an extensive literature 

review and field-based surveys. 

• The large scope of this study provides an unusually comprehensive analysis of the risks, 

mechanisms, and patterns of invasion in PWS. 

The project represents a cooperative and successful partnership of industry, citizen, 

agency, and scientific groups. This strong cooperative program addresses critical gaps in our 

understanding of invasion risks, as well as facilitates information exchange and participation 

among a broad spectrum of industry, citizen, agency, and scientific groups. 

• From a science perspective, this program results in a comprehensive analysis of invasion 

processes and risks for PWS, representing the first such study in the world for a high-latitude 

/ cold-water marine ecosystem.

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• From an industry and management perspective, this program assesses 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 disseminates findings and serves as a key source of 

information, especially through groups like the Smithsonian Environmental Research Center, 

the Regional Citizens’ Advisory Council of Prince William Sound, U.S. Fish & Wildlife 

Service, and NOAA Sea Grant. 

RESULTS 

Background 

Biological invasions of marine ecosystems in Alaska are a major environmental concern. 

• Biological invasions of coastal bays and estuaries are common throughout the world and are 

having significant ecological and economic impacts. 

• High-latitude / cold-water regions are also subject to biological invasions by many species 

with potential ecological and economic consequences similar to those reported for more 

temperate latitudes. 

• Transport of coastal planktonic organisms in ballast water of commercial ships appears to be 

the major source of new invasions worldwide in recent years. 

• Tankers arriving to Port Valdez release the third largest volume of ballast water of any U.S. 

port. 

BW Delivery Patterns and Biological Characteristics 

A large quantity of ballast water arrives to PWS in oil tankers. 

• For the past decade, tanker arrivals to Port Valdez have averaged 713 ships per year. 

• Tankers arriving to PWS in 1998 carried an estimated average of 65,775m 3 of total ballast 

water, including both segregated (non-oily) and nonsegregated (or oily) ballast water. 

• Segregated ballast water comprised an average of 54.7% of the total ballast water arriving to 

PWS in tankers. 

• Overall, an estimated 17,000,000 m 3 of segregated ballast water (an average of 32,715 m 3 per 

arrival) was discharged into PWS by oil tankers in 1998. 

Most ballast water delivered to PWS by crude oil tankers originates from U.S. domestic 

ports. 

• Tankers arriving directly from western U.S. ports accounted for 95.8% of the total tanker 

traffic, and 96% of the total segregated ballast water delivered by tankers, to PWS in 1998. 

• Arrivals from Puget Sound, San Francisco, and Long Beach comprised approximately 82.7% 

of all tanker traffic, as well as 86% of all segregated ballast water delivered by tankers, to 

PWS in 1998. 

• Most (69.6%) of the tankers arriving to Port Valdez from overseas came directly from Korea 

in 1998. 

• Tankers arriving from domestic ports transfer ballast water directly from that port to PWS, 

whereas foreign arrivals have replaced coastal ballast water with open-ocean exchange prior 

to their arrival.

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The voyage duration of tankers arriving to Port Valdez is relatively short compared to 

traffic arriving at other commercial ports, where invasions are common. 

• Ballast water spent an average of 6.6 days in the ballast tanks of oil tankers before arrival to 

Port Valdez, ranging between 4.8 to 10.2 days. 

A large quantity of planktonic organisms is released into PWS with segregated ballast 

water from oil tankers. 

• An average of 12,637 total organisms per m 3 (excluding chain-forming diatoms) was 

measured in our ballast water samples from 169 tanker arrivals, including those from both 

domestic and foreign source ports. 

• Overall, we estimate that roughly 264 billion organisms were delivered to PWS in the 

segregated ballast water of oil tankers during 1998. 

• Importantly, these estimates include only the largest plankton and miss many small 

planktonic organisms (e.g., bacteria, viruses, and other microorganisms), that would likely 

increase overall densities many fold. 

The abundance of planktonic organisms was greater in segregated ballast water from 

domestic source ports compared to that from foreign source ports. 

• Total density (across all taxonomic groups) of organisms was greatest on average in 

segregated ballast water from domestic arrivals compared to foreign arrivals. 

• Average densities of most taxonomic groups were 10- to 100-fold greater in segregated 

ballast water from domestic versus foreign sources. 

• The magnitude of density differences between domestic and foreign sources was much less 

for copepods and solitary diatoms. 

• Dinoflagellates were a notable exception to the general pattern, as average density was 

greatest in ballast water of the foreign arrivals. 

Significant variation existed in abundance of taxonomic groups in the segregated ballast 

water arriving from the major source ports. 

• Total density of organisms was lowest on average in ballast water from foreign arrivals 

compared to arrivals from each of the three major domestic ports (Puget Sound, San 

Francisco, and Long Beach) 

• Total density on average declined among the four major ports with increasing voyage 

duration. 

• In contrast, the greatest average densities for individual taxonomic groups (e.g., protozoans, 

brachyuran crabs, and bryozoans) did not always correspond to the shortest voyage duration. 

The abundance of plankton arriving in segregated ballast water from the major domestic 

ports varied both spatially and temporally. 

• The greatest densities occurred for all taxonomic groups, individually and combined, during 

the spring and summer months. 

• However, the timing of peak densities differed among taxonomic groups and among source 

ports.

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• The magnitude of seasonal variation in plankton densities also differed among source ports, 

being greatest for Puget Sound and San Francisco Bay compared to Long Beach. 

• Furthermore, significant annual variation also existed in the densities of plankton arriving to 

PWS from each of the major domestic source ports. 

NIS are present in the segregated ballast water released by oil tankers in PWS. 

• We identified 14 different nonindigenous species (13 crustaceans and 1 fish) arriving to Port 

Valdez in the ballast water of oil tankers. 

• To date, all of these identified NIS have been in ballast water from San Francisco Bay or 

Long Beach. 

• Importantly, these numbers are clearly underestimates, since only a subset of the plankton 

can be identified to species and only the largest fraction of planktonic organisms were 

included in our analyses. 

Organisms discharged in tanker ballast water, including known NIS, have high potential of 

initial survival in the salinity-temperature conditions of Port Valdez and PWS. 

• Seasonal cycles of salinities and temperatures in Port Valdez waters encompass the range of 

salinities and temperatures of arriving ballast waster, providing a good match between source 

ports and receiving waters. 

• Laboratory experiments indicate that a wide range of ballast water species (including some 

NIS) can survive the salinity and temperature conditions of Port Valdez upon initial 

discharge from tankers. 

Other Mechanisms of NIS Transport by Tankers 

Tankers also transfer organisms that are not in ballast water and that may become 

established in PWS. 

• Tanker hulls and sea chests sampled in dry dock sometimes carried a diverse array of fouling 

and nektonic organisms, including several NIS. 

• Sediment taken into ballast tanks during ballasting in shallow ports sometimes carried 

diverse and abundant bottom-dwelling organisms, including reproductive adult individuals. 

Ballast Water Exchange Experiments 

Preliminary experimental results suggest that ballast water exchange is as effective for oil 

tankers as for other vessel types. 

• Initial analyses suggest roughly 80-99% of the resident water is replaced per ballast water 

exchange event. 

• The efficacy of exchange appears to differ between exchange method, with Empty–Refill 

Exchange replacing the greatest proportion of water. 

• The efficacy of exchange also appears to differ among taxa. 

• Importantly, all analyses for the exchange experiments are still underway, and final results / 

conclusions are therefore pending project completion (anticipated in June 2000).

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Summary of NIS in Prince William Sound and Alaska 

A diverse array of NIS have been introduced into PWS and Alaska. 

• There are 24 species of NIS plants and animals in marine and estuarine ecosystems in 

Alaska, including 15 species recorded in PWS. 

• These NIS are taxonomically diverse and occupy a wide range of ecological niches and 

habitats, although there appear to be more NIS associated with boat harbors and with 

aquaculture activities. 

• Our focal taxonomic collections provided the first records of 7 NIS in Alaska, including 

some species that appear to be very recent introductions. 

• Many of the Alaskan NIS have larval stages which could be transported in ballast water. 

• None of the Alaskan NIS is clearly associated with ballast water of oil tankers as a primary 

mechanism of introduction, even though many NIS are frequently found in ballast water 

arriving to Port Valdez. 

• Instead, the transfer of NIS may have resulted from any one of multiple transfer mechanisms, 

including ballast water, ship fouling communities, and aquaculture. 

• Finally, it is important to note that many additional Alaskan marine species are cryptogenic 

(possibly introduced), as the historical baseline of biogeographic and taxonomic information 

is very limited for this biota. For example, we identified at least 29 cryptogenic species in 

Alaska (including 24 in PWS), exhibiting either wide global distributions often associated 

with spread by early shipping traffic or a variety of characteristics common to NIS. 

Biodiversity of Prince William Sound 

Taxonomy and biogeography of species in Alaskan marine ecosystems have received poor 

levels of study and understanding. 

• We discovered 10 new, previously undescribed species, as well as recorded range extensions 

for 74 other species from a diverse array of taxonomic groups. 

• It is now apparent that a large portion of many major groups remain undocumented, as well 

as cryptogenic in origin, due to limited surveys and historical analysis of the Alaskan biota. 

We have now initiated a biodiversity data base for marine species in PWS. 

• We have established a comprehensive data base for marine invertebrates in PWS. 

• The scope of this data base will be expanded to include algae, fish, mammals and birds in the 

next year. 

CONCLUSIONS 

Multiple risk factors exist that favor the establishment of NIS in PWS. 

• Approximately 550 tankers currently arrive per year to PWS and release an estimated 

17,000,000 metric tons of segregated ballast water. 

• Tankers repeatedly deliver ballast water from the same, limited source ports, providing 

repeated inoculations of the same species.

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• The voyage duration of these tankers is usually short (3-7 days), favoring high survivorship 

of transported plankton and resulting in the dense inoculation of competent organisms into 

PWS. 

• Environmental conditions of source ports match those in PWS for some portions of the year, 

and many organisms arriving in ballast water can tolerate conditions in receiving waters. 

• Most (95.6%) of arriving tankers do not undergo ballast water exchange, a process which can 

limit the transfer rate of NIS. 

• A large number (tens-to-hundreds) of NIS are known from the domestic ports that are the 

source of unexchanged ballast water arriving to PWS in oil tankers. 

• NIS are present in this domestic ballast water arriving to PWS in oil tankers. 

Ballast water exchange appears effective at reducing resident plankton on tankers, 

although a risk of invasion still exists. 

• Ballast exchange experiments suggest that tankers arriving to Port Valdez from foreign ports 

have reduced resident coastal organisms by > 90% through the current exchange practices. 

• Abundance of coastal organisms was 10-100 fold lower for oil tankers that were foreign 

arrivals (that underwent ballast water exchange) compared to domestic arrivals (that do not 

undergo exchange). 

• Although roughly equivalent to efficacy of exchange estimated for other vessel types, both 

data sets suggest that tens to hundreds of thousands of organisms/ship still arrive with 

exchanged ballast water. 

Alaskan waters, and those of PWS, are susceptible to invasion by NIS. 

• It is now evident that a diverse array of taxa have become established in Alaska and PWS. 

• These NIS occupy a broad range of marine and estuarine habitats. 

The number of marine NIS in Alaska appears to be significantly lower than other marine 

ecosystems at lower latitude. 

• Our surveys of PWS and Alaska were intensive and failed to detect many NIS known from 

the domestic source ports of oil tankers. 

• However, the limited historical record and scope of past surveys limits direct comparisons 

with low latitude marine ecosystems, for which extensive surveys and knowledge have been 

developed over decades to centuries of biological research. 

In general, the poor resolution of taxonomic and biogeographic data in Alaskan marine 

ecosystems is a substantial impediment for analysis of environmental impacts. 

• To date, we have been able to provide only a minimum estimate of NIS, as many species 

remain undescribed or cryptogenic until further analysis. 

• Assessment of other environmental impacts, such as oil spills, may also be limited without 

adequate baseline data on species composition and abundance. 

• We recommend a program of standardized surveys across multiple sites in PWS and Alaska 

to both improve the existing knowledge of NIS and provide a regional baseline of data.

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Acknowledgments 

Funding for this study was provided by Regional Citizens’ Advisory Council of Prince 

William Sound (RCAC), US Fish & Wildlife Service (USF&WS), and the NOAA National Sea 

Grant Program through Oregon State University (OSU) and University of Alaska Fairbanks 

(UAF). In-kind support and participation were provided by the Smithsonian Environmental 

Research Center (SERC), Maritime Studies Program/Mystic Seaport with Williams College, 

Alyeska Pipeline Service Company, and the oil shippers, particularly SeaRiver Maritime, ARCO 

Marine and British Petroleum (BP). Additional support for expanded ballast water exchange 

experiments was provided by the American Petroleum Institute, SeaRiver and ARCO, 

USF&WS, and U.S. Coast Guard. 

This project was stimulated and encouraged by the Nonindigenous Species Working 

Group of RCAC, chaired by Robert Benda (Prince William Sound Community College) and 

Gary Sonnevil (USF&WS). We thank Joel Kopp of RCAC for his able oversight and friendly 

assistance at all stages of this project, and Marilyn Leland and Linda Hyce of RCAC for their 

support. We are grateful to Rex Brown and the managers and staff of the Alyeska Pipeline 

Service Company and the Valdez Marine Terminal for access and help in sampling. The Masters, 

Officers, and crew members of the tankers gave willingly of their time and help; and we 

especially thank those of the ships SR Baytown, SR Long Beach, and SR Benicia, ARCO 

Independence, and ARCO Spirit for their extensive help on ballast water exchange experiments. 

Thanks also to W.P. (Pete) Rupp of SeaRiver Maritime and Victor Goldberg of ARCO Marine 

for their support of the exchange experiments. We especially thank the following agents in 

Valdez for their logistical assistance, advice and suggestions, and support: Kurt Hallier and 

Wayne Brandenburger of ARCO Marine; Bill Deppe, Phil Eichenberger, and John Poulos of 

SeaRiver Maritime, and Tom Colby of BP. The full cooperation of the oil companies operating 

these ships is gratefully acknowledged. 

Melissa Frey and George Smith of SERC provided key technical assistance for sampling 

ships, processing samples, conducting experiments, analysis, and many other crucial components 

at SERC’s laboratories in Valdez and in Maryland. Exchange experiments were assisted on 

board tankers by additional technicians of the SERC Invasions Biology Program: Linda 

McCann, Kim Philips, Safra Altman, Lynn Takata, Cathleen Coss, Kate Murphy, Dani Lipski, 

Laura Rodriguez and Brian Steves. Sorting of samples was assisted by Melissa Frey, George 

Smith, Sara Chaves, Kim Philips, Dani Lipski and Safra Altman. Data management and analysis 

were assisted by: Safra Altman, Melissa Frey, Midge Kramer, Kim Philips, George Smith, and 

Brian Steves. 

For assistance on the NIS cruises in Prince William Sound, special appreciation goes to: 

Todd Miller, for his assistance on the trip; Capt. Richard Vodicka who skippered the F/V 

Kristina in 1998 and helped in many ways large and small throughout the survey; Milos Falta, 

owner of the F/V Kristina and skipper during the 1999 surveys; Joel Kopp, RCAC, for loan of 

his zodiac, searching for a charter, and other logistical and support assistance; and the 

community of Tatitlek for housing the survey team, especially village leader Gary Kompkoff and 

boat operator/guide Steve Totemoff for logistical assistance in the survey of Valdez Arm and 

Tatitlek areas. Peter Jordan and Bryna Dunaway, high school students in Oregon, assisted in 

curating the dried plant collection. Mike Stekoll of the University of Alaska provided

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microscopes for the study. Lavern Weber, Director of the Hatfield Marine Science Center 

provided office and laboratory space for G.I. Hansen. These people all helped make the cruises a 

success. 

For assistance with the 1998 & 1999 Fouling Plate Surveys, we thank Gary Sonnevil (US 

Fish & Wildlife Service), Peter Armato (US Park Service and Alaska SeaLife Center), Chuck 

Pratt (Armin F. Koenig Fish Hatchery), Ken Morgan (Valdez Fish Hatchery), Andrea Tesch 

(Nuremburg Fish Hatchery), Stan Stephens (Stan Stephens Tours), and the pilots of Cordova Air 

Service for field assistance in deploying and retrieving plates. We appreciate use of facilities and 

further assistance from Alyeska Pipeline Service Company, Prince William Sound Science 

Center, and Prince William Sound Community College. In addition, we thank Conrad and 

Carmen Field for their help at Homer, especially for showing us “an unusual sea star” (Asterias 

amurensis). 

In addition to participation in the project by taxonomic experts (see cover page), species 

identifications were assisted by Klaus Ruetzler for sponges; Frank Ferrari, Olga Kalata and Chad 

Walter for copepods; Judy Winston for bryozoans; Lea Ann Henry for hydroids. Charles 

Lambert assisted with field collections of ascidians during the 1999 field survey. 

The cooperative support by all of these individuals and organizations made this project 

possible.

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Title Page 

Executive Summary 

Acknowledgements 

Table of Contents 

Table of Contents 

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1. Introduction 1-1 

1A. Project Goals 1-1 

1B. Structure & History of Project 1-1 

1C. Background 1-2 

1C1. Invasive Species & Ballast Water 1-2 

1C2. NIS in High Latitude/Cold Water Ecosystems 1-2 

1C3. Risk Factors for Prince William Sound 1-5 

2. Ballast Water Delivery Patterns 2-1 

3. Biological Characteristics of Ballast Water in Oil Tankers 3-1 

4. Predicting Initial Survival of Ballast Water Organisms 4-1 

5. Ballast Water Exchange Experiments 5-1 

6. Organisms in Sediments of Tanker Ballast Tanks 6-1 

7. Fouling Organisms on Hulls and in Sea Chests of Oil Tankers 7-1 

8. Summary of NIS in Prince William Sound and Alaska 8-1 

9. NIS Surveys 9A-1 

9A. Overview 9A-1 

9B. Rapid Community Assessments 9B-1 

9C. Focal Taxonomic Collections 9C1-1 

9C1. Marine Plants 9C1-1 

9C2. Planktonic Medusae, Ctenophores, and Pelagic Molluscs 9C2-1 

9C3. Polychaete Worms 9C3-1 

9C4. Peracaridan Crustaceans 9C4-1 

9C5. Copepod Crustaceans 9C5-1 

9C6. Decapod Crustaceans 9C6-1 

9C7. Shelled Molluscs 9C7-1 

9C8. Opisthobranch Molluscs 9C8-1

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9C9. Echinoderms 9C9-1 

9C10. Ascidians 9C10-1 

9D. Fouling Communities 9D-1 

9E. Museum, Voucher and Reference Specimens of Existing Collection 9E-1 

10. Biodiversity of Prince William Sound 10-1

Chapt 1. Introduction, page 1- 1 

Chapter 1. Introduction 

Anson H. Hines, Smithsonian Environmental Research Center 

Gregory M. Ruiz, Smithsonian Environmental Research Center 

1A. Project Goals 

The overall goal of this project was to assess the risk of biological invasion by 

nonindigenous species (NIS) introduced into Port Valdez / Prince William Sound, Alaska. 

Currently, ballast water is the major vector for introductions of NIS in coastal ecosystems, where 

ballast-mediated biological invasions are causing severe ecological and economic impacts. While 

the significance of ballast-mediated invasions has focused on temperate zone ports, little 

consideration has been given to NIS invasions at high latitudes, despite the volume of shipping 

and critical importance of certain cold-water ports to the world economy and especially US 

energy interests. Port Valdez is a high latitude-cold water port receiving the third largest annual 

volume of ballast water in the USA. Moreover, our recent review of NIS invasions at high 

latitude (Ruiz & Hines 1997) indicates that such cold water ecosystems have been invaded by a 

diverse array of marine and estuarine species. The specific objectives of the project were: 

• To analyze the delivery patterns, biological characteristics, and management practices of 

ballast water and other ships arriving to Port Valdez from coast-wise versus foreign voyages. 

• To assess viability of selected organisms arriving in tanker ballast water to Port Valdez. 

• To conduct experiments on the effectiveness of ballast water exchange procedures of tankers. 

• To evaluate organisms occurring in entrained sediments at the bottom of ballast tanks of 

crude oil tankers. 

• To evaluate fouling organisms on hulls and in sea chests of tankers as potential sources of 

NIS. 

• To analyze and search for NIS currently invading or already established in coastal waters of 

south central Alaska, using literature searches, an array of field sampling methods (field 

collections, fouling plates, and plankton sampling), and examination of existing preserved 

samples from Prince William Sound. 

The purpose of this report is to summarize the research conducted during 1997-1999 to 

assess the risk of biological invasions in Prince William Sound, especially with regard to oil 

tankers as a vector for transporting NIS into marine ecosystems. Progress during the project was 

reported in Ruiz & Hines, 1997 and Hines et al., 1998. Modified elements of the earlier reports 

are included in the present report, so as to provide a complete overview of the project within one 

document. Certain limited elements of research will be completed during 2000, including further 

analysis of existing collections at the University of Alaska Museum and Institute of Marine 

Sciences, and work-up of sample from ballast water exchange experiments conducted during 

summer 1999. These last elements will be reported separately upon completion. 

1B. Structure & History of Project 

This research project has built upon a Pilot Study conducted in 1997 (see Ruiz & Hines, 

1997), and includes an expanded scope of work conducted during 1998-1999. The multi-faceted 

approach to the research required a team of diverse CoPrincipal Investigators and subcontracted 

taxonomic experts. Drs. Anson Hines and Gregory Ruiz (SERC) have served as over-all project 

leaders, providing over-all administrative and scientific oversight for the team. In addition, Drs. 

Ruiz and Hines (SERC) had primary responsibilities for: analysis of ballast water delivery


patterns; biological characteristics of ballast water; experimental analysis of initial survival of 

ballast water organisms; ballast water exchange experiments; analysis of ballast tank sediments 

and tanker hull fouling; fouling community analysis, and management of most of the focal 

taxonomic studies; as well many aspects of field surveys of Prince William Sound. To sample 

and analyze ballast water of tankers arriving to Port Valez, two Biological Technicians (Melissa 

Frey and George Smith, of SERC) alternately rotated at about four month intervals between 

SERC’s temporary laboratory established in Valdez, Alaska and the SERC Biological Invasions 

Laboratory in Edgewater, Maryland. Ballast water exchange experiments were conducted with 

participation by several technicians and students from the SERC Biological Invasions Laboratory 

(see Acknowledgments). Co-PIs Nora Foster (UAF) and Dr. Howard Feder (UAF) had primary 

responsibility for analysis of existing samples in museum, reference and voucher collections in 

the UA Museum and UAF Institute of Marine Science. Nora Foster also participated actively in 

rapid community assessment surveys of Prince William Sound, and focal taxonomic analysis of 

molluscs. Dr. Howard Feder provided oversight to subcontracted focal taxonomic analysis of 

polychaetes. CoPI Dr. John Chapman (OSU) had primary responsibility to conduct rapid 

community assessment surveys of invertebrates of Prince William Sound, with focal taxonomic 

analysis of pericaridean crustaceans. CoPI Dr. Gayle Hansen (OSU) had primary responsibility 

for focal taxonomic field surveys of marine plants (especially macro-algae) of Prince William 

Sound. Dr. James Carlton (Mystic Seaport, Williams College) had primary responsibility for 

surveys of fouling communities of Prince William Sound. In addition, an array of systematic 

experts was subcontracted to analyze several focal taxonomic groups (see Chapter 9 below). 

Authorship of the chapters and subsections of this report indicate primary responsibilities for 

each major element. 

Throughout the 1997 Pilot Study and the 1998-1999 expanded phase, the project received 

guidance and comment from the Alaska NIS Working Group, which was organized by the 

RCAC of Prince William Sound and composed of academic scientists, resource managers from 

state and federal agencies, representatives of the oil and shipping industries, and concerned 

citizens of Alaska (see also Acknowledgments). 

Initial funding for the Pilot Study was provided by PWS RCAC, US Fish & Wildlife 

Service, and the US Coast Guard (Ruiz & Hines, 1997). Expanded research for the present 

project was extended with a proposal submitted in 1997 to the National Sea Grant Program, with 

co-funding from RCAC, USF&WS, Alyeska Pipeline Service Company, and in-kind support 

from the oil shipping companies (especially ARCO Marine, BP and SeaRiver Maritime). Funds 

awarded from the National Sea Grant Program were distributed to the Oregon Sea Grant 

Program for John Chapman at Hatfield Marine Science Center and to Alaska Sea Grant Program 

for Nora Foster and Howard Feder at University of Alaska Fairbanks and UA Museum. 

However, funding from Alaska Sea Grant was delayed for one year, so that funding was actually 

available beginning in 1999 and will carry through 2000. In 1998 SERC obtained supplemental 

funding from the American Petroleum Institute, supported by ARCO Marine and SeaRiver 

Maritime, to conduct ballast water exchange experiments on tankers. SERC also received 

further funding from USF&WS for these ballast water exchange experiments during 1999 

through a proposal submitted to the National Sea Grant Program. The work plan for the ballast 

water exchange experiments specifies that sample processing and analysis continue into 2000. 

SERC’s technical staff worked in close coordination with the shipping agents, masters, 

officers and crews of the oil tankers, and with Alyeska staff of the Valdez Marine Terminal. All


of these industry participants provided in-kind contributions and worked actively and 

cooperatively to assist project operations to sample ballast water and to conduct ballast water 

exchange experiments (see also Acknowledgments). 

Significantly more work than originally proposed was accomplished in nearly all 

components of the project. Moreover, we have been successful in gaining additional resources to 

support expanded elements of the project, including contributions in kind, and added external 

funds for ballast water exchange experiments. Most importantly, a hallmark of the project was 

the enthusiastic support of the project by the full array of private citizens, scientific institutions, 

governmental agencies, and industry, which served as cooperative partners. 

1C. Background 

1C1. Invasive Species & Ballast Water 

Aquatic nuisance species have invaded many, perhaps most, freshwater and marine ports 

around the world. 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 any of the plankton are viable and become 

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, 1995), the Great Lakes (Mills et al., 1993), Chesapeake Bay (Ruiz et al.,1999), Hawaii 

(Coles et al. 1999), and elsewhere (Ruiz et al., 1997). In San Francisco Bay, the rate of invasion 

has increased to about one new NIS invasion every 16 weeks, probably as a result of increased 

ballast water discharge (Cohen and Carlton, 1995). Whether the invasion is Eurasian zebra 

mussels in the Great Lakes, Asian clams in San Francisco Bay, or North American 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 that 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; 1999; 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). 

1C2. NIS in High Latitude/Cold Water Ecosystems 

Although there has been no significant analysis of NIS in polar marine ecosystems, there 

have been a limited number of NIS surveys in high temperate latitudes between 40 o - 60 o and a 

study of the Baltic Sea, which includes a major bay that extends substantially above 60 o . These 

studies include three regions in the northern hemisphere (Baltic Sea, Wadden Sea, and United 

Kingdom)(Reise et al., 1999, Lepapakoski 1984) and one region in the southern hemisphere 

(Tasmania/New Zealand)(Hayward 1997, R. Thresher, 1999 pers. comm.). Together, these 

studies clearly demonstrate that invasions are not limited to lower latitudes. The number of 

known NIS at these locations ranges between 32 and 80 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. As with most invasions, the actual 

impacts remain unmeasured (e.g., Ruiz et al., 1999). Nonetheless, based upon reported 

abundances and known ecology, species such as the green crab Carcinus 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 single invasion event (Cohen and 

Carlton, 1995). 

The numbers of NIS at high latitudes may be lower than those for temperate regions, 

although it is not clear whether low numbers of documented NIS reflect lack of invasion in high 

latitude ecosystems or lack of research focused on the invasion biology of these areas. At the 

outset of this study, the number of NIS 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 Valdez oil spill in Prince William Sound and 

other ecological research throughout the region (Ruiz & Hines, 1997). 

We advanced two hypotheses why NIS have not been as evident at high latitude as at 

mid-latitudes: 

(1) NIS are truly rare at high latitude. High latitude communities may be resistant to invasion 

(e.g., severe seasonal stress requires specialized evolutionary adaptations not possessed by 

non-native species). Transport patterns may not have been conducive to inoculation. 

Shipping/ballast water is major source of rapidly escalating invasions in temperate latitudes, 

but perhaps neither the relatively recent (20 yrs) surge in 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, 1995). 

(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 NIS in San 

Francisco Bay and Pacific northwest coast, but as of 1995 the number of NIS documented in 

San Francisco Bay was 212 species (Cohen & Carlton, 1995) and is now nearly 250 species 

(J.T. Carlton, personal communication). Three years ago, the number of NIS 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 >140 NIS, and the list is still growing 

with continuing research. Despite extensive biological/ecological assessments of coastal 

ecosystems associated with oil spills in Alaska, NIS probably remain inadequately studied. 

The existing and on-going surveys from oil spill work and other studies in Prince William 

Sound are probably not adequate because those surveys were designed for purposes other 

than detecting introduced species. Also, they mainly focused on rocky shores rather than on 

soft-bottom and fouling communities that are most invaded in other regions (e.g., Cohen & 

Carlton, 1995). Most introduced species have been discovered by taxonomic experts 

systematically examining specimens previously identified by non-specialists or conducting 

field surveys of their own.


1C3. Risk Factors for Prince William Sound 

Prince William Sound is a relatively pristine, cold water ecosystem at high latitude. 

Approximately 20% of US domestic oil production is shipped from Port Valdez at the head of 

the Sound (Fig. 1.1). Tankers arriving to Port Valdez come primarily from domestic source ports 

of the west coast of North America and Hawaii; but in the past three years tankers also have been 

traveling from foreign ports, especially in eastern Asia and rarely other locations (Fig. 1.2). 

Tankers arriving to Prince William 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, a quantity of 

domestic ballast water that greatly exceeds the volumes of foreign ballast water released in other 

U.S. West Coast ports and is the third largest volume for all U.S. ports (behind port systems of 

New Orleans and Chesapeake Bay). 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 NIS were 

unknown prior to our Pilot Study (Ruiz & Hines, 1997). The Pilot Study showed that nonsegregated 

ballast water contained few live planktonic organisms upon leaving tankers and 

entering the treatment plant, and that there is little risk of NIS in the discharge of water from the 

treatment plant (Hines et. al., in press). Accordingly, all of our analyses in subsequent research 

presented in this report focus on segregated ballast water. 

Figure 1.1.Map of Prince William Sound, Alaska, showing location of Valdez Marine Terminal. 

Segregated ballast water from tankers is discharged directly into the Sound/Port without 

treatment. This volume is many orders of magnitude greater than the ballast water released by 

other types of ships traveling to Prince William Sound. Release of ballast water into Prince 

William Sound increased markedly with the opening of the trans-Alaska Pipeline in 1977. 

Tankers have made more than 15,000 voyages through Prince William Sound to Port Valdez 

since the startup of the terminal in 1977. From 1987-1994, tanker arrivals to Valdez averaged


799 per year but have declined to less than 600 per year currently. Since 1996, oil shipping 

patterns authorized by the US Congress have changed to allow sale of crude oil on foreign as 

well as domestic markets. Tankers from foreign ports are required to conduct mid-ocean 

exchange of their coastal ballast water, which is expected to reduce numbers of organisms 

transported from foreign coastal ecosystems to Alaskan waters. However, the effectiveness of 

mid-ocean exchange is poorly measured. Moreover, the greatest volume of ballast water coming 

to Alaska derives from domestic ports of the west coast of North America, which themselves are 

highly invaded by NIS. Tankers on these domestic, relatively short coast-wise voyages are not 

required to exchange ballast water. 

To determine whether the known NIS 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. Huge 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. The 

tanker traffic to Port Valdez releases the third largest volume of ballast water into a US port. 

Other things being equal, larger ballast volumes mean larger inoculations of NIS. 

2. Short voyage 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; 1999). 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; 1999). 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 (see Chapt 2 and 3 below). 

3. Pattern of repeated delivery 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, Prince 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 (Fig. 1.2, Chapt 2 below). 

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. 

Temperature-salinity conditions in segregated ballast water of tankers arriving from several 

west coast source ports is shown to be similar to the waters of Port Valdez (see Chapt 4 

below). In the fjords of Prince William Sound, such as Port Valdez, heavy loads of


suspended sediment during summer snow/glacial melt also may be a major stress on marine 

organisms in surface waters. 

Figure 1.2. Major tanker routes to Port Valdez, Alaska. 

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., 1996). Exchange presumably limits the risk of invasion, as mid-ocean species are 

generally thought to be incapable of invading near-shore habitats. Delivery of ballast from 

coastal ports of the U.S. West Coast without oceanic exchange before release into Prince 

William Sound poses an elevated risk. Further experimental assessment of this role of 

mid-ocean exchange in reducing plankton abundance and diversity in ballast water is 

presented below in “Ballast Water Exchange Experiments”. While ballast water exchange is 

required for tankers from foreign ports, the National Invasive Species Act of 1996 considers 

tankers from U.S. west coast ports to be domestic, coast-wise traffic that does not require 

exchange. 

6. High frequency of known NIS - especially 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 of NIS. 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. These, in turn, form the sources donating much 

of the ballast water delivered to Prince William Sound. The 310+ known NIS of the west 

coast of North America vary considerably in abundance among 6 latitudinally separate 

regions (southern California, San Francisco Bay, northern California, Coos Bay Oregon, 

northwest region from the Columbia River estuary to British Columbia, and Alaska) (Ruiz & 

Hines, 1997). The number of known NIS varies from about 80+ species in southern 

California to about 40+ species in the northwest region of Washington and British Columbia, 

with the largest number of nearly 250 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, mollusks, and annelids comprising major fractions of NIS at most locations. 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 of NIS


probably reflects the level of study 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 in several locations along the west 

coast, indicating that invasions by the same species have occurred widely across latitudinally 

separate sites. The similarity of NIS at less studied sites may be expressed as a percent 

overlap with NIS in well-studied San Francisco Bay. The overlap of NIS at west coast 

source ports with those in San Francisco Bay is high, ranging from about 60-75%. For the 

limited sample known for Alaska, the overlap of NIS with San Francisco Bay is substantially 

lower at about 25%. However, it is not clear whether this lower overlap reflects reduced 

compatibility with the Alaskan region or the small sample size, or both. 

7. History of other vectors for NIS in Prince William Sound. Although ballast water from 

tankers is currently a major vector for introductions of NIS in Prince William Sound, several 

other vectors have been, and continue to be, active in Alaska for long periods of time and in 

the present. These transport mechanisms include: 

• Ballast water from other types of ships, especially bulk carriers of such products as wood 

(logs, wood chips), ore, and coal, which come from foreign or domestic ports to Alaska 

in ballast to load. These may provide inoculations in ports nearby Prince William Sound 

(e.g., Homer, Seward), that may be spread by coast-wise traffic. 

• Fouling of ship hulls, which was especially important historically in wooden ships and 

before anti-fouling paints. However, fouling continues to be common, and may be 

especially important in coast-wise transport to and within Alaskan waters. This vector 

may include all types of private, recreational and commercial vessels. 

• Intentional and accidental release from aquaculture activities. Both species that are 

cultured and species that may be coincident with the aquaculture (including disease 

organisms) may be released. Oyster culture, salmon culture, and cultured herring roe on 

kelp are especially common and potential sources of NIS in Prince William Sound. 

Mussel culture may be initiated in the area. 

• Fishery release has occurred commonly in the past through efforts of state and federal 

agencies to improve stocks. 

• Aquarium and pet trades have resulted in NIS invasions at many places around the world. 

• Large-scale changes in current patterns may transport NIS into Alaskan waters. During 

1998 the very strong El Niño along the eastern Pacific may have brought warm-water 

species much further north than their typical distribution. These shifts may also allow 

species transported by human activities to become established. 

In many ecosystems, NIS invade over time as a series of vectors shift in importance, and this 

accumulation of NIS can result in major changes in the diversity and function of coastal 

ecosystems (Cohen & Carlton 1995). 

1D. References 

Carlton, J.T. 1979. History, biogeography, and ecology of the introduced marine and estuarine 

invertebrates of the Pacific coast of North America. Ph.D. Thesis, Univ. Calif., Davis. 904 pp. 

Carlton, J.T. 1989. Man's role in changing the face of the ocean: biological invasions and the 

implications for conservation of near-shore environments. Conserv. Biol. 3(3): 265-273.


Carlton, J.T. and J. B. Geller. 1993. Ecological roulette: the global transport of nonindigenous 

marine organisms. Science 261: 78-82. 

Carlton, J.T., D. Reid and H. van Leeuwen. 1995. The role of shipping in the introduction of 

nonindigenous aquatic organisms to the coastal waters of the United States (other than the Great 

Lakes) and an analysis of control options. Report to U.S. Coast Guard, Marine Environment 

Protection Division, Washington, DC. 215 pp. 

Cohen, A.N. and J.T. Carlton. 1995. Nonindigenous aquatic species in a United States estuary: a 

case study of the biological invasion of San Francisco Bay and delta. Report to U.S. Fish & 

Wildlife Service, Washington, DC and National Sea Grant College Program, Connecticut Sea 

Grant. 246 pp. 

Coles, S. L., R.C. DeFelice, L.G. Eldredge and J.T. Carlton. 1999. Historical and recent 

introductions of non-indigenous marine species into Pearl Harbor, Oahu, Hawaiian Islands. Mar. 

Biol. 135(1): 147-158. 

Hallegraeff, G.M. and C.J. Bolch. 1992. Transport of diatom and dinoflagellate resting spores in 

ships’ ballast water: implications for plankton biogeography and aquaculture. J. Plankton Res. 

14: 1067-1084. 

Hayward, B.W. 1997. Introduced marine organisms in New Zealand and their impact in the 

Waitemata Harbour, Auckland. Tane 36:197-223. 

Hines, A.H., G.M. Ruiz, J. Chapman, J. Carlton and N. Foster. 1998. Biological invasions in 

cold-water coastal ecosystems: Ballast-mediated introductions in Port Valdez/Prince William 

Sound, Alaska. Progress Report to the Regional Citizens' Advisory Council of Prince William 

Sound. 

Leppakoski, E. 1984. Introduced species in the Baltic Sea and its coastal ecosystems. Ophelia, 

suppl 3: 123-135. 

Mills, E.L., J.H. Leach, J.T. Carlton and C.L. Secor. 1993. Exotic species in the Great Lakes: A 

history of biotic crises and anthropogenic introductions. J. Gt. Lakes Res. 19(1): 1-54. 

Reise, K., S. Gollasch and W.J. Wolff. 1999. Introduced marine species of the North Sea coasts. 

Helgol. Meeresunters. 52: 219-234. 

Ruiz, G.M., J.T. Carlton, A. H. Hines and E.D. Grosholz. 1997(a). Global invasions of marine 

and estuarine habitats by non-indigenous species: Mechanisms, extent, and consequences. Am. 

Zool. 37: 621-632 

Ruiz, G.M. and A.H. Hines. 1997. Patterns of nonindigenous species transfer and invasion in 

Prince William Sound, Alaska: Pilot Study. Report Submitted to the Prince William Sound 

Citizens’ Advisory Council. 80pp.


Ruiz, G.M., P. Fofonoff and A.H. Hines. 1999. Non-indigenous species as stressors in estuarine 

and marine communities: Assessing invasion impacts and interactions. Limnol. Oceanogr. 44(3, 

part 2): 950-972 

Smith, L.D., M.J. Wonham, L.D. MCann, D.M. Reid, G.M. Ruiz and J.T. Carlton. 1996. 

Biological invasions by nonindigenous species in United States waters: Quantifying the role of 

ballast water and sediments. Parts I and II. Final Report to the U.S. Coast Guard and the U.S. 

Department of Transportation. 246 pp. 

Smith, L.D., M.J. Wonham, L.D. McCann, G.M. Ruiz, A.H. Hines and J.T. Carlton. 1999. 

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

Invasions 1: 67-87. 

Williams, R.J., F.B. Griffiths, E.J. Van der Wal and J. Kelly. 1988. Cargo vessel ballast water as 

a vector for the transport of non-indigenous marine species. Est. Coast. Shelf Sci. 26: 409-420.

Chapt 2. Ballast Water Delivery Patterns, page 2- 1 

Chapter 2. Ballast Water Delivery Patterns 



George Smith, Smithsonian Environmental Research Center 

Melissa A. Frey, Smithsonian Environmental Research Center 

2A. Purpose 

The primary goal of this portion of the study was to characterize the traffic patterns and 

volumes of ballast water discharged into Port Valdez and Prince William Sound (PWS) by oil 

tankers. Since ballast water is a major mechanism for the transfer of NIS, we wished to describe 

the delivery patterns of the ballast water to the region by season, source region, voyage duration. 

Analysis of the biota associated with the tankers’ ballast water is discussed in the next chapter. 

2B. Methods 

We obtained data on ship arrivals and ballast water histories in two ways. First, we 

obtained information about the long-term (10-year) pattern of arrivals to Port Valdez from 

Alyeska and RCAC. Second, to characterize current patterns, we collected detailed data from 

vessels arriving to Port Valdez over the one-year period of 1998. Our goal in this latter approach 

was to collect comprehensive information on the origin (i.e., last port of call), date of arrival, and 

ballast water histories for as many arriving vessels as possible. Most of these data were collected 

by SERC staff, during interviews aboard vessels (see below). Additional data were sent to us by 

the ships’ personnel and shipping agents. 

Beginning December 1997, we implemented a sampling scheme to estimate the amount 

of segregated ballast water delivered to Prince William Sound and Port Valdez by source port 

and season. For tankers arriving to Port Valdez from each of the three primary domestic source 

port systems (Los Angeles, San Francisco Bay, Puget Sound), we boarded approximately 3 

tankers per month (i.e., 10 per quarter x 3 source ports = 30 per quarter). In addition, we 

attempted to board most tankers arriving to Port Valdez from foreign ports. 

Upon boarding, we conducted an interview of the ships’ personnel to collect information 

on the quantity, age, source region, and management of all ballast water. Following the 

interview, we proceeded to sample the segregated ballast water to characterize temperature, 

salinity, and resident biota (see Chapter 3). 

We excluded non-segregated ballast water from most of our current analyses. Although 

this can account for roughly 50% of the total ballast water aboard tankers arriving to Prince 

William Sound (see below), previous analyses indicated that very few viable organisms were 

present in this ballast water, which often includes some residual oil. Furthermore, the 

nonsegregated ballast water is pumped to an on-shore treatment facility at the Alyeska terminal. 

For review of previous results, as well as description of the treatment process, see Ruiz and 

Hines 1997. 

Since vessels with double bottoms are difficult to sample for biota, we focused our sampling 

effort primarily on vessels without double bottoms. Thus, to characterize the entire fleet (i.e., all


arrivals), we obtained additional data on ballast water histories of nearly all oil tankers arriving 

to Port Valdez in 1998. Ships’ personnel and shipping agents generously provided these data. 

2C. Results 

2C1. Number and Source of Tanker Arrivals to PWS 

Over the past decade (1989-98), tanker arrivals to Port Valdez have averaged 713 

(se=37.2) ships per year, ranging from 870 to 549 (Fig. 2.1). There has been a noticeable decline 

in arrivals since 1991, with each year having fewer arrivals than the previous one. 

Figure 2.1. Annual number of oil tankers arriving to Port Valdez, 1989-1998. Data as provided by Alyeska. 

1000 

NUMBER OF ARRIVALS 

800 

600 

400 

200 

0 

89 90 91 92 93 94 95 96 97 98 

YEAR 

Using 1998 to examine spatial and temporal patterns, tanker arrivals to Port Valdez were 

both distributed evenly among seasons and dominated by arrivals from U.S. domestic ports (Fig. 

2.2). An average of 137.3 (s.e.=2.98) vessels arrived each quarter, and 95.8% (s.e. = 0.82 %) of 

all arrivals came directly from a U.S. port. Of all tanker arrivals, 82.7% came from one of three 

domestic ports (Fig. 2.3): Puget Sound, Washington (43.0%); San Francisco Bay, California 

(28.8.%); and Long Beach, California (10.9%). The residual came from Oregon, Hawaii, 

Alaska, or foreign ports. Among arrivals from foreign ports, most (69.6%) came directly from 

Korea (Fig. 2.4). 

Figure 2.2. Number of oil tankers arriving to Port Valdez from foreign and domestic source ports by season 

in 1998. Seasons include: Winter (January-March), Spring (April-June), Summer (July-September), and Fall 

(October-December). Data based upon boarding interviews and reports from ships’ personnel (see text). 

160 

SOURCE PORT 

Foreign 

Domestic 

# VESSELS 

120 

80 

40 

0 

WINTER SPRING SUMMER FALL 

SEASON


Figure 2.3 Number of oil tankers arriving to Port Valdez from each domestic source port by season in 1998. 

Source ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); Columbia River, 

Oregon (OR); Cook Inlet, Alaska (AK); and Barbers Point, Hawaii (HI). Seasons include: Winter (January-March), 

Spring (April-June), Summer (July-September), and Fall (October-December). Data based upon boarding 

interviews and reports from ships’ personnel (see text). 

SOURCE PORT 

# VESSELS 

80 

60 

40 

20 

PS 

SF 

LB 

OR 

AK 

HI 

0 


SEASON 

Figure 2.4. Number of oil tankers arriving to Port Valdez from each foreign source port by season in 1998. 

Seasons Include: Winter (January-March), Spring (April-June), Summer (July-September), and Fall (October- 

December). Data based upon boarding interviews and reports from ships’ personnel (see text). 

# VESSELS 

6 

4 

2 

SOURCE PORT 

Korea 

China 

Taiwan 

Japan 

Singapore 

0 


SEASON 

2C2. Volume of Ballast Water delivered to PWS 

During 1998, oil tankers arriving to PWS carried an estimated average of 65,775m 3 

(s.e.=1,252; n=472) of total ballast water, the combination of segregated and nonsegregated 

ballast water. Segregated ballast water comprised an average of 54.7% (s.e.=2.1%; n=472) of 

the total among tankers. 

Across all vessels, tankers discharged an average of 32,715 m 3 (s.e.= 645; n=472) of 

segregated ballast water upon arrival to PWS (Table 2.1). Although there were no seasonal 

differences in the average amount of ballast water per tanker, there was a significant difference 

by source port (Fig. 2.5; 2-way ANOVA, F (3 (seasons), 5 (port source), 517 obs) =3.52, P = 0.004). 

Specifically, the mean volume was significantly greater for arrivals from foreign ports compared 

to arrivals from all other ports, and the mean volume was significantly lower for tankers from 

Hawaii relative to all other sources.


Table 2.1. Estimated volume of ballast water delivered by oil tankers to Port Valdez and PWS in 1998. 

Shown by source port and season are: (1) the estimated mean volume of ballast water arriving per tanker, including 

the standard error and sample size (n=number of vessels for which we have volume estimates), (2) the total number 

of tanker arrivals which is shown as N, (3) the total estimated volume of ballast water (calculated as mean volume X 

total arrivals). The bottom row (Overall) estimates the total ballast water volume and number of arrivals for all 

ships combined. Source ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); 

Foreign port with open-ocean exchange (EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI). 

Seasons include: Winter (January-March), Spring (April-May), Summer (July-September), and Fall (October- 

December). Source of data on volumes and arrivals as described in text. 

BW vol. (m³) 

Port/Source Season Mean(se) n N Total 

Winter 28163(1734) 56 64 1,802,432 

Spring 31421(2095) 51 60 1,885,260 

PS Summer 34448(1901) 53 59 2,032,432 

Fall 33894(2199) 46 53 1,796,382 

Grand total 206 236 7,516,506 

Winter 31841(2798) 31 34 1,082,594 

Spring 32809(2984) 40 41 1,345,169 

SF Summer 35765(2361) 40 44 1,573,660 

Fall 34371(2372) 37 39 1,340,469 

Grand total 148 158 5,341,892 

Winter 32526(3237) 18 18 585,468 

Spring 30399(2902) 12 13 395,187 

LB Summer 36045(3633) 14 14 504,630 

Fall 28850(1346) 15 15 390,180 

Grand total 59 60 1,875,465 

Winter 42153(4093) 9 9 379,377 

Spring 44294(3775) 6 6 265,764 

EX Summer 29856(3689) 5 5 149,280 

Fall 47056(5199) 3 3 141,168 

Grand total 23 23 935,589 

Winter 29182(6929) 4 9 262,638 

Spring 28250(2169) 4 4 113,000 

OR Summer 32429(1631) 2 3 97,287 

Fall 39138(5740) 6 7 273,966 

Grand total 16 23 746,891 

Winter 27142(2279) 6 7 189,994 

Spring 22229(1651) 4 5 111,145 

HI Summer 23766(4197) 6 6 142,596 

Fall 21729(1544) 4 5 108,370 

Grand Total 20 23 552,105 

Overall 32,715 472 523 16,968,448 

Note: Not included in the table are arrivals from Nikiski, AK (19) which provided no data, and other arrivals (7) for 

which port data are unavailable.


Figure 2.5. Mean volume of segregated ballast water per tanker arriving to Port Valdez and PWS by source 

port and season, 1998. The mean volumes are estimated for: (A) Each source port across all seasons; (B) Each 

season across all source ports. Standard error and sample size is shown above each bar; see Table 2.1 for further 

information. Source ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); 

Foreign port with open-ocean exchange (EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI). 

Seasons include: Winter (January-March), Spring (April-June), Summer (July-September), and Fall (October- 

December). Data based upon boarding interviews and reports from ships’ personnel (see text). 

23 

40 

206 

148 

59 

16 

30 

20 

CUBIC METERS (thousands) 

20 

10 

0 

40 

30 

PS SF LB EX OR HI 

PORT 

119 113 

117 

123 

20 

10 

0 


SEASON 

We estimated the total amount of segregated ballast water discharged into Prince William 

Sound during 1998 was approximately 17,000,000 m 3 (no. arrivals x average ballast water 

volume) for source port by season (see Table 2.1). The relative contribution of different source 

ports to the total varied greatly, reflecting variation in the number of arrivals (Fig. 2.6; Table 

2.1). As a result, Puget Sound contributed approximately 44% of the total, followed by San 

Francisco 31% and Long Beach 11%.


Figure 2.6. Cumulative volume of ballast water arriving to Port Valdez and PWS by source port and season, 

1998. The cumulative volumes are estimated for season by source port (see Table 2.1 for further detail). Source 

ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); Foreign port with openocean 

exchange (EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI). Seasons include: Winter 

(January-March), Spring (April-June), Summer (July-September), and Fall (October-December). Data based upon 

boarding interviews and reports from ships’ personnel (see text). 

CUBIC METERS (millions) 

2.0 

1.5 

1.0 

0.5 

0.0 


SEASON 

SOURCE PORT 

PS 

SF 

LB 

EX 

OR 

HI 

2C3. Age and Management of Ballast Water delivered to PWS 

The average age of ballast water arriving in tankers varied among source ports, ranging 

between 4.8 to 10.2 days (Fig. 2.7). The mean age among all arrivals was 6.6 days (s.e.= 0.2). 

For domestic source ports, the age of water was correlated with distance from Port Valdez to the 

source port, as ballast water came directly from the last port of call (the exception was for 

experiments conducted at our request, as described in Chapter 4). In contrast, all foreign arrivals 

exchanged their ballast water at sea, so the age of water was less than the voyage duration. Thus, 

for foreign arrivals, the actual source was considered open ocean exchange (EX) instead of the 

last port of call. 

Figure 2.7. Mean age (voyage duration) for ballast water arriving to Port Valdez by source port. The mean 

age is estimated for each source port based upon all boarding data (December 1997-July 1999). Standard error and 

sample size is shown above each bar. Source ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF); 

Long Beach, CA (LB); Foreign port with open-ocean exchange (EX); Columbia River, Oregon (OR); and Barbers 

Point, Hawaii (HI). 

BALLAST WATER AGE (Days) 

12 

8 

4 

0 

48 

50 

46 


SOURCE PORT 

2D. Discussion 

Over the past decade, Prince William Sound and Port Valdez together have received 

approximately 23.3 million m 3 (= 713 arrivals/yr x 32,610 m 3 /arrival) of segregated ballast water 

19 

2 

4


each year from oil tankers arriving to the Alyeska terminal. Although most of this water is 

released in Port Valdez, ships will sometimes begin discharging upon entering Prince William 

Sound en route for the Port. Thus, organisms released with this ballast water may experience a 

broader range of conditions outside of Port Valdez than we had originally considered. 

The total volume of ballast water delivered to Prince William Sound by tankers greatly 

exceeds the estimated quantity of ballast water arriving to other western U.S. ports (Carlton et al. 

1995). However, it is important to recognize that existing estimates for the other ports have 

included only the ballast water from foreign sources. In contrast, our estimates for Prince 

William Sound included both foreign and domestic sources, but were dominated by the latter. 

The amount of domestic ballast water released in other U.S. ports is only now being estimated. 

Nonetheless, even when we can include data on domestic sources for all ports, it appears likely 

that the total volume of ballast water released to PWS will still exceed that for the other western 

ports, due to the absence of extensive domestic tanker and bulker traffic (i.e., those vessels that 

discharge the greatest quantities of ballast water) at the other ports. Instead, the domestic traffic 

for other western U.S. ports is dominated by container ships, which release relatively small 

amounts of ballast water (Carlton et al. 1995; National Ballast Water Information Clearinghouse, 

unpubl. data). 

The total amount of ballast water arriving to Prince William Sound in tankers is also 

relatively large on a global scale. Within the U.S., PWS is third only to Chesapeake Bay and 

New Orleans in estimated ballast water discharge for 1991 (Carlton et al. 1995, Smith et al. 

1999). In a similar estimate of ballast water discharged to 46 Australian ports in 1991, only that 

for the port of Dampier exceeded the volume for PWS (Kerr 1994). As above, estimates for the 

both the U.S. and Australian ports were restricted to arrivals from foreign ports. Although these 

totals would clearly increase when including domestic arrivals, the overall patterns provide a 

useful context, suggesting PWS is on the extreme end of the spectrum for amount of ballast 

water discharge. 

It is also important to recognize the present level of tanker activity, and the magnitude of 

ballast water delivery, as a recent development in Port Valdez. The terminal began transporting 

oil via tankers in 1977. Based upon the arrivals rate and discharge volumes observed in this 

decade, we estimate over 700 million m 3 of segregated ballast water have been delivered over the 

past 3 decades of operation. This large cumulative volume underscores the potential importance 

of ballast water as a vector for the transfer of species. Unlike many other commercial ports, 

however, the volume of ballast water prior to oil exportation was virtually absent, as very few 

other vessels currently deliver ballast water to the Port (Ruiz et al., unpubl. data). 

Furthermore, the foreign export of oil from Port Valdez has only occurred since 1996, 

following authorization by U.S. Congress. Prior to this time, all oil export was only to domestic 

U.S. ports, which were therefore the source of ballast water delivery to PWS and Port Valdez. 

Although tankers now export oil to foreign ports from PWS, the delivery of ballast water from 

foreign traffic remains a small fraction (


2E. References 

Carlton, J.T., D. Reid and H. van Leeuwen. 1995. The role of shipping in the introduction of 

nonindigenous aquatic organisms to the coastal waters of the United States (other than the Great 

Lakes) and an analysis of control options. Report to U.S. Coast Guard, Marine Environment 

Protection Division, Washington, DC. 215 pp. 

Kerr, S. 1994. Ballast water ports and shipping study. Australian Quarantine and Inspection 

Service, Report No. 5, Canberra. 



Citizens’ Advisory Council. 80pp. 







Invasions 1: 67-87.

Chapt 3. Biological Characteristics of Ballast Water, page 3- 1 

Chapter 3. Biological Characteristics of Ballast Water in Oil Tankers 




Melissa A. Frey, Smithsonian Environmental Research Center 

3A. Purpose 

The overall goal of this research component was to characterize the biota associated with 

segregated ballast water of tankers arriving to Port Valdez. For this analysis, we designed a 

sampling program to measure temporal (seasonal, annual) and spatial (source port) variation in 

the biota associated with the ballast water. 

We focused primarily on the mid-large (>80 micron) zooplankton resident in the water 

column of ballast tanks, and present the results of this analysis here. We have included some 

information on the phytoplankton concentrations present in our samples, but the sampling 

methods (below) were not designed to characterize these organisms and many of the other taxa 

(e.g., bacteria, viruses, and other microorganisms) that are small in size. This choice does not 

imply that small organisms are not significant from an invasion standpoint, as the potential 

effects of toxic dinoflagellate blooms are very evident (see Introduction). Instead, we simply did 

not have the resources to include all taxonomic groups in our analyses. 

We chose to focus on the mid-large zooplankton for multiple reasons. First, the 

taxonomic resolution is relatively good compared to many of the other (smaller) organisms. 

Second, most known NIS that are established at the source ports of oil tankers, such as San 

Francisco Bay, occur (for some portion of their life history) in this zooplankton community. 

Third, we could readily gain access to the plankton community (as opposed to the bottom 

sediments). Finally, the sample analysis for larger zooplankton is not as technically difficult or 

time consuming as that necessary for the smaller organisms, allowing us to analyze samples from 

a large number of ships for statistical comparisons. 

The analysis of zooplankton presented here is one of the most comprehensive and 

quantitative studies of ballast water in the world. Additional data on the biota associated with 

ballast water also appear in other chapters. Our analysis of ballast water exchange (Chapter 5) 

includes survivorship of plankton during 8 separate voyages, effects of exchange on 

zooplankton, and some information on bacteria and ciliate protozoans. Biota associated with the 

bottom sediments of ballast tanks, as well as the hulls and seachests, are also examined (Chapters 

6 and 7, respectively). 

3B. Methods 

3B1. Source and Number of Sampled Ships 

For a 13-month period (December 1997 – December 1998, hereafter 1998), we conducted 

an intensive sampling program of ballast water on tankers that was stratified by source port and 

season. Most tankers and ballast water arriving to Port Valdez came from three U.S. domestic


port systems: Puget Sound, San Francisco Bay, and Long Beach (see Chapter 2). To characterize 

biota for these ports by seasons, we sampled a minimum of 3 tanker arrivals per month from 

each of the three domestic source port systems (i.e., 10 per quarter x 3 source ports = 30 per 

quarter). Although relatively few (23) tankers arrived from foreign ports in Port Valdez during 

this year, we sampled as many as possible (n=19) to compare the biota arriving from foreign 

versus domestic sources. We also sampled a limited number of arrivals from the other two 

domestic ports: Oregon and Hawaii (which comprised < 10% all arrivals; see Chapter 2). 

In June of three consecutive years (1997, 1998, 1999), we collected samples from 

approximately 10 tankers arriving to Port Valdez from the domestic ports of Puget Sound, San 

Francisco Bay, and Long Beach. Samples from 1997 were collected as part of a Pilot Study that 

we conducted for RCAC (Ruiz and Hines 1997), and the samples for 1998-1999 were collected 

in the present study. Together, these samples were used to characterize annual variation in the 

ballast water biota arriving to Alaska. 

3B2. Sample Collection and Analysis 

We boarded and sampled tankers immediately upon their arrival to Port Valdez. As 

described above, we sampled approximately 2-3 vessels per week over the 13-month period. 

Although we attempted to collect ballast water from every tanker boarded, vessels with double 

bottoms could not be sampled easily without disruption of ship operations and modification of 

our standard sampling protocol. In the present analysis, we therefore have included primarily 

(but not exclusively) vessels without double bottoms. 

We applied our established methods for qualitative and quantitative analysis of biota 

transported in ballast water, which evolved from methods developed by J.T. Carlton (e.g., 

Carlton and Geller, 1993; LaVoie et al.1999; Smith et al., 1999). Our protocol consisted of 

collecting the following information and samples: 

• Ship and ballast management information: Last port of call, number of tanks by type, 

capacity 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; 

• Physical variables of ballast water: Water temperature and salinity were measured (surface 

and 10m depth) for each tank sampled (as below), collecting ballast water with a Niskin 

bottle through the Butterworth hatches; oxygen (O 2 ) concentration was not measured 

because previous extensive analysis of ballast water tanks in other cargo ships indicated that 

O 2 concentrations rarely varied and were not appreciably lower than saturation (Smith et al., 

1996). 

• Biological samples of ballast water: Plankton samples were collected by towing a standard 

plankton net (80 micron mesh, 30 cm diameter) vertically through the entire height of the 

water column in each ballast tank; access to ballast tanks was obtained through the 

Butterworth hatches. A single tank was 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 10 cm. 

• Additional observations and opportunistic samples: Upon initiating sampling of ballast 

tanks, we routinely examined the surface waters to look for large, mobile biota (e.g., fish) 

and organisms attached to the sides of tanks; we often took opportunities to collect any such


organisms observed, as well as bottom sediments, since these are usually missed in our 

plankton tows. 

• Physical variables of port water: Shipside water temperature and salinity were measured 

(surface and 10m 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 at Valdez 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), to provide a 

qualitative assessment of plankton viability. 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, 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). 

We used two different methods to characterize the plankton samples, as follows: 

• Coarse Analysis. All samples were characterized by Coarse Analysis, consisting of a direct 

count of individuals according to general taxonomic groups, usually phyla (e.g., molluscs, 

crustaceans, echinoderms). The minimum of number of distinctly different taxa were also 

estimated in Coarse Analysis of each sample. 

• Fine Analysis. For a subset of samples (roughly 1/3 of the ships from domestic ports), Fine 

Analysis was used to enumerate all morphologically distinct taxa at the lowest taxonomic 

level possible. 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 of larvae 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. 

The two methods were selected to provide different types of information. The Coarse 

Analysis allowed us to test for patterns in the biota across all ships, increasing the statistical 

power of the analysis. Since many species were not present on each ship, such an approach was 

not feasible with finer taxonomic resolution. In contrast, the Fine Analysis allowed us to 

quantify the densities of particular taxa, usually crustacean groups with adult forms (see results), 

and test for the presence of nonindigenous species known from the source ports. These data also 

allow us to characterize the frequency and density of particular nonindigenous species in the 

ballast water. 

For both analyses, 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 desired taxonomic level 

(as above). For abundant taxa (> 100 individuals/sample), samples were split using a Folsom 

plankton splitter to achieve counts between 10-100 individuals per subsample (usually splits of 

1/8 to 1/32). For organisms in split samples, two subsamples were counted.


Taxonomic identification of plankton 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 (in Fine Analysis) for 

taxonomic verification and, wherever possible, species-level identification. These voucher 

specimens were sent to taxonomists at the Smithsonian Institution’s National Museum of Natural 

History and elsewhere for verification and identification. 

3B3 Data Analysis 

Throughout our analyses, we use “ship” as the level of replication within a class variable 

(e.g., source port, season, year), because multiple samples from the same ship are not 

independent of each other. Although our replicate plankton samples 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. Thus, we estimated density per ship as the mean of 

replicate tows. 

We derive most of the results reported from the Coarse Analysis. All enumeration is also 

completed for the Fine Analysis, but identification of only a portion of the voucher specimens 

has been finished to date. Although we cannot yet discuss the frequency and density of 

individual taxa (as described above), we confirm the presence of numerous nonindigenous 

species in the Fine Analysis, and these are reported here. We will include full results of the Fine 

Analysis, when completed, in a future publication and provide a copy to RCAC. 

Virtually all organisms collected in the ballast water samples were alive and appeared to 

be in good condition. Indeed, many of the organisms collected from these samples performed 

well in laboratory culture and experiments, as described in Chapter 4. Thus, we considered all 

organisms counted in fixed samples to be alive at the time of sampling. 

In most of the analyses, we have excluded the chain-forming diatoms. Although we 

enumerated these organisms to the full extent possible, quantitative counts are particularly 

problematic and unreliable, because the chains break apart during sample collection and 

processing. We have therefore included information on their prevalence and the counts made but 

excluded these data from most estimates of organism densities. 

Finally, the presentation of water temperature and salinity, for both the ballast water of 

oil tankers and Port Valdez, are presented in Chapter 4. 

3C Results 

Source and Number of Sampled Ships 

During this study, we sampled the ballast water of 169 tankers arriving to Port Valdez 

(Table 3.1). Our samples included 8-15 arrivals per quarter for each of the three major domestic


ports (Puget Sound, San Francisco Bay, and Long Beach) and 3-7 arrivals per quarter from 

foreign ports (primarily Korea; see Chapter 2). 

Table 3.1. Prevalence and densities of taxa in the ballast water arriving to Port Valdez for each source port. 

Shown for each taxon and source port are the prevalence, density among all ships, and density for ships only when 

taxon was present. Standard errors are shown in parentheses with each density measure. Source ports include: Puget 

Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); Foreign port with open-ocean exchange 

(EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI). The data include all sample dates. Sample 

sizes for each source port as follows: PS (n=48), SF (n=50), LB (n=46), EX (n=19), OR (n=2), HI (n=4). 

(%) (density/m³) 

Phylum Taxa source n Prevalence mean(se) all ships mean(se) when present 

DINOFLAGELLATA 

PS 48 88 775(223) 902(245) 

SF 50 54 66(21) 114(34) 

LB 46 65 71(30) 107(45) 

EX 19 84 729(658) 886(780) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 100 3.0(1.0) 3.0(1.0) 

DIATOMACEA 

PROTOZOA 

CNIDARIA 

CTENOPHORA 

PS 48 100 13866(3270) 13866(3270) 

SF 50 100 11683(4384) 11683(4384) 

LB 46 100 1170(241) 1170(241) 

EX 19 100 5346(2184) 5346(2184) 

OR 2 100 9409(6120) 9409(6120) 

HI 4 100 277(107) 277(107) 

PS 48 83 316(122) 361(139) 

SF 50 90 5506(3638) 6120(4037) 

LB 46 93 210(57) 220(59) 

EX 19 74 82(58) 110(78) 

OR 2 100 31(13) 31(13) 

HI 4 100 4.0(1.3) 4.0(1.3) 

PS 48 63 19.5(11.9) 55(32) 

SF 50 22 1.8(0.7) 8.3(2.3) 

LB 46 87 45(18) 53(21) 

EX 19 5 0.3(0.3) 5.7(0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

PS 48 17 0.2(0.1) 1.2(0.3) 

SF 50 4 0.01(0.008) 0.5(0.0) 

LB 46 37 1.6(0.6) 4.0(1.5) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0)


Table 3.1 continued 

Phylum Taxa source n Prevalence (%) mean(se) all ships mean(se) when present 

PLATYHELMINTHES 

PS 48 27 3.4(1.4) 13(4.3) 

SF 50 28 10.5(4.6) 37(14) 

LB 46 80 11(2.2) 13(2.4) 

EX 19 16 0.1(0.07) 0.9(0.1) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

NEMATODA 

ROTIFERA 

SIPUNCULA 

NEMERTEA 

PS 48 6 0.2(0.1) 2.5(0.9) 

SF 50 8 0.5(0.3) 6.5(2.4) 

LB 46 4 0.1(0.07) 2.5(0.5) 

EX 19 5 0.1(0.1) 1.4(0.0) 

OR 2 100 4.4(3.0) 4.4(3.0) 

HI 4 0 0(0.0) 0(0.0) 

PS 48 2 0.3(0.3) 12.5(0) 

SF 50 2 0.7(0.7) 33(4.5) 

LB 46 2 0.6(0.6) 27(0.0) 

EX 19 5 0.3(0.3) 5.6(0.0) 

OR 2 50 0.2(0.2) 0.4(0.0) 

HI 4 0 0(0.0) 0(0.0) 

PS 48 0 0(0.0) 0(0.0) 

SF 50 4 0.8(0.7) 19.8(11) 

LB 46 4 0.7(0.6) 15(4.1) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

PS 48 8 1.0(0.6) 12.4(5.3) 

SF 50 8 41(27) 516(275) 

LB 46 15 23(12) 150(66) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

ANNELIDA PS 48 90 370(136) 404(148) 

SF 50 80 199(55) 251(68) 

LB 46 100 33(6.5) 33(6.5) 

EX 19 2 2.1(1.3) 6.3(3.6 

OR 2 0 0(0.0) 0(0.0) 

HI 4 25 0.31(0.31) 1.2(0.0) 

MOLLUSCA 

Bivalvia PS 48 90 371(100) 396(106) 

SF 50 82 319(79) 389(93) 

LB 46 98 240(72) 246(74) 

EX 19 53 8.0(3.5) 15.2(5.5) 

OR 2 50 0.5(0.5) 1.0(0.0) 

HI 4 75 1.3(0.7) 2.0(0.5)




Gastropoda PS 48 71 139(39) 191(44) 

SF 50 54 34(18) 58(38) 

LB 46 91 35(7.6) 37(6.8) 

EX 19 37 26.6(17.6) 72(38) 

OR 2 50 0.2(0.2) 0.4(0.0) 

HI 4 0 0(0.0) 0(0.0) 

Other Mollusca PS 48 8 8.0(7.7) 105(83) 

SF 50 6 0.6(0.5) 10(7.6) 

LB 46 2 0.2(0.2) 9.0(0.0) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

ARTHROPODA/CRUSTACEA 

Amphipoda PS 48 29 1.9(1.3) 6.1(3.8) 

SF 50 30 0.61(0.16) 1.9(0.3) 

LB 46 48 0.7(0.2) 1.4(0.3) 

EX 19 19 0.07(0.05) 0.7(0.2) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

Anomura PS 48 29 0.91(0.64) 3.7(2.2) 

SF 50 24 0.2(0.08) 0.9(0.3) 

LB 46 63 3.8(1.4) 5.9(2.1) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

Brachyura PS 48 44 3.5(1.3) 9.2(2.8) 

SF 50 14 0.13(0.07) 1.1(0.5) 

LB 46 76 23.6(13.9) 30.2(18.6) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

Caridea PS 48 15 0.2(0.10) 1.3(1.2) 

SF 50 2 0.02(0.01) 0.2(0.0) 

LB 46 22 1.2(0.60) 3.8(1.5) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0)




Cirripedia PS 48 85 832(559) 951(637) 

SF 50 66 96(34) 108(47) 

LB 46 63 18(5.4) 37(7.2) 

EX 19 16 0.8(0.5) 6.9(1.7) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 50 3.1(1.9) 4.8(4.3) 

Cladocera PS 48 15 2.7(1.2) 21.7(5.1) 

SF 50 16 1.9(0.8) 11.7(3.7) 

LB 46 7 0.04(0.02) 0.6(0.1) 

EX 19 5 0.3(0.3) 5.7(0.0) 

OR 2 100 3.5(1.9) 3.5(1.9) 

HI 4 0 0(0.0) 0(0.0) 

Copepoda PS 48 100 2395(664) 2395(664) 

SF 50 100 9416(2060) 9416(2060) 

LB 46 100 5116(685) 5116(685) 

EX 19 100 2345(645) 2345(645) 

OR 2 100 43(6.0) 43(6.0) 

HI 4 100 8.2(4.4) 8.2(4.4) 

Cumacea PS 48 10 0.05(0.03) 0.6(0.2) 

SF 50 30 0.70(0.2) 2.1(0.5) 

LB 46 22 0.13(0.05) 0.6(0.2) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

Decapoda (misc.) PS 48 4 0.05(0.04) 1.3(0.7) 

SF 50 8 0.02(0.02) 0.5(0.2) 

LB 46 4 0.05(0.02) 2.5(0.0) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

Isopoda PS 48 25 0.70(0.40) 2.1(1.3) 

SF 50 18 0.34(0.26) 2.1(1.5) 

LB 46 9 0.06(0.03) 0.7(0.1) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

Mysidacea PS 48 2 0.007(0.007) 0.3(0) 

SF 50 54 2.48(0.6) 4.6(0.9) 

LB 46 78 3.52(0.8) 4.4(0.9) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 25 0.15(0.15) 0.6(0.1)




Ostracoda PS 48 25 0.35(0.16) 1.5(0.6) 

SF 50 8 0.40(0.20) 3.9(2.0) 

LB 46 17 0.28(0.14) 1.6(0.7) 

EX 19 16 0.93(0.54) 5.9(1.4) 

OR 2 50 0.6(0.6) 1.2(0.0) 

HI 4 0 0(0.0) 0(0.0) 

Tanaidacea PS 48 2 0.006(0.006) 0.3(0) 

SF 50 0 0(0.0) 0(0.0) 

LB 46 2 0.006(0.006) 0.3(0) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

BRYOZOA 

PHORONIDA 

CHAETOGNATHA 

ECHINODERMATA 

PS 48 35 16.1(5.4) 40.7(11.5) 

SF 50 40 59(33) 147(80) 

LB 46 22 1.5(0.5) 6.7(1.7) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

PS 48 2 0.01(0.01) 0.5(0.0) 

SF 50 2 0.01(0.01) 0.7(0.0) 

LB 46 15 0.5(0.4) 4.2(2.7) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

PS 48 33 0.7(0.4) 2.3(1.1) 

SF 50 12 0.5(0.3) 4.1(2.5) 

LB 46 72 5.8(2.2) 7.9(3.0) 

EX 19 21 0.2(0.1) 1.5(0.3) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

PS 48 33 32(15) 94(43) 

SF 50 12 6.3(4.8) 53(37) 

LB 46 26 5.8(3.5) 22(12) 

EX 19 5 0.3(0.3) 6.0(0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

CHORDATA 

Cephalochordata PS 48 0 0(0.0) 0(0.0) 

SF 50 0 0(0.0) 0(0.0) 

LB 46 9 0.1(0.01) 2.8(2.5) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0)




Fish PS 48 10 0.05(0.02) 0.3(0.1 

SF 50 6 0.02(0.01) 0.5(0.1 

LB 46 7 0.04(0.02) 0.5(0.1) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

Other Chordata PS 48 21 4.9(2.2) 23.8(8.4) 

(incl. larvacea) SF 50 8 5.2(3.5) 85(41) 

LB 46 63 59.1(30.6) 107(55) 

EX 19 5 0.06(0.06) 1.2(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

OTHER 

Eggs PS 48 73 87(39) 113(50) 

SF 50 66 39(16) 56(22) 

LB 46 61 141(41) 232(148) 

EX 19 21 6.3(4.1) 15(9.2) 

OR 2 100 12(11) 12(11) 

HI 4 20 7.3(2.4) 7.3(2.4) 

Trochophore 

PS 48 33 20(8.2) 65(23) 

SF 50 16 16(10) 115(65) 

LB 46 30 11(4.6) 36(13) 

EX 19 0 0(0.0) 0(0.0) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

Unidentified larvae PS 48 0 0(0.0) 0(0.0) 

SF 50 4 95(93) 2376(1388) 

LB 46 9 0.02(0.02) 0.4(0) 

EX 19 11 27(26) 253(178) 

OR 2 0 0(0.0) 0(0.0) 

HI 4 0 0(0.0) 0(0.0) 

For June of 1997-1999, we sampled the ballast water of 31 tankers arriving to Port 

Valdez from the domestic ports of Puget Sound (n=14), San Francisco Bay (n=9), and Long 

Beach (n=9). The data for 1998 and 1999 are included in the total 169 vessels sampled during 

this study, and the data for 1997 are derived from our previous work (as above). 

Figure 3.1. Number of oil tankers sampled upon arrival to Port Valdez. Shown are the number of ships from 

which ballast water samples were collected by source port (i.e., last port of call) and season. Source ports include: 

Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); Foreign port with open-ocean 

exchange (EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI). Seasons include: Winter (January- 

March), Spring (April-June), Summer (July-September), and Fall (October-December). Data were collected 

primarily from December 1997 – December 1998, and some additional data were collected in May-June 1999. 

# VESSELS SAMPLED 

15 

10 

5 

0 

PORT SOURCE 

PS 

SF 

LB 

EX 

OR 

HI 


SEASON


3C2. Abundance of Organisms in Ballast Water 

(a) Total Density by Source 

We measured an average of 12,637 (s.e. = 5,533) total organisms per m 3 in the ballast 

water for all 169 vessels from domestic and foreign source ports. This estimate excludes the 

chain-forming diatoms, which were detected in the samples from 16 domestic tankers. Although 

the chain-forming diatoms are difficult to quantify (as above), we estimated average densities in 

excess of one million organisms/m 3 , approximately 100 fold higher than the densities of solitary 

diatoms and all other taxa measured for either domestic or foreign tankers (Fig. 3.2). We have 

excluded the chain-forming diatoms from the subsequent analyses and discussion of total density 

or abundance (i.e., abundance of all organisms) below. 

Figure 3.2. Densities of organisms in ballast water arriving to Port Valdez from foreign and domestic source 

ports. The estimated mean densities (#/m³ and standard errors) are shown separately for chain-forming diatoms, 

solitary diatoms, and all other taxa in ballast water of ships arriving from each domestic ports and foreign ports. 

Chain-forming diatoms were only detected on domestic ships (n=16), whereas the other groups were present on all 

domestic (n=150) and foreign (n=19) arrivals that were sampled. The data include all sample dates. Since arrivals 

from foreign ports all underwent ballast water exchange in open ocean, the source is indicated as exchange. 

DENSITY (#m³) 

1.E+07 16 

1.E+06 10 6 

1.E+05 

10 4 

1.E+04 

1.E+03 

10 2 

1.E+02 

150 

150 

CHAIN DIATOMS 

SOLITARY DIATOMS 

OTHER TAXA 

19 

19 

1.E+01 

0 

1.E+00 10 

DOMESTIC EXCHANGE 

SOURCE 

The average total density was significantly greater in ballast water from domestic sources 

compared to that for foreign sources (Fig. 3.2; 1-way ANOVA, F (1,168) = 3.63, P = 0.048). The 

chain-forming diatoms were only evident in the ballast water of domestic arrivals, increasing the 

magnitude of density differences between foreign and domestic traffic. 

The total abundance of organisms in ballast water differed among domestic sources. Of 

the three major domestic ports, arrivals from Puget Sound and San Francisco Bay had 

significantly greater densities than those from Long Beach and foreign sources, whereas the 

latter two were not different (Fig. 3.3; ANOVA, F (2,143) = 3.71, P = 0.027). We excluded both 

Oregon and Hawaii from this comparison, due to the limited sample size and absence of data for 

some seasons. Although average density from Oregon arrivals was similar to that for Long 

Beach and foreign arrivals, density for Hawaii arrivals was over 10-fold lower than all others.


Figure 3.3. Densities of organisms in ballast water arriving to Port Valdez for each source port. The 

estimated mean densities (#/m³ and standard errors) are shown for all organisms by source port. The data include all 

sample dates (sample size indicated above bars) but exclude chain-forming diatoms. Source ports include: Puget 

Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); Foreign port with open-ocean exchange 

(EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI). 


1.E+05 

1.E+04 

1.E+03 

1.E+02 

1.E+01 

48 

50 

46 19 2 

4 

1.E+00 


SOURCE 

The average total densities among domestic source ports corresponded to voyage 

duration, with densities decreasing with voyage duration (see Fig. 2.7 of Chapt 2). Arrivals from 

Puget Sound and San Francisco Bay had the shortest voyage duration (average of 4.3 and 6.0 

days, respectively) and highest densities. Tankers arriving from Hawaii had the longest voyage 

duration (average of 10.2 days) and lowest densities, whereas arrivals from Oregon and Long 

Beach were intermediate to the other domestic ports in both respects. 

(b) Density by Taxonomic Group and Source 

All ballast water samples contained living organisms, but the prevalence and density 

varied among taxonomic groups (Table 3.1). Copepods and diatoms were detected in ballast 

water from 100% of the ships sampled, and protozoans (primarily tintinnids) were found in 

nearly all samples. These three groups also exhibited the highest densities, dominating the 

plankton community in ballast water (see below). 

As with the total density measures, most taxonomic groups also occurred at greater 

average densities in ballast water from domestic sources, when pooled, compared to that from 

foreign sources (Table 3.1 and Fig 3.4). For most groups, this difference was 10- to 100-fold. 

The magnitude of density differences between domestic and foreign sources were much less for 

crustaceans, primarily due to the presence of copepods (see Table 3.1) and solitary diatoms. 

Dinoflagellates were a notable exception to the general pattern, as average density was greatest 

in ballast water of the foreign arrivals.


Figure 3.4. Densities of major taxonomic groups in ballast water arriving to Port Valdez from foreign and 

domestic source ports. The estimated mean densities (#/m³ and standard errors) are shown separately for 10 

different major groups of organisms in ballast water of ships arriving from each domestic ports (n=150) and foreign 

ports (n=19). Eight groups are distinct phyla that were most abundant in the ballast water, and two are composed of 

multiple phyla, including eggs and trochophores (which were abundant but could not be classified by phylum) and 

all other invertebrates. The data include all sample dates. Since arrivals from foreign ports all underwent ballast 

water exchange in open ocean, the source is indicated as exchange. 

Echinodermata 

Chordata 

Other Inverts 

SOURCE 

Exchange 

Domestic 

Eggs & Trochophores 

TAXA 

Annelida 

Mollusca 

Dinoflagellata 

Protozoa 

Crustacea 

Diatomacea 

0 10 2 10 4 

1 10 100 1000 10000 10000 

DENSITY (#/m³) 

On a finer scale, significant variation existed in the abundance of taxonomic groups 

among specific source port systems (Fig. 3.5 and Table 3.1). More specifically, differences were 

present when comparing mean densities for each taxonomic group among Puget Sound, San 

Francisco Bay, Long Beach, and foreign arrivals (ANOVA, see Fig. 3.5 for statistically 

significant differences). Among these four ports, the densities of most taxa were relatively low 

for foreign arrivals, with the exception of diatoms and dinoflagellates. This pattern resulted from 

both the prevalence and densities of taxa among vessels. For example, the prevalence of most 

taxa was low in the ballast water of foreign arrivals, although the densities may not been 

particularly low for the few ships where a taxon was detected (Table 3.1). Across all arrivals, 

mean densities of the various taxa were generally lowest for both Oregon and Hawaii, although 

the limited sample size precludes any formal analysis. 

In contrast to total organism density, the greatest average densities for taxonomic groups 

did not always correspond to the shortest voyage duration (Fig. 3.5 and Table 3.1). For example, 

the highest average densities of protozoans, crustaceans, and bryozoans were measured for San 

Francisco Bay arrivals, and the brachyuran crabs were most abundant in samples from Long 

Beach. 

(c) Seasonal Variation in Density 

Significant differences among months were present in the total densities of organisms 

present in domestic ballast water (Fig. 3.6; ANOVA, F (11,168) = 1.98, P = 0.033). This resulted 

primarily from a relative increase during the spring and summer months.


Figure 3.5. Densites of major taxonomic groups in ballast water arriving to Port Valdez for each source port. 

The estimated mean densities (#/m³ and standard errors) are shown separately for 10 different major groups of 

organisms in ballast water of ships by source port. Eight groups are distinct phyla that were most abundant in the 

ballast water, and two are composed of multiple phyla, including eggs and trochophores (which were abundant but 

could not be classified by phylum) and all other invertebrates. Source ports include: Puget Sound, WA (PS); San 

Francisco Bay, CA (SF); Long Beach, CA (LB); Foreign port with open-ocean exchange (EX); Columbia River, 

Oregon (OR); and Barbers Point, Hawaii (HI). The data include all sample dates. Sample sizes for each source port 

as shown in Figure 3.3. Indicated by * are those taxa where mean density among ports, excluding HI and OR, is 

significantly different by ANOVA with confidence >95%. 

100000 

DINOFLAGELLATA* 

100000 

10000 

10000 

1000 

1000 

10100 

2 

10 100 

10 

10 

0 1 

01 


DIATOMACEA* 


100000 

10000 

1000 

10100 

2 

10 

0 1 

10 4 0 

PROTOZOA* 


100000 

10000 

4 

1000 

10100 

2 

10 

01 

ANNELIDA* 



100000 

MOLLUSCA 

10000 

1000 

10100 

2 

10 

1 


100000 

10000 

10 4 

10 4 0 

1000 

10 2 

100 

10 

1 

CRUSTACEA* 


100000 

10000 

ECHINODERMATA* 

100000 

10000 

10 4 

1000 

1000 

10100 

2 

100 

10 2 

10 

10 

01 

0 1 


OTHER INVERTS* 


100000 

100000 

CHORDATA* 

10000 

10 10000 

1000 

1000 

10100 

2 

10100 

2 

10 

10 

0 1 

0 1 


SOURCE 

EGGS AND TROCHOPHORES 

PS SF LB EX OR HI


Figure 3.6. Monthly densities of organisms in ballast water arriving to Port Valdez from domestic source 

ports. The estimated mean densities (#/m³ and standard errors) are shown for all organisms, except chain-forming 

diatoms, by month. The data for all domestic source ports are included in each month (sample size shown above 

bars). 

DENSITY (thousands/m³) 

80 

60 

40 

20 

0 

11 

10 

17 15 11 11 

10 

10 11 11 13 20 

Jan Mar May Jul Sep Nov 

SEASON 

The seasonal pattern in total density varied among arrivals from the 3 major domestic 

ports (Fig. 3.7). Arrivals from San Francisco Bay exhibited a strong spring peak in total 

plankton density, whereas the peak appeared later in arrivals from Puget Sound. In contrast, the 

density of organisms arriving from Long Beach was relatively stable throughout the year. 

Figure 3.7. Monthly densities of organisms in ballast water arriving to Port Valdez by domestic source ports. 

For each of the three major domestic source ports, the estimated mean densities (#/m³ and standard errors) are 

shown for all organisms, except chain-forming diatoms, by month. Sample size for each source port as follows, 

from left to right: Long Beach – 3,4,4,4,5,4,2,2,3,4,6,5; San Francisco –2,3,3,5,6,5,2,4,4,4,5,7; Puget Sound – 

5,3,3,2,5,6,5,4,4,3,2,6. 

DENSITY 

(thousands/m³) 

60 

40 

20 

0 

LONG BEACH 


DENSITY 


60 

40 

20 

0 

98.9 

SAN FRANCISCO 


DENSITY 


60 

40 

20 

0 

PUGET SOUND 


SEASON


For the individual taxonomic groups, densities varied both by source port and month (Fig. 

3.8). In general, peak densities occurred between spring and summer months for all taxonomic 

groups, but the timing of these peaks differed among groups. Summed across the three major 

domestic ports (Puget Sound, San Francisco Bay, and Long Beach; Fig. 3.8a): 

• Dinoflagellates, echinoderm larvae, chordates, and the combined eggs and trochophores 

exhibited peak mean densities in late summer; 

• Protozoans and diatoms exhibited a spring to early summer peak in mean density. 

• Molluscs and crustaceans for these combined ports were relatively high from spring through 

summer, exhibiting a bimodal distribution with spring and later summer peaks; 

• Annelids exhibited peaks in density during the summer months of June and August; 

• All other invertebrates, when combined, had relatively high densities from early spring 

through fall compared to the remainder of the year. 

The relative contributions of the three port systems to the overall temporal patterns varied 

significantly (Fig 3.8b-d). The general seasonal patterns (i.e., spring-summer peaks) in density 

were similar among ports, but clear differences existed in the magnitude and month of peak 

densities among ports. As noted previously for total density across the entire year (Fig. 3.5), the 

magnitude of peaks for the taxonomic groups did not correspond consistently to voyage duration. 

(d) Annual Variation in Density 

There were significant differences among years in the total density of plankton arriving 

from the three major ports for June 1997-1999 (Fig. 3.9). A 2-way ANOVA revealed differences 

among years (F (2,32) = 3.28, P = 0.055) but not ports (P>0.05), and the interaction was not 

significant (P>0.05). 

The magnitude of variation among years was more pronounced for the individual 

taxonomic groups in each of these ports (Fig. 3.10). Over half of the taxonomic groups exhibited 

significant differences among years, when analyzed individually for each port source (1-way 

ANOVA, see Fig 3.10 for statistically significant differences). As discussed above, some groups 

(e.g., echinoderm larvae and chordates) could not be compared statistically, due to low 

prevalence among ships. 

Interestingly, the changes among years were not consistent among port sources. For 

example, dinoflagellates increased in successive years for Puget Sound and Long Beach arrivals, 

but was greatest in 1998 and virtually absent in the other two years for San Francisco arrivals. 

While peak years in protozoan and crustacean densities were similar among the port sources, the 

peak years for all other taxa were highly divergent among the port sources.


Figure 3.8. Monthly densities of major taxonomic groups in ballast water arriving to Port Valdez from domestic source ports. The estimated monthly 

mean densities (#/m³ and standard errors) are shown separately for 10 different major groups of organisms in ballast water of ships arriving from domestic ports. 

Eight groups are distinct phyla that were most abundant in the ballast water, and two are composed of multiple phyla, including eggs and trochophores (which 

were abundant but could not be classified by phylum) and all other invertebrates. For each taxonomic group, the monthly mean densities are shown for: (a) Puget 

Sound; (b) San Francisco; (c) Long Beach; and (d) the three major ports combined. Sample size for each port as indicated in Figure 3.7.

Chapt 3. Biological Characteristics of Ballast Water, page 3- 18

Chapt 3. Biological Characteristics of Ballast Water, page 3- 19


Figure 3.9. Densities of organisms in the ballast water arriving to Port Valdez from domestic ports in June of 

three consecutive years, 1997-1999. The estimated mean densities (#/m³ and standard errors) are shown for all 

organisms, excluding chain-forming diatoms, by source port in each year (sample size indicated above bars). Source 

ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB). Densities are also 

shown by year for all three source ports combined (Overall). 

3 

5 

3 

10 

1.E+04 

2 

3 

4 

3 

4 

6 

12 

11 

1997 

1998 

1999 

Density (#/m³) 

1.E+02 

1.E+00 

LB SF PS Overall 

SOURCE 

Figure 3.10. Densities of major taxonomic groups in the ballast water arriving to Port Valdez from domestic 

source ports in June of three consecutive years, 1997-1999. The estimated mean densities (#/m³ and standard 

errors) are shown for 10 different major groups of organisms in ballast water of ships arriving by year from three 

different source ports. Eight groups are distinct phyla that were most abundant in the ballast water, and two are 

composed of multiple phyla, including eggs and trochophores (which were abundant but could not be classified by 

phylum) and all other invertebrates. Sample size for each source port and year as indicated in Figure 3.9. Indicated 

by * are instances where ANOVA showed significant differences in density among years (above bars) and among 

ports (below taxa labels) with confidence >95%. 

100000 

10000 

1000 

100 

LB (JUNE) 

1997 

1998 

* 

1999 

* * * 


10 

1 

100000 


10000 

1000 

100 

10 

Diatomacea 

* * 

Protozoa 

Annelida 

SF (JUNE) 

Mollusca 

Crustacea 

Echinodermata 

Other Inverts 

Chordata 

1997 

1998 

1999 

Eggs & Trocophores 

* 

* 

1 

100000 

10000 

1000 

100 

10 

1 

* 

* 

* 

PS (JUNE) 

1997 

1998 

1999 


Diatomacea 

Protozoa 

Annelida 

Mollusca 

Crustacea 


Echinodermata 

*Diatomacea 

Other Inverts 

*Protozoa 

Chordata 

Annelida 

Eggs & Trocophores 

Mollusca 

Crustacea 

Echinodermata 

Other Inverts 

Chordata 

*Eggs & Trochophores


3C3. Diversity of Organisms in Ballast Water 

In a preliminary analysis of our Fine Analysis data (see Methods), there was a significant 

difference in the minimum number of taxa, and/or species richness, detected among arrivals from 

the three main domestic ports (Fig. 3.11). The average species richness was greatest for Long 

Beach arrivals and lowest for San Francisco Bay arrivals, and it does not correspond to voyage 

duration. 

Figure 3.11. Minimum number of taxa detected in the ballast water arriving to Port Valdez by domestic 

source ports. Shown are the mean number (including standard errors and sample size, above bars) of distinctly 

different taxa observed in plankton samples of ships from each source port. Source ports include: Puget Sound, WA 

(PS); San Francisco Bay, CA (SF); Long Beach, CA (LB). All sample dates included. 

The season of peak species richness differed among port sources (Fig. 3.12). Arrivals 

from Puget Sound and San Francisco Bay exhibited peaks in mean species richness in the spring 

and summer, whereas those from Long Beach had their highest species richness in the fall and 

spring. Subsequent analyses indicated differences among source ports for each season, as well as 

differences among seasons for each source port (1-way ANOVA, Fig. 3.12 indicates statistically 

significant differences). 

To date, we have identified 14 different nonindigenous species arriving to Port Valdez in 

the ballast water of oil tankers (Table 3.2). One is a fish species and all the other species are 

crustaceans (copepods and amphipods, which have successfully invaded the respective source 

ports of arriving tankers. To date, all of these identified NIS have been in ballast water from San 

Francisco and Long Beach. 

We expect the cumulative list will increase, as final identifications are still underway for 

the Fine Analysis data. Upon completion, we will report the frequency and density of these NIS 

in ballast water arriving from the respective source ports.


Figure 3.12. Minimum number of taxa detected in the ballast water arriving to Port Valdez among domestic 

source ports and seasons. Shown are the mean number (including standard error above bars) of distinctly different 

taxa observed in plankton samples of ships from each source port and season. Source ports include: Puget Sound, 

WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB). Seasons include: Winter, (December – February); 

Spring (March – May); Summer (June – August); and Fall (September – November). Indicated by * are significant 

differences (ANOVA with confidence >95%) in diversity among seasons within port (see legend) and among ports 

within season (see x-axis labels). 

Minimum Number of Taxa 

40 LB 

PS 

SF* 

30 

20 

10 

0 

*Fall 

*Spring *Summer *Winter 

Season 

Table 3.2. Nonindigenous species identified in ballast water arriving to Port Valdez. The source of ballast 

water is indicated in which each species was detected; when two sources are indicated, the species was found in 

ballast water from each source port. Source ports are: San Francisco Bay, CA (SF); Long Beach, CA (LB). 

Broad Taxa Species Ballast Source 

Amphipoda Ampelisca abdita SF and LB 

Monocorophium acherusicum 

SF 

Sinocorophium heteroceratum 

LB 

Gammarus daiberi 

SF and LB 

Grandidierella japonica 

SF 

Copepoda Limnoithona tetraspina SF 

Oithona davisae 

LB 

Acartiella sinensis 

SF 

Pseudodiaptimus marinus 

SF 

Pseudodiaptimus forbesi 

SF 

Sinocalanus doerrii 

SF 

Tortanus dextrilobatus 

SF 

Mysidacea Acanthomysis bowmani SF 

Chordata Acanthogobius flavimanus SF 


Our analysis indicates that significantly greater numbers of organisms are discharged into 

Port Valdez and PWS in oil tankers arriving from domestic ports compared to those from foreign 

ports. This results from the number of arrivals and the density of organisms in their ballast 

water, as both are greatest for the domestic arrivals. Accounting for number of arrivals and 

density (by source port and season), Table 3.3 estimates the total supply of plankton that we


sampled to be roughly 264 billion organisms in 1998. Of this, approximately 3% arrived from 

foreign traffic. 

The differences observed in total density, as well as taxon-specific density, among arrivals 

from different source ports may result from a combination of multiple factors, including (a) 

differences in initial densities, (b) differences in survivorship, and (c) effects of ballast water 

exchange (conducted for foreign but not domestic arrivals). 

Table 3.3. Estimated number of large, planktonic organisms delivered in tankers’ ballast water to PWS and 

Port Valdez in 1998. Shown by source port and season are (1) the estimated total ballast water volumes, (2) mean 

densities of planktonic organisms, including standard errors and sample size, and (3) total number of planktonic 

organisms arriving in the ballast water of oil tankers. Total Volumes are derived from Table 2.1. Mean densities 

were estimated from analysis of plankton samples, which were collected by 80micron net, and exclude chainforming 

diatoms (see text for description). Where no samples were available for a season (e.g., Hawaii and Oregon), 

the grand mean across all samples of that source port was used. The bottom row (Overall) estimates the total ballast 

water volumes and total organisms delivered as a sum. Source ports include: Puget Sound, WA (PS); San Francisco 

Bay, CA (SF); Long Beach, CA (LB); Foreign port with open-ocean exchange (EX); Columbia River, Oregon (OR); 

and Barbers Point, Hawaii (HI). Seasons include: Winter (January-March), Spring (April-June), Summer (July- 

September), and Fall (October-November). 

Density organisms (#/m³) 

Phyto.+Zoopl. Zoopl. Phyto.+Zoopl. Delivered Zoopl. Delivered 

Port/Source Season Total BW (m³) Mean(se) Mean(se) n Billions Billions 

Winter 1,802,432 5963(1432) 879(343) 11 10.75 1.58 

Spring 1,885,260 17602(9543) 3427(1423) 11 33.18 6.46 

PS Summer 2,032,432 24116(8198) 8539(3062) 12 49.01 17.35 

Fall 1,796,382 8546(2636) 1269(384) 8 15.35 2.28 

Grand total 7,516,506 42 108.30 27.68 

Winter 1,082,594 24686(13263) 13435(9497) 8 26.72 14.54 

Spring 1,345,169 48504(23134) 29876(14848) 13 65.25 40.19 

SF Summer 1,573,660 14572(2914) 10696(1959) 10 22.93 16.83 

Fall 1,340,469 8840(1961) 7162(1488) 12 11.85 9.60 

Grand total 5,341,892 43 126.75 81.17 

Winter 585,468 4298(1094) 3426(954) 11 2.52 2.01 

Spring 395,187 4848(667) 3734(490) 10 1.92 1.48 

LB Summer 504,630 15951(3274) 15145(3175) 6 8.05 7.64 

Fall 390,180 6574(1613) 5508(1639) 12 2.57 2.15 

Grand total 1,875,465 39 15.05 13.27 

Winter 379,377 8791(3127) 1852(423) 7 3.34 0.70 

Spring 265,764 3466(2144) 1634(775) 3 0.92 0.43 

EX Summer 149,280 18447(14784) 3393(3141) 5 2.75 0.51 

Fall 141,168 3571(1398) 3179(1100) 2 0.50 0.45 

Grand total 935,589 17 7.51 2.09 

Winter 262,638 - - 0 2.49 0.02 

Spring 113,000 - - 0 1.07 0.01 

OR Summer 97,287 - - 0 0.92 0.01 

Fall 273,966 9474(6138) 65(13) 2 2.60 0.02 

Grand total 746,891 7.08 0.05 

Winter 189,994 772 17 1 0.15 0.00 

Spring 111,145 - - 0 0.05 0.00 

HI Summer 142,596 102(72) 14(2) 2 0.01 0.00 

Fall 108,370 - - 0 0.05 0.00 

Grand Total 552,105 3 0.26 0.01 

Overall 16,968,448 145 264.94 124.26


Among the domestic arrivals, many of the observed differences in prevalence and density 

probably result from initial differences at the locations where ballast tanks are filled. For 

example, this may explain the especially strong differences observed in densities of some 

organisms, such as dinoflagellates and protozoans (Fig. 3.8), among source ports. Although we 

have very limited data on the initial densities within ballast tanks at the start of the tankers’ 

voyages (see Chapter 5), the published literature indicates that significant variation in the density 

and diversity of plankton communities among these and other source ports should be expected. 

In this context, it is perhaps important to recognize some conspicuous differences that existed 

among the domestic source ports. Certainly there are many differences in the habitat 

characteristics (e.g., composition, extent, quality, proximity to ports, etc), which may influence 

what is initially entrained in the tankers’ ballast tanks. However, there are also two 

physical/chemical characteristics that are widely recognized to influence the composition and 

dynamics of biotic communities: temperature and salinity. Temperature clearly differed among 

domestic port systems, increasing from north to south. Among the major domestic ports, salinity 

was extremely low for San Francisco Bay compared to Puget Sound and Long Beach in 1998, in 

which rainfall was relatively high (due to El Nino Southern Oscillation) and had a 

disproportionately large effect on salinity in San Francisco Bay. 

Despite any initial differences in plankton communities, it is evident that survivorship 

during transit can contribute strongly to the observed differences in biota arriving from various 

source ports. A variety of studies have now shown a significant decline in the density of 

planktonic organisms in ballast tanks during voyages, and the magnitude of decline is timedependent, 

increasing significantly with voyage duration (Wonham et al. 1996, LaVoie et al 

1999, Smith et al. 1999; however see below for possible exceptions). We have obtained similar 

results aboard oil tankers arriving to Port Valdez (Chapter 5). For most taxa, the decline has been 

attributed to mortality. However, for a few groups included in our analysis, such as diatoms and 

dinoflagellates, it is possible for the organisms to develop dormant stages that can accumulate on 

the bottom of ballast tanks. 

Ballast water exchange undoubtedly had a significant effect on the plankton community 

associated with foreign arrivals, contributing to the major differences in biota between foreign 

and domestic arrivals. Exchange can significantly reduce the concentration of many organisms 

within ballast tanks, and it can also entrain additional organisms from the oceanic site of 

exchange (Ruiz et al. 1997, 1999; see also Chapter 5). In our study, the combination of ballast 

water exchange and voyage duration (which was relatively high for foreign ports) would both 

operate reduce initial densities of coastal plankton and contribute to the lower abundance of 

many taxonomic groups in ballast water of foreign arrivals compared to that from domestic 

arrivals. In contrast, the domestic arrivals did not undergo ballast water exchange and arrived to 

Port Valdez with the initial coastal water, following a relatively short voyage. 

We hypothesize that the combined effects of ballast water exchange and voyage duration, 

instead of initial densities, were responsible for observed differences in abundance of coastal 

organisms between domestic and foreign arrivals. More specifically, we suggest that these 

forces reduced the densities of predominantly coastal organisms such as cnidarians, flatworms, 

annelids, molluscs, chordates, echinoderms, bryozoans, barnacles, and many other crustacean 

groups (see Chapter 5 for further discussion).


The effects of ballast water exchange for some taxonomic groups, and its contribution to 

observed differences in their abundance between foreign and domestic arrivals, is not so well 

resolved. Unlike the low abundance of coastal organisms, foreign arrivals had relatively high 

densities of dinoflagellates, copepods, and solitary diatoms in their ballast water. Most of these 

organisms were probably oceanic in origin and were entrained during the exchange process. 

This is certainly the case for the copepods, for which the species were recognized as oceanic and 

the generation time is in excess of the voyage duration. However, there is some suggestion that 

an increase of phytoplankton can result from ballast water exchange, as generation times are 

relatively short and the organisms may respond rapidly to changes in water quality following 

exchange (Gollasch et al.1998, LaVoie et al. 1999). The extent to which populations of these 

taxa, either of coastal or oceanic origin, may have increased following exchange is uncertain. 

The temporal variation observed in plankton densities was largely expected. In general, 

the seasonal peaks in density corresponded to seasonal production and density variation 

measured for plankton in north temperate estuaries. The magnitude of variation observed among 

years also is evident in field studies, including especially an El Niño event such as that for 1998. 

The heavy rainfall in that year was associated with especially high densities of protozoans, 

solitary diatoms, and copepods in the arrivals from San Francisco Bay. 

Although we have identified 14 nonindigenous species in the ballast water arriving to Port 

Valdez, and have provided some comparative data on species richness, these results must be 

viewed with caution. Clearly these numbers are minimum estimates. Although both estimates 

will increase upon completion of the voucher identification (see results), the measures can only 

be applied to a subset of the taxa and will always represent a minimum value. More specifically, 

most of the larval invertebrates (e.g., molluscs, barnacles) include many different species, which 

cannot be readily distinguished as larvae. All bivalve larvae are therefore treated as one species 

in our analysis, masking the diversity that most certainly exists. Thus, this approach is useful 

primarily in describing minimum diversity of native and exotic species arriving in ballast water, 

and does not necessarily reflect actual diversity patterns in space or time. 

It is also important to recognize that our conclusions about patterns of abundance and 

diversity are focused on the large (>80 micron) segment of the plankton community within 

ballast tanks. We have provided some additional qualitative information about the macrofauna 

found on the bottom of tankers’ ballast tanks (Chapt 6, this report). Thus, our data do not 

address density or diversity of microorganisms and taxa missed by an 80 micron mesh. The 

dynamics of these groups are very much in debate, as few good data exist to discern the potential 

for population changes (either declines or increases), due especially to mortality, dormancy and 

cyst formation, or ballast water exchange. 

Beyond the variation in plankton delivery by time and source port, our data underscore 

that both the concentration and cumulative amount of plankton arriving in tankers’ ballast water 

to Port Valdez and PWS is relatively high compared to that estimated for other ports (e.g., 

Carlton and Geller 1993, Smith et al. 1999). This results from both the volume of water 

delivered (Chapter 2) and the concentration of plankton, as both values are relatively high. We 

hypothesize that the abundant plankton results from the short voyage duration for domestic 

traffic, accounting for approximately 97% of the total tanker arrivals at present. In contrast,


although Chesapeake Bay receives more total ballast water per year than PWS, most of the 

ballast water comes from Europe, arriving in 10-14 days with an average concentration of 200 

organisms / m 3 (Smith et. al. 1999; Ruiz et al., unpubl. data). Furthermore, it appears that 

ballast water arriving to the Chesapeake from domestic ports also has a lower density than that 

arriving to PWS from domestic ports. 

Finally, from an invasion perspective, there are three unusual features of our analysis that 

deserve explicit mention: 

• This is the most comprehensive analysis of domestic, coastwise ballast transfer. Most 

organisms that arrive in ballast water to PWS come from domestic source ports, which are 

themselves highly invaded. Thus, our study examines the opportunity for sequential 

invasions, which can “leapfrog” up the coast following initial colonization of North America. 

• The delivery of ballast water by tankers to Port Valdez is a relatively recent development, 

beginning in 1977. Although many features may influence the risk of invasion, it is often 

considered to increase with the frequency, density, and duration of inoculation. Our results 

indicate that the risk associated with the first two of these is relatively high. However, the 

operation of this transfer mechanism has only existed for three decades. Even at the current 

rates of organism delivery (see above), invasion success may be influenced strongly by 

duration. In contrast, many other ports have been receiving ballast water and ballast 

materials for a century or more. 

• Delivery of ballast water from foreign sources by tankers is even a more recent development, 

beginning in 1996. Although this accounted for only 3% of the total volume of ballast water 

delivered in 1998, all of the ballast water delivered by foreign arrivals had undergone 

exchange. Some coastal organisms remained in the exchanged tanks (see Chapter 5); 

however, the total supply of organisms from foreign ports is both relatively small and 

extremely recent compared to other port systems. 


Carlton, J.T. and J. B. Geller. 1993. Ecological roulette: the global transport of nonindigenous 


Gollasch, S., M. Dammer, J. Lenz and H.G. Andres. 1998. Non-indigenous organisms introduced 

via ships into German waters. Pp. 50-64 in Carlton, J.T. (ed.), Ballast Water: Ecological and 

Fisheries Implications. International Council for the Exploration of the Sea (ICES), Denmark. 

Lavoie, D. M., L.D. Smith and G.M. Ruiz. 1999. The potential for intracoastal transfer of nonindigenous 

species in the ballast water of ships. Est. Coast. Shelf Sci. 48: 551-564. 


Prince William Sound, Alaska: Pilot Study. Report, Prince William Sound Regional Citizens’ 

Advisory Council. 80pp. 

Ruiz, G.M., J.T. Carlton, A. H. Hines and E.D. Grosholz. 1997. Global invasions of marine and 

estuarine habitats by non-indigenous species: Mechanisms, extent, and consequences. Am. Zool. 

37: 621-632.


Ruiz, G.M., P. Fofonoff and A.H. Hines. 1999. Non-indigenous species as stressors in estuarine 

and marine communities: Assessing invasion impacts and interactions. Limnol. Oceanogr. 44: 

950-972. 







Invasions 1: 67-87. 

Wonham. M.J., W.C. Walton, A.M. Frese and G.M. Ruiz. 1996. Transoceanic transport of 

ballast water: Biological and physical dynamics of ballasted communities and the effectiveness 

of mid-ocean exchange. Final Report to the U.S. Fish and Wildlife Service and the Compton 

Foundation.

Chapt 4. Predicting Initial Survival of Ballast Water Organisms, page 4- 1 

Chapter 4. Predicting Initial Survival of Ballast Water Organisms 




Melissa Frey, Smithsonian Environmental Research Center 

4A. Purpose 

When ballast water is discharged into the receiving waters, the associated plankton 

encounters new conditions without time to acclimate. Survival may depend on short-term 

tolerances to accute variation in salinity-temperature combinations. If temperature-salinity 

conditions of ballast water closely match those of the receiving waters, then initial survival is 

predicted to be higher than when the conditions do not match closely. To determine if NIS 

arriving in ballast water can survive the initial exposure to temperature-salinity conditions in 

Prince William Sound, we tested the match of conditions between ballast water and ship-side 

water, and the short-term survival of ballast organisms in representative combinations. 

4B. Temperature & Salinity: Match of Source and Receiving Ports 

4B1. Methods 

Samples of ballast water were collected from segregated ballast tanks and from ambient 

waters adjacent to each ship sampled for ballast water plankton. A small niskin bottle lowered 

through hatch covers into the ballast tanks and lowered off the end of the ship’s berth to collect 

samples from the water surface and 10 m depth, which was determined to be below a potential 

thermocline or pycnocline. Salinity was determined to the nearest ppt with a refractometer and 

temperature to the nearest 0.5 o C with a hand-held thermometer. 

4B2. Results 

Temperature and salinity of the receiving waters of Port Valdez exhibit a distinct 

seasonal pattern (Fig. 4.1a, b). Water temperatures of Port Valdez at 10 m depth cycle 

seasonally from a low of 4 o C in February to a high of 13 o C in July. Surface water temperatures 

are more variable and 1-5 o C warmer than deep water in the spring. Salinity during December to 

April was about 31 ppt and the water column was well mixed. Water in Port Valdez was sharply 

stratified by depth as snow melted from late April to September, with salinities of surface waters 

dropping to 4-15 ppt while salinities at 10m depth declined only to about 25 ppt. 

Water in the segregated ballast tanks rarely exhibited much depth stratification. 

Temperatures of segregated ballast water varied seasonally with a winter mean low of about 7.5 

o C (+-3 o C) and a summer high of about 16 o C (+-3 o C) (Fig. 4.1c). Salinities of ballast water did 

not exhibit a seasonal pattern, but salinities fell into two distinct ranges, depending on the source 

port of the tanker (Fig. 4.1d). Most tankers delivered high salinity ballast water (ca. 30 ppt, 

range 20-36 ppt). In contrast, about 20% of the tankers throughout the year released ballast water 

of low salinity (ca. 4 ppt, range 0-14 ppt, mainly from Benicia in San Francisco Bay, especially 

during the heavy El Niño rains in 1998).


Figure 4.1 Seasonal Cycles of Temperatures and Salinities of Receiving and Ballast Water. Shown above are 

temperatures (A) and salinities of Port Valdez, Alaska at 0m and 10m depth. Shown below are average 

temperatures (C) and salinities (D) of ballast water discharged into Port Valdez by tankers. Averages are for two 

tank depths (0m and 10m depth) combined. 

A. Port Valdez 

C. Ballast Water 

Temperature (ºC) 

25 

20 

15 

10 

5 

0 

Dec Feb Apr Jun Aug Oct Dec 

0 m 

10 m 

Temperature (ºC) 

25 

20 

15 

10 

5 

0 


B. Port Valdez 

40 

D. Ballast Water 

40 

Salinity (‰) 

30 

20 

10 

0 m 

10 m 

Salinity (‰) 

0 

Dec Feb Apr Jun Aug Oct 

SEASON 

Dec 

30 

20 

10 

0 


SEASON 

Annual variation in temperature and salinity among source ports and receiving waters of 

Port Valdez was compared in June of 1997, 1998, and 1999 (Fig. 4.2). Temperature of ballast 

water from Long Beach (13-14 o C) and San Francisco(13-14 o C) was about a degree warmer than 

from Puget Sound (11-12 o C), but temperature of ballast water from Puget Sound was similar to 

the surface water of Port Valdez (11-12 o C), which was 1-3 o C warmer than water at 10 m depth 

(7-9 o C). However, there were no significant differences in temperatures among years. Salinity of 

ballast water from Long Beach (33ppt) was highest of the source ports, while that from San 

Francisco (10-14 ppt) was the lowest, and Puget Sound was intermediate (29-30 ppt). Surface 

salinity at the surface of Port Valdez (10-21ppt) was similar to ballast water from San Francisco 

Bay, while salinity at 10 m depth in Port Valdez (29-30 ppt) was similar to ballast water from 

Puget Sound. Salinity of ballast water from Long Beach and Puget Sound, as well as deep water 

at Port Valdez, did not differ among years. However, salinity of ballast water from San 

Francisco Bay and at the surface in Port Valdez was lowest in 1998, highest in 1997 and 

intermediate in 1999. 

Thus, there was often a good correspondence of physical characteristics between ballast 

water and receiving water, depending on the source port, time of year and water depth in Port 

Valdez. Temperatures of ballast water were a bit higher than of receiving waters, but the 

differences were not great, and there was considerable overlap between ballast and receiving 

water throughout the year. Higher salinities of ballast water from most source ports were similar 

to deeper water of Port Valdez throughout the year and similar to surface water during winter 

and early spring. During summer the vertical stratification in Port Valdez resulted in ballast 

water from both high and low salinities having good correspondence in major areas of the 

receiving waters. Based on these physical characteristics of ballast and receiving water, 

temperatures of Port Valdez would not appear to prevent survival of organisms from most source 

ports. Nonindigenous species from nearly fresh water, estuarine and full-strength sea water may 

also find corresponding salinities in Port Valdez.


Figure 4.2 Annual Variation in Temperature and Salinity of Ballast and Receiving Water. Shown are 

temperatures (top) and salinities (bottom) of ballast water arriving to Port Valdez in tankers from west-coast source 

ports (PS = Puget Sound, WA; SF = San Francisco, CA; LB = Long Beach, CA), and of recieving waters of Port 

Valdez at surface (V-0) and 10m depth (V-10). Bars indicate means and S.E. for June 1997, 1998, 1999. 

15.0 

June’97 

June’98 

June’99 

DEGREES (ºC) 

10.0 

5.0 

0.0 

PS SF LB V-0 V-10 

30.0 

SALINITY (‰) 

20.0 

10.0 

0.0 

PS SF LB V-0 V-10 

SOURCE 

4C. Temperature-Salinity Tolerance Experiments of Ballast Water Plankton 

4C1. Methods 

We conducted experiments at the SERC laboratory in Valdez to test for temperature x 

salinity tolerance of selected planktonic organisms arriving in segregated ballast water. Based on 

the two salinity categories of segregated ballast water released into Port Valdez (see above, Fig. 

4.1a, b), we grouped the experimental organisms into "freshwater taxa" and "seawater taxa" (Fig. 

4.3). 

Figure 4.3. Survivorship of Ballast Water Organisms in Salinity x Temperture Experiments. Survivorship of 

ballast water organisms at 96 hour exposure to 9 combinations of salinity and temperature in laboratory experiments. 

Three salinities (10, 20, 30 ppt) and three temperatures (3, 9, 15ºC) were tested to represent the range of seasonal 

variation in Port Valdez. (A) Trials with organisms from ballast water with fresh water sources (n = 9). (B) Trials 

with organisms from ballast water with seawater sources (n = 15). 

A 

B 

Mean Survivorship 

1.0 

0.8 

0.6 

0.4 

0.2 

0.0 

1.0 

0.8 

"Freshwater" Taxa 

3 C 

9 C 

15 C 

10 20 30 

"Seawater" Taxa 

0.6 

0.4 

0.2 

0.0 

10 20 30 

Salinity (ppt)


Nine combinations of three temperatures (3, 9, and 15 o C) and three salinities (10, 20, and 

30 ppt) were selected to represent the seasonal range of conditions for Port Valdez. Organisms 

used in the experiments are collected from the common species of plankton arriving in tanker 

ballast water. For each experiment, 10 individuals were placed in each of 3 replicate culture 

dishes at each of the 9 treatment combinations. Thus, there were 27 trials in each experiment (9 

treatments x 3 replicates). The test organisms were sorted in the lab and transferred directly to 

culture dishes maintained in incubators for 96 hrs, simulating release of ballast water into 

conditions of the Sound. Phytoplankton or brine shrimp nauplii were supplied as food to the 

cultures during the test period. 

Experiments (n = 24) were completed with organisms from the following taxonomic 

categories: 

Tintinnid protistan 

Nemertean worm larvae 

Spionid polychaete worm larvae 

Gastropod veliger larvae 

Copepods 

calanoid 

harpacticoid 

cyclopoid (Oithona spp.) 

Barnacle nauplii 

Crab zoea 

Mysid shrimp 

1 experiment 

2 experiments 

3 experiments 

1 experiment 

4 experiments 

1 experiment 

6 experiments 

3 experiments 

1 experiment 

2 experiments 

Nine of these were "freshwater taxa" and 15 were "seawater taxa". We intentionally selected 

copepods especially Oithona spp., for many of our experiments, because we recognized these as 

NIS arriving in apparently good condition from San Francisco Bay. 

4C2. Results 

Short-term survivorship of these ballast water organisms was high (>50%) for fresh water 

taxa at 10 ppt, and for seawater taxa at 20-30 ppt. These short-term experiments also showed that 

the ballast organisms had distinct, but quite broad tolerances that clearly overlap conditions of 

temperature and salinities in Port Valdez. For example, although there was considerable 

variation in survivorship among individual experiments, mean survivorship of calanoid copepods 

varied from about 20-80% for each test salinity, but survivorship of calanoids in most 

experiments increased with salinity (Fig. 4.3). Oithona spp. (which include known NIS 

copepods) were able to tolerate salinity-temperature conditions they would encounter in the 

receiving waters of Port Valdez. 

The freshwater and seawater taxa differed substantially in their patterns 

of temperature x salinity tolerance (Fig. 4.3). The survivorship of seawater taxa at any of the 3 

test temperatures generally increased with increasing salinity, and there were not great 

differences in survivorship among temperatures. However, survivorship of freshwater taxa at all 

temperatures generally declined sharply with increasing salinity, and survivorship at 3 o C was 

markedly lower than at 9 or 15 o C.


4D. Conclusions 

Planktonic species arriving to Port Valdez in ballast water have high potential of 

surviving the salinity-temperature conditions that they encounter during initial discharge from 

the ship. Although some taxa will not tolerate some salinity layers in the seasonally stratified 

conditions in the Port, the overlap of ballast water with Port conditions at some strata is high. 

Plankton in the ballast water, including known NIS such as Oithona spp., should be able to 

tolerate these conditions. Conditions other than initial salinity-temperature combinations 

probably determine whether or not these organisms survive to become established within Prince 

William Sound.

Chapt 5 Ballast Water Exchange Experiments, page 5- 1 

Chapter 5. Ballast Water Exchange Experiments on Tankers 




Melissa Frey, Smithsonian Environmental Research Center 

Safra Altman, Smithsonian Environmental Research Center 

Kimberly Philips, Smithsonian Environmental Research Center 

Tami Huber, Smithsonian Environmental Research Center 

Sara Chaves, Smithsonian Environmental Research Center 

5A. Purpose 

The primary objective of this research component is to measure the efficacy of ballast 

water exchange in removing various types of taxa from ballast tanks of oil tankers. There are very 

few quantitative studies that have measured the effects of exchange, and these are restricted to 

just a few vessel types and measure the effect on a small subset of entrained taxa. It is likely, 

however, that the efficacy of exchange varies by vessel type, tank design, and organism type. To 

date, there have been no measures of ballast water exchange for oil tankers. 

Ballast water exchange is the most widely used national and international management 

strategy to limit new invasions associated with ships’ ballast water (Hallegraeff 1998, Zhang & 

Dickman 1999, Dickman and Zhang 1999,. Moreover, exchange is currently the only treatment 

method available for commercial ships to reduce the quantities of non-indigenous coastal plankton 

in ballast water (National Research Council 1996). This practice is recommended by the 

International Maritime Organization (IMO) to reduce the risk of invasion by shipping. 

Furthermore, the U.S. Congress passed the National Invasive Species Act of 1996 (NISA) to 

encourage ballast water exchange. Specifically, NISA requests that vessels arriving from outside 

of the Exclusive Economic Zone (EEZ) voluntarily conduct open-ocean exchange of ballast tanks 

to be discharged in U.S. ports 

Commercial ships practice two basic types of ballast water exchange to replace coastal with 

oceanic water. Flow-Through (FT) Exchange is conducted by pumping oceanic sea water 

continuously through a ballast tank to flush out the ballast water originating from a coastal source 

port. Empty-Refill (ER) Exchange is performed by emptying a ballast tank of its coastal water and 

refilling it with oceanic water. 

Each exchange method may vary in efficacy due to the amount and circulation of water 

being removed, independent of any tank- or vessel-specific effects on efficacy. For example, FT 

Exchange initially has the effect of dilution but not complete replacement of ballast water. 

Alternatively, organisms may differ in their distribution or response to water turbulence. Some 

taxa may swim against currents or always reside near the bottom of tanks, which could greatly 

influence the effect of ballast water exchange on removal. 

To maximize the degree of exchange, multiple exchanges are often recommended. The 

current IMO standard recommendation is 300% exchange for Flow-Through, while 100-200%


exchange is common for Empty-Refill. These recommended standards provide a theoretical level 

of at least 90% replacement of coastal water by oceanic water, but this is largely untested among 

the broad range of vessel types and tank configurations. Specifically, there are almost no 

experimental analyses which quantify the efficacy of alternative exchange methods and multiple 

tank exchanges, even though (a) this is the present national and international management strategy 

being implemented and (b) the cost of such exchanges is substantial in ship fuel and operations 

time. 

We initiated a rigorous quantitative comparison of ballast water exchange methods on oil 

tankers arriving to PWS. Across two years, we conducted replicated exchange experiments, 

allowing us to measure the effects of both exchange methods on reduction of entrained organisms 

and standard physical and biological tracers. 

We hypothesize that (1) Empty-Refill exchange will have the highest efficiency, (2) 

relatively little reduction in density occurs after the first exchange event, and (3) a significant 

difference exists among taxa on the effect of ballast water exchange. Our experiments were 

designed to directly test these hypotheses and provide needed quantitative data on this management 

practice. 

This work was initiated in two phases, extending the duration of the analyses and 

allowing us to increase the replication (and therefore strengthen the statistical power and value of 

this analysis). The first phase was initiated in summer of 1998. Through the cooperation and 

financial support from the American Petroleum Institute, SeaRiver and ARCO, we conducted the 

ballast water exchange experiments on four separate voyages of tankers to Alaska in June/July of 

that year. The second phase was initiated in spring of 1999, when we received additional 

funding from U.S. Fish and Wildlife Service to conduct similar experiments (with increased 

measurements) on another four voyages in summer. In addition to analysis of these experiments, 

we agreed (with additional funding from phase two) to provide a review of existing data on the 

efficacy of ballast water exchange, allowing us to examine our results from oil tankers to those 

reported for other vessel types. 

All of the experiments have been completed, but we have not yet completed the full 

analysis of all samples. We report here on the experiments conducted, including the status of 

sample analysis and initial results. We will provide a comprehensive report of the results across 

both phases upon completion of our analyses. We anticipate that these results will be available 

by June 2000. 

5B. Methods 

Although the overall goal of this research was to measure the efficacy of ER and FT 

methods of ballast water exchange in removing coastal plankton from ballast water, our 

experimental design allowed us to address three specific objectives: 

• Compare the efficacy of ER and FT exchange methods in removing a range of different 

materials (biotic and abiotic); 

• Measure the effect of repeated exchanges: comparing 100, 200, and 300% exchange of the 

tanks.


• Measure the survivorship of organisms in ballast water over the course of routine voyages. 

Since the density of organisms can change during a voyage (see Chapter 3 for discussion), it 

was important to control for such changes in experimental tanks that were independent of the 

exchange treatment. For this purpose, we included identical measures for an unexchanged 

control tank on every vessel. Although key to the experimental analysis, this also provided 

an opportunity to address this third objective. 

All experiments were conducted aboard oil tankers during regular operations, en route to 

Port Valdez. Each tanker served as an experimental platform. Each ship was boarded by a pair of 

SERC staff at a domestic source port (San Francisco, Puget Sound, or Long Beach), where ballast 

water used to fill the segregated ballast tanks just in advance of departure for Port Valdez. The 

tanks underwent various treatments and were sampled repeatedly during the voyage (below). 

Experimental Design 

The experiment consisted of a replicated, factorial, and paired design. On each ship, we 

sought to use 3 different ballast tanks that were each subjected to a different treatment: No 

exchange (=Control), ER Exchange, and FT Exchange. Each Treatment tank was sampled as 

many as 5 time points, coinciding with: initial ballast loading, 100% exchange, 200% exchange, 

300% exchange, and final at arrival to PWS. 

Ballast water was loaded in accordance with standard operating procedures at dockside. All 

exchanges occurred in open, oceanic conditions well outside the influence of coastal waters (>75 

miles offshore). Exchanges of tanks were managed by ships’ crews in coordination with the desired 

sampling schedule. For FT Exchange, sea water was pumped into the ballast tanks, causing ballast 

water to overflow through the top of the tanks and onto the deck. After a volume of water equal to 

the volume of the tank was pumped, the exchange was interrupted and samples were collected. For 

ER Exchange, ballast tanks were drained initially by gravity and then by pumping before refilling 

with sea water. The process required approximately 12 hours for each multiple of exchange. 

The specific details of implementation and sampling are described below in various 

sections. 

Biotic and Abiotic Tracers. 

In addition to quantifying the effect of ballast water exchange on entrained plankton 

communities in the ballast tanks, we used four different types of tracers for parallel measures of 

efficacy. One of these (salinity of the resident water) simply involved collecting and measuring 

attributes of the resident water. The other three involved materials that we added directly to the 

ballast tanks, including: Rhodamine dye to trace the fate of the initial water; 1 um Fluorescent 

Microspheres that simulate passive particles such as cysts; and newly hatched Artemia (brine 

shrimp) nauplii, native to San Francisco Bay, as a living particle. 

Each tracer can provide information about different components of the ballast tank 

environment during exchange (as indicated above). Moreover, we were interested in developing 

some standardized measures for comparisons across ships. Since the resident community within 

each ship’s ballast tanks may differ considerably (see Chapter 3), the tracers could provide a


common currency for comparing exchange performance among many ships in a way that is simply 

not possible for the entrained plankton communities. 

Tracers were added to ballast tanks in a standardized way, following approval for use in 

these experiments by the U.S. Environmental Protection Agency. The quantity of tracer was 

chosen to produce desired concentrations for the specific volumes of each tank, such that 

anticipated dilution during exchanges would allow us to detect at least a 100-fold reduction in 

measurable concentrations of each tracer. 

Tracers were added to at least two locations in each tank during early stages of ballast tank 

filling (i.e., before the tank was 25-50% full), so as to increase the opportunity for mixing 

throughout each tank. Rhodamie and microspheres were added directly to all tanks. In contrast, 

Artemia were initially cultured (i.e., the cysts were added to salt water and hatched in buckets in 

advance of boarding the ship), and the resulting organisms were used to inoculate ballast tanks. 

Sample Collection. 

Replicate samples were collected at 2 –3 different locations (i.e., tank access points) from 

each tank, for up to 5 different sampling periods (as above). Sampling procedures for the plankton 

community followed our established protocol for characterization of ballast water (see Chapter 3 

for description); Artemia abundance was measured as a component of the plankton (see below). 

Replicate whole water samples were collected from two depths (0m and 10m), using a Niskin 

bottle. Whole water samples were used to measure salinity, temperature, and concentrations of dye 

and microspheres. 

For each sampling location and period, we collected 2 replicate samples for all measures. 

Thus, for each tank and sampling period, we obtained at least: 4 plankton samples (2 locations x 2 

samples); 8 rhodamine samples (2 locations x 2 depths x 2 samples); 8 mircosphere samples (2 

locations x 2 depths x 2 samples). Temperature and salinity measures were made immediately 

upon all replication Niskin samples (at least 8 per tank and sampling period, as above). 

Although we followed the same general sampling protocol for all voyages (in both years), 

we collected additional whole water samples during the 1999 experiments to measure changes in 

the abundance of total bacteria and ciliate protozoans. For both measures, we collected at least 8 

samples per tank and sampling period (2 locations x 2 depths x 2 samples). Samples were 

collected from all experiments in 1999 to measure total bacteria. However, we only included 

samples from one vessel for the protozoans, due to the time-intensive nature of analysis for this 

group. 

Sample processing. 

Water temperature and salinity were measured immediately, using a hand-held 

thermometer and refractometer, respectively. Plankton/Artemia samples were examined aboard 

ship initially to assess general condition (live/dead, active, lethargic) soon after collection, using 

dissecting microscopes; these samples were then preserved in 5% buffered formalin. The preserved 

plankton samples were sorted and enumerated in the laboratory as described in Chapter 3 for Fine


Analysis. Thus, densities were estimated for each taxon, and voucher samples were sent to experts 

to verify the taxonomic identity. 

The tracers in whole water samples are being quantified in the laboratory at SERC (dye 

concentration with a fluorometer, micro-spheres with direct counts under a fluorescent compound 

microscope). Total bacteria are also being estimated by direct count with a compound microscope, 

using standard techniques. The protozoan have been sent to a colleague (Dr. Richard Pierce, 

expert in ciliate protozoa) for direct counts by taxon. 

5C. Results 

The experiments were conducted on 8 different voyages, which were divided evenly 

between the two years (Table 5.1). All experiments were conducted from June to mid July, to 

control for seasonal variation and to occur during a period of high plankton abundance (see 

Chapter 3). Six of the 8 ships included all three treatments: ER exchange, FT exchange, and 

control. These were SeaRiver ships, departing from the ports of San Francisco Bay and Puget 

Sound. However, the large ARCO tankers were not able to perform ER exchange; experiments 

aboard these remaining 2 ships included only FT exchange and control tanks. 

Table 5.1. Overview and status of ballast water exchange experiments conducted aboard oil tankers arriving to 

Port Valdez, 1998-1999. For each of 8 replicate experiments: (A) The upper table indicates the vessel, start date, 

source port, exchange methods, and number collected samples (physical/chemical and biological); (B) The lower table 

indicates the status of the respective samples. Physical/chemical tracers include salinity, rhodamine, and flourescent 

microspheres (shown in lower table). Biological tracers include resident zooplankton and brine shrimp (Artemia) in 

both years, as well as total bacteria in1999 only. . Source ports: Puget Sound, WA (PS); San Francisco Bay, CA (SF); 

Long Beach, CA (LB). Exchange types: Empty-Refill (ER) and Flow-Through (FT). See text for experimental 

design. 

A. 

# of samples # of samples 

Ship Date Port Source Exchange type(s) Physical tracers Biol. Tracers 

S/R Baytown 27-Jun-98 SF ER+FT 120 60 

S/R Benicia 01-Jul-98 SF ER+FT 120 60 

S/R Long Beach 08-Jul-98 SF ER+FT 120 60 

ARCO Independence 18-Jul-98 LB FT 80 40 

S/R Baytown 11-Jun-99 PS ER+FT 60 144 

ARCO Spirit 12-Jun-99 LB FT 64 164 

S/R Baytown 08-Jul-99 PS ER+FT 60 144 

S/R Long Beach 19-Jul-99 SF ER+FT 120 252 

B. 

status of proccessing (ip=in progress) 

Ship Date Salinity Rhodamine Microspheres Zooplankton/Artemia Bacteria 

S/R Baytown 27-Jun-98 done done ip done - 

S/R Benicia 01-Jul-98 done done done done - 

S/R Long Beach 08-Jul-98 done done done done - 

ARCO Independence 18-Jul-98 done done done done - 

S/R Baytown 11-Jun-99 ip done done done ip 

ARCO Spirit 12-Jun-99 ip done done done ip 

S/R Baytown 08-Jul-99 ip done done ip ip 

S/R Long Beach 19-Jul-99 ip done done ip ip


We distributed our experiments among the three source ports to maximize the range of 

conditions (e.g., taxonomic groups, voyage duration, vessel types, and salinity), allowing us to test 

for general patterns among oil tankers. For example, diversity (and salinity) was generally greatest 

for ships from Long Beach and Puget Sound, and voyage duration differed among source ports 

(see Chapters 2 and 3). However, the traffic from each source port presented some unique 

constraints to the overall design: 

(1) Ships from Long Beach could not perform ER Exchange; 

(2) Ships from San Francisco contained low salinity waters, especially during the 1998 El Nino 

year, creating a possible physiological stress for organisms when exposed to exchange (not 

present at the other ports); 

(3) Ships from Puget Sound were not able to complete as many exchanges during the short voyage 

duration, limiting the total exchange volume to 100% or 200% (instead of the 300% possible 

for the longer voyages from other source ports). 

For all experiments on all 8 vessels, we have measured the effect of at least one full (100%) 

exchange for the exchange methods and tracers indicated in Table 5.1. In addition, for the majority 

of vessels we have also measured the effect of multiple exchange events. 

Table 5.1 also indicates the number of samples taken for each voyage and the status of 

these samples. The analysis for physical tracers is actually twice the number shown, as the same 

sample is used for analysis of rhodamine and microspheres. 

Although the samples are now at various stages of analysis (Table 5.1), our initial 

analyses suggest a significant difference between ER and FT exchange in the reduction of 

rhodamine dye. For example, Figure 5.1 shows the average change in concentration of 

rhodamine for the respective treatments across multiple exchange events in 1998. The 

concentration of dye was reduced by 80 and 99% (for FT and ER exchange, respectively) 

compared to the initial concentrations. Interestingly, the concentration of dye in the control tank 

increased between the first and second measures, and this change is attributed to inadequate 

mixing at time T o for some ships (as evidenced by vertical stratification that was present in our 

raw data). Similar patterns exist for the rhodamine data collected in 1999. 

Despite the rhodamine results, demonstrating relatively high levels of exchange, it is 

premature to draw conclusions about the efficacy of exchange to remove organisms. We are 

now analyzing the samples to measure removal rates for both biological and physical tracers 

(i.e., microspheres), and we expect to complete these analyses by June 2000. At present, it is 

evident that some taxa declined in abundance (following exchange) to the same extent as 

rhodamine dye. Figure 5.2 shows a decline in the abundance of Limnoithona sp. for both ER and 

FT exchange on one vessel. The density actually increased in the control tank, and this was due 

most likely to growth of copepodite stages instead of mixing. Changes in the abundance of other 

taxa appear to be much less striking, although analysis of the overall pattern (including variation 

among taxa) must await completion of all quantitative counts and taxonomic verification.


Figure 5.1. Effect of ballast water exchange on rhodamine dye concentrations. Data are from exchange 

experiments conducted in the ballast tanks of oil tankers arriving to Port Valdez. Shown for 1998 voyages (n=4 

vessels) is the mean percent change in rhodamine dye concentration (compared to the initial time measure) at each 

of four successive time points for 3 different treatments: Control – ballast tanks that did not undergo exchange; ER 

Exchange – ballast tanks that underwent Empty-Refill Exchange; FT Exchange – ballast tanks that underwent Flow- 

Through Exchange. See text for experimental design. 

200% 

CONTROL 

ER EXCHANGE 

FT EXCHANGE 

DYE REMAINING 

150% 

100% 

50% 

0% 

T0 T1 T2 T3 Tf 

TIME POINT 

Figure 5.2. Effect of ballast water exchange on Limnoithona sp.density. Data are from exchange experiments 

conducted in the ballast tanks of an oil tanker arriving to Port Valdez. Shown for one voyage is the mean density of 

the copepod SPECIES at each of five successive time points for 3 different treatments: Control – ballast tanks that 

did not undergo exchange; ER Exchange – ballast tanks that underwent Empty-Refill Exchange; FT Exchange – 

ballast tanks that underwent Flow-Through Exchange. See text for experimental design. 


4000 

3000 

2000 

1000 

TREATMENT 

CONTROL 

ER EXCHANGE 

FT EXCHANGE 

0 

T0 T1 T2 T3 Tf 

TIME POINT 

Changes in the density of entrained biota within the control tanks of each vessel will 

measure survivorship over time (i.e., during transit). This will allow us to test our hypothesis 

(above) about the effect of voyage duration on survivorship, and whether differences among port 

sources are due to such time-dependent survivorship. 


This is the first study to compare the relative efficiency of exchange methods (ER and FT 

exchange) for any vessel type or taxon.


Quantitative and experimental analyses of ballast water exchange have been very limited to 

date, and these can be classified into 3 general types: 

1. Comparison of ballast water in ships that have or have not exchanged ballast water. 

These data indicate that, compared to ships that have not conducted mid-ocean ballast water 

exchange, ships with exchanged ballast water have reduced abundance of plankton. However, 

with this approach, it is not possible to (a) compare directly methods of exchange (FT vs. ER) , 

(b) control for initial plankton densities or the percentage of water exchanged (as below). Thus, 

the data are highly variable and interpretation is limited (e.g., Smith et al. 1996). 

2. Comparison of ballast water in tanks of the same ship that have not exchanged ballast 

water, with measurements made only after exchange is complete and upon arrival to port. 

These data suggest a reduction of roughly 90% occurred, but interpretation is also limited with 

this design (Ruiz and Hines 1997; see below). Initial variation among tanks can be 

considerable, depending upon the timing (e.g., day vs. night) and sequence of ballasting, which 

creates potentially large differences among tanks independent of exchange treatment. 

Furthermore, it is not possible to compare efficiency between methods of exchange, or for 

multiples of exchange, because ships usually only perform one method and volume of 

exchange. 

3. Comparison of ballast water in tanks of the same ship before and after exchange of ballast 

water, with measurements made on board ship at various stages of the exchange process. 

These data provide a clear measure of efficiency within a single tank, and we have conducted 

this analysis on approximately 5 military vessels and 1 commercial bulk carrier (Ruiz et al. 

1999, Wohnam et al. 1996). However, the sample size is small (and taxa included in ships to 

date are limited), and comparison between exchange methods or multiples of exchange (on the 

same ship) has not been included or possible to date. 

The 1997 Pilot Study provided initial data comparing the end result of FT Exchange (300% 

and 100%) on plankton abundance. These data suggested that approximately 70- 90% of coastal 

plankton was removed by FT exchange, compared to control tanks from the same source. 

Interestingly, it was not clear that an increased level of exchange (100 vs. 300 %) produced a 

parallel reduction in key taxonomic groups. 

In both the Pilot Study and the current study (Chapter 3), it was evident that abundance of 

coastal organisms was 10-100 fold lower in tankers from foreign ports (that underwent ballast 

water exchange) compared to domestic arrivals (that do not undergo exchange). Although this 

difference may result from the exchange, it is confounded by differences in the initial 

concentrations (i.e., source ports) and voyage duration that can also have a strong influence. 

The results of this study – the most comprehensive and rigorous to date - will 

significantly advance our understanding of the strengths and limitations of ballast water 

exchange, providing multiple quantitative measures for the two exchange methods, both for oil 

tankers specifically but for commercial ships more generally.


Importantly, when completed, our study will also provide a set of standards for 

evaluating ballast water management in two ways. First, we have developed and tested a 

standard set of assays to measure exchange efficiency across vessel types, vessel tanks, and 

under various conditions. This will be useful in comparing efficiency among studies. Second, 

the results obtained by this and future studies will provide a benchmark against which to assess 

the efficacy of emerging technologies. 


Carlton, J.T. and J.B. Geller. 1993. Ecological roulette: The global transport of nonindigenous 


Dickman, M. and F. Zhang. 1999. Mid-ocean exchange of container vessel ballast water. 2: 

Effects of vessel type in the transport of diatoms and dinoflagellates from Manzanillo, Mexico, 

to Hong Kong, China. Mar. Ecol. Prog. Ser. 176: 253-262. 

Hallegraeff, G.M. 1998. Transport of toxic dinoflagellates via ships’ ballast water: bioeconomic 

risk assessment and efficacy of possible ballast water management strategies. Mar. Ecol. Prog. 

Ser. 168:297-309. 

National Research Council. 1996. Stemming the Tide: Controlling Introductions of 

Nonindigenous Species by Ships’ Ballast Water. National Academy Press, Washington, D.C. 




Ruiz, G.M., L. S. Godwin, J. Toft, L.D. Smith, A.H. Hines and J.T. Carlton. 1999. Ballast water 

transfer and management by U.S. Navy vessels. Final Report to the U.S. Dept. of Defense. 24pp. 

Smith, L.D., M.J. Wonham, L.D. McCann, D.M. Reid, G.M. Ruiz and J.T. Carlton. 1996. 


ballast water and sediments. Parts I and II. Final report to the U.S. Coast Guard and the U.S. 

Department of Transportation. 

Wonham, M.J., W.C. Walton, A.M. Frese and G.M. Ruiz. 1996. Transoceanic transport of 

ballast water: Biological and physical dynamics of ballasted communities and the effectiveness 

of mid-ocean exchange. Final Report to the U.S. Fish & Wildlife Service and the Compton 

Foundation. 

Zhang, F. and M. Dickman. 1999. Mid-ocean exchange of container vessel ballast water. 1: 

Seasonal factors affecting the transport of harmful diatoms and dinoflagellates. Mar. Ecol. Prog. 

Ser. 176: 243-251.

Chapt 6. Organisms in Sediments of Tanker Ballast Tanks, page 6- 1 

Chapter 6. Organisms in Sediments of Tanker Ballast Tanks 



6A. Purpose 

At certain times and source ports, appreciable quantities of bottom sediment are taken up 

by tankers during ballasting. The entrained sediment potentially includes bottom dwelling 

organisms, which may be discharged and introduced into a receiving port (Smith et al. 1996). 

Few studies of such entrained sediment exist, but our samples of bulk carriers in Chesapeake Bay 

revealed that the bottoms of ballast tanks often hold a wide variety of large crabs, fish, shrimp, as 

well as many small organisms. To determine whether tankers arriving to Prince William Sound 

transported organisms associated with sediment in the bottoms of segregated ballast water tanks, 

we sampled a subset of ships traveling between Port Valdez and west coast ports, and between 

west coast ports and Asian ports. 

6B. Methods 

During 1998-99 we supplied 13 ships with “sediment sampling kits”, which we 

developed in cooperation with the shipping agents. The ships’ mates collected core samples and 

evident organisms in the sediment during routine cleaning operations, which usually occurred on 

voyages from Valdez to west coast ports, when ballast tanks were empty and open for 

maintenance, and in Asian ports, when ships were in dry-dock. The samples were preserved in 

10% formaldehyde sea water, labeled and returned to Valdez. Samples were then sent to the 

SERC lab in Maryland for processing and identification. Subsamples of whole sediment were 

sent to Mary McGann in USGS, Menlo Park for identification of foramenifera. Remaining 

sediment was washed through a 0.5 mm mesh sieve and identified under a dissecting 

microscope. 

6C. Results 

Sediment samples of the 13 tankers contained a diverse array of taxa, including fish, 

polychaete worms, mollusks, adult crabs and other crustaceans, cnidarians, and other 

invertebrates (Fig. 6.1,Tables 6.1, 6.2). The ships averaged 2.8 taxa per ship, ranging from 0-6 

taxa, with annelid worms occurring in about 90% of the ships. The number of individuals per 

sample varied widely from 1-147 individuals, with a mean of 47 individuals. Small crustaceans 

(particularly cumaceans) were the most abundant taxa, however polychaete worms were the most 

prevalent. The sediments also contained several species of Foraminfera, including Trochammina 

hadai, an NIS that has invaded many west coast ports and is very common in San Francisco Bay 

(McGann, pers. comm.), and which is reported from Prince William Sound in samples collected 

from deep sediments following the ExxonValdez oil spill. Organisms were abundant in 

sediments taken up in both San Francisco Bay (Benicia) and in Long Beach, where the ship 

intakes are near the port bottom. Ships sampled in dry dock in Asia tended to have few 

organisms, perhaps as a result of longer voyage time across the Pacific. However, the diversity 

of higher taxonomic groups present in sediments of ballast tanks did not show any obvious 

pattern by source port.


Table 6.1. Taxa Recovered From Ballast Tank Sediments of 12 Tankers. 

Foraminifera Annelida Mollusca Crustacea Chordata Other inverts. 

Ammonia hadai Capitellidae Mytilidae Alpheidae Engraulidae Bryozoa 

Bulimina sp. Nereis sp. Nudibranchia Amphipoda Sciaenidae Sipuncula 

Elphidium sp. Oligochaeta Balanus balanoides Turbellaria 

Globigerina sp. Spionidae Calanus 

Haglophragmoides sp. Syllidae Canuellidae 

Jadammina macrescens Caridea 

Lagena sp. Cirripedia 

Rosalina globularis Crangonidae 

Trochammina hadai Cumacea 

Trochammina inflata Grapsidae 

Trochammina pacifica Harpacticoida 

Hyperiidae 

Majidae 

Ostracoda 

Tanaidacea 

Table 6.2. Presence/Absence of Taxa in Ballast Tank Sediment Samples Presented by Source Region(s). 

Source(s) Diatomacea Foraminifera Annelida Mollusca Crustacea Chordata Other Inverts n 

Korea P P P A A A A 1 

LB & Korea P P P P P P P 2 

PS A A A P P A P 1 

PS & SF P P P P P P P 1 

SF A P P P P P P 6 

SF&China A A P A P A A 1


Figure 6.1. Prevalence (A) and numbers (B) of organisms recovered from sediments of tanker ballast tanks. 

Bars indicate means for 12 tankers. 

A 

Cnidaria 

Chordata 

Diatomacea 

Other Inverts 

Foraminifera 

Mollusca 

Crustacea 

Annelida 

0 20 40 60 80 

PERCENT SHIPS 

n=12 

B 

Cnidaria 

Chordata 

Diatomacea 

Other Inverts 

Foraminifera 

Mollusca 

Annelida 

Crustacea 

NA 

NA 

0 20 40 60 80 

# OF ORGANISMS RECOVERED/SHIP 


Sediment that accumulated in the bottom of ballast tanks often contained organisms from 

a diverse array of taxa. Many of these were adults in full reproductive condition. At least one 

NIS (the foraminiferan Trochammina hadai) found in these samples appears to be established in 

Prince William Sound, although the current status of this invasion is not known (McGann, pers. 

comm. 1999). 

In future work, it would be valuable to sample sediment in ballast tanks that have 

undergone mid-ocean exchange. It is not clear if bottom-dwelling organisms will be affected by 

exchange in the same ways as planktonic organisms in the water column. 


McGann, M. 1999. Personal communication. 

Smith, L.D., M.J. Wonham, L.D. McCann, D.M. Reid, G.M. Ruiz and J.T. Carlton. 1996. 


ballast water and sediments. Parts I and II. Final report to the U.S. Coast Guard and the U.S. 

Department of Transportation.

Chapt 7. Organisms Fouling Hulls and Sea Chests, page 7- 1 

Chapter 7. Organisms Fouling Hulls and Sea Chests of Tankers 



7A. Purpose 

Historically, fouling organisms on ships have been a major source of introduced species 

(Carlton 1979a, 1979b, 1987, 1989). Modern anti-fouling paints and high ship speeds greatly 

reduce the amount of fouling today. However, fouling is often common in sea chests and at 

certain points on the bottom. We sampled tankers during routine maintenance in dry dock, 

selected to estimate the potential range of fouling and diversity of fouling organisms. 

7B. Methods 

We sampled the fouling communities of two ships in dry dock: the S/R Baytown (in San 

Francisco Bay), which had not been cleaned in dry dock for approximately 2 years; and the S/R 

Benicia (in Portland), which had been cleaned in dry dock within about 6 months. The S/R 

Baytown had remained within San Francisco Bay for several months without making an ocean 

voyage prior to haul out, providing time for further accumulation of fouling organisms. 

Representative patches of fouling communities were scraped from the bottoms of the ships 

within 6 hours of haul out and before any cleaning had commenced. The sea chests and strainers 

of the ambient water intakes were also sampled. All samples were preserved in 10% 

formaldehyde and returned to the laboratory for sorting and identification using a dissecting 

microscope. 

7C. Results 

The two ships exhibited divergent extremes in the quantity and diversity of organisms 

(Table 7.1). The ship that had not been in dry dock for approximately 2 years exhibited 

extensive fouling communities, with abundant mussels and associated worms, crustaceans, and 

sediments. At least one NIS for the west coast (the mussel Musculista senhousia) was identified 

specifically on this ship. In contrast, the ship that had been hauled recently had a relatively 

sparse number of organisms, with most of the hull completely clean of fouling communities, and 

only organisms present in the sea chest. However, even this ship had organisms that are NIS for 

the west coast (e.g., the striped bass Morone saxatilis) in its water intake strainers. 


We hypothesize that these two vessels represent the extremes in fouling communities, 

corresponding to the length of time since the last entry into dry dock for bottom cleaning. 

However, there are two other features that may contribute to these overall patterns. First, the S/R 

Baytown had been resident in San Francisco Bay for over 6 months, and may have developed an 

unusually rich fouling community. Second, the other vessel entered relatively fresh water of the 

Columbia River that may have had an adverse effect on the resident community of fouling 

organisms. To distinguish the effect of dry dock schedule on fouling community structure (from 

these other confounding variables), it would be valuable to sample more ships which differ in 

time since last haul out, but preferably sampled at the same dry dock to control for potential 

effects of different salinity. Nevertheless, both ships carried NIS, which indicates that this is an 

active mechanism of transport and introduction.


Table 7.1. Organisms from hulls and sea chests of two oil tankers in dry dock (*=NIS). 

S/R Baytown 

S/R Benicia 

Algae 

Cnidaria 

Ulva sp . Garveia franciscana * 

Diatomacea 

Mollusca, Bivalvia 

Protozoa Mytlilus sp. 

Folliculina sp. 

Crustacea, Cirripedia 

Cnidaria Balanus sp. 

Cordylophora caspia * 

Crustacea, Amphipoda 

Garveia franciscana * Corophium sp. 

Nematoda 

Pisces 

Unidentified sp. Morone saxatlis * 

Nemertea Sardinopsis sagax 

Unidentified sp. 

Polychaeta 

Neries sp . 

Ophelidae, unidentified sp. 

Polydora sp. 


Musculista senhousia * 

Mytilus sp . 

Crustacea/Copepoda 

Cyclopoida, unidentified sp. 

Harpacticoida, unidentified sp. 

Crustacea/Amphipoda 

Corophium sp. 

Gammaridae, unidentified sp. 

Crustacea/Isopoda 


Crustacea/Brachyura 


Bryozoa 

Bowerbankia 

Membrenipora 

Victorella sp. 


Carlton, J.T. 1979a. History, biogeography, and ecology of the introduced marine and estuarine 

invertebrates of the Pacific coast of North America. Ph.D. Thesis, Univ. Calif., Davis. 904 pp. 

---. 1979b. Introduced invertebrates of San Francisco Bay. Pp. 427-444 in Conomos, T. J. (ed.), 

San Francisco Bay: The Urbanized Estuary. California Academy of Sciences, San Francisco. 

---. 1987. Patterns of transoceanic marine biological invasions in the Pacific Ocean. Bull. Mar. 

Sci. 41(2): 452-465. 

---. 1989. Man’s role in changing the face of the ocean: biological invasions and the implications 

for conservation of near-shore environments. Conserv. Biol. 3(3): 265-273.


Table 7.1. Organisms from hulls and sea chests of two oil tankers in dry dock (* = NIS). 

S/R Baytown 

S/R Benicia 

Diatomacea 

Cnidaria 

Protozoa Garveia franciscana * 

Folliculina sp. 

Crustacea, Cirripedia 

Cnidaria 

Balanus sp. 

Garveia franciscana * 

Crustacea, Amphipoda 

Cordylophora caspia * 

Corophium sp. 

Hydroid – unident sp. 


Bryozoa 

Mytilus sp. 

Bowerbankia sp. 

Pisces 

Canopeum sp. Morone saxatilis * 

Victorella sp. 

Sardinopsis sagax 

Nemertea 


Nematoda 

Unidentified spp. 

Polychaeta 

Ophellidae, unidentified sp. 

Polydora sp. 

Nereis sp 

Crustacea/Copepoda 

Harpacticoida, unidentified sp. 

Cyclopoida, unidentified sp. 

Crustacea/Amphipoda 

Gammaridae, unidentified sp. 

Corophium sp. 

Crustacea/Isopoda 


Crustacea/Brachyura 



Musculista senhousia * 

Mytilus sp. 

Algae 

Ulva sp.

Chapt. 8. Summary of NIS, page 8- 1 

Chapter 8. Summary of NIS in Prince William Sound and Alaska 



Paul W. Fofonoff, Smithsonian Environmental Research Center 

8A. Purpose 

To summarize our knowledge of marine NIS in Prince William Sound specifically and 

Alaska generally, we extracted information obtained from the literature, our field surveys, focal 

taxonomic research by systematists, and analysis of existing specimens in museum and reference 

collections (see detailed reports in Chapt 9). Because prior ecological and systematic work in 

Alaska has not focused on NIS, we also wish to establish a baseline for the status of NIS in 

Alaskan waters, against which future introductions may be measured. We partitioned the species 

records into 5 categories: 

(1) Definite & probable NIS, along with particularly suspicious cryptogenic species; 

(2) Cryptogenic species; 

(3) New, undescribed species discovered by this study. 

(4) Species with range extensions into south central Alaska discovered by this study; and 

(5) Species that were reported/suspected as NIS, but which we dismissed upon further analysis. 

The sudden appearance of apparently new or undescribed species in an ecosystem is 

often a good indicator of a biological invasion. Similarly, analysis of species’ range extensions 

is an important tool in detecting NIS, which may be introduced from distant biogeographic 

provinces or from adjoining provinces. However, where the native biota is as poorly studied as 

in Alaska, it may be difficult to distinguish NIS from native species that are new to science, or 

from new records of species within their normal range. Discovery of undescribed species and 

range extensions needs to be evaluated in the context of other indicators of biological invasions, 

such as association with sites of human activities and particular transport mechanisms. Thus, 

designating species as native or NIS requires a series of graded criteria (see Methods below), but 

the origin of many species may remain unknown, i.e., “cryptogenic”. These cryptogenic species 

may be further categorized into species that have particular, suspicious attributes in some 

criteria, or species that have not received adequate research to evaluate their origin. In other 

cases, species initially may be designated or suspected as NIS, but further consideration by 

experts may refute the initial concern. 

8B. Methods 

The graded criteria (derived from J.T. Carlton, e.g., Carlton 1979a, Chapman & Carlton 

1991) 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 unknown origin (Carlton 1996). In Alaska, the marine biota in many 

groups have received little systematic and biogeographic analysis, and a large portion of species 

in these groups may be cryptogenic in origin due to lack of study without particular suspicions of 

invasive characteristics. 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. In several 

cases, we have indicated cryptogenic species that have some suspicious characteristics of NIS. 

• Paleontological - NIS are absent from fossil record even though they are present 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 studied by early trained observers. 

• Biogeographic - NIS exhibit grossly disjunct patterns of distribution (we took care to 

evaluate artifacts of the distribution of biologists/taxonomists); native 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 natural dispersal 

mechanisms and have documented human-mediated mechanisms which could effect their 

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 distribution. 

• Evolutionary/Genetic - NIS have isozyme or DNA frequencies which match distant proposed 

source populations and are significantly different from adjacent natural populations; native 

species have population genetics which blend with adjacent natural populations; cryptogenic 

species have not been studied with molecular techniques. 

We also researched all published and anecdotal reports of NIS or range extensions of species that 

we were able to find in the scientific and informed popular literature for the region. We use 

these reports interactively with our field and museum work, both to direct our field surveys and 

re-examination of existing collections, and to determine the history of suspicious species that we 

collected in the field. 

8C. Results 

A diverse array of 24 species of plants and animals has been introduced into Alaskan 

waters, with 15 of these species being recorded in Prince William Sound (Table 8.1; see also 

Species Notes below). Of these definite/probableNIS, we collected 12 species in our Focal 

Taxonomic Analyses (Chapt 9), including 5 species of algae (Ceramium sinicola, Croodactylon 

ramosum, Fucus cottoni, Macrocystis integrifolia, Codium fragile tomentosoides), 1 species of 

sponge (Cliona thosina), 1 hydroid at Homer (Garveia franciscana),1 polychaete worm 

(Heteromastus filiformis), 2 molluscs (Mya arenaria, Crassostrea gigas), 1 bryozoan 

(Schizoporella unicornis), and 1 tunicate (Botrylloides violaceus). Our findings include 7 “first


TABLE 8.1 Definite/Probable NIS for Alaska 

* Found by this project 

Species Common Name Probable AK Regions Date 1st Invasion Population Ecological References 

Region of Origin Record Status Status Impacts 

Rhodophyta 

* Ceramium sinicola a red alga NE Pacific (CA) Prince William Sound 1998 Probable Established Fouling Hansen 1998 

* Chroodactylon ramosum a red alga NW Pacific Prince William Sound 1998 Probable Established Fouling Hansen 1998 

Phaeophyta 

* Fucus cottoni (=muscoides) 

Probable Established Unknown Hansen 1998; South and 

a rockweed NE Atlantic Prince William Sound; 

Kenai Peninsula 

Tittley 1986 

* Macrocystis integrifolia a kelp NE Pacific (SE AK) Prince William Sound 1979 Definite 

Not 

reproducing NIS vector Hansen 1998 

* Microspongium globosum a brown alga NW Pacific Prince William Sound 1998 Probable Established Fouling Hansen 1998 

Sargassum muticum 

Chlorophyta 

Japanese brown 

alga 

NW Pacific SE Alaska


TABLE 8.1 continued 



Chordata- Ascidiacea 

* Botrylloides violaceus (=Botrylus 

aurantius ) 

a tunicate NW Pacific Prince William Sound 1999 Definite Established Fouling G.Lambert 1999pers.comm. 

Chordata-Osteichthyes 

Alosa sapidissima American Shad NW Atlantic N to Cook Inlet; Kodiak I. 1896 Definite Migrant Predator on Chapman 1942; McPhail 

salmonid 

fry 

and Lindsey 1986; USGS 

1999 

Dallia pectoralis (FW) Alaska Blackfish Arctic Slope (FW) Anchorage area (FW) 1950s Definite Established Predator on Morrow 1980; USGS 1999 

salmonid 

fry 

Esox lucius (FW) Northern Pike Northern N. 

America (FW) 

Salmo salar Atlantic Salmon N Atlantic 

(Anadromous) 

Salvelinus fontinalis (FW) Brook Trout Eastern N. America 

(FW) 

Anchorage area (FW) 1970s Definite Established Predator on 

salmonid 

fry 

SE Alaska-Prince William 

Sound 

1990 Definite Unknown Predator/ 

competitor 

of salmonids 

SE Alaska 1920 Definite Established Predator on 

salmonid 

fry 

Morrow 1980; USGS 1999 

Wing et al. 1992; Freeman 

1998 pers. comm.; USGS 

1999 

Morrow 1980; Alaska 

Department of Fish and 

Game 1994; USGS 1999


records” for NIS in the region. Our Rapid Community Assessment (Chapt 9) found 1 NIS 

species (the soft-shelled clam Mya arenaria ) to be widely distributed in intertidal sediments 

throughout Prince William Sound and the Kenai Peninsula. Two species (the oyster Crassostrea 

gigas, and the kelp Macrocystis integrifolia) are not established as self-sustaining, reproducing 

populations within the Sound; but these aquaculture introductions are being sustained by ongoing 

inputs that serve as a potentially important mechanism of transport for many other 

associated species. Further notes on each of these NIS are provided below. 

The literature reports 11 other NIS species, including 1 algal species (Sargassum 

muticum), 1 marsh plant (Cotula coronopifolia), 1 foraminiferan (Trochammina hadai), an 

amphipod crustacean (Jassa marmorata), 1 bryozoan (Cryptosula pallasiana), 1 brittle star 

(Ophiothrix koreana), and 5 species of fish (Alosa sapidissima, Dallia pectoralis, Esox lucius, 

Salmo salar, Salvelinus fontinalis). Several of these fish species were intentionally introduced in 

fresh water to augment fisheries, and we have included them here because they potentially have 

important impacts on native salmonid species in the region. Further notes on each of these NIS 

are provided below. 

In addition, we consider two cryptogenic species to be particularly suspicious as NIS, 

because of their new appearance at harbor areas (i.e., Homer, Cordova)(Table 8.2). These 

species include a sea star (Asterias amurensis) that is native to Alaska in the Bering Sea, but 

which has a history of invading other regions (probably via ballast water transport), and which 

appears to have suddenly extended its range to Homer in south central Alaska. Despite surveys 

of the area by good naturalists, this large animal has not been recorded at Homer/Katchemak Bay 

until now. We also discovered a new, undescribed species of ascidian (Dispalia sp. nov.) in the 

fouling communities of Homer and Cordova, but it was not present at other locations with rich 

fouling communities but lacking intense boat/ship traffic. Further notes on each of these 

suspicious species are provided below. 

A large portion of Alaskan marine species is cryptogenic in origin due to inadequate 

biogeographic and taxonomic study. However, many cryptogenic species also either exhibit wide 

distributions that may reflect global spread by early shipping traffic (Carlton 1996) or have other 

suspcicious traits of NIS (Table 8.3). During this project we collected at least 24 such species in 

Prince William Sound, and identified at least 5 others found elsewhere in Alaska. 

During our study we discovered several apparently new/undescribed species (Table 8.4) 

and documented range extensions for many other species. (Table 8.5), which also highlights the 

need for more analysis of Alaskan marine biodiversity. We found specimens of 10 apparently 

new/undescribed species in Prince William Sound, including 1 brown alga, 6 polychaete worms, 

2 molluscs, and 1 tunicate (Table 8.4). Formal species description of the tunicate species 

(Distaplia sp. nov.) is proceeding. We documented range extensions or first records for Prince 

William Sound or Cook Inlet (although some are known in the Bering Sea and further north) for 

74 species (4 algae, 11 hydrozoan cnidarians, 2 ctenophores, 24 polychaete worms, 20 molluscs, 

7 crustaceans, 2 bryozoans, 1 echinoderm, 2 tunicates, and 1 fish) (Table 8.5).


TABLE 8.2 Highly Suspicious Cryptogenic Species 

* Found by this project 



Echinodermata - Asteroidea 

* Asterias amurensis 

Asian Sea Star 

NW Pacific; Bering 

Sea 

Homer Spit 1999 Suspicious 

Range 

extension 

Established 

Predator on Baranova 1976; Ward and 

molluscs & Andrew 1995; Foster et al. 

other inverts 1999 (Chapt 9, this report) 

Chordata - Ascidiacea 

* Distaplia sp. nov. 

a tunicate unknown Homer, Prince William 

Sound (Cordova) 

1998 Suspicious 

New 

Established Fouling G.Lambert 1999 pers. comm.


TABLE 8.3 Examples of Cryptogenic Species 



Rhodophyta 

Porphyra miniata a red alga Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998 

Phaeophyta 

Demaraeaea attenuata a brown alga NW Pacific Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998 

Punctaria latifolia NE Pacific Prince William Sound 1998 Range 

Hansen 1998 

extension 

Punctaria plantaginea a brown alga Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998 

Heterokontophyta-Xanthophycaeae 

Vaucheria longicaulis a golden-brown alga NE Pacific Prince William Sound 1998 Range 

Hansen 1998 

extension, 

overlooked 

Chlorophyta 

Blidingia marginata a green alga NE Pacific Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998 

Caposiphon fulvescens a green alga NE Pacific Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998 

Halochlorococcum moorei a green alga NE Pacific Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998 

Kornmannia leptoderma non zostericola a green alga NE Pacific Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998 

Angiospermophyta 

Atriplex patula (=A. p. var. littoralis) Orach; Spearscale Eurasia SE Alaska 1883 Cryptogenic Established Unknown Meehan 1884; Hulten 1968 

Atriplex prostrata (=A. patula var. Halberd-Leaved Orach Eurasia SE Alaska Cryptogenic Established Unknown Hulten 1968 

hastata) 

Cnidaria 

Protohydra sp. a worm-like hydroid Cosmopolitan 

(CA-BC ) 

Prince William Sound Cryptogenic Established Unknown Kozloff 1987; Foster 1999 pers. 

comm.; UAF collections 

Annelida- Polychaeta 

Barantolla (americana species complex) a capitellid polychaete Circumboreal Prince William Sound 1988 Cryptogenic Established Unknown Kozloff 1987; Kudenov 1998, 

pers. comm. 

Amphitrite (cirrata species complex) a terebellid polychaete Circumboreal Prince William Sound 1998 Cryptogenic Established Unknown Kozloff 1987; Kudenov 1998, 

pers. comm. 

Capitella (capitata species complex) a capitellid polychaete Cosmopolitan Prince William Sound 1980 Cryptogenic Established Unknown Jewett 1998; Cohen and Carlton 

1995; Kudenov 1998, pers. 

comm. 

Decamastus sp. a capitellid polychaete Cosmopolitan 

(WA-BC) 

Prince William Sound 1998 Cryptogenic Established Unknown Jewett 1998; Kozloff 1987; Cohen 

and Carlton 1995 

Eteone (longa species complex) a phyllodocid polychaete Circumboreal Prince William Sound 1980 Cryptogenic Established Unknown Pettibone 1963; Kozloff 1987; 

Kudenov 1998, pers. comm. 

Eumida (sanguinea species complex) a phyllodocid polychaete Circumboreal Prince William Sound 1998 Cryptogenic Established Unknown Kozloff 1987; Kudenov 1998, 

pers. comm. 

Harmathoe (imbricata species complex) a polynoid polychaete Circumboreal Prince William Sound 1980 Cryptogenic Established Unknown Kozloff 1987; Cohen and Carlton 


comm. 

Mediomastus sp. a capitellid polychaete Cosmopolitan Prince William Sound 1988 Cryptogenic Established Unknown [Jewett 1998]; Cohen and Carlton 

1995 

Pholoe (minuta species complex) a sigalionid polychaete Circumboreal Prince William Sound 1979 Cryptogenic Established Unknown Jewett 1998; Cohen and Carlton 


comm. 

Polydora quadrilobata a spionid polychaete NE Pacific 

(British 

Columbia) 

Prince William Sound Cryptogenic Established Unknown Kozloff 1987; Foster 1999 pers. 

comm.; UAF collections





Crustacea- Copepoda 

Leimia vaga a harpactacoid copepod NW Atlantic Prince William Sound 1999 Cryptogenic Established Unknown Cordell 1999 pers. comm. 

Mollusca- Bivalvia 

Macoma balthica Baltic Clam Northern 

oceans, NW 

Atlantic cryptic 

Bryozoa 

Alcyonidium "polynoum" or "mytili" a bryozoan Unknown 

(Pacific, NW 

Atlantic) 

Alaska Pacific coast 

before 

1924 

Cryptogenic Established Likely Carlton 1979; Meehan et al. 1989; 

Cohen & Carlton 1995 

Kachemak Bay Cryptogenic Established Unknown Carlton 1979; Cohen & Carlton 

1995; Winston 1999 pers. comm. 

Callopora lineata a bryozoan Unknown Prince William Sound Cryptogenic Established Unknown Foster 1999 pers. comm.; 

Winston 1999 pers. comm. 

Celleporella hyalina a bryozoan Unknown Resurrection Bay Cryptogenic Established Unknown Foster 1999 pers. comm.; 


Cellepora craticula a bryozoan Unknown Prince William Sound Cryptogenic Established Unknown Foster 1999 pers. comm.; 


Cribilina corbicula a bryozoan Unknown Prince William Sound Cryptogenic Established Unknown Foster 1999 pers. comm.; 


Parasmittina trispinosa a bryozoan Unknown Prince William Sound Cryptogenic Established Unknown Foster 1999 pers. comm.; 


TABLE 8.4 New or Undescribed Species 

Species Common Name Probable AK Regions Date 1st Invasion References 

Region of Origin Record Status 

Phaeophyta 

Coilodesme n. sp. a brown alga NE Pacific Prince William Sound 1998 New species Hansen 1998 


Eumida sp. a phyllodocid polychaete Unknown Prince William Sound 1998 undescribed species Kudenoff 1998 pers. comm. 

Exogone sp. a syllid polychaete Unknown Prince William Sound 1998 undescribed species Kudenoff 1998 pers. comm. 

Glycera sp. a glycerid polychaete Unknown Prince William Sound 1998 undescribed species Kudenoff 1998 pers. comm. 

Nephtys sp. a nephtyid polychaete Unknown Prince William Sound 1998 undescribed species Kudenoff 1998 pers. comm. 

Polygordius sp. an archiannelid polychaete Unknown Prince William Sound 1998 undescribed species Foster 1999 pers. comm.; 

UAF collections 

Scolopos sp. an orbiniid polychaete Unknown Prince William Sound 1998 undescribed species Kudenoff 1998 pers. comm. 

Mollusca - 

Gastropoda 

*Adalaria sp 1. a nudibranch, Adalaria sp. 1 Unknown Prince William Sound 1999 undescribed species Goddard 1999 pers. comm. 

of Behrens (1991) 

Adalaria sp. 2 a nudibranch Unknown Prince William Sound 1999 Unidentified species Goddard 1999 pers. comm. 

Chordata - Asciiacea 

*Diastaplia n. sp. a tunicate Unknown Homer, Prince William 

Sound (Cordova) 

1998 New species G. Lambert 1999 pers. comm. 

* This species has been known from West coast of US for several years, but is not yet described (Goddard, 1999 pers. comm.)


TABLE 8.5 Species Range Extensions or First Records for Cook Inlet / Prince Wiliam Sound (sc AK) 



Rhodophyta 

Polysiphonia senticulosa a red alga NE Pacific Prince William Sound 1998 Range extension Hansen 1998 

Phaeophyta 

Ectocarpus acutus a brown alga NE Pacific Prince William Sound 1998 Range extension Hansen 1998 

Ectocarpus dimorphus a brown alga NE Pacific Prince William Sound 1998 Range extension Hansen 1998 

Chlorophyta 

Codium fragile spp. fragile a green alga NE Pacific Prince William Sound 1998 Range extension Hansen 1998 

Cnidaria-Hydrozoa 

Aequorea aequorea a hydromedusa NE Pacific (SE AK) Prince William Sound, 

Bering Sea north 

1999 First Record sc AK Mills, Chapt 9C2, this report 

Aequorea victoria a hydromedusa NE Pacific (SE AK) Prince William Sound 1999 First Record sc AK Mills, Chapt 9C2, this report 

Clytia gregaria (=Phialidium a hydromedusa NE Pacific (SE AK) Prince William Sound, 1999 First Record sc AK Mills, Chapt 9C2, this report 

gregarium) 


Eperetmus typus a hydromedusa NE Pacific (SE AK) Prince William Sound, 



Euphysa sp. a hydromedusa NE Pacific (SE AK) Prince William Sound, 


Gonionemus vertens a hydromedusa NE Pacific (SE AK) Prince William Sound, 


Halitholus sp. a hydromedusa NE Pacific (SE AK) Prince William Sound, 


Melicertum octocostatum a hydromedusa NE Pacific (SE AK) Prince William Sound, 


Proboscidactyla flavicirrata a hydromedusa NE Pacific (SE AK) Prince William Sound, 


Sarsia spp. a hydromedusa NE Pacific (SE AK) Prince William Sound, 


Tiaropsis multicirrata a hydromedusa NE Pacific (SE AK) Prince William Sound, 


Ctenophora: 

Bolinopsis infundibulum a ctenophore NE Pacific (SE AK) Prince William Sound, 










Pleurobrachia bachei () a ctenophore NE Pacific (SE AK) Prince William Sound, 1999 Range extension N Mills, Chapt 9C2, this report 

Dutch Harbor 


Chaetozone senticosa a cirratulid polychaete NE Pacifc Prince William Sound 1980 Range extension N Kudenov 1998, pers. comm. 

Cirratulus cirratulus a cirratulid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 

Dodecaria sp. a spionid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 

Drilonereis falcata minor a lumbrinereid polychaete NE Pacific (BC Canada) Prince William Sound 1980 Range extension N Kozloff 1987; Foster 1999 pers. comm.; 

Drilonereis minor () a lumbrinereid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 

Flabelligera mastigophora a lumbrinereid polychaete NW Pacific (Chukchi Sea) Prince William Sound 1980 Range extension S Foster 1999 pers. comm.; UAF 

Hesperonoe complanata a polynoid polychaete NE Pacific Prince William Sound 1980 Range extension S Kozloff 1987; Foster 1999 pers. comm.; 

Lumbrineris limicola a lumbrinereid polychaete NE Pacific Prince William Sound Range extension Kozloff 1987; Foster 1999 pers. comm.; 

Lumbrineris luti a lumbrinereid polychaete NE Pacific (BC Canada) Prince William Sound 1988 Range extension N Kozloff 1987; Kudenov 1998 pers. comm. 

Magelona berkleyi a magelonid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 

Magelona hobsoni a magelonid polychaete NE Pacific (BC Canada) Prince William Sound 1988 Range extension N Kozloff 1987; Foster 1999 pers. comm.; 

Magelona sacculata a magelonid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 

Mesochaetopterus taylori a chaetopterid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 

Microphthalmus sczelkowi a hesionid polychaete NE Pacific (CA) Prince William Sound 1980 Range extension N Foster 1999 pers. comm.; UAF 

Mysta barbata a hesionid polychaete NW Pacific (Chukchi Sea) Prince William Sound Range extension S Foster 1999 pers. comm.; UAF 

Onuphis (=Nothria, an onuphid polychaete NE Pacific Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 

Oriopsis sp. a sabellid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.;





Nemidia sp. a polynoid polychaete Bering Sea Prince William Sound Range extension S Foster 1999 pers. comm.; UAF 

Nemidia tamarae a polynoid polychaete Bering Sea Prince William Sound Range extension S Foster 1999 pers. comm.; UAF 

Phyllodoce medipalpa a phyllodocid polychaete NE Pacific Prince William Sound Range extension N Kozloff 1987; Kudenoff 1998 pers. comm. 

Rhynchospio gluteae a spionid polychaete Unknown Prince William Sound Range extension Kudenov 1998 pers. comm. 

Syllis (Typosyllis) harti a syllid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Foster 1999 pers. comm.; UAF 

Syllis (Typosyllis) harti a syllid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Foster 1999 pers. comm.; UAF 

Tharyx secundus a cirratulid polychaete NE Pacific (BC Canada) Prince William Sound 1980 Range extension N Kozloff 1987; Foster 1999 pers. comm.; 


Mollusca- Gastropoda- Prosobranchia 

Barleeia acuta Acute Barleysnail NE Pacific (BC Canada) Prince William Sound Range extension N Kozloff 1987; Turgeon et al. 1988; Foster 

1999 pers. comm.; UAF collections 

Mollusca- Gastropoda- Opisthobranchia 

Acanthodoris nanaimoensis Wine-Plumed Spiny Doris NE Pacific (BC Canada) Prince William Sound 1999 Range extension N Kozloff 1987; Turgeon et al. 1988; Foster 

1999 pers. comm.; UAF collections; 

Goddard 1999 pers. comm. 

Adalaria jannae Janna’s Adalaria NE Pacific (BC Canada) Prince William Sound 1999 Range extension N Kozloff 1987; Goddard 1999 pers. comm 

Adalaria sp.1 of Behrens (1991) Armed Adalaria NE Pacific (SE AK) Prince William Sound 1999 Range extension N Goddard 1999 pers. comm 

Alderia modesta Modest Alderia NE Pacific (BC Canada) Prince William Sound 1999 Range extension N Kozloff 1987; Turgeon et al. 1988; 

Goddard 1999 pers. comm 

Ancula pacifica Pacific Ancula NE Pacific (SE AK) Prince William Sound 1999 Range extension N Kozloff 1987; Turgeon et al. 1988; 


Cuthona albocrusta White-Crust Cuthona NE Pacific (BC Canada) Prince William Sound 1999 Range extension N Kozloff 1987; Turgeon et al. 1988; 


Cuthona pustulata NE Pacific (BC Canada) Homer 1999 Range extension N Kozloff 1987; Turgeon et al. 1988; 


Eubranchus olivaceus Green Balloon Aeolis NE Pacific (BC Canada) Prince William Sound 

and Cook Inlet 

Range extension N 

Kozloff 1987; Turgeon et al. 1988; Foster 

1999 pers. comm.; UAF collections; 


Geitodoris heathi Heath’s Dorid NE Pacific (SE AK) Prince William Sound 1999 Range extension N Goddard 1999 pers. comm 

Janolus fuscus NE Pacific (SE AK) Cook Inlet 1999 Range extension N Foster 1999 pers. comm.; Goddard 1999 

pers. comm 

Albatross Aglaja NE Pacific (SE AK) Prince William Sound 1980 Range extension N Kozloff 1987; Turgeon et al. 1988; Foster 


Melanochlamys diomedeum NE Pacific (SE AK) Prince William Sound 1998 Range extension N Kozloff 1987; Foster 1999 pers. comm.; 

Melanochlamys ocelliger Arctic Odostome Bering Sea Prince William 

Sound,Shumagin Is., 

Range extension S Berh 1894; Lee & Foster 1985; Foster 


Kodiak Is. 

Odostomia arctica Hansine Seaslug NE Pacific (BC Canada) Prince William Sound Range extension N Kozloff 1987; Turgeon et al. 1988; 


Olea hansineensis Banded Polycera NE Pacific (N to Hawkins 

Island, Prince William 

Sound) 

Prince William Sound 

Range extension W Foster 1999 pers. comm.; Goddard, pers. 

comm., 1999 

Palio zosterae Arctic Barrel-Bubble Bering Sea Prince William Sound Range extension S Turgeon et al. 1988; Foster 1999 pers. 

comm.; UAF collections 

Retusa obtusa 

Mollusca- Bivalvia Glacial Mussel Bering Sea Prince William Sound Range extension S Foster 1999 pers. comm.; UAF 

collections 

Musculus glacialis 

Crustacea- Copepoda Unidentified copepod NE Pacific (subtropical) Prince William Sound Range extension S Ted Cooney 1998 pers. comm. 

Unidentified Calanoid 

Crustacea- Leptostraca 

Nebalia sp. a nebaliacean NE Pacific (BC Canada) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 


Crustacea- Isopoda 

Gnathia tridens an isopod NE Pacific (CA) Prince William Sound Range extension N Foster 1999 pers. comm.; UAF 

collections





Munna chromocephala an isopod NE Pacific (WA) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 


Munna ubiquita an isopod NE Pacific (WA) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 


Pleurogonium sp. an isopod NE Pacific (WA) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 


Synodidotea ritteri an isopod NE Pacific (CA) Prince William Sound Range extension N Smith and Carlton 1975; Foster 1999 

pers. comm.; UAF collections 

Brachiopoda 

Terebratalia crossei a brachiopod NW, NE Pacific Prince William Sound Range extension N, Foster 1999 pers. comm.; UAF 

NE 

collections 

Bryozoa 

Cribilina annulata a bryozoan Bering Sea Prince William Sound Range extension S Kozloff 1987; Foster 1999 pers. comm.; 


Filicrisia smithi a bryozoan Bering Sea Prince William Sound Range extension S Foster 1999 pers. comm.; UAF 

collections 

Echinodermata-Asteroida 

Asterias amurensis Asian Sea Star NW Pacific; Bering Sea Homer Spit 1999 Range extension REFS 

Chordata- Ascidiacea 

Chelysoma columbianum a tunicate NE Pacific (BC Canada) Prince William Sound Range extension N Kozloff 1987; Foster 1999 pers. comm.; 


Halocynthia hilgendorfi igaboja a tunicate NE Pacific (BC Canada) Prince William Sound Range extension Kozloff 1987; Foster 1999 pers. comm.; 


Chordata- Osteichthyes 

Sphyraena argentea Pacific Barracuda NE Pacific (BC Canada) Prince William Sound 1998 Range extension Valdez Vanguard newspaper, 1998


We also considered several reports and specimens that were initially considered as 

possible NIS, but which we reject primarily as misidentifications of similar native species (Table 

8.6). 

TABLE 8.6 Species/Specimens Misidentified as NIS 

Putative Species Common Name Probable AK Regions Date 1st Invasion References 

Region of Origin 

Record Status 

Rhodophyta 

Porphyra redidiva a red alga NE Pacific Prince William Sound 1998 Misidentified earlier Hansen 1998 

Chlorophyta 

Monostroma fractum a green alga NE Pacific Prince William Sound 1998 Misidentification, overlooked Hansen 1998 

Angiospermophyta 

Myriophyllum spicatumEurasian Watermilfoil Eurasia SE Alaska (FW) Probable misidentification USGS 1998 

Cnidaria-Hydrozoa 

Halitholus sp. a hydromedusa NE Pacific Prince William Sound 1998 Not identifiable to species Mills, Chap9C2,this rept 

Leuckartiara sp. a hydromedusa NE Pacific Prince William Sound 1998 Not identifiable to species Mills, Chap9C2,this rept 


Anaspio boreas a spionid polychaete Gulf of Alaska Prince William Sound Uncertain identification Foster 1999 pers. 

comm.; UAF 

collections 

Polydora cf. P. 

brachycephalata 

a spionid polychaete 

NE Pacific 

(Oregon) 

Prince William Sound Misidentification Kozloff 1987; Foster 

1999 pers. comm.; 


There are few over-arching ecological traits that characterize marine NIS in Alaska. NIS 

were variable in their local distributions in the region, with distributions of most species 

apparently limited to particular sites, but with many sites having some NIS. Although NIS were 

frequently associated with harbor areas and aquaculture sites, some species (e.g., Mya arenaria) 

occurred widely wherever the appropriate habitat was present. NIS occurred in a wide range of 

habitats from coastal marshes (Cotula coronopifolia) and the high intertidal zone (Fucus 

cottonii) to deep subtidal waters (Trochammina hadai), and from variable and low salinity areas 

(Mya arenaria, Heteromastus filiformis) to stenohaline high salinities (Botryloides violaceus). 

NIS included species inhabiting hard and soft substrates. NIS also include species from a wide 

range of motility, from migratory fish to sessile plants and invertebrates; and they included a full 

range of trophic modes from autotrophs (algae) to suspension feeders to predators. While many 

of the NIS have life cycles with a dispersal stage (especially echinoderms and bivalves with 

long-lived planktonic larvae), others had little motility (e.g., sessile tunicates with short-lived 

planktonic larval stages). Thus, although NIS were most common in habitats most impacted by 

human activities, there were few sites or habitats within the region and few ecological niches that 

were immune from invasion. 

Although oil tankers transport great quantities of abundant and diverse plankton 

(including known NIS) into Prince William Sound, we have not identified any established NIS 

that is clearly attributable to introduction via tanker ballast water. However, analysis of probable 

transport mechanisms for species introductions is difficult in most regions where multiple 

transfer agents have been active. In south central Alaska, NIS were commonly found at harbor 

areas (e.g., Homer), where ballast water and hull fouling associated with cargo ships and the full 

range of fishing and other vessels are potential vectors. Bulk carriers like wood chip and log 

ships arriving in ballast to Homer, as well as tankers arriving to Port Valdez, are sources of the 

largest volumes of ballast water (Smith et al., 1999). Fishing and recreational vessels often have 

extensive fouling communities which may be transported coastwise within the region. NIS were 

also found at sites of associated with aquaculture (e.g., Tatitlek) and with fishery introductions


(e.g., Atlantic salmon). Oyster (Crassostrea gigas) culture imports spat from Washington and 

Oregon hatcheries as “clean” seed for grow-up in the field. However, associated parasitic, 

commensal and fouling organisms frequently could be transported unintentionally with the spat 

(Carlton, 1992). Similarly, transfer of kelp (Macrocystis integrifolia), however “clean” in 

appearance, from Oregon and Washington (in the past) and southeast Alaska (in the present) 

could also serve as a vector for many fouling species, epiphytes, or organisms hiding in 

holdfasts. Several species of fish have been introduced intentionally into Alaskan freshwaters, 

where they may impact salmonids at key stages of their migratory life cycle. Also, escapes of 

Atlantic salmon from pen culture have resulted in established populations of Salmo salar in 

British Columbia, as well as in increasingly frequent instances of this NIS fish being caught in 

Prince William Sound and throughout south central to southeast Alaska. 

The number of marine NIS in Alaska appears to be significantly lower than other marine 

ecosystems along the west coast of North America, where numbers of NIS range from about 50 

species in Puget Sound (Cohen et al., 1998) to 250 species in San Francisco Bay (Cohen & 

Carlton, 1995; Carlton, pers. comm.). 

Species Notes: 

RHODOPHYTA 

Ceramium sinicola- This red alga was found as an epiphyte of Codium fragile (tomentosoides) 

near Green Island. It has not been found previously north of southern California, and is strongly 

suspected of being an introduction (Hansen 1998). 

Chroodactylon ramosum- This microscopic, primitive, red alga was found growing on oyster 

floats at Tatilek. This species is previously known from Japan, Australia, and southern California 

(and the Great Lakes, where it was introduced, Mills et al. 1993) but has not been found in the 

well-studied waters of British Columbia and Washington. It may have been introduced with 

oysters (Hansen 1998). 

PHAEOPHYTA 

Fucus cottoni (=muscoides)- This brown seaweed is known from European coasts from northern 

Spain to Scandinavia (South and Tittley 1986). In the northeast Pacific, it was first found by G. 

I. Hansen on Vancouver Island in 1981, and subsequently found to be abundant in high marsh 

and mudflat areas along Prince William Sound (Hansen, 1998; Chapt 9 Hansen). Its status as a 

separate species has been questioned by Fletcher (1987), who considers this species to be an 

ecotype of F. vesiculosus adapted to marsh and mudflat habitats. Specimens from Prince 

William Sound have been sent to Esther Serrao, Portugal, who is studying the phylogenetic 

relationships of Fucus using molecular techniques. The widespread distribution of this plant in 

British Columbia and Alaska, suggests that it is not a recent introduction (Hansen 1998; Chapt. 9 

Hansen). 

Macrocystis integrifolia- This giant kelp is found from California to southeast Alaska. Since 

1979, this kelp had been transported by plane from southeast Alaska to Prince William Sound to 

be used as substrate for the Herring-Roe-on-Kelp fishery. Blades of kelp are placed in 

impoundment nets with gravid herring, which deposit their eggs on the kelp. The egg-laden


blades are then harvested and shipped to Japan, as a delicacy. Blades and holdfasts of kelp are 

commonly found in Prince William Sound, but attached plants have not been found, indicating 

that this kelp has not become established. While “clean” kelp blades are selected for the fishery, 

the practice represents a potential vector for transport of microscopic developing stages of algae 

and invertebrates into Prince William Sound (Jay Johnson, Alaska Fish and Game, pers. comm. 

to G. I. Hansen ; Hansen 1998). Our examination of several large plants including blades, stipes 

and holdfasts at Knight Island in the Sound during June 1998 showed that a variety of 

gastropods, ophiuroids, amphipods and bryozoans were present. It was not clear whether these 

associated organisms colonized the plants or were present at the time of release into the Sound. 

Microspongium globosum- This tiny brown alga is known previously from the North Atlantic 

and Japan, but it has not been found in the waters of British Columbia and Washington. It grows 

epiphytically on the cryptogenic brown alga Demaraeaea attenuata, attached to oyster floats at 

Tatilek (Hansen 1998). 

Sargassum muticum- This Japanese seaweed was first observed on the U.S. west coast in 1947, 

in Coos Bay, Oregon. By 1986, it was well established from southern California to southeast 

Alaska. It was probably transported across the Pacific on the shells of Pacific Oysters from 

Japan, and then transported along the coast by currents, shipping, and oyster transplants (Scagel 

1956; Scagel et al. 1986; Cohen and Carlton 1995; US Fish & Wildlife Service, Nonindigenous 

Aquatic Species Database 1999). 

CHLOROPHYTA 

Codium fragile (ssp. tomentosoides), Dead Man’s Fingers- The green algal species Codium 

fragile occurs on the West Coast as a species complex consisting of several unnamed subspecies, 

presumably native (Cynthia Trowbridge, pers. comm. to G. I. Hansen), as well as the introduced 

C. f. tomentosoides. The latter is native to the Northwest Pacific, and now widely introduced in 

temperate waters (Farnham 1980; Carlton and Scanlon 1985; Trowbridge 1995). On the West 

Coast, this seaweed has previously been known only from San Francisco Bay, where it was first 

collected in 1977, and probably was introduced on ship fouling (Cohen and Carlton 1995). A 

form of Codium nearly identical to C. f. tomentosoides was found in 1998, at Green Island, in 

Prince William Sound, together with a more typically native Codium. According to experts on 

the genus consulted by Hansen, both forms lie within the morphological range of the native 

populations, but molecular studies will be needed to determine their identity and relationships. 

In any event, the occurrence of Codium in Prince William Sound represents a range extension 

from southeastern Alaskan waters, and a possible introduction. 

ANGIOSPERMOPHYTA 

Cotula coronopifolia, Brass Buttons- This attractive flowering plant of the aster family is native 

to South Africa. It was first reported on the Pacific Coast in 1878, along San Francisco Bay and 

now occurs in coastal marshes from southern California to southeast Alaska (Hultén 1968; 

Cohen and Carlton 1995). Brass Buttons was probably transported in the dry ballast of ships to 

San Francisco Bay and other Pacific ports, as well as to scattered sites on the Atlantic coast of 

North America and Europe (Hultén 1968). Seeds of this plant (Cohen and Carlton 1995) are a 

favorite food of waterfowl, which may be how this species reached Alaska.


Additional flowering plants, identified by Hultén (1968) as “introduced weeds” of “waste 

places” and roadsides, probably occur at the edges of seashores and salt-to-fresh tidal marshes on 

the Pacific coast of Alaska, based on their habits and distribution elsewhere in North America. 

The following species are likely to occur in tidal marsh and shore habitats, especially disturbed 

ones: Agrostis gigantea (Redtop); Polypogon monspeliensis (Beard Grass); Puccinellia distans 

(Alkali Grass); Rumex crispus (Curly Dock); Rumex obtusifolius (Round-Leaved Dock)); Rumex 

maritimus (Golden Dock); Polygonum prolificum (Prolific Knotweed); Spergularia rubra (Sand 

Spurrey); Plantago major (English Plantain) (e.g. Fernald 1950; Gleason and Cronquist 1991; 

Cohen and Carlton 1995). Polygonum prolificum is native to eastern North America; the other 

species are of Eurasian origin (Hultén 1968). Many of these species were present on the coast of 

southeast Alaska by 1883 (Meehan 1884), and may have been introduced in ship’s ballast. 

PROTOZOA- FORAMINIFERA 

Trochammina hadai- This foraminiferan is native to Japan, and was first found in North America 

in San Francisco Bay in 1990-1993 (Cohen and Carlton 1995; McGann and Sloan 1996). It was 

subsequently found in many Pacific Coast estuaries, from San Diego Bay to Puget Sound (Cohen 

et al. 1998; McGann 1998 pers. comm.). In San Francisco Bay it forms very dense populations 

and it processes large amounts of carbon in the benthic communities throughout the estuary. T. 

hadai was also found in EVOS samples taken from deep (300 ft) water of Prince William Sound 

(McGann 1998 pers. comm.). This benthic protozoan inhabits the sediments (preferably muddy) 

of brackish-marine estuaries (Matsushita and Kitazato 1990, Kitazato and Matsuchita 1996). It 

probably has been introduced in ballast water, and was common in sediments in the ballast tanks 

of oil tankers travelling between west coast ports and Port Valdez (McGann and Sloan 1996; 

McGann 1998 pers. comm.). However, sediment samples collected from low intertidal to 

shallow subtidal zones throughout Prince William Sound during 1998-1999 did not contain T. 

hadai, so the extent of this population in the Sound remains unclear (Hines & McGann, pers. 

comm.). 

PORIFERA 

Cliona thosina- This boring sponge was originally described in 1888 using specimens on oyster 

shells from an unknown locality (possibly France or Mexico). C thosina was found boring in 

field cultured oysters (Crassostrea gigas) in Prince William Sound in 1998 (Hines, 1998; 

Ruetzler pers. comm. 1998). Its boring activities weaken oyster shells and can cause shell 

deformation, breakage and increase vulnerability to predators (such as crabs). The larval stages 

of C. thosina are short lived (1-2 days), limiting its ability to be transported in ballast water. 

Oysters cultured in the Sound arrive as “clean” spat derived from laboratory cultures in Oregon 

and Washington. However, oyster spat is not always as “clean” as the suppliers claim, and many 

associated species may be found in these types of aquaculture sources (Carlton, 1992). Cliona is 

common in oysters of the lower west coast, so it is possibly derived from these populations. 

CNIDARIA- HYDROZOA 

Garveia franciscana (Rope Grass Hydroid)- This hydroid has been found in many estuaries 

around the world, but its origin is uncertain. The Indo-Pacific and the Black--Caspian Sea basin 

have been suggested as possible native regions (Cohen and Carlton 1995; Calder 1997 pers. 

comm.) It was first described from San Francisco Bay in 1902, which was its only known 

location on the west coast of North America (Cohen and Carlton 1995), until we found it near


Homer in 1999 (Lee-Anne Henry pers. comm. 1999; Chapt 9 Fouling Communities). In other 

regions of the world, this hydroid has been an economically important fouling organism, 

adversely affecting ships, power plants and fishing gear (Simkina 1963; Andrews 1973; McLean 

1972). 

ANNELIDA- POLYCHAETA 

Heteromastus (filiformis)- This sediment-dwelling, free-burrowing polychaete, of the family 

Capitellidae, was first described from Europe, but it is now widely distributed in coastal waters 

around the world. On the west coast of North America, H. filiformis was first reported in 1936, 

from San Francisco Bay, and subsequently has been found north to British Columbia and Prince 

William Sound. Its introduction to the Pacific Coast could have occurred with Atlantic or Pacific 

oysters, or in the ballast water of ships (Carlton 1979; Cohen and Carlton 1995). Heteromastus 

“filiformis”, as with some other capitellid species, may constitute a complex of several 

morphologically similar species (Cohen and Carlton 1995). H. filiformis was collected 

commonly in Port Valdez in 1971-1972, 7 years after the 1964 earthquake that disrupted the 

benthic system, indicating that it was established well before initiation of tanker traffic to the 

Port (Feder et al. 1973). 

Lumbrineris heteropoda- This infaunal and polychaete is known from the Sahkalin and Japan, 

and from two Alaskan specimens, one from Resurrection Bay, and another from Glacier Bay. 

The wide gap between the known range and the Alaska records is suggestive of an introduction 

(Nora Foster, 1999 pers. comm.). 

MOLLUSCA- BIVALVIA 

Crassostrea gigas (Pacific Oyster; Japanese Oyster)- The Pacific Oyster was first planted in 

North American waters in 1902, in Puget Sound. By 1939, it was cultivated in Ketchikan, 

Alaska, and it is now reared in Prince William Sound, Katchemak Bay and other locations. 

Alaskan waters are too cold for natural reproduction of C. gigas, so spat must be transferred from 

southern waters (Quayle 1969; Carlton 1979;R. Piorkowski 1999 pers. comm.). 

Mya arenaria (Softshell Clam)- The Softshell Clam has a complex biogeographical history. This 

species evolved in the Pacific, in the Miocene Period, and subsequently invaded the Atlantic, but 

became extinct in the Eastern Pacific (Strasser 1999). Living populations of Mya arenaria 

remain in the Bering Sea, but on the Eastern Pacific Coast, shells of softshell clams are absent 

from subfossil deposits and shell middens, including those recently examined for M. arenaria 

(Foster, Chapt 9C7, this report). Mya arenaria was re-introduced to the Pacific Coast in San 

Francisco Bay in 1874, probably with plantings of Eastern Oysters (Crassostrea virginica). It 

was soon widely transplanted along the coast, reaching Alaska by the 1960s-1970s (Carlton 

1979). The clam has been widely established for decades in Prince William Sound and Port 

Valdez (Feder et al. 1973, Feder and Paul 1973), and was heavily impacted by benthos upheaval 

in the 1964 earthquake (Baxter 1971). 

CRUSTACEA-AMPHIPODA 

Jassa sp.; Jassa marmorata- A tube-dwelling amphipod of the genus Jassa from Prince William 

Sound was found in University of Alaska collections (Nora Foster pers. comm.). Specimens are 

being examined by John Chapman, but are not yet identified to species. Jassa marmorata, native


to the northwest Atlantic, has been widely introduced in the world’s oceans, and has been 

collected from Alaska waters (Point Slocum, Conlan 1989; Cohen and Carlton 1995; Chapman 

1998 pers. comm.). Amphipods of this genus build tubes on hard surfaces, including ship hulls, 

but also have been collected from ballast water (Cohen and Carlton 1995). 

BRYOZOA 

Cryptosula pallasiana- This bryozoan is apparently native to the Atlantic Ocean, but is now 

widely distributed in the Pacific. An early (1925) record of C. pallasiana from Homer, Alaska 

was a misidentified specimen of C. okadai, but in 1944-46 it was found in Sitka (U. S. Navy 

1951; Carlton, pers. comm.), as well as San Francisco Bay, and Newport Harbor, California. It 

was probably transported in ship fouling (Cohen and Carlton 1995). 

Schizoporella (unicornis)- This northwest Pacific bryozoan was first collected in the Eastern 

Pacific in 1927, in Puget Sound (Carlton 1979; Cohen and Carlton 1995). Its first Alaska 

collection was made between 1944 and 1949, in Kodiak (U. S. Navy 1951; Powell 1970; Dick 

and Ross 1988; Carlton, pers. comm.). Schizoporella unicornis may have been introduced in ship 

fouling or with plantings of Pacific Oysters (Cohen and Carlton 1995). In 1999, it was found in 

Tatitlek. (This form, while definitely introduced to the Pacific coast, may actually be a complex 

of several species (Winston 1999 pers. comm.). 

ECHINODERMATA- OPHIUROIDEA 

Ophiothrix koreana- A single brittlestar from Southeast Alaska (Juneau) has been tentatively 

identified as O. koreana (Kyte 1998, pers. comm.). If this identification is correct, this collection 

would be the first record of this northwest Pacific ophiuroid from the eastern Pacific. Since only 

a single specimen has been collected, the existence of established populations is unknown. Most 

brittlestars have long-lived planktonic larvae, so ballast-water transport is likeliest, but transport 

with oysters or ship fouling can not be ruled out. 

CHORDATA- ASCIDIACEA 

Botrylloides violaceus (=Botryllus aurantius)- This colonial tunicate is native to the northwest 

Pacific (Japan), and may have been first found on the West Coast in 1973, in San Francisco Bay 

(Cohen and Carlton 1995). It is now widespread, from southern California to British Columbia 

(Cohen et al. 1998; Lambert and Lambert 1998). Botrylloides violaceus was abundant on fouling 

plates in Prince William Sound in 1999 (G. Lambert 1999 pers. comm.). 

CHORDATA-OSTEICHTHYES 

Alosa sapidissima (American Shad)- This anadromous fish, native to the Atlantic coast of North 

America, was introduced in 1871, to the Sacramento River. It rapidly spread along the Pacific 

coast, and was first collected in Alaska in the Stikine River in 1896. Shad spawn in freshwater 

rivers from San Francisco Bay, north to the Columbia River, but feeding adult and juvenile fish 

wander as far north as Cook Inlet and the Kamchatka Peninusla (Chapman 1942; Cohen and 

Carlton 1995). A specimen of this species from Port Moller, Alaska Peninsula resides in the 

University of Alaska Museum collections (Foster 2000, pers. comm.). American Shad have been 

captured by seines and gill nets in Southeast Alaska during strong El Niño years, e.g., 1969 and 

1983 (J. Karinen, 2000 pers. comm.)


Dallia pectoralis (Alaska Blackfish)- This small freshwater fish is native to the North slope and 

Yukon-Kuskakwim Delta of Alaska and eastern Siberia, but it was introduced in 1950 to Hood 

and Spenard Lakes in Anchorage, in the Susitna River drainage, and has spread to other lakes in 

the vicinity (Morrow 1980). We are unaware of records of this fish in brackish or tidal waters, 

but we are including it here because of concerns of adverse impacts on Rainbow Trout 

(Oncorhynchus mykiss) and other salmonid populations in the Anchorage area (Morrow 1980). 

Esox lucius (Northern Pike)- This large predatory, freshwater gamefish is native to most of the 

glaciated regions of North America and Eurasia (Scott and Crossman 1973). In Alaska, the 

native range includes the Bering Sea drainage, and North Slope, but not Pacific watersheds. 

Northern Pike were illegally introduced to the Susitna River valley in the 1970s (Morrow 1980). 

This species is known to enter brackish waters (Scott and Crossman 1973), though we are 

unaware of estuarine occurrences in Alaska. Pike have a reputation as predators of salmonids, so 

their introduction has long been discouraged on the West Coast (Lampman 1946; Dill and 

Cordone 1997). 

Salmo salar (Atlantic Salmon)- Atlantic Salmon are native to both sides of the North Atlantic, 

and spawn in rivers of Europe and eastern North America. Many unsuccessful attempts were 

made to stock this species on the West Coast, beginning in the Sacramento River in 1874 (Dill 

and Cordone 1997), but the extensive use of S. salar in net-pen aquaculture has again raised the 

possibility of its establishment in Pacific waters. Rearing of Atlantic Salmon is illegal in 

Alaskan waters, but occurs in British Columbia and Washington (USGS, Nonindigenous Aquatic 

Species Database 1999). In Alaska, the Atlantic Salmon was first caught off Cape Cross, in 

southeastern waters, in 1990 (Wing et al. 1992). Since then, many S. salar have been caught in 

the state’s marine waters, and in 1998, the first one was caught in Alaska freshwater (Freeman 

1998 pers. comm.; USGS, Nonindigenous Aquatic Species Database 1999), and some have been 

caught in Prince William Sound with landings reported at Port Valdez and Cordova (Benda 1997 

pers. comm., Freeman 1998 pers. comm.). Many escaped cultured fishes are in poor condition 

(USGS, Nonindigenous Aquatic Species Database 1999), but successful reproduction has been 

documented on Vancouver Island (Volpe 1999), and could well occur in Alaska waters. 

Salvelinus fontinalis (Brook Trout)- This eastern North American fish was introduced into 

southeast Alaska in the 1920s, and continued to be stocked into the 1950s. In its native range, 

the Brook Trout has anadromous populations in coastal regions, from Massachusetts to Labrador 

(Morrow 1980). We have not found documentation of sea-running fish in Alaskan waters, but 

estuarine occurrences of this trout are possible. However, few populations occur in coastal lakes, 

and none are known from streams or rivers (Alaska Department of Fish and Game 1994) The 

Brook Trout may hybridize with the native Dolly Varden, but the impact of this crossing on 

native populations is unknown (Morrow 1980). 

Suspicious Species 

ECHINODERMATA- ASTEROIDEA 

Asterias amurensis (Common Asian Sea Star)- This sea star is native to the Northwest Pacifc, 

including the coasts of Japan and Russia north to the Tatarskii Inlet and the southern Kuril 

Islands, and to the Bering Sea Coasts of Russia and Alaska (Baranova 1976; Ward and Andrew


1995, Jewett and Feder 1981). Its recent appearance at Homer in Cook Inlet could represent a 

natural range extension. However, this species has long-lived planktonic larvae and could also 

be carried in ship ballast water. Asterias amurensis has successfully invaded the coast of 

Tasmania, where it poses a threat to shellfisheries (Ward and Andrew, 1995). Shipping traffic 

into Homer by bulk carriers of logs and wood chips increased markedly recently as spruce trees 

killed by a beetle outbreak have been forested. These ships probably bring large quantities of 

ballast water to Homer from Asian ports within the established of range A. amurensis. The 

recent appearance of A. amurensis in the low intertidal zone at the tip of Homer Spit was noted 

as a sudden arrival by experienced naturalists (C. & C. Field, 1999 pers. comm.), who have been 

studying the area for several years. This large conspicuous sea star would not be overlooked 

easily and was not recorded previously in the many benthic trawl surveys of the Gulf Alaska (N. 

Foster, UA Museum 1999 pers. comm.). Specimens of A. amurenis are not represented in the 

University of Alaska Museum’s collections for trawl surveys in either the Gulf of Alaska (Foster, 

1999 pers. comm.) or Cook Inlet (Feder and Paul 1981, Feder et al. 1981). The survey of Cook 

Inlet also included scuba surveys. It would have been very unusual for such collections to miss 

a large conspicuous sea star (30 cm from ray tip to ray tip). We consider this species to be 

cryptogenic in Cook Inlet, with characteristics that are very suspicious of an NIS. Because it is a 

voracious predator, it could have a major impact on benthic communities. 

ASCIDIACEA 

Distaplia n. sp.- This tunicate is a new, undescribed species (G. Lambert, 2000, Chapt 9C10, this 

report), which is very abundant in fouling communities on floats and man-made substrates in 

marinas at Homer and Cordova. It was first collected in 1998 in Homer and was found in both 

Homer and Cordova in 1999. It was not found at other sites within Prince William Sound where 

other, native species of tunicates were common in fouling communities but lack similar 

shipping/boating traffic (e.g., Tatitlek, Chenega, Port Chalmers). These sites were sampled by 

the same expert taxonomists and systematists who found the species at Homer and Cordova. 

Also, sites with and without the new species were sampled at the same times with an equivalent 

sampling effort (fouling plates, Chapt 9D, this report). Its appearance is also suspicious, because 

it was not identified in 1901 when tunicates were collected in the region at nearby sites (Ritter 

1903), but this ascidian taxonomist tended to “lump” species of Distaplia (G. Lambert, 1999 

pers. comm.). This tunicate could be a formerly rare native species that has taken advantage of 

the newly created marina habitat (G. Lambert 1999 pers. comm.), or a recent introduction. 

CRYPTOGENIC SPECIES 

PHAEOPHYTA 

Demaraeaea attenuata- This is a possible introduction from the Northwest Pacific, but G. 

Hansen treated this alga as cryptogenic, although she considered its epiphyte Ceramium sinicola 

to be a more likely introduction (Chapt 9C1, this report). 

ANGIOSPERMOPHYTA 

Atriplex patula & A. prostrata- These flowering plants commonly occur on marshes and 

beaches, but Hultén (1968) refers to them (lumped as A. patula) as an “introduced weed”. 

Botanists are divided on their status in North America, and on the East Coast they can be traced 

back to the early 1700’s. Here, we designate them “cryptogenic”.


ANNELIDA-POLYCHAETA 

Capitella capitata and other polychaete species complexes- Many cosmopolitan polychaete 

“species” are believed to represent groups of sibling species, with little morphological 

differentiation, but possibly with differing life histories and environmental adaptations. This has 

been shown for the pollution-tolerant worm Captitella capitata (Grassle and Grassle 1976), and 

is suspected for many others. Cryptic invasions by foreign sibling species could be common for 

species with planktonic larvae, especially in newly polluted harbors, where less-tolerant natives 

could be replaced by better adapted invaders. Such invasions could only be detected by genetic 

methods, or by very exacting morphological studies. 

Polydora quadrilobata- This spionid polychaete has a wide distribution, including both sides of 

the North Atlantic, the Northwest Pacific, and the Northeast Pacific coast from California to 

Puget Sound (Blake 1971; Kozloff 1987). At least 7 species of spionids have been introduced to 

the Northeast Pacific (Cohen and Carlton 1995; Cohen et al. 1998). Foster (N. Foster, UA 

Museum, 1999 pers. comm.) considers P. quadrilobata’s wide distribution to be suspicious, in 

view of the numerous introductions of this group: “The suspicious designation results from my 

perception that Spionidae do seem to make up a large proportion of the NIS listed by the Puget 

Sound expedition.” 

BRYOZOA 

Alcyonidium (polyoum) Native and introduced cryptic species are presumed to exist on the 

Pacific Coast. However, Alaska animals may be more likely to represent native forms, while 

San Francisco Bay bryozoans are more likely to be introduced (Cohen and Carlton 1995). 

MOLLUSCA- BIVALVIA 

Macoma balthica- Native and introduced cryptic species are presumed to exist on the Pacific 

Coast. However, Alaska animals may be more likely to represent the native forms (Meehan et 

al. 1989; Cohen and Carlton 1995). 

CRUSTACEA-COPEPODA 

Leimia vaga- This benthic harpacticoid copepod was first described from Nova Scotia, but has a 

limited distribution in the North Atlantic and is found in several Oregon and Washington 

estuaries (Chapt 9C5; Jeff Cordell 1999 pers. comm.). In 1999, our surveys found it in Prince 

William Sound (Chapt 9C5, this report). Its disjunct distribution is suggestive of transport 

between coasts, but the origin of this species is unknown. 


We have identified 24 species of plants and animals comprising the NIS of marine related 

ecosystems in Alaska, including 15 species recorded from Prince William Sound. In addition, 2 

cryptogenic species have highly suspicious characteristics of NIS. These species represent a 

diverse array of taxa that occupy a wide range of ecological niches and habitats, although there 

appear to be more NIS associated with boat harbors and with aquaculture activities. Several of 

these NIS are first records for Alaska. We also recorded several new, previously undescribed 

species as well as numerous range extensions for species, which probably reflect the poor level 

of study and understanding of taxonomy and biogeography in Alaskan marine ecosystems. Many 

of the Alaskan NIS have larval stages which could be transported in ballast water; however,


other vectors (including intentional and incidental release for fisheries and aquaculture) are 

obvious possibilities. None are clearly associated with ballast water of oil tankers as a primary 

mechanism, even though many NIS are frequently found in ballast water arriving to Port Valdez 

(see Chapt 3, this report). The number of marine NIS in Alaska appears to be significantly lower 

than other marine ecosystems along the west coast of North America, where numbers of known 

NIS range from about 50 to 250 species. The complexities and uncertainties of the native biota 

and the history of vectors in the south-central Alaskan region will inevitably result in an evolving 

analysis, typically revealing previously hidden importance and impacts of NIS. 


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Chapt 9A. Overview of NIS Surveys, page 9A- 1 

Chapter 9A. Overview of NIS Surveys 



9A1. Purpose 

A central goal of this project was to determine whether NIS have been, or are becoming, 

established within Prince William Sound. Because of the diversity of potential vectors and 

changing patterns of transport mechanisms, our purpose was to provide as broad and 

comprehensive a search for NIS as possible with our resources. Since previous ecological and 

systematic work in Alaska has not focused on NIS, we also wished to provide a baseline for the 

status of NIS in Alaskan waters, against which future introductions could be measured. Recent 

and on-going work on NIS along the temperate west coast of North America indicates that many 

invasive species appear to be spreading northward from a peak of NIS diversity in San Francisco 

Bay, as well as other highly invaded source ports for ballast water arriving to Prince William 

Sound. Some of these NIS (e.g., Carcinus maenas) have moved rapidly from central California 

to Washington and British Columbia, and may be expected to reach Alaskan waters in coming 

years. We focused on Prince William Sound for our field surveys and analysis of existing 

samples; but because NIS often spread coast-wise, we also sampled ports on the adjacent Kenai 

Peninsula, and we considered scientific reports broadly from Alaskan waters. 

9A2. Approach 

To detect recent or well-established NIS in Port Valdez / Prince William Sound and 

adjoining areas of risk for invasion, we used several methods, including: 

• Rapid assessment field surveys of estuarine and marine invertebrates and plants for Port 

Valdez, Prince William Sound, Seward and Homer. The objective was for experienced 

general ecologists to survey major habitats and communities, especially for large NIS plants 

and animals detectable in the field by experienced naturalists. 

• Focal taxonomic field collections in Prince William Sound, Seward and Homer. The 

objective was for taxonomic experts to sample and analyze key taxonomic groups that have 

known NIS but which are difficult for generalists to identify, providing definitive 

identification and careful, authoritative assessments of the native, invasive and cryptogenic 

status. Whenever possible, we also wished the taxonomic experts to have the opportunity to 

sample the sites using their specialized methods and knowledge for collecting the focal 

taxon. 

• Fouling plate surveys in Prince William Sound, Seward and Homer. The objective was to 

provide a replicated standard sampling method of assay for NIS in a community that is prone 

to invasions, but which has received little prior ecological analysis in Alaska. 

• Re-examination of museum and reference collections for Prince William Sound. The 

objective was to re-examine extensive collections already available in the University of 

Alaska Museum and vouchers samples from Exxon Valdez Oil Spill (EVOS) and other 

ecological studies, developing a screening method of screening for potential NIS. 

Our approach of utilizing this array of methods served to maximize spatial, temporal, taxonomic, 

and habitat coverage, while still focusing our limited resources upon elements known to be of 

highest risk of invasion. The field surveys provided broad coverage of Prince William Sound, as


will as Anchorage, Homer and Seward as important ports in neighboring Cook Inlet and the 

Kenai Peninsula (Fig. 9.A.1).


In any ecosystem, the ability to detect established NIS varies in space and time, because 

most aspects of species distribution and abundance are probabilistic. Like any population, NIS 

populations are typically patchily distributed across the full range of habitats, and may undergo 

marked seasonal and annual fluctuations. In high latitude ecosystems like Prince William 

Sound, this variation is extremely pronounced, due to rapid changes in photoperiod and 

fluctuations in surface salinities resulting from warm season runoff. By using an array of 

sampling at several times throughout the growing season, we increased the probability of 

detecting NIS. 

In addition to the stochastic aspect of detecting NIS, detection of NIS in Prince William 

Sound is difficult because the biota is not well described by taxonomists and biogeographers. 

Despite the extensive sampling in Prince William Sound for EVOS and other ecological 

programs, there are no comprehensive keys or field guides to the biota of the region (a notable 

exception is the guide to Alaskan molluscs; Foster 1991). In our Pilot Study (Ruiz & Hines 

1997) of Port Valdez alone, we found a surprisingly high percentage of new species records, and 

our literature analysis showed that 20-50% of the species are cryptogenic in origin. 

It was necessary to sample several habitats of Prince William Sound that have received 

little scientific study. For example, there are almost no publications on the fouling community of 

floats and pilings, yet NIS are very common in this habitat in west coast source ports, where 

some of these NIS have been very destructive. Soft-bottom habitats of Prince William Sound 

have received less study than rocky substrates. 

Also, in much of the previous work associated with the Exxon Valdez Oil Spill, 

taxonomic identifications were not carried out to species but were reported at relatively high 

taxonomic levels (e.g., Family, Order). Without careful identification to species by expert 

taxonomists, NIS are often confused with similar native species. Therefore, we brought 

systematic experts for several focal taxonomic groups to Alaska for field collections or 

contracted them to identify selected subsets of samples. 

9A3. Site selection and sampling design 

In addition to a survey of Port Valdez and Sawmill Bay during our Pilot Study in 

June 1997, we conducted two broad expeditions to survey Prince William Sound for NIS during 

low tide series of 20-28 June 1998 and 8-16 August 1999. Fouling Plate Surveys were 

conducted during 7-17 September 1998 and in 8-16 August 1999. 

The surveys in September 1998 and August 1999 included sampling stations at Homer 

and Cordova on the Kenai Peninsula adjacent to Prince William Sound. The survey sampling 

design focused on invertebrates and plants in a variety of habitats of shallow water, the intertidal 

zone, and accessible man-made surfaces (e.g., floats, pilings, buoys). We selected sampling sites 

that were judged to be most susceptible to invasion by NIS: 

• areas most likely to be in the path of ballast water discharged from tankers (as estimated by 

circulation models and the path of the Exxon Valdez Oil Spill); 

• ports and sites of sustained disturbance by human activities (especially Port Valdez, Cordova, 

and Whittier, as well as Homer and Seward); 

• habitats associated with previously reported NIS and cryptogenic species;


• warmer water areas; 

• marinas, floats, buoys, and pilings with accessible fouling communities; and 

• sites of active aquaculture for the Japanese Oyster Crassostrea gigas. 

Together, these habitats comprise a broad area of the shallow, nearshore margins and islands of 

Prince William Sound. The survey attempted to gain broad coverage of these habitats, including 

46 sites in June 1998, 9 sites for fouling plates in September 1998, and 33 sites including a 

subset of 7sites for fouling plates in August 1999, which were spread throughout the major 

regions of the Sound and the port sites of the Kenai Peninsula (see map Fig. 9A.1, Table 9A.1). 

The survey sampled three major habitats: intertidal and shallow rocky substrates; intertidal and 

shallow soft sediments; and fouling communities on floats, buoys, pilings and oyster culture 

structures. The salinity of the array of sites ranged from fresh to fully marine areas of the Sound. 

The rapid assessment survey methods were similar to those employed in NIS surveys of 

San Francisco Bay (Cohen & Carlton 1995) and Puget Sound (Cohen et al. 1998). The approach 

utilizes a team of experienced naturalists and general ecologists to sample as great a diversity of 

organisms at as broad an array of sites as possible within the region of concern. The team for the 

surveys consisted of: 

• John Chapman (OSU), general NIS of northeast Pacific and peracaridan crustacea; 

• Nora Foster (UAF), marine invertebrates of Alaska, especially mollusca; 

• Anson Hines (SERC), barnacles and decapod crustaceans; and 

• Todd Miller (Hatfield Marine Science Center), technical assistance and peracaridan 

crustaceans. 

The survey teams utilized a variety of transportation modes to travel among sites 

throughout Prince William Sound, including vans, ferries, float planes, small boats, vessels of 

Stan Stephens Tours, and the Fishing Vessel Kristina. Following collecting, samples were 

processed in temporary laboratories provided by USF&WS Refuge in Homer, the Seward 

Marine Science Center, University of Alaska in Seward, Prince William Science Center in 

Cordova, the SERC Invasions Biology Laboratory in Valdez, the Prince William Sound 

Community College in Valdez, and several hotels. The F/V Kristina also served as laboratory 

platform for processing of samples while in transit among some of the sites. For each sampling 

site, the diversity and relative abundance of species were recorded. Field notes included GPS 

readings, sketches of the sites, salinity and temperature readings, and notes on common or 

abundant species identified in the field. Samples were collected by hand, by scraper to remove 

fouling organisms, and by trowel to collect soft sediment. Samples of sediment, algalinvertebrate 

turf, and scrapings of fouling communities were washed and sieved on 5mm, 1mm 

and 0.5 mm mesh. Each sample was “rough sorted” immediately after collection to aid in 

identification of large or delicate specimens and to preserve voucher specimens for subsequent 

work-up in the laboratory. Voucher samples were preserved in either 70% EtOH or 10% 

formalin (as appropriate to the type of organism). Voucher samples from the survey have been 

distributed to appropriate taxonomic experts for definitive identification in the laboratory using 

microscopes.


Map 

No. 

TABLE 9A.1. Collecting Sites for NIS Surveys 

(1997,1998,1999). 

Site Station 98 Cruise 

Station 

No. 

Latitude Longitude 1997 

Pilot 

Study 

Rapid Community 

Assessments 

1998 

Surveys 

1999 

Survey 

Fouling Plate 

Surveys 

1998 1999 

1 Anchorage Port Anchorage 61º14'N 149º45'W X 

Westchester Lagoon 61º13'N 149º50'W X 

2 Potter Potter flats 61º05'N 149º40'W X 

3 Katchemak Bay Homer small boat harbor X X X 

Homer spit X X X 

Homer spit mudflat X X X 

4 Sadie Cove X 

5 Seward Seward small boat harbor X X X 

Lowell Point 

X 

6 Whittier Whittier small boat harbor 60º46'37"N 148º41'24"W X X X 

Whittier ferry dock 60º46'25"N 148º40'55"W X X 

7 Shotgun Cove Shotgun Cove 60º47'26"N 148º32'30"W X 

8 Esther Island Lake Bay buoy 28 60º47'37"N 148º05'01"W X 

Lake Bay oysters 30 60º48'00"N 148º05'24"W X 

9 Squaw Bay Squaw Bay oysters 36 60º50'00"N 147º49'20"W X 

10 Eaglek Bay Eaglek Bay oysters 37 60º51'00"N 147º45'36"W X 

11 Fairmont Bay Fairmont Bay oysters X X 

12 Growler Island Growler mudflat 38 60º54'15"N 147º07'48"W X X 

Growler dock 39 60º54'13"N 147º07'48"W X X 

13 Valdez Arm, Sawmill Bay Sawmill Bay shore 9 61º03'15"N 146º47'24"W X X 

Sawmill Bay mudflat 10 61º03'23"N 146º47'24"W X X 

Navigation buoy 40 61º03'16"N 146º41'39"W X 

14 Valdez Arm Potato Point X 

15 Port Valdez Anderson Bay X 

16 Valdez Duck Flats low intertidal 44 61º07'28"N 146º18'00"W X X 

Duck Flats high intertidal 45 61º08''24"N 146º19'30"W X X 

Floating cargo dock 43 61º07'25"N 14618'36 "W X X 

SERVS dock 61º07'25"N 146º21'15"W X X X 

Small boat harbor 46 61º07'25"N 146º21'15"W X X X X X 

USCG dock " " X X 

Ferry dock " " X 

17 Port Valdez, Dayville flats Dayville flats 4 61º04'54"N 146º19'00"W X X 

18 Valdez Marine Terminal Alyeska small boat ramp 1 61º05'12N 146º23'30"W X X 

Alyeska small boat harbor 2 61º05'10"N 146º22'28"W X X 

Alyeska entrance 3 61º05'10"N 146º21'55"W 

Terminal floats 42 61º05'20"N 146º24'09"W X X


Map 

No. 

TABLE 9A.1. (continued) Collecting Sites for NIS 

Surveys (1997,1998,1999). 

Site Station 98 Cruise 

Station 

No. 

Latitude Longitude 1997 

Pilot 

Study 

Rapid Community 

Assessments 

1998 

Surveys 

1999 

Survey 

Fouling Plate 

Surveys 

1998 1999 

19 Valdez Arm, Rockey Point Rockey Point 11 60º57'36"N 146º45'36"W X 

20 Busby Island South reef 5 60º52'55"N 146º46'29"W X 

Busby Island 6 60º52'54"N 146º46'24"W X 

21 Tatitlek Tatitlek Narrows oysters 8,12 60º52'06"N 146º43'30"W X X X 

Village dock 60º52'06"N 146º43'30"W X X X 

Ferry dock 60º52'06"N 146º43'28"W X X 

22 Bligh Island Cloudman Bay 7 60º50'11"N 146º43'15"W X 

23 Sheep Bay Upper Sheep Bay 13 60º52'12"N 146º43'48"W X 

Middle Sheep Bay 14 60º40'21"N 145º57'06"W X 

24 Cordova Small boat harbor 15,19 60º32'30"N 145º46'28"W X X X 

Mudflat S of Small Boat Harbor 18 60º32'28"N 145º46'28"W X X X 

Marine Science Center 17 60º32'48"N 145º46'27"W X X 

Fish Dock & Flats 16 60º32'27"N 145º46'26"W X 

Ferry dock 60º32'27"N 145º46'24"W X 

25 Hawkins Island Windy Bay 20 60º33'54"N 145º58'38"W X 

26 Orca Bay Channel Buoy 21 60º32'22"N 146º55'55"W X 

27 Hinchinbrook Island Constantine Harbor X 

28 Montague Island Port Chalmers X X 

29 Green Island Green Island 22 60º18'19"N 145º58'38"W X 

30 Evans Island Sawmill Bay 23 60º03'31"N 147º59'47"W X 

Port San Juan 24 60º04'2"N 148º03'36"W X 

31 Chenega Island Chenega dock X X 

32 Eleanor Island Northwest Bay middle arm 25 60º32'57"N 147º34'48"W X 

33 Knight Island Passage Knight Island Passage buoy 26 60º33'52"N 147º49'11W X 

34 Main Bay Main Bay 27 60º31'58"N 148º04'41"W X 

Main Bay fish hatchery 28 60º31'16"N 148º05'35"W X


The experts for the Focal Taxonomic Collections included: 

• Gayle Hansen (Hatfield Marine Science Center, Oregon State University), phycologist with 

special expertise in Alaskan macroalgae; 

• John Chapman (Hatfield Marine Science Center, Oregon State University), peracarid 

crustacea; 

• Jeff Cordell (School of Fisheries, University of Washington), copepod crustaceans; 

• Nora Foster (University of Alaska Museum), molluscs and other marine invertebrates of 

Alaska; 

• Jeffery Goddard (University of California, Santa Barbara), opisthobranch molluscs 

• Jerry Kudenov (University of Alaska, Anchorage), polychaete worms; 

• Gretchen Lambert (Friday Harbor Laboratories, University of Washington), ascidians; 

• Charles Lambert (Friday Harbor Laboratories, University of Washington), ascidians; 

• Claudia Mills (Friday Harbor Laboratories, University of Washington), cnidarian medusae 

and ctenophora; 

• Lise Schickel (University of California, Santa Barbara), decapod crustaceans and parasitic 

crustaceans; 

• Anson Hines (SERC), barnacles and decapod crustaceans; 

• Judith Winston (Virginia Natural History Musem), bryozoans; and 

• Lea Ann Henry (University of Toronto), hydrozoan cnidarians. 

At numerous locations in Port Valdez / Prince William Sound, sediment samples were collected 

for identifications of foraminiferans, particularly the Asian NIS Trochammina hadai, which is 

extensively introduced in San Francisco Bay and other west coast source ports. This NIS was 

reported from deep-water samples taken for the Exxon Valdez Oil Spill study. Samples were 

processed and sent to Mary McGann of US Geological Survey, Menlo Park, CA, for 

identification. 

Other details of the methods are provided below within the individual chapters for each 

Focal Taxonomic Collection, the Fouling Community Analysis and the Re-examination of the 

Museum, Reference and Voucher Collections. 

9A4. References 






J. Cordell, L. Harris, T. Klinger, A. Kohn, C. Lambert, G. Lambert, K. Li, D. Secord, and J. Toft. 

1998. Puget Sound expedition: A rapid assessment survey of non-indigenous species in the 

shallow waters of Puget Sound. Washington State Department of Natural Resources. Olympia, 

Washington. 

Foster, N.R. 1991. Intertidal Bivalves: A Guide to the Common Marine Bivalves of Alaska. 

University of Alaska Press, Fairbanks. 152 pp.




Citizens’ Advisory Council. 80pp

Chapt 9B. NIS Surveys: Rapid Community Assessments, page 9B- 1 

Chapter 9B. Rapid Community Assessment 

John Chapman, Hatfield Marine Science Center, Oregon State University 

Collections of some taxonomic groups from the 1998 survey were transferred to 

systematic experts for focal taxonomic analysis provided in other sections of this report- see 

sections below for Focal Taxonomic Collections. 

Other data and analysis to be provided by John Chapman have not been forthcoming.

Chapt 9C1. Marine Plants, page 9C1- 1 

Chapter 9C1. Focal Taxonomic Collections: Marine Plants in Prince William Sound, 

Alaska 

Gayle I. Hansen, Hatfield Marine Science Center, Oregon State University 

Background 

Several NIS marine plants with potential for invasion of Alaskan waters have been 

reported on the west coast of North America. For example, the pervasive algae Sargassum 

muticum, Lomentaria hakodatensis, and the Japanese eelgrass Zostera japonica are thought to 

have been introduced with the aquaculture of oysters by the importation of spat from Japan. At 

least 5 oyster farms occur in Prince William Sound, and all have imported spat. For the herringroe-on-kelp 

(HROK) pound fishery, the giant kelp Macrocystis integrifolia is transported to 

Prince William Sound via plane from southeast Alaska (the northern limit of this species) to be 

used as a substrate for herring roe. Although the giant kelp cannot recruit in Prince William 

Sound, it seems likely that other species, accidentally co-transported with Macrocystis, could 

become established. Our Pilot Study (Ruiz and Hines 1997) also considered several NIS algal 

species reported from Alaskan waters, including a report of a cosmopolitan species Codium 

fragile tomentasoides from Green Island. 

Methods 

Sample Period. Marine benthic algae, seagrasses, and intertidal lichens were sampled as a part 

of the cruise aboard the F/V Kristina during 20-28 June 1998, described above for invertebrates. 

Site Information. A subset of 19 of the 46 sites selected for invertebrate sampling were chosen 

for the plant study, including 13 intertidal sites (4 within Port Valdez and 9 in Prince William 

Sound) and 6 off-shore float sites. Site abbreviations (for tables and figures to follow), 

coordinates, temperature, and salinity are given in Table 9C1.1. Please note that the site numbers 

in Table 9C1.1 for plants do not correspond to the site numbers on the map (Fig. 9A2) or Table 

9A1 for invertebrates. The substratum types, listed for each site, are only those sampled for 

algae and seagrasses. For analysis and discussion, the 19 plant sites have been grouped into 5 

basic habitat types: harbors, mud bays, rocky headlands and reefs, rocky bays, and floats. These 

will be discussed in greater detail in the Results section below. 

Surveying Techniques. At each site, intertidal areas accessible by foot within the time period 

provided were sampled. Since introduced species could potentially occur in any of the marine 

plant taxonomic groups, it was important to sample the entire range of species present from as 

broad an area as possible. Marine algal populations are well-known for being extremely patchy 

in distribution, caused primarily by narrow species requirements (and tolerances) for substratum, 

tidal height (exposure), salinity, nutrients, and sunlight. Since the species were patchily 

distributed, they were encountered and collected sporadically, not uniformly over time. For this 

study, abundance was noted only when unusually large patches of a particular species were 

encountered; it was not documented uniformly for entire sites. 

Time Allotment. As shown in Table 9C1.s, sampling times at major sites varied from 10 

minutes to 2 hours. At low diversity sites, such as Cloudman Bay, the time provided was 

sufficient for complete algal collection; at other sites, such as Green Island, the time was often


TABLE 9C1.1. General Site Information, 1998 Collections* 

ABBR. DATE LOCATION LAT LON SUBSTR. T SAL 

(ºC) (0/00) 

Port Valdez (Val) 

al-sbr Jun 20 Alyeska, small boat ramp, 61º 05' 12''N 146º 23' 30"W br, co 9 0 

Port Valdez 

al-pil Jun 20 Alyeska, small boat harbor, 61º 05' 10"N 146º 22' 28"W pi 9 0 

Port Valdez 

sough Jun 20 Slough, near Alyeska gate, 61º 04' 54"N 146º 19' 00"W mu 11 0 

Port Valdez 

duckflat Jun 28 Duckflat, Port Valdez 61º 07' 28''N 146º 18' 00''W mf 17-22 0-4 

Other Harbors 

Cor Jun 23 Cordova, Orca Inlet 60º 32' 28''N 145º 46' 28''W dm, mf 10-16 5-28 

Whit Jun 26 Whittier, Passage Canal 60º 46' 25''N 148º 40' 55''W dm, bo, gr, mu 4-10 8-14 

Mud/Cobble Bays 

CB Jun 21 Cloudman Bay, Bligh I. 60º 50' 11''N 146º 43' 15''W mu, co 10 3 

SMB Jun 22 Sawmill Bay, Valdez Arm 61º 03' 15''N 146º 47' 24''W mu, co 8 5 

Gro Jun 27 Growler I. 60º 54' 15''N 147º 07' 48''W mu, co 22 11 

Rk Headlands and Reefs 

RP Jun 22 Rocky Point, Valdez Arm 60º 57' 36''N 146º 45' 36''W br, co 11 15 

Bus Jun 21 Busby I., south reef 60º 52' 55''N 146º 46' 29''W br, co 11 23 

Green Jun 24 Green I., northwest reef 60º 18' 19''N 147º 23' 47''W bo, br 11 30 

Rk Bays 

NW Jun 25 Northwest Bay, middle arm, 60º 32' 57"N 147º 34' 48''W gr, co 12 10-27 

Eleanor Island 

Floats and Buoys 

TAT Jun 22 Tatitlek Narrows, Bligh I. 60º 52' 12''N 146º 43' 48''W oy 12 26 

WBF Jun 23 Windy Bay, Hawkins I. 60º 33' 54''N 145º 58' 38''W oy 14 28 

MBF Jun 25 Main Bay 60º 31' 58''N 148º 04' 41''W bb 14 19-20 

EIF Jun 25 Lake Bay, Esther I. 60º 48' 00''N 148º 05' 24''W bb 12 16 

SBF Jun 26 Squaw Bay 60º 50' 00''N 147º 49' 20''W oy 14 24 

EBF Jun 26 Eaglek Bay 60º 51' 00''N 147º 45' 36''W oy 14 24 

Abbreviations: 

*= coordinates, temperature, and salinity provided by T. Miller lon=longitude 

abbr.=abbreviations 

mf=mudflat 

bb=barrier buoy 

mu=mud 

bo=boulders 

oy=oysterfloats 

br=bedrock 

pi=wood pilings 

co=cobble 

rk=rocky 

dm=docks/marina 

sal=salinity 

gr=gravel 

subst=substratum 

lat=latitude 

t=temperature 

seriously inadequate to sample fully the algal diversity present. Differences among sites in 

amount of time for collecting were not factored out or corrected after sampling was completed; 

however, the sites which were judged to be undercollected are designated with an “*” in Table 

9C1.2. 

TABLE 9C1.2. Collection Efficiency Records 

Data Type 

Major Collection Sites (without off-shore floats) 

Val al-sbr al-pil slough duckflat Cor Whit CB SMB Gro RP Bus Green NW 

New Records 5 1 1 1 4 5 2 2 5 4 2 2 6 1 

Total Species 47 35 7 7 24 41 56 10 45 63 61 59 71 69 

Collection Time 165 40* 10 15 100 120 120 30 45* 105 70* 75* 65* 136 

Correlation R R 2 * = undercollected sites 

Total Species:Time** 0.896 43% ** = both without Val (Total Valdez) included 

New Records:Time** 0.498 7%


Field Sampling & Processing. All algal sampling was done by hand or with a chisel. Collected 

specimens were then placed in plastic bags for transport back to the boat for processing. On 

board, the samples were sorted to species and then either pressed in a plant press or preserved in 

5% formalin/seawater. Site notes and preliminary species lists were made in the field, and some 

final identifications were done on board. However, for most species final determinations were 

not made until returning to the Hatfield Marine Science Center in Newport, Oregon, where 

compound microscopy was available. The smaller marine algae that require microscopic 

examination while still alive for identification were necessarily excluded from this study. After 

identification, both liquid and dried specimens were curated, labeled, and deposited in the 

herbarium at OSU/HMSC for reference in future Alaskan marine algal studies. 

Identifications. Since no marine flora (identification guide) of Prince William Sound exists, the 

algae collected during this study were identified using a wide variety of literature. For common 

species, the most important references utilized were Abbott and Hollenberg (1976), Gabrielson et 

al. (1993), Perestenko (1994), and Sears (1998). For more obscure species, much of the world 

taxonomic literature on temperate/arctic marine algae was employed. To confirm the 

identification of particularly difficult or important species, some specimens were sent out to 

colleagues for identification using molecular techniques. These taxa are designated with a "#" in 

the species charts. Due to the costs of these tests, these results will not be presented here, but 

instead will be presented at a later date as part of the papers prepared by these experts. 

Distributions, Residency Status, and New Records. In determining if species were introduced, 

the local and global distributions had to be determined from the literature. Some of the 

references used for this process were: Scagel et al. (1993), Sears et al.(1998), Selivanova and 

Zhigadlova (1997), Lee (1980), Guiry (1998), Rueness (1977), Phillips and Menez (1988), 

Yoshida et al. (1995), Adams (1983, 1994), Womersley (1984, 1987, 1994, 1996), Lindstrom 

(1977), Hansen et al. (1981), and Hansen (1997). These distributions, summarized in the 

abbreviated form explained below, are shown under range (Ra) in the first column of the species 

site lists (Tables 9C1.3 – 9C1.5). The ranges provided the basis for determining the Residency 

Status (St) of the species. Residency status rankings include the following 5 categories: 

• E (Endemic) = species known only from Alaska 

• N (Native) = species native to the North Pacific, including species with ranges limited to the 

northeast Pacific (nep) and those that occur in all other areas around the northern Pacific rim 

(np). 

• C (Cryptogenic) = species with extremely broad distributions that occur circumboreally (cb) 

and/or extend to the southern hemisphere (ws). 

• I (Introduced) = species that appear to have been introduced to the area. 

• F (Failed Introduction) = deliberately introduced species that have failed to colonize the 

area


TABLE 9C1.3. Marine and Estuarine Plants Collected in Port Valdez, Alaska 

NIS ANALYSIS 

PORT VALDEZ 

TAXA 

…………..June 1998 Collections……..…. Total 

Ra St NR So Val al-sbr al-pil slough duckflat Checklist 

RHODOPHYTA, Rhodophyceae 

ws C Ahnfeltia fastigiata O 

nep N Ahnfeltiopsis gigartinoides O 

nep N Antithamnionella pacifica O 

ws C Audouinella purpurea O 

ws C Bangia atropurpurea X X X 

nep N# Ceramium gardneri O 

np N Constantinea subulifera O 

np N Corallina frondescens O 

np N Corallina vancouveriensis O 

np N Cryptonemia borealis O 

np N Cryptonemia obovata O 

nep N Cryptosiphonia woodii X* X* O 

cb C Devaleraea ramentacea X X O 

cb C Dumontia contorta O 

np N Dumontia simplex O 

nep N Endocladia muricata O 

ws C Erythrotrichia carnea O 

np N Glioipeltis furcata X* X* O 

np,ar N Halosaccion firmum X X O 

np N Halosaccion glandiforme X* X* O 

ws C Hildenbrandia rubra O 

np N Leachiella pacifica O 

np N Lithophyllum dispar O 

nep N Mastocarpus papillatus complex X* X* O 

np N Mastocarpus cf. pacificus X* X* X 

nep N Mazzaella heterocarpa O 

np N Mazzaella phyllocarpa X X X 

nep N Mazzaella splendens O 

nep N Microcladia borealis O 

ws C Nemalion helminthoides O 

np N Neorhodomela aculeata X X O 

np N Neorhodomela larix X X O 

np N Neorhodomela oregona X* X* O 

nep N Odonthalia floccosa O 

np N Odonthalia kamtschatica O 

np N Odonthalia setacea (drift) O 

nep N Palmaria hecatensis X* X* O 

cb C Palmaria mollis/palmata O 

np N Phycodrys riggii O 

ws C Polysiphonia brodiaei O 

nep N Polysiphonia hendryi v. deliquescens X X O 

nep N Polysiphonia hendryi v. hendryi O 

nep N Polysiphonia hendryi v. luxurians O 

nep N Polysiphonia pacifica v. pacifica O 

nep N Porphyra cuneiformis O 

nep N Porphyra mumfordii O 

np N Porphyra perforata O 

cb C# NR NAT Porphyra purpureo-violacea O


Table 9C1.3. continued 

nep N NR Wa Porphyra rediviva X* X* X 

np N Pterosiphonia bipinnata X* X* O 

np N Ptilota filicina O 

cb C Ptilota serrata (incl. pectinata ) O 

cb C Rhodomela lycopodioides O 

cb C Scagelia americana O 

np N Tokidadendron kurilensis X X O 

nep N Weeksia coccinea X* X* X 

HETEROKONTOPHYTA, Phaeophyceae 

Val al-sbr al-pil slough duckflat Checklist 

cb C Agarum clathratum (cribrosum) O 

cb C Chordaria flagelliformis Xun X un O 

np N Chordaria gracilis O 

cb C Coilodesme bulligera O 

np N Costaria costata O 

cb C Desmarestia aculeata O 

cb C Desmarestia viridis O 

cb C Dictyosiphon foeniculaceus X* X* X un O 

nep N Ectocarpus parvus O 

ws C Ectocarpus siliculosus O 

nep N Elachista lubrica X* X* O 

cb I NR NAT Fucus cottonii X* X* X 

cb C Fucus gardneri/distichus/evanescens X* X* X* un X* O 

cb C Fucus spiralis X X X X 

np, a N Laminaria "groenlandica"/bongardiana X* X* O 

cb C Laminaria saccharina X* X* X O 

np N Laminaria yezoensis X* X* O 

ws C Leathesia difformis O 

cb C Melanosiphon intestinalis X X O 

ws C Petalonia fascia O 

ws C Pilayella littoralis/washingtonensis X* X* X X* O 

ws C Scytosiphon simplicissimus X* X* X un O 

np N Soranthera ulvoidea X* X* O 

ws C Sphacelaria rigidula O 

cb C Spongonema tomentosum O 

HETEROKONTOPHYTA, Xanthophyceae 


ws C NR BC Vaucheria longicaulis () mats X* X* X 

CHLOROPHYTA, Chlorophyceae 


cb C Acrosiphonia arcta X* X* X* O 

nep N Acrosiphonia coalita O 

np N Acrosiphonia saxatilis O 

cb C Blidingia chadefaudii O 

ws C NR BC Blidingia marginata X* X* X* X 

ws C Blidingia minima X* X* X* O 

cb C Blidingia subsalsa X* X* X* X* O 

cb C Chaetomorpha capillaris/cannabina O 

nep N NR Wa Chaetomorpha recurva O 

ws C Cladophora albida O 

ws C Cladophora sericea O 

ws C Enteromorpha clathrata O 

ws C Enteromorpha in compressa O 

ws C Enteromorpha intestinalis X* X* X* O 

ws C Enteromorpha linza O 

ws C Enteromorpha prolifera/torta X* X* X* X* X



cb C Gayralia oxyspermum X* X* X 

cb C NR BC Halochlorococcum moorei X* X* X* X 

cb C Kornmannia zostericola (epiphytic) O 

cb C Monostroma grevillei/arcticum O 

ws C Rhizoclonium implexum X* X* X* X* O 

ws C Rhizoclonium riparium X* X* X* X* O 

ws C Rhizoclonium tortuosum O 

ws C Ulothrix implexa (non flacca ) X* X* X* X X* O 

np C# Ulva fenestrata /expansa/lactuca X* X* X* O 

cb C Ulvaria obscura X* X* X* O 

ws C Urospora penicilliformis X* X* X 

SEAGRASSES 


cb C Zostera marina X* X* O 

LICHENS 


cb C Verrucaria maura O 

cb C Verrucaria mucosa O 

Ra St NR So Val al-sbr al-pil slough duckflat Checklist 

TOTALS: 

Species 47 35 7 7 24 112 

New Records 5 1 1 1 4 7 


! = abundant or common 

# = currently being examined with molecular techniques 

= uncertainty of identification 

al-sbr=Alyeska boat ramp and vicinity 

al-pil=Alyeska small boat harbor pilings 

BC=British Columbia 

C=cryptogenic 

cb=circumboreal 

Checklist=total records for Port Valdez including literature and the present study 

duckflat=Mudflat east of the town of Valdez 

N=native to North Pacific 

NAT=North Atlantic 

nep= northeast Pacific 

np= North Pacific 

NR = new record to Alaska 

nr = northward range extension within Alaska 

O= records from the literature and pilot study 

slough=Slough about 1 mile from Alyeska gate 

So=Closest source to PWS 

Stat=NIS Status (native, cryptogenic, introduced, etc) 

un= living unattached 

Val=Total records for Port Valdez for the present study 

Wa=Washington 

ws= widespread, occurring in North Pacific, North Atlantic, and Australia or New Zealand 

X*= current record noted with specimen 

X= current record noted in the field


TABLE 9C1.4. Marine and Estuarine Plants Collected at Shore*** 

Sites in Prince William Sound 

NIS ANALYSIS 

HARBORS MUD BAYS HEADLANDS RK 

TAXA 

AND REEFS BAYS 

Ra St NR So Val Cor Whit CB SMB Gro RP Bus Green NW 


ws C Ahnfeltia fastigiata X* X*! 

nep N Antithamnionella pacifica X* X* X* 

ws C Antithamnionella spirographidis X X 

ws C Audouinella purpurea X* X* 

ws C Bangia atropurpurea X X* X* X* 

np N Bossiella cretacea X* X* 

nep N Bossiella plumosa X* 

nep N Callithamnion acutum X* 

nep N Callithamnion pikeanum v. laxum X* 

nep N Callithamnion pikeanum v. pikeanum X* 

cb C# Ceramium cimbricum X* X* 

nep I# NR Cal Ceramium sinicola (on Codium ) X* 

nep N# Ceramium gardneri X* 

nep N# Ceramium pacificum/washingtonensis X* X* X* X* X* X*! 

ws C# Ceramium rubrum/kondoi X* X* 

np N Constantinea subulifera X* X X* X*! X*! 

np N Corallina frondescens X* X* X* X 

np,ch N Corallina officinalis v. chilensis X* X* X*! 

nep N Cryptosiphonia woodii X* X* X* X* X* X* X*! X* 

nep N Delesseria decipiens X 

cb C Devaleraea ramentacea X X* X* 

cb C Dumontia contorta X* X* X* X* X X* 

np N Dumontia simplex X 

nep N Endocladia muricata X* X* 

ws C Erythrotrichia carnea X* X* X* X* 

np N Glioipeltis furcata X* X X* X* X* X* X*! X*! 

np,ar N Halosaccion firmum X X* X X X 

np N Halosaccion glandiforme X* X X* X X* X X* 

np N Leachiella pacifica X* X* X* X* 

nep N Mastocarpus papillatus complex X* X* 

np N Mastocarpus cf. pacificus X* X X* X* X X* 

np N Mazzaella phyllocarpa X X* X* X*! X* X*! X! 

nep N Mazzaella splendens X 

nep N Microcladia borealis X 

np N Neorhodomela aculeata X X X* X* X* X*! X*! X! 

np N Neorhodomela larix X X* X* X* 

np N Neorhodomela oregona X* X*! X X* X* X X*! X*! X*! 

np N Neoptilota asplenioides X* X X* X! X 

nep N Odonthalia floccosa X! X X*! X* X*! X*! 

np N Odonthalia setacea (drift) X* 

nep N Opuntiella californica X* 

np N Palmaria calophylloides/stenogona X X X X** X*! 

nep N Palmaria hecatensis X* X*! X* X* X 

cb C Palmaria mollis/palmata X* X! X* X X* X* X! 

np N Phycodrys riggii X X* X* X*



nep N Platythamnion pectinatum X X X* 

np N Pleonosporium cf vancouverianum X* 

nep N Polysiphonia eastwoodae X* 

ws C Polysiphonia brodiaei X 

nep N Polysiphonia hendryi v. deliquescens X X 

nep N Polysiphonia hendryi v. hendryi X* X* 

nep N Polysiphonia pacifica v. determinata X* 

nep N Polysiphonia pacifica v. pacifica X* X* X* X* 

np,nz N nr SeA Polysiphonia senticulosa X* X* X* X* X* X* 

cb C Polysiphonia stricta (urceolata) X* X* X* 

nep N Porphyra cuneiformis X* X* X 

cb C NR Com Porphyra miniata X* 

nep N Porphyra nereocystis X* 

nep N NR Wa Porphyra rediviva X* 

np N Porphyra torta/abbottae X* 

np N Pterosiphonia bipinnata X* X* X* X* X* X* X* X*! 

cb C Ptilota serrata (incl. pectinata ) X* X* X* X* 

np,ar N Rhodymenia pertusa X X* X X* X! 

cb C Scagelia americana X* X* X* X* X* X* 

nep N Smithora naiadum X* 

np N Tokidadendron kurilensis X X* X*! X*! 

nep N Weeksia coccinea X* X X* 


Val Cor Whit CB SMB Gro RP Bus Green NW 

np N# Alaria taeniata/angusta/crispa X* X X 

np N# Alaria tenuifolia/pylaii//membranacea X* 

np N# Alaria praelonga/marginata X X* X* X*! 

cb C Agarum clathratum (cribrosum) X X* X* 

np N Analipus japonicus X X* X* X* X* X 

cb C Chorda filum X* X* X* X 

cb C Chordaria flagelliformis Xun X*! X* X* X* X* X* X* 

cb C Coilodesme bulligera X* 

ak E NR Coilodesme n. sp. X* 

np,nz N Colpomenia bullosa X* 

ws C Colpomenia peregrina X* X* X* X* 

np N Costaria costata X* X* 

np N Cymanthera triplicata X* 

np N Cystoseira geminata X* 

cb C NR Com Delamarea attenuata X* X* 

cb C Desmarestia aculeata X X X X X* X 

cb C Desmarestia viridis X X X* 

cb C Dictyosiphon foeniculaceus X* X*! X*! X* X* X* X* X* X* X 

ws C Ectocarpus siliculosus X 

cb C Elachista fucicola X* X X X* 

nep N Elachista lubrica X* X* X* 

cb C Eudesme virescens X* X* X* X* 

cb I NR NAT Fucus cottonii X* X* X* X* 

cb C Fucus gardneri/distichus/evanescens X* X*! X*! X X X* un X* X X*! X* 

cb C Fucus spiralis X X* X X X X! X! 

np,ar N Laminaria "groenlandica"/bongardiana X* X* X* X* X* X* X* X*! 

cb C Laminaria saccharina X* X X*! X* X! 

np N Laminaria yezoensis X* X*! X*



ws C Leathesia difformis X X X* X! X 

nep N Leathesia nana X* X* X* X* 

nep F Macrocystis integrifolia X* drift 

cb C Melanosiphon intestinalis X X*! X* X* X* X* X* X* 

ws C Pilayella littoralis/washingtonensis X* X* X* X* X* X* X X X* 

ws C nr SeA Punctaria latifolia X* X* 

ak E Punctaria lobata X* 

cb C NR Jap Punctaria plantaginea* X X* X X X* 

cb C Punctaria tenuissima X* 

cb C Ralfsia fungiformis X* X* 

np N Saundersella simplex X* X* 

ws C Scytosiphon simplicissimus X* X* X* X* X X* X* X* 

np N Soranthera ulvoidea X* X* X X* X X* X* X* 

np N Soranthera ulvoidea f. difformis X* X* 

cb C Sphacelaria racemosa X* X* 

ws C Sphacelaria rigidula X X* X* X* 

HETEROKONTOPHYTA, Xanthophyceae 


ws C NR BC Vaucheria longicaulis () X* X*! 



cb C Acrosiphonia arcta X* X* X*! X X* X* X* X X* 

np N Acrosiphonia saxatilis X* X* X* 

ws C NR BC Blidingia marginata X* 

ws C Blidingia minima X* X* X* X* X X* X* X* 

cb C Blidingia subsalsa X* X* X* 

cb C NR BC Capsosiphon fulvescens X* 

ws C Cladophora albida X* X* X* X* X X* X* 

cb C Cladophora hutchinsiae X* 

ws C Cladophora sericea X X* X X* X* X* X* X* X*! 

np N Cladophora stimpsonii X* X* 

nep N nr SeA Codium fragile subsp. fragile X* 

ws I# NR Wa Codium fragile subsp. tomentosoides X* 

ws C Enteromorpha intestinalis X* X* X X X X X 

ws C Enteromorpha linza X*! X X* X* X 

ws C Enteromorpha prolifera/torta X* X X* X* X* X X* 

cb C Gayralia oxyspermum X* X X X* 

cb C NR BC Halochlorococcum moorei X* X* 

cb C Kornmannia zostericola (epiphytic) X* 

cb C NR NAT Kornmannia leptoderma (epilithic) X* 

nep N NR Wa Monostroma fractum X* X* 

cb C Monostroma grevillei/arcticum X X X* X* 

ws C Percursaria percursa X* X* 

ws C Rhizoclonium implexum X* 

ws C Rhizoclonium riparium X* 

ws C Rhizoclonium tortuosum X* X* X* X* X 

ws C Ulothrix implexa (non flacca ) X* X* X 

ws C# Ulva fenestrata /expansa/lactuca X* X* X X* X* X* X* X* 

cb C Ulvaria obscura X* X X* X X* 

np N Ulvella setchellii X* 

ws C Urospora penicilliformis X* X


Table 9C1.4. Continued 

SPERMATOPHYTA, Seagrasses 


cb C Zostera marina X* X* X* X X* X X* 

nep N Phyllospadix scouleri X X* 

nep N Phyllospadix serrulatus X 

LICHENS 


cb C Verrucaria maura X X X 

47 41 56 

TOTALS: Species (total =146) 

47 41 56 10 45 63 61 59 71 69 

New Records (total =17) 

5 5 2 2 5 4 2 2 6 1 

GROUP TOTALS (with overlap excluded): 

Harbors Mud Bays Headlands Rk Bays 

Species in Merged Groups 87 

78 

96 

69 

New Records in Merged Groups 8 

7 

8 

1 


Site Abbreviations and Dates: 

! = abundant or common Bus=Busby Island south reef, 21 June 

# = currently being examined with molecular techniques CB=Cloudman Bay, East Bleigh I, 21 June 

***=shore sites include both shore and marina sites 

Cor=Cordova, 23 June 

= uncertainty of identification Green=Green Island, northwest point, 24 June 

ak=Alaska 

Gro=Growler Island, near resort, 27 June 

ar=arctic 

NW=Northwest Bay, Knight Island, 25 June 

BC=British Columbia 

RP=Rocky Point headland, 22 June 

C=cryptogenic 

SMB=Saw Mill Bay, 22 June 

Cal=California 

V-Ck=Port Valdez checklist (all records known) 

cb=circumboreal Val=all Port Valdez collections, June 1998 

Ch=Chile 

Whit=Whittier, 26 June 

Com=Commander Islands, Russia 

Cor=Cordova 

drift= dying unattached 

E=endemic to Alaska 

F=failed introduction 

I=possible introduction 

Jap=Japan 

N=native to North Pacific 

NAT=North Atlantic 

nep= northeast Pacific 

np=North Pacific 

NR = new record to Alaska 

nr = northward range extension within Alaska 

nz=New Zealand 

O=Presence known from the Pilot Study and literature 

Ra=Distribution Range 

RK=rocky 

SEA=Southeast Alaska 

So=Closest source to PWS 

St=range status (see N, C, E, and I) 

un= living unattached 

Wa=Washington 

ws= widespread, occurring in North Pacific, North Atlantic, and Australia or New Zealand 

X*=Presence known from the current study; specimens available 

X=Presence known from the current study; no specimen taken


TABLE 9C1.5. Marine Algae Collected from Off-Shore*** Floats 

in Prince William Sound, June 1998 

NIS ANALYSIS 

TAXA Float FLOATS 

Only 

Range Stat NR So Cklist TAT WBF MBF EIF SBF EBF on floats 

CYANOPHYTA, Cyanophyceae 

ws C Calothrix crustacea X X* ** 

ws C Rivularia atra X X* ** 


nep N Antithamnionella pacifica X X* 

cb I NR SD Chroodactylon ramosum X X* ** 

nep N nr SeAk Polysiphonia senticulosa X X* 

ws C Polysiphonia urceolata X X* ** 

cb C Scagelia americana X X* 


nep N Coilodesme californica X X* ** 

ak E NR Coilodesme n. sp. X X* 

np N Cystoseira geminata X X* 

cb C NR Com Delamarea attenuata X X* 

nep N nr BC Ectocarpus acutus X X* ** 

nep N nr BC Ectocarpus dimorpha X X* ** 

nep N Ectocarpus parvus X X* **, V-CK 

Ectocarpus sp. (Acinetospora ) X* 

Giffordia sp. X* 

np N Laminaria groenlandica X X* 

ws C Laminaria saccharina X X* X* 

np N Laminaria yezoensis X X* 

cb C Melanosiphon intestinalis X X* 

cb I NR Jap Microspongium globosum X X* ** 

ws C Pilayella littoralis X X* 

Pilayella sp. (elongate X* 

intercalary structures) 

cb C NR Jap Punctaria plantaginea X X* 

cb C nr SeAk Punctaria latifolia (Desmotrichum ) X X* 

ws C Scytosiphon simplicissima X X* 


ws C Cladophora albida X X* X* 

ws C Cladophora sericea X X* X* X* 

ws C Enteromorpha prolifera/torta X X* 

ws C Percursaria percursa X X* 

TOTALS: 27 17 1 4 1 4 7 9 

New Records 9 4 1 1 0 0 3 4 


*=Specimen available LJ=LaJolla, California Float Sites and Dates (Coord. with JC) 

**=Only on floats in this study N=native EBF=Eaglek Bay floats, 26 June 

***=Sites accessed by boat nep=northeast Pacific EIF=Ester Island float, 25 June 

BC=British Columbia np=North Pacific MBF=Main Bay barrier buoy, 25 June 

C=cryptogenic nr=new record from neighboring area SBF=Squaw Bay float, 26 June 

cb=circumboreal NR=new record from remote area TAT=Oyster floats near Tatilek, 21 & 22 June 

Com=Commander Islands, SeAk=Southeast Alaska WBF=Windy Bay floats, 23 June 

Jap=Japan V-Ck=also known from the Valdez Checklist Float Cklist=species used in this study 

ws=widespread


Species were then categorized as to whether they were new distribution records (NR) to the area. 

These included species that had never before been reported from Prince William Sound (nr) or 

Alaska (NR). In each of these cases, the closest known records to the area were given as the 

source (So). Since the new records seemed to be the most likely category in which to find 

recognizable NIS, they are highlighted in gray throughout the charts and tables. 

This preliminary quantification of marine plant species and NIS in Prince William Sound 

required a number of lengthy, detailed steps. After gathering, identifying, and curating all of the 

species, site lists had to be prepared and both local and global biogeographic information 

compiled. Then, with this information in hand, the residency status and new distribution records 

of each species were determined. Only after all of this was completed could NIS begin to be 

recognized. Since many of the steps in this process revealed important data that characterized not 

only NIS but the marine flora in general, this report presents the site lists in their entirety and 

then summarizes the results for comparative purposes in tables and graphs. Since the results are 

lengthy, they have been organized into the following 7 major parts that are presented below: 

• The Species Lists by Site, including Port Valdez, Shore, and Floats. 

• The Total Species Numbers and Composition of the Individual Sites. 

• Total Species Numbers and Composition in each Habitat Type. 

• Native, Cryptogenic, and Introduced Species and their Taxonomic Composition. 

• Native, Cryptogenic, and Introduced Species in the Habitat Types. 

• New Species Records and Probable Introductions. 

• Comments on the Five Probable Introductions and One Important Failed Introduction. 

Results 

During our 9-day search for NIS in Port Valdez and Prince William Sound, 489 plant 

samples were processed (Table 9C1.6). These samples contained 155 different species 

dominated by the red (Rhodophyceae), brown (Phaeophyceae), and green (Chlorophyceae) 

algae, in that order. Among these species, 21 were found to be new records to the area, and, of 

these, at least 5 appear to be introduced. In addition, 70 species were found to be cryptogenic, 

some of which have suspicious characteristics of NIS. 

TABLE 9C1.6. Collection Data* 

TAXONOMIC SAMPLES 

TOTAL NEW 

GROUP Total Herb. Form. SPECIES RECORDS 

Rhodophyceae 199 135 64 69 5 

Phaeophyceae 162 120 42 49 8 

Chlorophyceae 117 99 18 30 7 

Xanthophyceae 2 2 0 1 1 

Seagrasses 7 4 3 3 0 

Lichens 0 0 0 1 0 

Cyanophyceae 2 1 1 2 0 

Total, June 1998 489 361 128 155 21 


* =Samples and species counts are only for the June 1998 trip. 

Herb.=Pressed herbarium sheets 

Form.=Bottles of preserved specimens


The total species and new species records/collecting site were correlated with collection 

time (Fig. 9C1.1). Longer collecting periods yielded more species at an R 2 value of 43%. New 

records, on the other hand, appear to be almost unaffected by collection time, showing an R 2 of 

only 7%. 

FIG. 9C1.1. Collection Efficiency 

80 

180 

Total Species 

70 

60 

50 

40 

30 

20 

Val 

al-sbr 

al-pil 

slough 

duckflat 

Cor 

Whit 

10 

0 

Major Collection Sites 

CB 

SMB 

Gro 

RP 

Bus 

Green 

NW 

New Records 

Total Species 

Collection Time 

160 

140 

120 

100 

80 

60 

40 

20 

0 

Collection Time (minutes) 

The Site Species Lists. Species of marine and estuarine plants identified at all sites sampled 

during our June 1998 survey are listed in Tables 9C1.3, 9C1.4, and 9C1.5. The plants sampled 

were predominantly macrobenthic marine algae of the Rhodophyceae, Phaeophyceae, 

Chlorophyceae, Xanthophyceae, and Cyanophyceae along with several species of seagrasses and 

marine lichens. Species occurrence at the various sites is designated with an X in the lists. If 

samples were taken and curated for identification purposes, the species are listed with an X*, 

indicating that vouchers are available in the OSU/HMSC herbarium for study. In addition, each 

species is categorized for several biogeographic features that are necessary for the NIS Analysis, 

explained in the Methods section above. 

• Species of Port Valdez. Samples from 4 sampling sites in Port Valdez included 47 algal 

species and 5 new records (Table 9C1.3). The sites covering the largest areas (the Alyeska 

small boat ramp and the duckflat) contained the majority of the species. The highest species 

count occurred at the Alyeska boat ramp where the greatest amount of hard substratum was 

available for algal settlement. The highest number of new records occurred in the duckflat. 

A few of these species are good candidates for NIS status. However, their lack of earlier 

discovery may have an obvious explanation. Mudflats, like the duckflat, are not only 

notoriously poor habitats for most marine algae, but they can be dangerous in Alaska. 

Therefore, earlier phycologists avoided many of these areas. Knowing this to be true, it was 

possible to predict the occurrence of some new records (e.g., Fucus cottonii and Vaucheria 

longicaulis) in the mudflats and sloughs. Also shown in Table 9C1.3 is a Checklist of Algal 

Species for Port Valdez, which includes the species collected from the 1998 sampling, as


well as those found previously during the 1997 Pilot Study (Ruiz & Hines, 1997) and the 

literature (Calvin and Lindstrom, 1980, and Weigers et al., 1997). In addition to our summer 

sampling, this list includes year-round collections taken by the earlier investigators. The 

total count for the entire Port Valdez area, including these earlier records, amounts to 112 

species. 

• Species of Shore Sites. Shore sites include all intertidal areas and marinas sampled during 

the June 1998 cruise, along with the combined records for Port Valdez (Table 9C1.4). These 

10 sites covered a wide range of habitats, which were grouped into 4 major habitat types: 

Harbors, Mud Bays, Headlands and Reefs, and Rocky Bay (presented in more detail below). 

The overall species count for all of the shore areas was 146 species with 17 new records. 

The highest species diversity occurred at Green Island, Northwest Bay, and Growler Island, 

all of different habitat types; while the highest number of new records occurred at Green 

Island, Saw Mill Bay, Cordova, and Port Valdez, also a mixture of habitat types. Only two 

species (Dictyosiphon foeniculaceus and Fucus gardneri ) were found at all shore sites 

sampled. Four others (Cladophora sericea, Acrosiphonia arcta, Pilayella littoralis, and 

Neorhodomela oregona) were found at all but one site (and were possibly overlooked there). 

Numerous species (31) were common to all of the habitat types, but there were also an 

extraordinary number of species that appeared to be limited to only 1 habitat type (17 were 

found only in harbors, 9 only in mud bays, 19 only on headlands and reefs, and 5 only in 

rocky bays). Two species restricted to harbors are new records to the area: Porphyra 

rediviva, a newly discovered free-floating marsh plant that could be easily transported by 

ships, and Vaucheria longicaulis , a species unique to high mudflats that is common to many 

southern west coast harbors. Although not a new record, another interesting harbor species is 

Antithamnionella spirographidis. This species is reported to be common to harbors in British 

Columbia and is thought to be introduced to that area (Lindstrom in DeWreede 1996). 

However, its circumboreal and Australian existence leads me to categorize it as cryptogenic 

in this paper. 

• Species from Floats. Marine plants were sampled from five oyster floats and one barrier buoy 

(MBF) (Table 9C1.5). A total of 27 different algal species were identified from the floats. 

Of these, 9 were not collected at any of the other sites during our trip. Most of these unique 

species are small and could have been overlooked in other areas, but several are species that 

probably could only find suitable habitat on the floats. Over half the 27 species collected are 

well-known fouling organisms (e.g., Cladophora sericea, Pilayella littoralis, and 

Polysiphonia urceolata). Nine new species records, the highest habitat number in our 

survey, were also found on the floats. This may be related to the fact that most of the floats 

sampled are used in aquaculture, which could be a source of introductions. The highest 

counts for both species and new records occurred on the floats at Tatitlek Narrows (TAT) 

and Eaglek Bay (EBF) used in active oyster culture. Two of the new records found at these 

sites (Chroodactylon ramosum and Microspongium globosum) and possibly more are thought 

to be introduced. One species (Polysiphonia senticulosa) is considered to be a range 

extension from southeast Alaska, but it is already widespread in Prince William Sound. This 

species is presumably native in Washington to Southeast Alaska, and was recently reported 

to be introduced and pervasive in New Zealand (Nelson and Maggs, 1996).


The Total Number of Species and Species Composition of the Individual Sites. The overall 

number of species was 155 for all sites, but the numbers of species per site ranged from only 10 

to 71 species, indicating that there is considerable variation in species composition among 

habitats (Table 9C1.7, Fig. 9C1.1, 9C1.2). The 21 new records across all areas ranged from 1 to 

9 at the individual sites and was highest on floats. The highest species count (71 species) 

occurred at Green Island, the most exposed and highly saline site. At this site, the proportion of 

red algal species was nearly 2 times that of the brown algae and 4 times that of the greens. The 

lowest species count (10 species) occurred at Cloudman Bay, a sheltered, estuarine mud bay. At 

this site there were almost no red algae, and the brown algae were more abundant than the 

greens. 

Table 9C1.7. Total Species Numbers and Composition at Each Site, June 1998 

Taxonomic Group Val Cor Whit CB SMB Gro RP Bus Green NW Floats Total* V-Total** 

Rhodophyceae 18 12 23 1 15 26 27 27 40 37 5 69 84 

Phaeophyceae 13 11 14 5 20 24 23 21 21 22 16 49 52 

Chlorophyceae 14 16 17 3 9 12 9 8 10 9 4 30 36 

Xanthophyceae 1 1 0 0 0 0 0 0 0 0 0 1 1 

Seagrasses 1 1 1 1 1 1 1 3 0 0 0 3 3 

Lichens 0 0 1 0 0 0 1 0 0 1 0 1 2 

Cyanophyceae 0 0 0 0 0 0 0 0 0 0 2 2 2 

TOTALS 47 41 56 10 45 63 61 59 71 69 27 155 180 

NEW RECORDS 5 5 2 2 5 4 2 2 6 1 9 21 23 

Abbreviations: Sites as in Table 1; Val=the Port Valdez collections combined; Floats=the float collections combined; 

Total*=with overlap and earlier collections excluded; V-Total**=Total* with the Port Valdez Checklist species included. 

Figure 9C1.2. Total Species Numbers and 

Composition at each Site, June 1998 

80 

70 

60 

Total Species 

50 

40 

30 

20 

10 

0 

Val Cor Whit CB SMB Gro RP Bus Green NW Floats 

Collecting Sites 

Rhodophyceae Phaeophyceae Chlorophyceae 

Xanthophyceae Seagrasses Lichens 

Cyanophyceae 

Total Species Numbers and Composition in each Habitat Type. Since several sampling sites 

had mixed habitats, categorizing the sites into distinct habitat types had some weaknesses. 

However, it increased the number of species sampled for each category of habitat, providing 

more power to the data analysis (Table 9C1.8, Fig. 9C1.3a, b).


TABLE 9C1.8. Habitat Type and Species Composition 

Taxonomic Group Valdez Harbors Mud Bays Rk Bays Headlands Floats Total 

Rhodophyceae 18 37 31 37 48 5 69 

Phaeophyceae 13 22 31 22 30 16 49 

Chlorophyceae 14 25 15 9 14 4 30 

Xanthophyceae 1 1 0 0 0 0 1 

Seagrasses 1 1 1 0 3 0 3 

Lichens 0 1 0 1 1 0 1 

Cyanophyceae 0 0 0 0 0 2 2 

Total 47 87 78 69 96 27 155 

Fig 3a. Habitat Type and 

Species Composition 

120 

100 

Total Species 

80 

60 

40 

20 

0 

Valdez 

Harbors 

Mud Bays 

Rk Bays 

Headlands 

Floats 

Habitat Type 



Cyanophyceae 

Fig. 3b. Habitat Type and 

Percentage Composition 

100% 

80% 

Percentage Composition 

60% 

40% 

20% 

0% 

Valdez Harbors Mud Bays Rk Bays Headlands Floats 

Habitat Type 



Cyanophyceae


The 5 habitat types with their features and included sites are: 

• Harbors* (Val, Cor, and Whit). Sheltered areas with variable salinities (0-28 ppt) and 

variable substrates including mud, cobble, and wood (the pilings). Heavily influenced by 

boat traffic and other human activities. [* Note that Port Valdez (Val), also included in the 

Harbor group, is included separately in several of the tables to show the comparable diversity 

of this targeted site.] 

• Mud Bays (CB, SMB, Gro). Sheltered bays with salinity ranges from 3-11 ppt with a 

substratum of primarily mud, although cobble and bedrock was often available. 

• Rocky Bays (RB). One semi-sheltered bay with a salinity ranging from 10-27 ppt and a 

substratum varying from gravel to cobble to bedrock. 

• Headlands and Reefs (RP, Bus, Green). Very exposed habitats with salinity ranges from 15- 

30 ppt and a substratum consisting almost totally of bedrock and cobble. 

• Floats (TAT, WBF, MBF, EIF, SBF, EBF). Exposed to semi-sheltered off-shore habitats. 

Salinities ranged from 16-28 ppt and substrata included 5 plastic oyster floats and line and 1 

cement buoy (MBF). 

Headlands and reefs with high exposure and high salinity had the greatest species 

diversity. As would be expected for temperate zones, they are dominated in descending order by 

red, brown, and green algae. Surprisingly, the next largest diversity of species occurred in the 

harbors. Although the red algae also predominated there, harbors had a large number of green 

algae. In the mud bays and on off-shore floats, there was a tendency for increase in the 

percentage of brown algae. However, since total species number varies among habitat types, 

composition of algal groups may be partly an artifact of small number of species at the low 

diversity sites. 

The numbers of new records among the various habitat types were fairly uniform except 

in rocky bays where the sample size (1 bay) was small (Table 9C1.9). The slight increase in 

numbers on floats may be significant, but overall, the data indicate that habitat type has little to 

do with the discovery of new species records. 

TABLE 9C1.9. Habitat Type and New Species Records 

Valdez Harbors Mud Bays Headlands Rk Bays Floats (pvc/ Totals* 

TAXONOMIC (mixed) (mixed) (mud/cob) and Reefs (gravel/cob) concrete) 

GROUP SP NR SP NR SP NR SP NR SP NR SP NR SP NR % 

Rhodophyceae 18 1 37 2 31 1 48 2 37 1 5 2 69 5 7 

Phaeophyceae 13 1 22 2 31 4 30 4 22 0 16 7 49 8 16 

Chlorophyceae 14 2 25 3 15 2 14 2 9 0 4 0 30 7 23 

Xanthophyceae 1 1 1 1 0 0 0 0 0 0 0 0 1 1 

Seagrasses 1 0 1 0 1 0 3 0 0 0 0 0 3 0 

Lichens 0 0 1 0 0 0 1 0 1 0 0 0 1 0 

Cyanophyceae 0 0 0 0 0 0 0 0 0 0 2 0 2 0 

Total 47 5 87 8 78 7 96 8 69 1 27 9 155 21 

% Habitat Total 11 9 9 8 1 33 14 

% NR Total (21) 24 38 33 38 5 43 100 

* = excludes overlapping records 

cob= cobble


Native, Cryptogenic, and Introduced Species and their Taxonomic Composition. Of the 155 

total algal species found, 52% are native, 45% are cryptogenic, and 3% (5 species) appear to be 

introduced (Table 9C1.10, Fig. 9C1.4). The taxonomic composition of these groups parallels the 

findings in the Pilot Study survey for Port Valdez. The native species contain a very large 

percentage of red algae, about 64% of the total. The brown algae make up 27% of the natives, 

and the greens only 6%. The composition of the cryptogenic forms is almost the reverse. The 

red algae are only about 23% of the total count, while the browns and the greens both average 

about 35%. 

Table 9C1.10. Native, Cryptogenic, and Introduced 

Species and their Taxonomic Composition 

Taxonomic 

Group Total N 

Status** 

C I NR 

Rhodophyceae 69 51 16 2 5 

Phaeophyceae 49 22 25 2 8 

Chlorophyceae 30 5 24 1 7 

Xanthophyceae 1 0 1 0 1 

Seagrasses 3 2 1 0 0 

Lichens 1 0 1 0 0 

Cyanophyceae 2 0 2 0 0 

Totals 155 80 70 5 21 

% of Total 100 52 45 3 14 

NR 21 7 9 5 

** = For simplification, 2 endemics and 1 failed introduction are included with the natives. 

N=native, C=cryptogenic, I=potentially introduced, NR=new records 

Fig. 9C1.4. Residency Status and 

Taxonomic Composition 

180 

160 

Total Species 

140 

120 

100 

80 

60 

40 

Cyanophyceae 

Lichens 

Seagrasses 

Xanthophyceae 

Chlorophyceae 

Phaeophyceae 

Rhodophyceae 

20 

0 

Total N C I NR 

Status 

Of the 21 new species records across all habitats, 5 were red algae, 8 brown, 7 green, and 

1 was a Xanthophyte (Table 9C1.9), reflecting a fairly uniform distribution of new records across 

at least the 3 major taxonomic groups. However, the percentage of new species records by group 

increased dramatically from red to brown to green algae. It is possible that this increase relates, 

in part, to our overall level of taxonomic understanding in each of these 3 major classes. Since 

stable morphological features usable in taxonomy decrease as one moves from the red to the 

brown to the green algae, ease of accurate identification likewise decreases. Hence, it is likely


that our knowledge of the Alaskan flora is correspondingly most complete for the reds, then the 

browns, and lastly the greens. 

Native, Cryptogenic and Introduced Species by Habitat Types. In all habitat types except the 

floats, the native and cryptogenic species were fairly evenly distributed and ranged from 44 to 

55% of the species (Table 9C1.11, Fig. 9C1.5a, b). However, there were some predictable 

reversals of dominance. In the harbors and mud bays and on the floats, the cryptogenic species 

were the most abundant, while on the headlands and reefs and in the rocky bays, the native 

species predominated. This reversal reflects the confounding effect of variation in groups among 

habitats (Figs. 9C1.3a, b). Since red algae predominated on the reefs and rocky bays (and green 

algae are relatively low in numbers), native species, consisting mostly of red algae, were also 

predominant there. On the other hand, in harbors and mud bays red algae were not as common 

(and green algae are more abundant); hence the numbers of native species were lower in those 

habitats. The reefs and rocky bays were under-collected in most cases, weakening the 

conclusions about algal species in these areas. On floats the cryptogenic forms consisted of 

55% of the species while the native species consisted of only 37%. Since most of the floats 

sampled were from oyster farms, it is likely that they are periodically cleaned. Each cleaning of 

the floats would provide cleared primary substrata for ephemeral (opportunistic) species that are 

quick to colonize and reproduce. Since ephemeral species are most often cryptogenic, their 

higher percentage may be understandable. 

Table 9C1.11. The Native, Cryptogenic, and Introduced 

Species Occurring in Each Major Habitat 

Residency 

Status Harbors 

Habitat Types 

Mud Bays Headlands Rk Bays Floats Totals 

Native* 38 35 52 36 10 80 

Cryptogenic 48 42 42 33 15 70 

Introduced 1 1 2 0 2 5 

Totals 87 78 96 69 27 155 

New Records 8 7 8 1 9 21 

*2 endemics and 1 failed introduction (under rk bay browns) are included in natives. 

Total Species 

Fig 9C1.5a. Species Residency Types 

Occurring in Each Major Habitat 

120 

100 

80 

60 

40 

20 

0 

% of Habitat 

Fig 9C1.5b. Major Habitats and Residency 

Type Composition 

100% 

80% 

60% 

40% 

20% 

Harbors 

Mud Bays 

Habitat Type 

Headlands 

Rk Bays 

Floats 

Introduced 

Cryptogenic 

Native* 

0% 

Harbors 

Mud Bays 

Habitat Types 

Headlands 

Rk Bays 

Floats 

Introduced 

Cryptogenic 

Native*


New Species Records and Probable Introductions. The 21 new species records for the June 

1998 trip are the most likely candidates to be NIS; however, several factors should be considered 

further before the status of the species can be determined definitively (Table 9C1.12). Since all 

of the new records appeared to have been overlooked for at least some period of time, the 

obvious questions related to why this oversight occurred are: 

TABLE 9C1.12. New Records of Benthic Marine Algae to Prince William Sound 

(species overlooked, misidentifications, range extensions, and possible introductions) 

Justification Stat Taxon Location Type Ra So C Comments 

Rhodophyta, Rhodophyceae 

gi I Ceramium sinicola Green Exp nep Cal 3 epiphyte, MB in progress 

gi, sol I Chroodactylon ramosum TAT F cb S. Cal. 1 microscopic 

rex N Polysiphonia senticulosa RP, Cor, Whit, Bus, All np,nz SeAk 1 easy to recognize 

Green, NW, TAT ex M invasive 

mid C Porphyra miniata Gro M cb Com. 2 MB in progress 

mid N Porphyra redidiva Val (duckflat) M nep Wa 1 recently described 

Heterokontophyta, Phaeophyceae 

mid E Coilodesme n. sp. Green, TAT Exp, F ak 1 epiphyte, morph. needed 

mid C Delamaraea attenuata SMB, Bus, EBF Exp, F cb Com 1 recently illustrated 

rex N Ectocarpus acutus MBF, EBF F nep BC 1 

rex N Ectocarpus dimorphus EBF F nep BC 1 

gi I Fucus cottonii Val (slough), SMB, M cb N. Atl 1 common in marshes 

CB, Gro 

MB in progress 

gi, sol I Microspongium globosum EBF F cb Jap, N. Atl 1 microscopic 

rex C Punctaria latifolia TAT, SMB, RP Exp, F cb SeAk 1 

mid C Punctaria plantaginea SMB, Cor, Whit, All cb N. Atl. 1 some think cold water 

Green, Gro ex F form of latifolia 

Heterokontophyta, Xanthophyceae 

reproductive material 

sol, rex C Vaucheria longicaulis Val, Cor M, H ws BC 3 needed to confirm sp. 

Chlorophyta, Chlorophyceae 

rex C Blidingia marginata Val M ws BC 1 

sol, rex C Capsosiphon fulvescens CB M cb BC 1 microscopic 

rex N Codium fragile* (NE Pacific form) Green Exp nep SeAk 1 epi=C. sinicola 

gi I Codium fragile 

subsp. tomentosoides Green Exp ws Wa. 2 MB needed for subsp. 

sol, rex C Halochlorococcum moorei Val, Cor H cb BC 1 microscopic, endophytic 

mid, sol C Kornmannia leptoderma non 

zostericola (epilithic) Cor H cb N. Atl. 2 Culture work needed 

mid, sol N Monostroma fractum Gro M nep Wa. 2 Culture work needed 

Abbreviations: (see earlier charts) 

* = Recently also reported in O’Clair et al. , 1996, morph.=morphological studies 

from my earlier EVOS collections 

rex=range extension 

Category=preliminary decisions based on 

sol=species overlooked 

morphological and distributional features 

Stat=residency status 

and the literature available 

Type=habitat type 

ex=except 

Exp=exposed cobble 

C=Certainty of Idenfication 

F=on floats 

1=absolute certainty 

H=harbor, on cobble 

2=Morphological identity but additional 

gi=geographic isolation study (eg, MB or cultures) needed 

I=likely introduction 

3=Vegetative morphology similar, but reproductive 

ID=identification or MB data needed for positive identification 

M=mud 

MB=molecular biological study 

mid=earlier misidentification


• Are any of the species taxonomically problematic Such problematic species often end up in 

new records lists; and, indeed, several of the new records are problematic species that require 

further study for positive identification. Investigations are currently in progress for 5 of these 

species. 

• Could the species have been mistaken for other similar species in the past Species that 

resemble one another can be confused easily. Often these mistakes are not revealed until a 

species is newly illustrated or described. In these cases, misidentifications and distributions 

could easily be corrected with herbarium searches. In the list, at least 6 species fall into this 

category, including at least 1 undescribed species. 

• Has small size or habitat restriction influenced the species discovery Microscopic species 

are frequently overlooked as are species from unusual habitats. On the list, 4 species are 

microscopic, and 1 occurs in the unlikely habitat of a high marsh. 

• Is the species new to the area through range extension or through an actual introduction For 

marine plants, historical (baseline and fossil) information, geographic isolation, and 

molecular data are appropriate for proving the latter. 

The final justification for categorizing a species of marine plant as introduced (Table 9C1.12) 

was based on many of these factors, but remains tentative. All 5 species listed as introduced are 

geographically isolated. Nine other species are northward range extensions from southeast 

Alaska or British Columbia, and these species are tentatively identified as native. However, 

these range extensions could be caused by either natural dispersal, possibly caused by El Niño 

events of the past few years, or they could be introduced with aquaculture transports. 

Of the 21 new records, at least 5 are at this time very strongly supported for NIS status 

based on their geographic isolation, and this is a very conservative estimate. To further confirm 

the status of these, molecular biological proof of identifications are currently in progress. 

Comments on the 5 Probable Introductions and on 1 Important Failed Introduction. 

Additional description of each of the most probable introduced species and their habitats and 

distribution are provided below: 

Chroodactylon ramosum. Chroodactylon is a microscopic primitive red alga that is typically 

bright bluegreen in color. Its uniseriate, dichotomously branched filaments are unmistakable 

under the microscope. Although common to the North Atlantic in both Europe and North 

America, in the Pacific it is only known from Japan, southern Australia, and southern California. 

Because this alga generally occurs in estuarine or freshwater habitats (Vis and Sheath 1993), its 

occurrence in the turf algae of the oyster floats at Tatitlek was a surprise, except that it could 

have been brought in with oysters. The lack of records for this species in the well-worked 

marine and estuarine environments of British Columbia and Washington indicates that it is truly 

an isolated population and in all likelihood introduced. 

Codium fragile subsp. tomentosoides and the northeast Pacific complex*. The normal range of 

the native species complex of Codium fragile in the northeast Pacific is from Baja California to 

southeast Alaska. This complex appears to consist of several unnamed subspecies (C. 

Trowbridge, pers. comm.). Separate from this is an alien subspecies called tomentosoides that 

has been reported to occur in San Francisco Bay. This alien subspecies is differentiated from the


native complex by having a different branching frequency and more rounded and mucronate 

utricle tips. At Green Island, two different subspecies of Codium appear to occur. The low 

intertidal form appears to be identical to the native complex. Its utricle tips are sharply pointed 

and it is fairly tightly branched. The second form occurs in the mid to upper intertidal and is 

more loosely branched with very short mucronate tips, nearly identical to subsp. tomentosoides. 

However, experts in the field (Silva and Max) have told me after considerable hesitancy, that 

neither are truly subsp. tomentosoides, and that both fall clearly within the native complex. This 

indicates that the Green Island Codium is probably a range extension or an introduction from 

southeast Alaska or from Washington, Oregon, or California. Perhaps studies on its epiphyte 

(discussed below) will enable us to detect its true source. (* Recently O’Clair et al., 1996, also 

noted that Codium occurs on Green Island. This record appears to be based on G.I. Hansen’s 

earlier EVOS project collections, now located in Juneau.) 

Ceramium sinicola. This Ceramium species was an epiphyte of Codium fragile at Green Island. 

The species, unlike Ceramium codicola, does not have bulbous rhizoids. It is completely 

corticated except for some slightly broken cortication near the tips like in the southern California 

species Ceramium sinicola. The morphology of the plants most closely matches the descriptions 

of Dawson (1950) and Setchell and Gardner (1924) for C. sinicola, and male, female, and 

tetrasporic specimens have all been observed. Its occurrence in Alaska is extremely unlikely 

unless it is a recent introduction. This past year I have been working with Mr. Tae Oh Cho, who 

is monographing the world species of Ceramium with both morphological and molecular 

techniques. He has agreed to look at this material for me and will also look at my Alaskan 

material of Ceramium rubrum which he feels may actually be C. kondoi, a Japanese/Korean 

species, and possibly another introduction to Alaska. 

Fucus cottonii. This species is unrecorded for the North Pacific, and yet it occurred in nearly all 

of the high mudflat/marsh areas visited during the June 1998 cruise of Prince William Sound. 

The plant was first observed in unpublished notes by G.I. Hansen on Vancouver Island in 1981 

and then at several Prince William Sound and Kenai sites during the EVOS studies. In some 

areas it dominated the supralittoral zone extending even into the terrestrial. At Cloudman Bay it 

occurred 100 meters away from the bay on stream banks intermixed with mosses and vascular 

plants. The plants in Alaska are mat-forming and either loose-lying on mud, entangled with 

other algae (such as Fucus gardneri), or intermixed with terrestrial plants. They range from 1-5 

cm in height. The blades are dichotomously branched and often terete and only 1-3 mm in 

diameter. In some habitats, they become flattened, still without a visible midrib, and up to 5 mm 

in diameter. No receptacles were found during the June trip, but during the EVOS studies G.I. 

Hansen found a number of plants with relatively small (up to 2 cm long), elongate, somewhat 

pointed receptacles with conceptacles and oogonia bearing 8 eggs. In some areas, the extent of 

the mats of this small fucoid makes me question its form of reproduction. Since receptacles are 

so uncommon, propagation of the mats must be by fragmentation and vegetative growth, an 

advantageous feature for dispersal. 

There is some question as to the use of Fucus cottonii (= F. muscoides) as a valid species. 

Fletcher (1987) considers the species as a high marsh ecad of Fucus vesiculosis, an Atlantic and 

Arctic species, but others have accepted F. cottonii as a distinct species (Guiry, 1998). To 

confirm the validity of the species and my designation of the Prince William Sound material,


samples are being sent to Esther Serrao in Portugal, who is studying the phylogenetic 

relationships of the genus with molecular techniques. 

Microspongium globosum. This tiny brown alga was found growing epiphytically on 

Delamaraea attenuata on the floats at Tatitlek. The thalli were abundant and bore plurilocular 

sporangia that clearly match the diagrams for this species in Fletcher (1987). Known only from 

the North Atlantic and Japan, the species makes a surprising appearance in Alaska. It also has 

not been reported from the well investigated areas to the south. Its occurrence on the oyster 

floats at Tatitlek as an epiphyte on another new record to Alaska indicates to me that this species, 

possibly along with its host, is another new introduction to the area. However, its vector could 

also have been oysters from Japan. 

Macrocystis integrifolia. Since 1979 (Jay Johnson, Alaska Fish and Game, pers com.) 

Macrocystis has been imported (by plane) from southeast Alaska to Prince William Sound to be 

used as substrate for herring eggs in the lucrative Herring-Roe-On-Kelp (HROK) fishery. 

Normally only blades and fronds of the giant kelp are transported northward for the fishery. 

These are then placed in impoundment nets which house both the kelp and the fish. The eggladen 

blades are then harvested and sold primarily to Japan as a gourmet food item. 

Theoretically, the blades that are brought up to the Sound are clean (the most desirable for 

HROK) and are all harvested for later sale. However, during our June trip and during many of 

Hansen’s earlier trips to the area, blades and holdfasts of the kelp that had escaped were found 

adrift in Prince William Sound. Perhaps due to the climate, none of these plants appear to have 

propagated in the area since none have ever been found attached anywhere north of southeast 

Alaska. Hence, in the site list the species is listed as a failed introduction. However, even with 

the transport of “eye-clean” blades, it is likely that numerous small algal and animal species are 

co-transported accidentally from southeast Alaska to Prince William Sound every year with this 

kelp. This may account, as much as the current El Niño, for many of our new range extensions. 

Discussion 

During the search for marine plant NIS in Prince William Sound, it was important to 

characterize the flora at each of the sites so that the probable introduced species and their impacts 

on the community could be recognized. In addition, information on the taxonomic and residency 

status composition of these communities was absolutely essential to be able to determine 

vulnerable sites for future invasions. Although 155 species of plants collected during the 1998 

cruise is probably only about half that of the actual flora of Prince William Sound, the data 

compiled reveal important trends in community composition. In addition, the final lists of new 

records and probable introductions give valuable insight into the difficulties of recognizing NIS. 

Limitations of the data. Although 19 sites were visited during the June 1998 survey, the 

time allowed for sampling was inadequate at many of the beaches. This had a substantial impact 

on the overall data. Moreover, the lack of year-round collections for the area limits the results in 

ways that cannot even be predicted. In terms of numbers, this can be shown clearly by 

comparing the total count of 112 species shown for Port Valdez in the Checklist (which includes 

seasonal collections) with the count of 47 for the area obtained during this short trip.


Information derived from additional collections and herbarium specimens from our sites would 

help to overcome these difficulties and to improve the resolution of the data. 

Although temperature and salinity influenced the total species counts for marine algal 

species, this could not be demonstrated clearly with the data on hand. Prince William Sound has 

regions heavily effected by rain, snow and ice melt, and marked changes in temperature and 

salinity occur throughout the year. During summers, salinity is the lowest as runoff produces a 

freshwater lens on the surface of many of the bays, including Port Valdez and Whittier. In these 

areas intertidal species are subjected to wide salinity fluctuations with the tidal cycle. Since these 

physical factors were measured only during the limited sampling periods, they do not reflect the 

range of conditions encountered over time by the intertidal species sampled. 

Limited historical knowledge of the flora and new records. Only two floristic papers 

on the marine algae (and plants) of Prince William sound have ever been published (Calvin and 

Lindstrom, 1980; Wiegers et al., 1997). In addition, an overall identification guide to the marine 

algae does not exist for Prince William Sound or even for Alaska, and we are left with using an 

assortment of references from neighboring areas to identify species. This lack of both taxonomic 

information and baseline data for Prince William Sound is clearly evident in the discovery of 21 

new records to the area amounting to 13.5% of the species collected during our short 9 day 

cruise. During our earlier Pilot Study, an additional 3 new records were found in Port Valdez 

alone. Though fairly evenly distributed among the 3 major taxonomic groups, there were a few 

more new records among brown and green algae than among the red, and the majority of the 

species appeared to be cryptogenic. In addition, new record species were slightly more abundant 

at certain sites. Green Island, the most diverse site in the study, bore 6 new records, while the 

float sites combined bore 9. Both of these areas (probably along with many others in Prince 

William Sound) appear to have been understudied in the past. Since, for this study, our probable 

NIS were derived from the new records, it is understandable that each of these 2 sites (or site 

types) also bore 2 of the 5 probable NIS designated in the study. 

Taxonomic composition. In nearly all temperate outer-coastal habitats, the red algal 

species are the highest in numbers followed by the brown and then the green algae forming a 

R>B>G hierarchical pattern of dominance. Proceeding from open coasts into protected bays and 

estuaries, the ratio changes to reflect a reduction in the number of red algae. For instance, off the 

coast of Oregon, the R:B:G ratio is 61:22:17. In Prince William Sound, the overall ratio of 

R:B:G in the species surveyed was 47:33:20, a ratio probably indicating the influence of 

sheltered and less saline water. However, the overall composition pattern was still R>B>G 

(Table 9C1.13). The R>B>G dominance pattern in Prince William Sound occurred only at 

Rocky Headlands and Reefs and in Rocky Bays, all areas of moderate to high water movement 

(exposure) and relatively uniform salinity and temperature supporting established communities 

with numerous annuals and perennials. In Harbors, the proportion of green algae increased and 

the pattern became R>G>B, reflecting the tolerance of green algae for lower salinities found in 

this habitat. In addition, since many of the green and brown algae are ephemeral (opportunistic), 

they can survive the wide fluctuations in temperature and salinity. Moreover, since ephemeral 

forms are often fouling organisms, many are repeatedly brought in to seed these areas by boat 

traffic. In the mud bays and on the floats, the proportion of brown algae increased. In mud bays, 

the frequent shifts in mud level smothers many of the species, providing niches primarily for


ephemerals and unattached forms. On the oyster floats, the early successional ephemeral forms 

are also encouraged due to the periodic cleaning of the habitat. The higher and generally more 

uniform salinity of both of these habitats appears to enable the ephemeral browns to outcompete 

the ephemeral greens. 

Table 9C1.13. Summary of the Hierarchical Composition 

and Physical Features of each Habitat Type 

Observed during the June 1998 Survey 

Habitat Types 

Harbors Mud Bays Rk Headlands Rk Bays Floats 

Composition 

and Reefs 

Taxonomic* R>G>B B=R>G R>B>G R>B>G B>R>G 

variable variable variable 

Residency Status C>N C>N N>C N>C C>N 

Exposure low low high low med-high 

Salinity variable low-med high variable high 

Temperature variable variable uniform uniform uniform 

Substratum variable soft hard hard hard 

* = includes only the 3 major taxonomic groups sampled. 

Resident type composition. Cryptogenic species predominated in the more disturbed 

and variable habitats of the Harbors, Mud Bays, and Floats, while the native species 

predominated in the less disturbed and more uniform habitats provided by Rocky Headlands and 

Reefs, and Rocky Bays. Cryptogenic algal species appear to contain a high percentage of 

ephemeral forms. Hence, their ability to survive in fluctuating environments and perhaps in 

ballast water and on ship bottoms is high. 

Introduced species and their impact. The 5 probable plant NIS discovered during our 

survey are all isolated (and probably young) populations. Although four of these species do not 

appear to have wide distribution in Prince William Sound, Fucus cottonii does appear to have an 

expanding range. It was found at 4 of our sites and appears to be prevalent in the supra-littoral of 

all of these areas. Unique to sloughs and the marsh area of mudflats, this species does not seem 

to be replacing any of the known marine or estuarine species. However, in Cloudman Bay, it 

may actually be out-competing some terrestrial plants. Fortunately, none of the probable NIS 

plants found in Prince William Sound appear to be hazardous to the environment. None are as 

toxic or as invasive as the Mediterranean introduction Caulerpa taxifolia (Lemee et al., 1993; 

Verlaque and Fritayre, 1994). 

The transport mechanisms of these introductions is only partially clear. The two species 

(Chroodactylon ramosum, Microspongium globosum ) found on oyster floats could have been 

brought into the area with the transplantation of oysters for aquaculture purposes. The vector for 

Codium and its epiphyte Ceramium is more debatable. The subspecies Codium fragile fragile 

was potenitally transported up from southeast Alaska with Macrocystis for the HROK industry. 

But the only method of transport for the subspecies Codium fragile tomentasoides would have to 

be either ballast water or as fouling on the hulls of ships. The importation mechanism of Fucus 

cottonii is even less clear. Its relatively widespread occurrence in Prince William Sound (and in 

patchy spots along the west coast) indicates that it is probably not a recent introduction. 

However, since it is a predominantly unattached species, it is also an excellent candidate for 

transport by ballast water.


Other potential introductions and their significance. What of the other 70 cryptogenic 

species which are possibly introductions, but which have less obvious characteristics of 

invasion These species are, by definition, wide-ranging and many are abundant, often heavily 

impacting the communities in which they occur. Proof of the NIS status of these prominent 

species is possible, but it will require detailed comparative morphological study and world-wide 

molecular biological tracking of their distributions. Furthermore, knowledge of the impacts of 

these species on community structure will demand complex physiological and ecological studies 

of the species in both their introduced and native habitats. These studies are important projects 

for future investigators who are concerned about the conservation of our native biodiversity. 

References 

Abbott, I. A., and G. J. Hollenberg. 1976. Marine Algae of California. Stanford University 

Press, Stanford. xii+827 pp. 

Adams, N. M. 1983. Checklist of marine algae possibly naturalized in New Zealand. N. . J. 

Bot. 21: 1-2. 

Adams, N. M. 1994. Seaweeds of New Zealand, an illustrated guide. Canterbury University 

Press Publ., New Zealand. 360 pp. 

Calvin, N. I., and S. C. Lindstrom. 1980. Intertidal algae of Port Valdez, Alaska: species and 

distribution with annotations. Bot. Mar. 23: 791-797. 

Carlton, J. T. 1996. Biological invasions and cryptogenic species. Ecology 77 (6): 1653-1655. 

Chapman, J., and G. Hansen. 1997. Surveys of nonindigenous aquatic species for Port Valdez, 

Alaska. In: Ruiz, G.M. and A.H. Hines. 1997. Patterns of nonindigenous species transfer and 

invasion in Prince William Sound, Alaska: Pilot Study. Report, Prince William Sound Regional 


Dawson. E. Y. 1950. A review of Ceramium along the Pacific coast of North America with 

special reference to its Mexican representatives. Farlowia 4: 113-138. 

DeWreede, R. E. 1996. The impact of seaweed introductions on biodiversity. Global 

Biodiversity 6: 2-9. 

Fletcher, R. L. 1987. Seaweeds of the British Isles. Vol. 3. Fucophyceae (Phaeophyceae). Part 

1. British Museum (Natural History), London. 359 pp. 

Gabrielson, P. W., R. F. Scagel, and T. B. Widdowson. 1989. Keys to the benthic marine algae 

and seagrasses of British Columbia, southeast Alaska, Washington and Oregon. Phycological 

Contribution Number 4. Dept. of Botany, University of British Columbia, Vancouver. vi+187 

pp. 

Guiry, R. 1998. An internet accessible “taxonomic database” on “seaweeds” (primarily of 

Europe). http://seaweed.ucg.ie.


Hansen, G. 1997. A revised checklist and preliminary assessment of the macrobenthic marine 

algae and seagrasses of Oregon. Pp. 175-200 in Kaye, T., A. Liston, R. Love, D. Luoma, R. 

Meinke, and M. Wilson (ed.). Conservation and Management of Native Flora and Fungi. Native 

Plant Society of Oregon, Corvallis. 

Hansen, G. I., D. J. Garbary, J. C. Oliveira, and R. F. Scagel. 1981. New records and range 

extensions of marine algae from Alaska. Syesis 14: 115-123. 

Lee, R. K. S. 1980. At catalogue of the marine algae of the Canadian Arctic. Publications in 

Botany, No. 9. National Museums of Canada, National Museum of Natural Sciences, Ottawa, 

ON. 82 pp. 

Lemee, R., D. Pesando, M. Durand-Clement, A. Dubreuil, A. Meinesz, A. Guerriero, and F. 

Pietra. 1993. Preliminary survey of toxicity of the green alga Caulerpa taxifolia introduced into 

the Mediterranean. J. Appl. Phycol. 5:485-493. 

Lindstrom, S. C. 1977. An annotated bibliography of the benthic marine algae of Alaska. 

ADF&G Technical Data Report No. 31. Juneau. 172 pp. 

Nelson, W. A., and C. A. Maggs. 1996. Records of adventive marine algae in New Zealand: 

Antithamnionella ternifolia, Polysiphonia senticulosa (Ceramiales, Rhodophyta), and Striaria 

attenuata (Dictyosiphonales, Phaeophyta). N. Z. J. Mar. Freshwat. Res. 30: 449-453. 

O’Clair, R. M., S. C. Lindstrom, I. R. Brodo. 1996 [1997]. Southeast Alaska’s Rocky Shores: 

Seaweeds and Lichens. Plant Press, Auke Bay. 

Perestenko, L. P. 1994. Red Algae of the Far-Eastern Seas of Russia. Komarov Botanical 

Institute, Russian Academy of Sciences, St. Petersburg. 331 pp. [In Russian]. 

Phillips, R. C., and E. G. Menez. 1988. Seagrasses. Smithsonian Contributions to the Marine 

Sciences 34. v+104 pp. 

Rueness, J. 1977. Norsk Algeflora. Universitetsforlaget, Oslo. 266 pp. [In Norwegian]. 

Ruiz, G. M., and A. H. Hines. 1997. The risk of nonindigenous species invasion in Prince 

William Sound associated with tanker traffic and ballast Water Management: Pilot Study. 

Regional Citizens’ Advisory Council of Prince William Sound RFP Number 632.97.1. 47 pp + 

52 pp tables and graphs. 

Scagel, R. F., P. W. Gabrielson, D. J. Garbary, L. Golden, M. W. Hawkes, S. C. Lindstrom, J. C. 

Oliveira, and T. B. Widdowson. 1989 [1993]. A Synopsis of the Benthic Marine Algae of 

British Columbia, southeast Alaska, Washington and Oregon. Phycological Contribution 

Number 3. Dept of Botany, University of British Columbia, Vancouver, BC. 535 pp.


Sears, J. R. 1998. NEAS Keys to the Benthic Marine Algae of the Northeastern Coast of North 

America from Long Island sound to the Strait of Belle Isle. NEAS Contribution Number 1, 

Dartmouth, MA. xi+161pp. 

Selivanova, O. N., and G. G. Zhigadlova. 1997. Marine algae of the Commander Islands: 

preliminary remarks on the revision of the flora. Bot. Mar. 40: 1-24. 

Setchell, W. A., and N. L. Gardner. 1924. Expedition of the California Academy of Sciences to 

the Gulf of California in 1921. Proc. of the Cal. Acad. Sci., 4 th Series, 12: 12-88. 

Verlaque, M., and P. Fritayre. 1994. Mediterranean algal communities are changing in face of 

the invasive alga Caulerpa taxifolia (Vahl) C. Agardh. Oceanol. Acta 17: 659-672. 

Vis, M. L., and R. G. Sheath. 1993. Distribution and systematics of Chroodactylon and 

Kyliniella (Porphyridiales, Rhodophyta) from North American streams. Jap. J. of Phycology 41: 

237-241. 

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regional multiple-stressor ecological risk assessment for Port Valdez, Alaska. IETC No. 9701 

and RCAC 1033.102. Inst. of Environmental Toxicology and Chemistry, Western Washington 

Univ., Bellingham, WA. 

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Government Printer, South Australia, 329 pp. 

Womersley, H. B. S. 1987. The marine benthic flora of southern Australia, Part 2. Australian 

Government Printing Division, Adelaide, 484 pp. 

Womersley, H. B. S. 1994. The marine benthic flora of southern Australia. Part 3A. Australian 

Biological Resources Study, Canberra, 508 pp. 

Womersley, H. B. S. The marine benthic flora of southern Australia. Part 3B. Australian 

Biological Resources Study, Canberra. 392 pp. 

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Phycol. 43: 115-171. [In Japanese].

Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 1 

Chapter 9C2. Focal Taxonomic Collections: Planktonic Cnidaria, Ctenophora, and 

Pelagic Mollusca 

Claudia Mills, Friday Harbor Laboratories, University of Washington 

Methods 

Medusae, ctenophores, and pelagic molluscs were collected at sites in both Prince William 

Sound and Cook Inlet from August 8–14, 1999 using small plankton nets and a water scoop attached 

to a long handle. Specimens were examined in the field, relaxed, and fixed for transport to the 

laboratory. Specimens were reexamined microscopically at the Friday Harbor Laboratories in 

September 1999, in order to verify or assign species names. 

Results 

All pelagic Hydrozoa, Scyphozoa, Ctenophora and Mollusca are identified by site in Table 

9C2.1. Separate species lists for these groups follow for Prince William Sound (Table 9C2.2) and 

Cook Inlet (Table 9C2.3). An annotated species list follows (Table 9C2.4), including all species on 

both lists. In the time allotted to this project, I do not feel that I completed a comprehensive search 

of the literature for species previously collected in Prince William Sound and Cook Inlet, but no 

other papers came to mind. I also did not search for unpublished data at the University of Alaska. 

A few coelenterates were conspicuously missing from the region. We saw no 

stauromedusae, no Epiactis anemones on eelgrass, no Chrysaora or Phacellophora 

scyphomedusae, and no Anthopleura elegantissima or A. xanthogrammica. 

No known nonindigenous species of planktonic Cnidaria or Ctenophora were collected in 

Prince William Sound or Cook Inlet by our scientific teams in either 1998 or 1999. In the 1999 

expedition, 15 species of Hydrozoa were collected (including 3 hydroids [see section 9D. Fouling 

Communities) for more thorough hydroid work-up] and 14 species of hydromedusae), two 

scyphomedusae and unidentified scyphozoan polyps (scyphistomae), and two species of 

ctenophores. Two molluscan species were also taken in the water column. 

The following species appear to be new records in the Prince William Sound region: 

Hydromedusae 

*Aequorea aequorea 

*Aequorea victoria 

*Clytia gregaria (=Phialidium gregarium) 

Eperetmus typus 

Euphysa sp. 

Gonionemus vertens 

Halitholus sp. 

*Melicertum octocostatum 

*Proboscidactyla flavicirrata 

Sarsia spp. 

Tiaropsis multicirrata


Ctenophora: 

*Bolinopsis infundibulum 

*Pleurobrachia bachei 

New NAME for common Scyphomedusa 

Aurelia labiata 

* indicates common species whose presence in PWS may be known, but I have not seen reports 

in print. Dr. Jennifer Purcell (Horn Point Laboratory, University of Maryland) is working with 

some of these, but her results are unpublished as yet. 

Following the annotated species list is a list and discussion of nonindigenous cnidarian 

species already present in some west coast estuaries that might be positioned to ultimately invade 

locations in Alaska. This list is accompanied by an Appendix (following the report) titled 

“Commentary on species of Hydrozoa, Scyphozoa and Anthozoa (Cnidaria) sometimes listed as 

non-indigenous in Puget Sound”, reprinted from Cohen et al. (1998). References are given at the end 

of the main report as well as the Appendix. 

References 

Arai, M. N. and A. Brinckmann-Voss, 1980. Hydromedusae of British Columbia and Puget Sound. 

Can. Bull. Fish Aquat. Sci., 204: 192 pp. 

Bigelow, H. B. 1912. The ctenophores. Bull. Mus. Comp. Zool., 56: 369-404, 2 pls. 

Bigelow, H. B. 1913. Medusae and siphonophorae collected by the U. S. Fisheries steamer 

“Albatross” in the Northwestern Pacific, 1906. Proc. U. S. Nat. Museum, 44: 1-119, 6 pls. 

Bigelow, H. B. 1920. Medusae and Ctenophora. Rep. Canadian Arctic Exp. 1913-18, Southern Party 

1913-16, Volume VIII: Mollusks, Echinoderms. Coelenterates, etc., Part H: 3H-22H, 2 pls. 

Calder, D. R. 1988. Shallow-water hydroids of Bermuda: the Athecatae. Royal Ontario Museum Life 

Sciences Contributions, 148: 1-107. 

Cohen, A., C. Mills, H. Berry, M. Wonham, B. Bingham, B. Bookheim, J. Carlton, J. Chapman, J. 

Cordell, L. Harris, T. Klinger, A. Kohn, C. Lambert, G. Lambert, K. Li, D. Secord, and J. Toft, 

November 1998. Report of the Puget Sound Expedition, September 8–16, 1998: a Rapid 

Assessment survey of non-indigenous species in the shallow waters of Puget Sound. Washington 

State Department of Natural Resources, Olympia, Washington, 37 pages. 

Greenberg, N., R. L. Garthwaite and D. C. Potts, 1996. Allozyme and morphological evidence for 

a newly introduced species of Aurelia in San Francisco Bay. Marine Biology, 125: 401-410. 

Harbo, R. M. 1999. Whelks to Whales: Coastal Marine Life of the Pacific Northwest. Harbour 

Publishing, Madeira Park, B.C., Canada.


Kramp, P. L. 1961. Synopsis of the medusae of the world. J. Mar. Biol. Assoc. U.K., 40: 1–469. 

Mills, C. E. 1981. Seasonal occurrence of planktonic medusae and ctenophores in the San Juan 

Archipelago (NE Pacific). Wasmann J. Biol., 39: 6-29. 

Mills, C. E. 1998 to present. Web Site: http://faculty.washington.edu/cemills/ 

Mills, C. E. and F. Sommer, 1995. Invertebrate introductions in marine habitats: two species 

of hydromedusae (Cnidaria) native to the Black Sea, Maeotias inexspectata and Blackfordia 

virginica, invade San Francisco Bay. Marine Biology, 122: 279-288. 

Murbach, L. and C. Shearer. 1903. On medusae from the coast of British Columbia and 

Alaska. Proc. Zool. London 2:164-192, pls. 17-22. 

Purcell, J. E. 1998 Project report 98163S - Jellyfish as competitors and predators of fishes. 

On the web at http://www.uaa.alaska.edu/enri/apex/98163S.html. 

Ricketts, E. F. and J. Calvin, 1939. Between Pacific Tides. Stanford University Press, Stanford. 

Wrobel, D. and C. Mills, 1998. Pacific Coast Pelagic Invertebrates: a Guide to the Common 

Gelatinous Animals. Sea Challengers and the Monterey Bay Aquarium, Monterey, California.


Table 9C2.1. Hydrozoa, Scyphozoa, Ctenophora and pelagic Mollusca by site, 1999. 

Collections and identifications by Claudia E. Mills. 

1999 PWS EXPEDITION 

C.E. MILLS REPORT 

Homer 

Sadie 

Cove 

Seward 

Marina 

Lowell 

Pt 

Seward 

south of 

Esther 

Is. 

Whittier 

Fairmoun 

t Bay 

HYDROZOA 

X 

Aequorea aequorea v, albida X X X X X X 

Aglantha digitale 

X 

Bougainvillia superciliaris 

x 

Clytia gregaria 

X X X X X 

(=Phialidium gregarium) 

Eperetmus typus X X 

Euphysa sp. 

X 

Eutonina indicans 

X 


X 


X 

Leuckartiara sp. 

X 

Melicertum octocostatum X X X 

Mitrocoma cellularia 

X 

Obelia hydroids X X X 

Proboscidactyla flavicirrata X + 

polyps 

X X X 

Sarsia/Coryne sp. hydroids X 

Sarsia spp. medusae X X X X 

SCYPHOZOA 


X 

Cyanea capillata X X X X X X X 

unidentified scyphistomae 

(probably Aurelia sp.) 

X 

X 

Valdez 

Mar 

ina 

Tatitlek 

Bus 

-by 

Is. 

Cord 

ova 

CTENOPHORA 

Bolinopsis infundibulum 

Pleurobrachia bachei 

X 

X 

MOLLUSCA 

Melibe leonina X X X 

Clione limacina X


Table 9C2..2. Prince William Sound species list. See Table 1 for specific locations. 

Collections and identifications by Claudia E. Mills, unless otherwise noted. 

HYDROZOA 

REFERENCE 

Aequorea aequorea v. albida PWS 99; Purcell, 1998 

Aequorea victoria/ A. aequorea v. aequorea Purcell, personal communication 1999 

Catablema multicirrata Bigelow, 1913 

Clytia gregaria PWS 99 


Eperetmus typus PWS 98, PWS 99 

Euphysa sp. PWS 99 

Gonionemus vertens PWS 99 

Halitholus sp. PWS 99 

Melicertum octocostatum PWS 99 

Obelia longissima PWS 98 

Obelia spp. hydroids PWS 99 

Proboscidactyla flavicirrata PWS 99 

Sarsia spp. medusae PWS 99 

Staurophora mertensii Bigelow, 1913 

Tiaropsis multicirrata PWS 98 

SCYPHOZOA 

Aurelia"aurita" (Mills quotes) Purcell, 1998 

Aurelia labiata PWS 99 

Cyanea capillata PWS 99; Purcell, 1998 

Unidentified scyphistomae PWS 99 


CTENOPHORA 

Bolinopsis infundibulum PWS 99 

Pleurobrachia bachei PWS 99; Purcell, 1998 

MOLLUSCA 

Melibe leonina PWS 99 

* PWS 98 refers to specimens collected by Ruiz et al., June 1998 in Cook Inlet. 

PWS 99 refers to specimens collected by Greg Ruiz et al., August 1999 in Cook Inlet.


Table 9C2.3. Cook Inlet species list. 

Collections and identifications by Claudia E. Mills, unless otherwise noted. 

HYDROZOA 

*REFERENCE AND LOCATION 

Aequorea aequorea v. albida PWS 99 - Homer Marina 


PWS 99 - Sadie Cove, Katchemak Bay 

Bougainvillia superciliaris PWS 99 - Sadie Cove, Katchemak Bay 

Clytia gregaria 






PWS 99 - Homer Marina 



Melicertum octocostatum 




Obelia sp. hydroids 


Proboscidactyla flavicirrata hydroids PWS 99 - Homer Marina 

Sarsia/Coryne sp. hydroids PWS 99 - Homer Marina 

Sarsia spp. medusae 

PWS 99 - Homer Marina, Sadie Cove 

SCYPHOZOA 

Cyanea capillata 

Unidentified scyphistomae 


PWS 99 - Homer Marina, Sadie Cove 


CTENOPHORA 

(none) 

MOLLUSCA 

Clione limacina PWS 99 - Homer marina 

Melibe leonina 

PWS 99 - not collected but told of site at Jakalof Bay 

by Carmen Field 

* PWS 99 refers to specimens collected by Ruiz et al., August 1999 in Cook Inlet.


Table 9C2.4 

PRINCE WILLIAM SOUND ANNOTATED SPECIES LIST 

(combines both Cook Inlet and Prince William Sound locations) 

HYDROZOA 

Aequorea aequorea var. albida 

Distribution. Most of the Aequorea medusae that we saw were beached. Such specimens were seen at the 

Homer Marina, Lowell Point in Seward, the Whittier Marina, and in Cordova and Valdez. A few were 

seen in the water while underway south of Esther Island, along with Cyanea capillata. 

Remarks. This name was applied by Bigelow (1913) to Aequorea specimens measuring 120 mm and 

165 mm bell diameter, collected in Dutch Harbor. Such very-large Aequorea occur throughout southern 

Alaska, and are accompanied in some places by smaller specimens that seem very similar to Aequorea 

victoria at Friday Harbor (called Aequorea aequorea var. aequroea by Bigelow, 1913). Whether they 

are different sizes of the same species or 2 different species has still not been resolved (even the modern 

use of "A. victoria" as species name for Friday Harbor medusae is controversial). Only large-sized 

specimens (most 120-160 mm diameter) were seen in Prince William Sound in August 1999. Similar 

large Aequoreas in Prince William Sound were called A. victoria by Purcell (1998). 

Aequorea victoria or Aequorea aequorea var. aequorea 

Remarks. We did not collect any smaller specimens of Aequorea, but I am told by Dr. Jennifer Purcell, 

who has been doing a recent plankton study in Prince William Sound that small Aequoreas that look like 

those at Friday Harbor are also present. Bigelow (1913) calls these A. aequorea var. aequorea. Arai 

and Brinckmann-Voss later applied the name A. victoria to the same animals. It is not clear to me that 

A. victoria is not a junior synonym to A. aequorea. 


Distribution. Several Aglantha digitale medusae were collected at the head of Sadie Cove, 

Katchemak Bay, on August 8, 1999, by dipping from a small boat in about 8 feet of water. Most were 

within a layer of fresher water that occupied the upper 15" of the water column and were dead and 

decomposing. Many others were seen, but not collected. 

Remarks. There is no question about the identification of this material, although why these medusae 

were in the layer of low salinity water is not clear. This circumpolar species is well known in the 

North Pacific, North Atlantic and Arctic Oceans, including the Bering Sea. 

Bougainvillia superciliaris 

Distribution. Five Bougainvillia superciliaris medusae were collected at the head of Sadie Cove, 

Katchemak Bay, on August 8, 1999, by dipping from a small boat in about 8 feet of water. All were 

below a layer of fresher water that occupied the upper 15" of the water column. Several others were 

seen, but not collected. 

Remarks. These 6-12 mm high specimens best correspond with the description of Bougainvillia 

superciliaris, having its characteristic prominent peduncle above the manubrium. The Sadie Cove 

specimens had 36-40 tentacles on each of the four marginal bulbs, which is quite a bit higher than the


10-22 tentacles described for B. superciliaris in Kramp (1961). Bigelow (1913) found a single specimen 

of B. superciliaris of the same size off Attu Island in the Aleutians, but also with less than 20 tentacles 

in each group. This tentacle number discrepancy leads to the question about species identification for the 

Sadie Cove material. 

Catablema multicirrata 

Distribution. This species (2 medusae) was collected by Bigelow (1913) off Orca in Prince William 

Sound on July 19, 1906. 

Remarks. We did not find it in August 1999, but did not sample at that location. 

Clytia gregaria (=Phialidium gregarium) 

Distribution. Many individuals of this species were collected in Sadie Cove, Katchemak Bay and in 

Fairmount Bay, Tatitlek, off Busby Island and in the Cordova Marina. 

Remarks. Most of the specimens correspond well to the description of Clytia gregaria (as Phialidium 

gregarium) in Kramp (1961), with about 40 tentacles and a few rudimentary bulbs alternating with 

marginal vesicles in 12 mm diameter medusae; the gonads were on the distal 1/2 of the radial canals. 

Some smaller medusae (7 mm diameter), with a few less tentacles and shorter gonads may be C. 

gregaria, or could be C. lomae, looking very similar to specimens collected in September 1998 in Puget 

Sound by Claudia Mills and Erik Thuesen. Seasonal morphological variation with changes in 

zooplankton prey availability have not been described, so it is difficult to be positive about the species 

name in some cases. The genus name Clytia has typically been applied only to the hydroid form, but it is 

an older genus name than Phialidium, and should be applied to both phases of the life cycle. 


Distribution. One young Eperetmus typus medusa was collected at the head of Sadie Cove, 

Katchemak Bay, on August 8, 1999, by dipping from a small boat in about 8 feet of water. It was 

below a layer of fresher water that occupied the upper 15" of the water column. Three more Eperetmus 

typus medusae were collected in Fairmount Bay, Prince William Sound, in vertical plankton tows 

taken off the side of the Kristina with Jeff Cordell's 130 µm mesh plankton net in about 70- 90 feet of 

water. 

Remarks. These 8-12 mm diameter immature specimens were very lively swimmers, that sank rapidly 

when they were not swimming. Both locations were protected coves and in both cases the animals may 

have been fairly near the bottom, but not enough is known to verify whether this species is indeed 

typical of protected coves and associated with the bottom in the same way as Gonionemus or 

Polyorchis. 

Euphysa sp. 

Distribution. Three small Euphysa sp. medusa were collected in Fairmount Bay, Prince William 

Sound, in vertical plankton tows taken off the side of the Kristina with Jeff Cordell's 130 µm 

mesh plankton net in about 70- 90 feet of water on August 10, 1999. 

Remarks. These 1.5–3.5 mm high medusae were found in the lab by Jeff Cordell in his plankton tow 

material. The two smaller specimens clearly had 3 larger tentacles and either one small tentacle or one


bare bulb. The larger specimen had 4 tentacles. There is insufficient information to assign them to 

species. 


Distribution. Six Eutonina indicans medusae were collected in the Homer Marina, Katchemak Bay, 

Cook Inlet on August 8, 1999. 

Remarks. I am surprised that we did not find more of this species. 


Distribution. More than 30 Gonionemus vertens medusae were seen in a dense eelgrass bed at low tide 

in front of Tatitlek, near the small town marina, on August 12, 1999. 

Remarks. These medusae emerged from within blades of eelgrass in the low intertidal as the tide came 

in. They were abundant. The pigmentation was less colorful and more brownish that specimens in the 

San Juan Islands. This species was previously not known north of Sitka (where it is mentioned by 

Ricketts and Calvin, 1939), except that probably the same species (as G. agassizii) was collected early 

this century by Trevor Kincaid in a salt lake on Unalaska Island, in the Aleutians (Murbach and 

Shearer, 1903). The same or a very similar species also occurs in Japan and the Russian Far East. 


Distribution. Three Halitholus sp. medusae were collected at the Tatitlek commercial (ferry) dock on 

August 11, 1999, in vertical plankton tows taken off the side of the Kristina with Jeff Cordell's 130 µm 

mesh plankton net in about 50 feet of water. Two more of these medusae were seen using a flashlight, 

but not collected, later the same evening from the small Tatitlek town marina at about midnight. 

Remarks. These 7-8 mm high specimens cannot be referred to any described species of Halitholus. 

They are very similar to both Halitholus sp. I and Halitholus sp. II of Arai and Brinckmann-Voss (1980, 

pp. 48-52), which were previously known from British Columbia and Washington State. 


Distribution. Four Leuckartiara sp. medusae were collected at the head of Sadie Cove, 

Katchemak Bay, on August 8, 1999, by dipping from a small boat in about 8 feet of water. All 

were below a layer of fresher water that occupied the upper 15" of the water column. Several 

others were seen, but not collected. 

Remarks. These approximately 15 mm-high specimens cannot be referred to any described 

species of Leuckartiara. They bear some resemblance to Leuckartiara foersteri of Arai and 

Brinckmann-Voss (1980, pp. 52-53), which is known from British Columbia and Washington 

State. They had 8 large tentacles and 12 small tentacles, with no additional rudimentary marginal 

bulbs. 

Melicertum octocostatum 

Distribution. About ten Melicertum octocostatum medusae were found in the Homer Marina, 

Katchemak Bay, Cook Inlet, Fairmount Bay and the Cordova Marina.


Remarks. This species is well known elsewhere in Alaska as well as in the North Pacific and Atlantic; it 

probably occurs throughout Prince William Sound. These specimens were relatively small, all being 

under 12 mm in bell height. 


Distribution. Homer Marina, Katchemak Bay, Cook Inlet on August 8, 1999. 

Remarks. Only a single small (15 mm diameter) specimen was collected. This specimen was only 

provisionally identified as M. cellularia until it was compared with a comparable-sized living M. 

cellularia in Friday Harbor. The small and large tentacles on the margin are the same, confirming the 

species identification. 

Obelia longissima (Pallas, 1766) 

Distribution. Various sites in Prince William Sound, summer 1998. 

Remarks. John Chapman sent all of the hydroids he collected in 1998 to Claudia Mills, who passed 

them on to Dr. Wim Vervoort of the Natural History Museum at Leiden, the Netherlands. Dr. Vervoort 

identified all of the hydroids that he saw as Obelia longissima. John Chapman has both the hydroids and 

their specific collection information. 

Obelia spp. hydroids 

Distribution. These hydroids were collected at least at the Seward Marina and at Lowell Point on 

August 10 and 11, 1999. 

Remarks. In my inexpert opinion, the blackened portions of some of the stems imply that these were 

probably Obelia longissima. I originally guessed that they might be Garveia franciscana, but they are 

not. They should be inspected by a hydroid specialist. 

Proboscidactyla flavicirrata 

Distribution. The hydroid of P. flavicirrata was found at the distal tips of several, 6 cm-long sabellid 

worm tubes in the Homer Marina, Katchemak Bay, Cook Inlet on August 8, 1999. This hydroid was 

actively producing medusa buds although no medusae were seen in this marina. Several P. flavicirrata 

medusae were collected in Fairmount Bay, Tatitlek and the Cordova Marina. 

Remarks. These medusae are very small; they probably occur throughout Prince William Sound. 

Sarsia/Coryne sp. hydroids 

Distribution. One or more clumps of hydroids that looked like Sarsia were collected by Jeff Goddard in 

the Homer Marina, Katchemak Bay, Cook Inlet on August 8, 1999. 

Remarks. I did not look carefully at this material. If it was reproductive and making medusa buds, it can 

be assigned to Sarsia; if it was reproductive and bearing fixed gonophores, it could be assigned to 

Coryne. Sarsia hydroids cannot usually be identified to species without their mature medusae. 

Sarsia spp. medusae


Distribution. About ten Sarsia medusae were collected from Sadie Cove and the Homer Marina, 

(Katchemak Bay, Cook Inlet) and Fairmount Bay and Busby Island. 

Remarks. Sarsia is a typical north-boreal hydrozoan genus. Many conspecific Sarsias are known from 

the Puget Sound / Strait of Georgia region and the entire life cycle - both hydroid and mature medusa - is 

usually needed for identification to species. Most of the medusae collected had the apical canal above 

the manubrium that is seen in Sarsia princeps and looked quite a bit like those pictured as S. princeps by 

Bigelow (1920), although they were rather small for that species. A second species seemed to also be 

present. 

Staurophora mertensii 

Distribution. This species (5 medusae) was collected by Bigelow (1913) in Prince William Sound - no 

further site description or date given. 

Remarks. We did not find it in August 1999. 

Tiaropsis multicirrata 

Distribution. Several Tiaropsis multicirrata were collected at station PWS 98-21 on June 24, 1998. John 

Chapman identified this site as Green Island, near Montague Island, on the south side of Prince William 

Sound. 

Remarks. Specimens sent to Claudia Mills for identification, summer 1998. 

SCYPHOZOA 


Distribution. Only two Aurelia labiata were seen, in the Cordova Marina, on August 13, 1999. 

Remarks. I would have expected to see this species or its northern congener Aurelia limbata (similar, but 

with a brown rim and tentacles) in many more locations. Aurelia tends to occur in dense aggregations at 

the surface - such "swarms" were described to me by resident kayakers as present in Sheep Bay near 

Cordova and at Long Bay off Culross Passage near Whittier, but we did not see them. See Wrobel and 

Mills (1998) for a discussion of the differences between A. aurita and A. labiata. Purcell (1998) refers to 

the Prince William Sound species as A. aurita, but probably without knowing about the recently 

rediscovered A. labiata name. 

Cyanea capillata 

Distribution. Cyanea capillata was probably the most common medusa in Prince William Sound 

in August 1999. We saw it in the Homer Marina and Sadie Cove in Cook Inlet, as well as at the 

Whittier Marina, en route at the south end of Esther Island and south of Eaglek Bay, in 

Fairmount Bay, at Tatitlek and in the Cordova Marina. A.J. Paul told me that it is common in 

Resurrection Bay, but further out than the town of Seward. 

Remarks. In Prince William Sound this species comes in a range of colors, from red to pink or 

lilac, to yellowish, to a colorless "white". The size ranged from a couple of cm to about 30-40 cm 

in bell diameter. It was abundant in open water.


Unidentified scyphozoan polyps 

Distribution. Jeff Goddard observed scyphistomae on the docks at both Homer and Whittier. 

Remarks. Most scyphistomae on docks on the west coast have proven to be those of Aurelia spp. 

Other species of scyphozoan scyphistomae have not been observed in the field and I assume that 

they select other types of habitats. If these were Aurelia, they may have been either Aurelia 

labiata or perhaps Aurelia limbata, which is likely to also occur in the region. 

CTENOPHORA 

Bolinopsis infundibulum 

Distribution. Only one Bolinopsis specimen was seen on the PWS 1999 trip. It was collected on 

August 11, 1999, at the Tatitlek commercial (ferry) dock, in vertical plankton tows taken off the 

side of the Kristina with Jeff Cordell's 130 µm mesh plankton net in about 50 feet of water. 

Remarks. I do not hesitate to call this specimen Bolinopsis infundibulum, which I have also 

collected at Dutch Harbor. 

Pleurobrachia bachei 

Distribution. Only one Pleurobrachia specimen was seen on the PWS 1999 trip. It was collected on 

August 11, 1999, at the Tatitlek commercial (ferry) dock, in vertical plankton tows taken off the side of 

the Kristina with Jeff Cordell's 130 µm mesh plankton net in about 50 feet of water. 

Remarks. Examination of this preserved ctenophore left some question about its species identity. 

This animal was fairly contracted in its preserved state, at which point the funnel canal appeared 

to be shorter than the pharynx, which is indicative of Pleurobrachia pileus (see Bigelow, 1912). 

This species name should not be applied lightly, however, to a North Pacific specimen, since all 

those collected from British Columbia to California have been identified as Pleurobrachia 

bachei. Comparison with 3 living P. bachei of the same size (about 7 mm) from Friday Harbor, 

revealed that the pharynx/canal ratios in that species to be similar to the preserved specimen 

from PWS, so that name is applied here. 

MOLLUSCA 

Clione limacina 

Distribution. A young pteropod collected in the Homer Marina (Cook Inlet) on August 8, 1999 was 

probably Clione limacina. 

Remarks. The identification was not confirmed by careful microscopic examination. This species is 

found in boreal and temperate regions worldwide. 


Distribution. Several Melibe leonina was observed either swimming in the water column or 

attached to kelp at each of: Fairmount Bay, Tatitlek and Cordova. In addition, Carmen Field


informed me that this species is common in Jakalof Bay within Katchemak Bay, although we did 

not confirm that location. 

Remarks. This species was seen swimming well up in the water column at Fairmount Bay and over 

eelgrass at Tatitlek. It was attached to laminarian kelp in the marina at Cordova.


NONINDIGENOUS CNIDARIA, (MOST) ALREADY PRESENT IN SOME 

WEST COAST ESTUARIES 

that might be positioned to ultimately invade locations in Alaska. 

CNIDARIA 

HYDROZOA 

Bougainvillia muscus (Allman, 1863). Hydroid known on the west coast only from Friday 

Harbor, Washington (Mills, 1981, as B. ramosa); possibly the same species of Bougainvillia that 

is a pest/contaminant in some aquariums in California. Temperature tolerances not known. This 

may actually be a complex of cryptic species rather than one species (Calder, 1988). 

Blackfordia virginica Mayer, 1910. Hydroids and medusae on the west coast known from north 

San Francisco Bay and Coos Bay (J. T. Carlton, personal communication); has a wide salinity 

and temperature tolerance. Also reported from the Chesapeake Bay and several European and 

Asian harbors, and its apparent point of origin, the Black Sea. Full temperature tolerances not 

known, but most of Alaska is north of its known distribution. 

Cladonema radiatum Dujardin, 1843. Hydroids and tiny medusae abundant in eelgrass community 

in Padilla Bay, Washington, not far from Anacortes and Cherry Point oil terminals. Temperature 

tolerances not known, but this species is found in numerous locations worldwide, including northern 

and Mediterranean Europe, its putative natural range. 

Cordylophora caspia (Pallas, 1771). Hydroid known from the mouth of the Samish River in 

Samish Bay, not far south of the Cherry Point oil terminals. Is also found in very low salinity 

tributaries to north San Francisco Bay and elsewhere on the west coast. Requires very low 

salinity, temperature tolerances not known; assumed to be of Ponto-Caspian origin. 

Ectopleura crocea (L. Agassiz, 1862). This Atlantic hydroid is probably established in at least 

California and British Columbia (see photo attributed to this species in Harbo (1999, p. 32). 

Species of Tubularia/Ectopleura cannot be positively identified without examining the 

reproductive medusoids, which is rarely done by non-specialists. 

Maeotias inexspectata Ostroumoff, 1896. Medusae on the west coast known from low 

salinity tributaries to north San Francisco Bay, seemingly always in salinities less than 15 psu, 

maybe to as low as 1-2 psu (Mills and Sommer, 1995). Also known intermittently from the 

Chesapeake Bay and several European estuaries, and its apparent point of origin, the Black 

Sea and Sea of Azov. Temperature tolerances not known, but most of Alaska is north of its 

known distribution. This species was newly collected in the Baltic Sea in Estonia in August 

1999 (Risto Vainola, pers. comm.). 

Moerisia spp. Several species in this genus have been described from a variety of widely 

separated locations worldwide, including some rivers emptying into north San Francisco Bay 

(J.T. Rees, personal communication), in which both polyps and medusae of this genus have been 

found. Also known from the Chesapeake Bay. It is not clear how many species are involved


worldwide. Some populations are known to be single-sexed, inplying a single introduction. 

Temperature and salinity tolerances not known. 

SCYPHOZOA 

Aurelia aurita (Linnaeus, 1758). The most commonly-reported species of Aurelia worldwide. With 

its seemingly highly-transportable sessile polyp, there is some reason to assume that this species was 

carried early to many additional locations, although its home range is so-far not defined - genetic 

studies are currently underway by several researchers. All Aurelia that I have inspected carefully in 

Alaska appear to be Aurelia labiata Chamisso and Eysenhardt, 1821 or Aurelia limbata Brandt, 1835 

(the latter species known from the Aleutians and the Bering Sea). A. labiata was originally described 

from central California, but seems to range all the way up the North American Pacific coast. It 

would not be too surprising to find A. aurita also living on the west coast. A report of a geneticallydifferent 

population of Aurelia (Greenberg et al., 1996) in San Francisco Bay is likely to be such. 

Because it is more amenable to culture, most public aquariums on the west coast have Aurelia aurita 

on display, providing a possible source of introduction. The name Aurelia aurita has been rather 

indiscriminately applied in the literature, including on the west coast of North America, without 

careful morphological inspection. 

ANTHOZOA 

Diadumene lineata Merrill, 1870. Cryptogenic sea anemone known from several locations in 

Washington and California, as well as worldwide. We searched in seemingly appropriate habitat for 

this species in several locations including the intertidal at Lowell Point, Seward, but none were found. 

Temperature tolerances are not known and might be an issue in Alaska. 

Nematostella vectensis Stephenson, 1935. Cryptogenic sea anemone known from quiet, lowsalinity 

lagoon habitats in most coastal American states, as well as numerous locations 

worldwide. This species was not found during the 1999 Prince William Sound Expedition, but 

we may not have encountered the right kind of habitat. Temperature tolerances are not known 

and might be an issue in Alaska. 

CTENOPHORA 

Mnemiopsis leidyi A. Agassiz, 1865. This is apparently the only species of ctenophore known to have 

invaded a marine habitat outside of its home range (the Black Sea). This genus, whose 3 putative 

species are not entirely resolved taxonomically, is native to the eastern coast of North America, 

extending from New England well into Argentina. It has not been found yet in the Pacific Ocean. 

NOTE: Many ctenophores are assumed to have very broad global distributions, and it is not known at 

this time to what extent, if any, their ranges have been extended artificially by man.

Chapt 9C3. Polychaete Worms, page 9C3- 1 

Chapter 9C3. Focal Taxonomic Collections: Polychaete Worms 

Jerry Kudenov, Department of Biology, University of Alaska, Anchorage 

Summary 

Nearly all of the Prince William Sound samples of polychaetes collected during the 1998 

Expedition have been examined. Certain taxa, such as the Spirorbinae, have not yet been 

identified. Excluding the latter subfamily, 61 species have been tentatively identified and 

partially described. In essence, there appear to be no clearly defined species that could be listed 

as a NIS. However, at least five species may be new to science (species of Eumida, Scoloplos, 

Exogone, Nephtys and Glycera). At least three new range extensions may be noted for 

Phyllodoce medipapillata, Chaetozone senticosa and Rhynchospio glutaea. Finally, six species 

that have widespread distributions in the northern hemisphere are represented in the present 

material, including: Pholoe minuta, Eteone longa, Barantolla americana, Harmothoe imbricata, 

Capitella capitata and Amphitrite cirrata. The systematics of each of these species is terribly 

confused and precise identifications are impossible to render presently. For example, Eteone 

longa was originally described from Greenland (1780), and has since been reported from 

numerous localities in the Arctic, Atlantic and Pacific Oceans where it appears to be 

phenotypically “identical” wherever it occurs. Of course, this is likely not true since the 

distributions of most species are restricted spatially and temporally. Resolving such dilemmas 

falls outside the scope of this study, and these six species are therefore identified as above, 

pending future revisions. Although all identifications are reasonably precise and non-indigenous 

species are represented in these samples, all results are based on literature descriptions and are 

preliminary; present materials must be more carefully compared to known reference specimens. 

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Acad. nat. Sci. Phila., 61: 235-295. 

Moore, J.P. 1911. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of 

southern California in 1904. Euphrosynidae (sic) to Goniadidae. Proc. Acad. Nat. Sci. Phila., 63: 

234-318. 

Müller, O.F. 1771. Von Würmern des sussen und salzigen Wassers. Copenhagen, Heinich Mumme 

und Faber. 200 pp. 

Müller, O.F. 1776. Zoologica Danicae Prodromus seu Annimalium Daniae et Norvegiae 

indigenarum characters, nomine et synonyma imprimis populaium. Havniae. 274 pp.


Oersted, A.S. 1843. Groenlandiae Annulata dosibranchiata. K. Danske Videns. naturw. math-Afh. 

Copenhagen, 10: 153-216. 

Pettibone, M.H. 1957. North American genera of the family Orbiniidae. Journal of the Washington 

Academy of Sciences 47: 159-167. 

Rathke, H. 1843. Beitrage zur Fauna Norwegens. Nova Acta Acad. Leop. Carol. Nat. Cur. Halle, 

20: 1-264. 

Schmarda, L.K. 1861. Neue wirbellose Thiere beobachet und gesammelt auf einer Reise um die 

Erde 1853 bis 1857. a. Turbellarien, Rotatorien und Annelidien. Part 2: 1-164. 

Zachs, I.G. 1933. Polychaeta of the North-Japanese Sea. Explorations of the Seas of the 

USSR 19:125-137. (In Russian). 

Table 9C3.1 POLYCHAETA, PRINCE WILLIAM SOUND, SUMMER 1998 

SIGALIONIDAE 

Pholoe minuta Fabricius, 1780 

PHYLLODOCIDAE 

Eteone longa (Fabricius, 1780) 

Eulalia bilineata (Johnston, 1840) 

Eumida species A (new species) 

Phyllodoce medipapillata Moore, 1909 

Phyllodoce species 

NEREIDIDAE 

Chelonereis cyclurus Harrington, 1897 

Platynereis species (bicanaliculata) 

CAPITELLIDAE 

Barantolla americana Hartman, 1963 

GONIADIAE 

Glycinde picta Berkeley 1927 

GLYCERIDAE 

Glycinde armigera Moore, 1911


ORBINIIDAE 

Scoloplos species A New Species 

POLYNOIDAE 

Harmothoe imbricata (Linnaeus, 1767) 

Harmothoe extenuata (Grube, 1840) 

Harmothoinae 

Lepidonotinae 

CHRYSOPELATIDAE 

Chrysopetalum occidentale Johnson, 1897 

SYLLIDAE 

Exogone cf. dwisula Kudenov & Harris, 1995 

Sphaerosyllis cf. californiensis Hartman, 1961 

Trypanosyllis gemmipara Johnson, 1901 

Eudontosyllis species A 

Tyosyllis alternata (Moore, 1908) 

Typosyllis hyalina Grube 1863 

Typosyllis pulchra Berkeley & Berkeley, 1938 

Typosyllis stewarti Berkeley & Berkeley, 1942 

Autoylus (Procerea) cornutus Agassiz, 1863 

LUMBRINDERIDAE 

Lumbrineris latrielli Audouin & Milne-Edwards, 1834 

ORBINIIDAE 

Leitoscoloplos pugettensis (Pettibone, 1957) 

Naineris dendritica (Kinberg, 1867) 

SPIONIDAE 

Spio filicornis (Müller, 1776) 

OPHELIIDAE 

Armandia brevis Hartman, 1938 

Ophelia limacina (Rathke, 1843) 

NEREIDIDAE 

Nereididae (postmetamorphic juvenile) 

SPIONIDAE 

Prionospio steenstrupi Malmgren, 1867


CIRRATULIDAE 

Cirratulus cingulatus Johnson, 1901 

Chaetozone senticosa Blake, 1996 

CAPITELLIDAE 

Capitella capitata (Fabricius, 1780) 

NEPHTYIDAE 

Nephtys species A (N. ciliata) 

Nephtys species A (juvenile) 

Nephtys species B (juvenile) 

ARENICOLIDAE 

Abarenicola pacifica Healy & Wells 1959 

OWENIIDAE 

Owenia fusiformis della Chiaje, 1841 

GLYCERIDAE 

Glycera cf. nana Johnson, 1901 

SPIONIDAE 

Rhynchospio glutaea (Ehlers, 1887) 

Prionospio sp. 

Dipolydora cf. socialis (Schmarda, 1861) 

Dipolydora sp. A (near bidentata) Zachs, 1933 

Dipolydora sp. B 

Dipolydora sp. C (near giardi (Mesnil, 1896)) 

Polydora limicola Annenkova, 1934 

Diplydora quadrilobata (Jacobi, 1883) 

Polydora sp. 

MALDANIDAE 

Nicomache personata Johnson 1901 

PECTINARIIDAE 

Pectinaria granulata Johnson 1901 

AMPHARETIDAE 

Ampharete species A 

TEREBELLIDAE 

Amphitrite cirrata Müller. 1771 

Polycirrus species III Hobson & Banse, 1981


SABELLIDAE 

Laonome cf. kroyeri (Malmgren, 1866) 

Schizobranchia insignis Bush 1904 

SERPULIDAE 

Crucigera zygophora (Johnson, 1901) 

Serpula vermicularis Linnaeus, 1767 

Spirorbis species 

Table 3. Polychaeta Collected in 1998 PWS Expedition 

Note: Materials listed below as PWS NIS 1998 include Stations followed by the number of 

specimens in parentheses. 

Pholoe minuta Fabricius, 1780 

PWS NIS 1998: Sta 1(1 specimen); Sta 5(1); 6 (fragments); Sta 7 (2); Sta (3); Sta 10(4); Sta 

11(1); Sta 41(3). 

Based on specimens, these are not Pholoe minuta sensu Fabricius. Original taxon described as 

having papillae disbursed over entire ventral and parapodial surfaces. Specimens all have small, 

close-set, short papillae over entire ventral surface; parapodia with conspicuous digitiform 

papillae. Whatever “Pholoe minuta” represents, it must be a polyphyletic species at the very 

least. It has a recorded distribution in both the Arctic and south Atlantic Oceans. This species is 

correctly identified to a single taxon. However its actual identity is questionable in view of its 

widespread distribution. 

Eteone longa (Fabricius, 1780) 

PWS NIS 1998: Sta 3(1 specimen ); Sta 15(2); Sta 16(3); Sta 17(8); Sta 18(9); Sta 36(1); Sta 

41(1). 

Technically, the Eteone longa/flava group is in severe disarray and is undoubtedly represents a 

complex taxonomic assemblage of closely related species. Whatever taxon is represented by 

PWS specimens of “E. longa” must remain obscure until a definitive study is published. 

Eulalia bilineata (Johnson, 1840) 

PWS NIS 1998: Sta 21(1 specimen). 

Keys out according to Blake (1996) and Pleijel (1991). Only one specimen, which is poorly 

preserved used for this identification; therefore it is considered to be tentative. 

Eumida species A (new species) 

PWS NIS 1998: Sta 21(2 specimens). 

Genus identification correct. Prostomium small, with 4 distal and one median unpaired antenna. 

Dorsal cirri triangular, pointed, lanceolate. Ventral cirri subtriangular, subtly pointed. Segment 

1 highly reduced, not fused to segment 2; in largest specimen it actually extends onto 

prostomium (although this may be artifactual); tentacular cirri lateral to prostomium. Segment 2


with two pairs of tentacular cirri, ventral pair shortest; lacking setae. Segment 3 with one pair 

tentacular cirri, with fascicle of setae in neuropodium. 

Parapodia all distally rounded, without hint of dorsal lobe; all about the same length throughout 

the body. Ventral cirri asymmetrical, subquadrangular, longest anteriorly and gradually 

decreasing in length, size posteriorly. 

Pygidium lacking appendages (lost). 

Phyllodoce medipapillata Moore, 1909 

PWS NIS 1998: Sta 21(1); 36(1). 

Two beautiful specimens, complete, well preserved with proboscides everted. Key out according 

to Blake (1994). Present record is a range extension, and potentially also an introduced species 

to PWS, assuming there are no other records of it between here and central to southern California 

(0-300 m) where it seems to be restricted. 

Phyllodoce species 

PWS NIS 1998: Sta 15(2 specimens) 

Juvenile individuals, one of which has proboscis everted. Both extremely small, unidentifiable. 

Chelonereis cyclurus Harrington, 1897 

PWS NIS 1998: Sta 3(3 specimens); Sta 5(4); Sta 6(3); Sta 9(1); Sta 10(1); Sta 11(32); Sta 

12(33); Sta 14(1); Sta 16(2); Sta 19(7); Sta 21(65); Sta 22(7); Sta 23(28); Sta 24(14); Sta 24N(2); 

Sta 25(10); Sta 28(57); Sta 30(8); Sta 31(2); Sta 32(3); Sta 34(35); Sta 35(106); Sta 36(2); Sta 

40(1); Sta 41(2); Sta 44(2). 

Characteristic species. Notosetae homogomph spinigers. Neurosetae heterogomph spinigers and 

falcigers. Largest specimen lacks homogomph falcigers in neuropodia. 

Platynereis species (bicanaliculata (Baird, 1863)) 


Juvenile lacking tentacular cirri. This identification is highly tentative, based on a single 

specimen! 

Nereididae (postmetamorphic juvenile) 


Identified as “Platynereis” but the specimen is a postmetamorphic juvenile that is not 

identifiable to genus. 

Glycinde picta Berkeley 1927 

PWS NIS 1998: Sta 4(1 specimen ); Sta 7(1); Sta 15(1); Sta 16(2); Sta 41(1). 

All with ventral arc of micrognaths, characteristic of Glycinde picta along Pacific coast of North 

America. 

Glycinde armigera Moore, 1911 

PWS NIS 1998: Sta 13(1 specimen ); Sta 23(1).


Identification tentative in light of a dissection performed previously and prior to the present 

examination that damaged the critical region of macro- and micrognaths. 

Harmothoe imbricata (Linnaeus, 1776) 

PWS NIS 1998: Sta5(2 juveniles); Sta6(2 juveniles); Sta 10(3 specimens); Sta 11(10); Sta 

14(2); Sta 19(2); Sta 21(8); Sta 23(4); Sta 24(3) Sta24(1); Sta 28(7); Sta 30(2); Sta 34(7); 

Sta 35(17); Sta 36(9). 

Another widespread species that must be re-examined critically. Tentatively assigned to 

Harmothoe imbricata. 

Harmothoe extenuata (Grube, 1840) 


Seems to key out well. Some of the specimens included as H. imbricata likely identical to this 

taxon. 

Harmothoinae 

PWS NIS 1998: Sta 19(1 specimen); Sta 24N(2). 

These are most likely “H. imbricata “ juveniles, and should be referred to above as “” 

Lepidonotinae 

PWS NIS 1998: Sta 23(1 specimen ); Sta 24N(1); 34(2). 

Note that Station 34(2 specimens) contains the best specimens, which seem to key out to 

Parhalosydna, which seems somewhat of a stretch. Most of the specimens are not well 

preserved, nearly all lack elytra, and identification can not be made positively. 

Chrysopetalum occidentale Johnson, 1897 

PWS NIS 1998: Sta 23(1 specimen); 24(1). 

Highly characteristic species. 

Exogone cf. dwisula Kudenov & Harris, 1995 

PWS NIS 1998: Sta 19(1 specimen); Sta 21(1); Sta 23(10); Sta 32(1). 

This species is very closely allied to E. dwisula, and also to E. gemmifera. Antennae closely set, 

laterals about .67-.75x length of median. Pharynx extends through 1.5-2 segments, with anterior 

unpaired middorsal tooth. Proventriculus extends through 2 segments, with 15 rows muscle 

cells. Peristomial antennae small, inconspicuous, not visible in dorsal view. Setae number 4-5 

per parapodium, of 3 kinds: a) falcigers with deeply incised blades, confined to anterior setigers; 

b) stout awl-shaped spinigers, numbering 1-2 per anterior parapodium, 1 per median and 

posterior parapodia; c) dorsal and ventral simple seta, the former present in all setigers, the latter 

in the last few setigers. Aciculae numbering 1 per parapodium, all terminating in distally 

enlarged heads (blunt or beaked) 

Sphaerosyllis cf. californiensis Hartman, 1961 

PWS NIS 1998: Sta 10(1 specimen); Sta 19(1); Sta 19(11); Sta 21(2); Sta 21(2); Sta 23(65); 

Sta 23(5). 

One difference between these specimens and those examined by Kudenov & Harris (1995) is the 

presence of an additional pair of conspicuous papillae on distal parapodial surfaces; one is


anterior, the other posterior. A most unusual aspect to the setal morphology is the fact that the 

cutting teeth on blades of compound falcigers are set in 2 rows, members of one row alternating 

with those of the other row. 

This has not been reported for S. californiensis. Then again, I don’t believe anyone has ever 

looked closely enough! These specimens will represent a new species if S. californiensis lacks 

these alternating rows of teeth on blade cutting surfaces. 

Trypanosyllis gemmipara Johnson, 1901 


One small specimen, 61 segments. Bidentate falcigers. Trepan with 10 teeth; middorsal tooth 

absent. 

Eudontosyllis species A 


Only 2 specimens, both anterior fragments. Specimen of Sta 19 in 2 pieces, with dorsal cirri; Sta 

23 in 1 piece, lacking dorsal cirri. Specimens key out to Eudontosyllis Knox 1960, which 

according to Fauchald (1977) is represented by a single species. 

Essential descriptive elements include: Prostomium reduced, with 2 pairs of large lenticulate; 

eyes Palps reduced, fused only basally. Paired occipital nuchal organs extending over setiger 1, 

not fused to dorsum; Antennae very long, smooth basally, terminating in a few distal moniliform 

elements, each long and cylindrical; Peristomial tentacles long, number 1 pair; Notoacicula 

present, each conspicuous, with distally bent tips; Notosetae as multispinose capillaries in small 

inconspicuous tufts. Neurosetal fascicles with bidentate compound falcigers. 

The one specimen with notosetal fascicles may be epitokous (Sta 23). Need to check out the 

specimen from Sta 19 for comparison. 

Tyosyllis alternata (Moore, 1908) 

PWS NIS 1998: Sta 5(1 specimen); 9(1); Sta10(1, juvenile). 

Typosyllis hyalina Grube 1863 


Typosyllis pulchra Berkeley & Berkeley, 1938 


Typosyllis stewarti Berkeley & Berkeley, 1942 

PWS NIS 1998: Sta 21(5 specimens); 21(1); 23(1). 

Characteristic increase in thickness of falcigers in posterior segments. Many of these have lost 

their blades. 

Autoylus (Procerea) cornutus Agassiz, 1863 

A.cornutus Okada, 1933:645-647, figs. 3,4; Pettibone, 1963:144, fig. 37e. 

A.cornatus Hartman, 1944:338, pl. 13, fig. 5.


A. (Regulatus) cornutus, Imajima, 1966:49-51, Text-fig. 13a-i. 

PWS NIS 1998: Sta 19(1 specimen); Sta 20(2); Sta 25(1). 

Only two specimens include in these samples. Specimen (Sta. 19) is small, lacking tentacular 

and dorsal cirri. Dorsal cirri of setiger 1 longest; those from setiger 2 all shorter and about the 

same size. Both asexual forms exhibiting stolons between segments 13-14. Trepan with 18 

teeth: 9 larger and 9 smaller. Dorsal simple setae thick, distally truncate and serrated. Nuchal 

organs restricted to posterolateral regions of prostomium; not extended to posterior margin of 

setiger 1. 

The species was originally reported from Atlantic habitats (Labrador to Chesapeake Bay; 

Plymouth) and has also been reported from Japan to British Columbia-Washington. 

Specimens (Sta. 20) are sexual forms for which only the genus is a certain identification. 

Swarming or sexually swimming stages have not been related to asexual phases, unfortunately, 

along the Pacific coast of North America (or most other places, except see Gidholm 1965, 1966). 

Nephtys species A (N. ciliata) 

NIW NIS 1998: Sta 10(1 specimen); Sta11(1); Sta 15(3); Sta 16(1); Sta 16(4); Sta 36(5). 

This appears to be a new species. It does not key out to anything in Banse & Hobson (1974) 

where, in the key, this species drops out of the key at couplet 6 (page 73). The couplet provides 

a choice between large, postsetal notopodial lobes without a middorsal proboscideal papilla 

versus medium-sized postsetal notopodial lobes with out without a dorsal median proboscideal 

papilla. 

To couplet 9, the next choice is interramal cirri, proboscis with unpaired dorsal papilla, which 

leads to a choice between N. ciliata or N. caecoides. 

It is not N. caecoides. Key leads to N. ciliata which lacks a dorsal pigment pattern. Notopodial 

postsetal lobe partly covered by acicular lobe, which, in N. caeca is large, but is relatively small, 

compared to the postsetal notopodial lobe. Proboscis proximally with small warts. 

In all, this appears to be N. ciliata. One principal difference appears to be the size of the 

postsetal notopodial lobes. 

Specimen 11(1) poorly preserved; assignment tentative. 

Nephyts species A (juvenile) 

NIW NIS 1998: Sta 15(7 specimens); Sta 17(1); Sta 17(1). 

All specimens are postmetamorphic or young juveniles and are unidentifiable to species. Note 

that all have conical acicular lobes and poorly defined postsetal lamellae. Specimen (17(1)) 

obviously a newly postmetamorphic juvenile; assigned to this taxon for convenience. 

Nephyts species B (juvenile) 

NIW NIS 1998: Sta 11(3 specimens).


Interramal cirri short, almost straight except for distally curved tip, hanging almost straight 

down. Interramal cirri beginning from setiger 5. Acicular lobes generally rounded although 

notopodial lobe slightly bilobed; neuropodial lobe more evenly rounded. 

Glycera cf. nana Johnson, 1901 

NIW NIS 1998: Sta 10(1 specimen); Sta 11(1); Sta 16(1); Sta 24(3); Sta 36(1). 

These specimens are mighty peculiar! Postsetal lobes are rounded, with biramous parapodia as 

per Glycera. Ailerons winged as per Glycera. Proboscis with 3 kinds of papillae including long, 

slender and shorter tapering forms plus spherical papillae. Inferior presetal lobe pointed, 

appearing rather different from that for Glycera nana. 

Hilbig (1994) describes Glycera nana in terms that, compared to the present materials, intimates 

that the PWS specimens are sufficiently different to represent a new species. 

Lumbrineris latrielli Audouin & Milne-Edwards, 1834 

PWS NIS 1998: Sta 5(2 specimens); Sta 11(1). 

Keys out according to both Banse & Hobson (1974) and also Ruff (1995), particularly in view of 

the latter’s comments. Specimen (Sta. 11) with dental formula: 1+1, 5+4, 2+2, 1+1. Yellow 

aciculae. Compound falcigers in anterior segments. Posterior pre- and postsetal lobes not 

elongate. No obvious pigmentation patterns in preserved specimens. 

Scoloplos species A New Species 

As Scoloplos armiger PWS NIS 1998: Sta 4(2 specimens); Sta 10(4); Sta 15(2); Sta 36(1); Sta 

41(22). 

As Scoloplos species PWS NIS 1998: Sta 7(1 specimen); Sta 10(3). 

As Orbiniidae PWS NIS 1998: Sta 10(4 specimens); Sta 41(1). 

All of these individuals represent a new taxon. There are no subpodial lobes present whatsoever. 

Number of thoracic segments numbering 14-15. Branchiae from posterior thoracic segments. 

Thoracic neurosetae with distally smooth, transparent hoods. Abdominal neurosetae include 

both capillaries and delicate spines. Neuropodial lobes in larger specimens digitiform; smaller 

specimens notched. These lobes are clearly different from those of both S. armiger and S. 

acmeceps. 

Leitoscoloplos pugettensis (Pettibone, 1957) 


Specimens correctly identified to species. No abdominal subpodial lobes present. 

Naineris dendritica (Kinberg, 1867) 


Agrees well with descriptions. Originally identified as N. quadricuspida on label, however, only 

one record can be assigned to this species, and even then, Hartman (1961) noted distinct and 

significant differences between her material compared to those described by Fabricius. In other 

words, N. quaricuspida does not occur on this coast!


Spio filicornis (Müller, 1776) 

PWS NIS 1998: Sta 5(2 specimens); Sta 10(1); Sta 15(12); Sta 17(1). 

Keys out according to Blake (1996). 

Armandia brevis Hartman, 1938 

PWS NIS 1998: Sta 5(3 specimens); Sta 7(3); Sta 10(5); Sta 11(12); Sta 11(1*); Sta 15(4). 

Everything keys out extremely well to this taxon following Hartman (1969). Need to check out 

the validity of this genus based on Colin Herman’s comments a few years ago. Specimen (Sta. 

11(1*)) is poorly preserved, has a pair of prostomial eyespots, and hints of lateral eyespots, and 

is taken here to represent Armandia brevis. 

Ophelia limacina (Rathke, 1843) 


This is a supposedly cosmopolitan species. It has 37 setigers compared to 39 originally 

described. First 10 setigers abranchiate. Ventral groove present from around setiger 10-11. 

Prionospio steenstrupi Malmgren, 1867 


Specimens poorly preserved, and trashed in most cases. Gills not well intact, and in a few 

specimens (Sta. 11) they are of variable lengths. The neuropodial lamella of setiger 2 with the 

characteristic ventral protuberance; those of setiger 3 squarish to ventrally pointed also. 

Identification tentative pending additional specimens. 

Cirratulus cingulatus Johnson, 1901 

PWS NIS 1998: Sta 6(1 specimen); Sta 11(1); Sta 16(2); Sta 17(8) + Sta 17(3); Sta 24(3); Sta 

24N(1). 

Specimens agree fairly well with description provided by Blake (1996:350-351). Neurosetal 

spines begin from setigers 23-26; notosetal spines from setigers 35-37. Specimen from Sta 24N 

may be a juvenile, with neurosetal spines from setiger 9. and notosetal spines from setiger 16. 

Eyes present in all specimens as line of 4-6 individual eyespots. The three specimens from Sta 

17 are very large and show size-dependent morphology concerning transverse band of 

tentacles/cirri. 

Chaetozone senticosa Blake, 1996 

PWS NIS 1998: Sta 15(9 specimens); Sta 17(57). 

Keys out according to Blake (1996), although the final identification needs to be confirmed 

based on methyl green. Specimens with around 60-70 setigers, hooks beginning from around 

setiger 35-40. Prostomium short, triangular, with a single achaetous annulus. 

Originally reported from Central and Northern California. This may be a range extension, 

assuming the identification is valid. 

Barantolla americana Hartman, 1963 

PWS NIS 1998: Sta 3(1 specimen); Sta 4(1); Sta 9(2); Sta 15(3); Sta 19(1); Sta 41(1).


This is a dubious taxon. Originally described has having capillary setae only in notosetiger 6 

and neurosetiger 7; mixed capillaries in notosetiger 7 and neurosetiger 8; hooks only in 

notosetigers 8-11 and neurosetigers 9-11. In contrast, Fauchald (1977) lists Barantolla as 

having 6 setigers with capillaries followed by 1 mixed capillaries and hooks, and then 4 more 

with hooks only. 

The present specimens are at odds with the above descrepancies. Specimen 3(1) with mixed 

notosetae on setiger 5; 7(2) with capillaries only in notosetigers 1-5 and neurosetigers 1-6, 

mixed setae in notosetiger 6 and neurosetiger 7, and hooks only thereafter in notosetigers 7-11 

and neurosetigers 8-11; specimen 41(1) with capillaries only in both noto- and neurosetigers 1-6, 

and hooks only in both noto- and neurosetigers 7-11 (setiger with mixed setae apparently 

absent). 

Capitella capitata (Fabricius, 1780) 

NIW NIS 1998: Sta 7(3 specimens); Sta 9(1); Sta 15(9); Sta 41(17). 

Whatever Capitella capitata is, these specimens can be assigned to the stem species. But this is 

one the “cosmopolitan” species that can be almost anything. 

Abarenicola pacifica Healy & Wells 1959 

NIW NIS 1998: Sta 7(1 specimen); Sta 17(3); Sta 18(9). 

One large specimen (Sta. 7), intact; all others are juveniles. No question concerning identity. 

Owenia fusiformis della Chiaje, 1841 

NIW NIS 1998: Sta 7(1 specimen); Sta 41(1). 

Only two specimens. Have a collar as per Owenia collaris. Setae appear to have configuration 

found in Owenia fusiformis. Refer to recent paper on the family from IP4 (Paris). 

Rhynchospio glutaea (Ehlers, 1887) 

PWS NIS 1998: Sta 10(1 specimen); Sta 10(1). 

New record for Alaska. Not particularly surprising. 

Prionospio sp. 


Recheck. One specimen appeared to have gills on middle body segments. These are not 

polydorids as noted on label. 

Dipolydora cf. socialis (Schmarda, 1861) 


Tentative identification, but not Dipolydora socialis. Falcate spines of setiger 5 strongly falcate, 

with hint of flange; bristle absent. Setiger 1 postsetal lamellae poorly developed. Gizzard-like 

structure present around setigers 17-18, but not as portrayed by Blake (1996). 

Dipolydora sp. A (near bidentata) Zachs, 1933 

PWS NIS 1998: Sta 17(18 specimens); Sta 19(1); Sta 23(2). 

This taxon may be a shell borer. Spines of setiger 5 without distal bristles, with flange-tooth on 

lateral surface (not on convex surface as fas as I can see). Caruncle to posterior setiger 4.


Notosetae present on setiger 1. Prostomium deeply incised, bifurcate. Posterior notosetae 

capillaries; spinous packets, modified setae absent. Neurosetae without manubrium, from setiger 

7, bidentate to end of body. 

Dipolydora. sp B 


Prostomium incised, strongly bilobed. 4 pairs of eyes. Notosetae setiger 1 present, lobe reduced 

to papillar lobe. Caruncle to posterior setiger 3. Setiger 5 strongly modified; heavy spines 

distally falcate, heavy triangular tooth on concave surface, bristles present in notch between 

tooth and tip of spine. Bidentate neurosetae without manubria, from setiger 7. Branchiae from 

setiger 7. 

Does not key out using Blake 1996…falls out at couplet 10 (10B where choice is presence of 2 

accessory teeth) since this specimen has only 1 visible accessory tooth, and appears to lack a 

cowling. 

Dipolydora sp. C (near giardi (Mesnil, 1896)) 

PWS NIS 1998: Sta 21(>100 specimens); Sta30(1). 

Numerous specimens. Prostomium incised, bilobed. No eyes. Setiger 1 complete, notosetae 

present, notopodium reduced to digitiform lobe. Caruncle to setiger posterior margin setiger 3. 

Setiger 5 modified; spines with accessory tooth on concave surface, with partial cowling on 

opposite side of concave surface extending to convex surface; bristles absent. 

Branchiae from setiger 9. Neurosetal hooks without manubria, from setiger 7. 

Dipolydora quadrilobata (Jacobi, 1883) 


Incomplete specimen, and identification is tentative. 

Polydora limicola Annenkova, 1934 


Incomplete specimen, and identification is tentative. 

Polydora sp. 


This is a juvenile specimen. It is small and slender. Prostomium entire. Eyes numbering 8. 

Setiger 5 is not modified. Pygidium 4-lobed. Probably not assignable to any known taxon. 

Nicomache personata Johnson 1901 


Identification certain to this species, although specimen is incomplete. Pigmentation pattern 

characteristic of species.


Pectinaria granulata Johnson 1901 

PWS NIS 1998: Sta 11(5 specimens); Sta 16(3); Sta 17(1); Sta 36(22). 

Correct identification. 

Laonome cf. kroyeri (Malmgren, 1866) 


Avicular uncini with short bases; companion setae absent. Both capillary and spatulate setae 

present. Radioles lacking external stylodes; collar bilobed. Need to reconfirm identity. 

Schizobranchia insignis Bush 1904 

PWS NIS 1998: Sta 19(4 specimen); Sta 23(2) + Sta 23(>20). 

Identification correct. Smallest specimen placed into shell vial (Sta. 23) is perhaps a juvenile, 

with all the setal features consistent. Note that large vial (Sta. 23) mislabeled as Terebellidae 

(instead of Sabellidae). 

Amphitrite cirrata Müller, 1771 

PWS NIS 1998: Sta 19(2 specimens); Sta 23(3); Sta 24(2); Sta35(1); Sta 36(1). 

This is another “cosmopolitan” species…at least in the northern hemisphere. A taxonomic black 

hole similar to Phole minuta and Eteone longa!! Specimen (Sta. 35) is juvenile terebellid, 

probably Amphitrite cirrata; it is not Ampharetidae)! 

Polycirrus species III Hobson & Banse, 1981 


This keys out as per Hobson & Banse (1981). However, as a general comment, the key is 

extremely poor and relies on imprecise terminology that overlaps features used to describe 

notosetae! In any case, it would seem appropriate to attach names instead of roman numerals. 

Ampharete species A 


The generic identification is correct. Single specimen lacks a tail, and cannot be identified to 

species. It is not Ampharete labrops, which has eyespots on upper buccal lip; present specimen 

lacks eyespots in corresponding region. 

Crucigera zygophora (Johnson, 1901) 

PWS NIS 1998: Sta 19(6 specimens); Sta 23(>20); Sta 24(1). 

Identification correct. Present specimens are textbook examples. 

Serpula vermicularis Linnaeus, 1767 


Identification correct. Tentacles solid red or banded red and white, at least in preservative. 

Spirorbis species 

PWS NIS 1998: Sta 17(>50 specimens). 

Dextrally spiraled tubes, 3 thoracic setigers. Species identifications pending examination of 

remaining materials.

Chapt 9C4. Peracaridan Crustaceans, page 9C4- 1 

Chapter 9C4. Focal Taxonomic Collections: Peracaridan Crustaceans 

John W. Chapman, Department of Fisheries & Wildlife, Hatfield Marine Science Center, Oregon 

State University 

Summary 

No clear peracaridan NIS were discovered among the scores of species collected in three 

surveys of 72 sites in 21 general areas of Prince William Sound and south central Alaska 

between 1997 and 1999. No NIS were found in UAF samples from the area collected 

previously. Two peracaridan species previously considered to be introduced are likely to be 

misidentified. Five species of NIS gammaridean amphipods were found in ballast water of 

tankers travelling to Prince William Sound, indicating that this is an active mechanism of NIS 

transport to Alaska, even though they do not appear to have invaded or become established there. 

Invasions of Alaskan estuaries and marine waters by a broad diversity of peracaridan species 

have not occurred. The diversities of peracaridan NIS invasions in the northern hemisphere vary 

with climate, as do invasions by other taxa noted previously. Most marine and estuarine 

peracaridan NIS thus appear to be incapable of invading Alaska from lower latitudes due to the 

extreme climate. The risk of invasions by high diversities of NIS of pericaridans thus appears to 

be extremely low. 

These findings do not indicate whether a few NIS could be present at ecologically 

catastrophic abundances, however. Eight peracaridans that are prominent members of either 

fouling or benthic communities sampled in the survey, have unclear origins or cannot yet be 

clearly distinguished from species that are nonindigenous to the northeast Pacific. They are 

therefore classified as cryptogenic. These cryptogenic peracaridan species occur in the same 

areas as the soft shell clam, Mya arenaria Linnaeus, 1758, which is one of the most clearly 

documented NIS in south central Alaska. If proven to be NIS, these cryptogenic peracaridan 

species, would be evidence that even a few NIS capable of invading Alaskan estuaries can 

increase to ecologically catastrophic densities. They would indicate that surveys of peracaridan 

NIS diversity, such as this one, are an insufficient basis for estimates of risk. Whether these 

peracaridan crustaceans are, in fact, native to the region therefore should be tested by analyses of 

morphological variation, molecular genetics and by crossbreeding viability tests with their 

presumed original populations. 

Introduction 

A major objective of the south central Alaskan NIS survey was to determine whether 

introductions of marine or estuarine species have already occurred. An ultimate objective of the 

overall risk analysis is to predict whether Alaskan waters are vulnerable to NIS invasions. The 

survey results and comparisons of climate effects on peracaridan NIS diversity over the northern 

hemisphere provide a basis for this prediction. 

Predicting which nonindigenous species (NIS) can be introduced, where, and the factors 

that control their survival are major objectives of invasion ecology. These predictions require 

knowledge of the interactions between dispersal and processes that determine NIS survival. The


mechanisms of NIS dispersal among estuaries are becoming well known (e.g., Cangelosi 1999, 

Cohen 1998, Frey et al.1999, Draheim and Olson 1999, Miller and Chapman 2000, Moy 1999, 

Ruiz et al. 1999, Thresher 1999), while the processes limiting NIS survival and production 

among estuaries remain poorly known. The distributions of NIS reveal how survival varies as 

dispersal occurs and thus indicate the interactions of dispersal, ecology, and survival. 

Interpreting the geography NIS distributions is thus a necessary part of the search for factors 

controlling NIS invasions. 

NIS are particularly diverse and abundant in estuaries of the northeastern Pacific, 

including San Francisco Bay, California (Carlton and Geller, 1993, Cohen and Carlton 1995, 

1997, Ruiz et al. 1997a, 1997b) and in Europe (Leppakoski, 1994, Leppakoski and Olenin, 1999, 

Eno, et al. 1997). The majority of these NIS have origins from western ocean coasts (Cohen and 

Carlton 1995, Leppakoski and Olenin, 1999) and progressively fewer NIS are known with 

increasing latitudes (Carlton 1979, Cohen et al. 1998, Mills et al. 2000). These geographical 

patterns do not appear to result entirely from the mechanisms of dispersal or patterns of endemic 

species diversity. Other processes controlling NIS distributions warrant consideration. 

Potential climate effects on east and west and north and south patterns of NIS diversity 

among estuaries and coastal waters of the northern hemisphere are of paramount concern in any 

NIS risk analysis for Alaska. The cold temperate climate of Alaska is at the extreme northern 

range of many northeast Pacific intertidal species (O’Clair 1977, O’Clair and O’Claire 1998). 

Climate effects are therefore considered in this section. Salinity and temperature variations in 

estuaries are dominant processes of climate that limit NIS survival. Most estuarine species 

survive within narrow temperature and salinity ranges. Most chemical, biological, and 

hydrological processes that also limit the abundances and distributions of estuarine organisms are 

also controlled by, or closely correlated with, salinity and temperature (e.g., Southward 1969, 

Green 1971, Ebbesmeyer et al. 1991, Cohen and Carlton 1995 1999, Chapman 1998, Thompson 

1998). NIS distributions are therefore influenced by salinity and temperature within local 

estuaries. In turn, salinity and temperature are affected by climate (Ebbesmeyer et al. 1991, 

Cayan 1993). Precipitation and air temperature variations (Ebbesmeyer et al. 1991, Cayan 1993) 

can be interpreted to infer salinity and temperature variations in local estuaries even though 

direct, long-term measures of these parameters are lacking in most cases. 

Amphipods recovered from 34 ballast water samples taken from tankers during 1998 

were examined to determine whether amphipods are transported by ballast water traffic from 

west coast ports and harbors. 

Methods 

Rapid assessment survey 

Prince William Sound, Seward and Homer, Alaska (60 o 00' - 61 o 00' N) survey results are 

compared to results of NIS surveys of the native, cryptogenic and introduced peracaridans from 

Puget Sound, Washington (47 o 10' - 49 o 00' N) and San Francisco Bay, California (37 o 30' - 38 o 

10' N) to resolve how peracaridan NIS invasions are distributed over a broad range of latitudes.


Collections were made from three sites in Port Valdez in early spring of 1997, 46 sites 

throughout Prince William Sound in June 1998, and from 23 sites in Prince William Sound, 

Seward and Homer in 1999 (Ruiz and Hines 1997, Hines and Ruiz 1998) (Figure 9C4.1). 

Twenty six sites were surveyed in Puget Sound, Washington in September 1998 (Cohen et al. 

1998). San Francisco Bay was surveyed in early fall, late spring or summer of 1993, 1994, 1996 

and 1997 at 25 regular sites plus several irregular sites (Cohen 1998, Cohen and Carlton 1995, 

1997, 1998). The three systems are excellent for comparison because they have all received and 

have been interconnected by significant aquaculture and shipping activities that are vectors of 

NIS dispersal in at least the last century. 

Figure 9C4.1. Prince William Sound, Alaska and south central Alaska rapid assessment sites with open boxes 

indicating all 46 sampling sites of 1998, inset indicating the five port areas sampled in 1999 and the shaded boxes 

indicating the eleven general sampling areas of the sound in 1999. (Latitudes longitudes and site descriptions are in 

appendix table 9C4.5). 

Each area was surveyed by the author in the same fashion. Survey samples were 

collected by hand, scrapings, cores, or dredge as necessary to remove biological communities or 

substratum from floats, intertidal pilings rocks and intertidal or shallow subtidal mudflats 

accessible at each collection site. These samples were washed on an 0.5 mm mesh sieve directly 

or decanted onto an 0.5 mm mesh sieve and washed following vigorous sloshing in buckets of


seawater, to suspend organisms from the removed substratum. Harbor float, rock and piling 

substratums were emphasized in all three survey areas but other available habitats were sampled 

extensively as available. Organisms were picked directly from substratums during sample 

collection or from the sieves after washing or from voucher samples of substratums and 

examined under a stereomicroscope. All collected organisms were fixed in 10% formalin before 

transfer to 70% ETOH for long-term preservation. All specimens were identified to lowest 

possible taxonomic category. 

Voucher specimens will be deposited in the Los Angeles County Museum, the California 

Academy of Sciences and the Smithsonian National Museum of Natural History. The precise 

locality records and notes for each collection site are available from the author. Temperature and 

salinity was measured at each collection site. Surface salinties ranged between 0 and 33 o / oo in 

all three survey areas. Surface water temperatures ranged between 8 and 20 o C in Prince William 

Sound, between 10 and 21 o C in Puget Sound and between 12 and 30 o C in San Francisco Bay. 

San Francisco Bay is a well mixed estuary. Low surface salinities and clear stratification occur 

in both sounds in summer and were apparent during the surveys. 

All species from the three surveys were collected and examined directly by the author 

and thus are assumed to be a more standardized sample than would be likely from comparisons 

of different surveys and sampling methods employed by different investigators. The indigenous 

origins of species are inferred from previously published records or herein using the criteria of 

Carlton (1979) and Chapman and Carlton (1991, 1994). The criteria used for cyptogenic species 

(species that are not clearly native or introduced) are adopted from Carlton (1996a). Only 

populations of species that have been moved by human activities to new locations, that are 

reproductive there, and that satisfy the criteria for nonindigenous species are considered here to 

be NIS. 

Alaskan Climate and Peracaridan NIS Invasion Risks 

Sources of NIS to Alaskan estuaries are available from a global population. The 

peracaridan Crustacea of the northern hemisphere considered here are as a sample of that 

population. Crustacea comprise approximately 25% of the 250 NIS reported from San Francisco 

Bay (Cohen and Carlton 1995, J. T. Carlton, personal communication), where they are the most 

diverse NIS taxon. The majority of these crustacean NIS are peracaridans. The Peracarida 

consist of relatively small, short-lived species that are primarily mysids, amphipods, isopods, 

tanaidaceans and cumaceans. The Peracarida are prominent in most North Pacific and North 

Atlantic marine and estuarine NIS communities (Bowman et al. 1981, Chapman 1988, 1999, 

Chapman and Carlton 1991, 1994, Mees and Fockedey 1993, Leppakoski 1994, Cohen and 

Carlton 1995, 1997, Eno et al. 1997, Toft et al. 1999). Both native and nonindigenous 

Peracarida are diverse, taxonomically well known, and ubiquitous in aquatic environments (e.g., 

Barnard and Barnard 1983, Barnard and Karaman 1991, Chapman 1988, Cohen and Carlton 

1995, Chapman 2000). Peracarida develop directly, without larval dispersal stages or unique life 

history traits that complicate identifications and interpretations of their geographical 

distributions. Peracaridans may thus provide clear indications of the patterns of diversity within 

and among broad geographical regions.


The species selected for the east to west geographical analysis of northern hemisphere 

peracaridan NIS are common or abundant where they occur and documented either in the 

literature or by personal observations. Species that are poorly documented, not examined 

directly, cryptogenic, or that are not introduced across the North Atlantic or North Pacific, were 

not included. For instance, the amphipod Chelicorophium curvispinum (Sars, 1895), which 

spread from the Black and Caspian Sea to northern Europe (Eno et. al. 1997), and the introduced 

mysid Acanthomysis bowmani Modlin and Orsi, 1997 in San Francisco Bay, which has unknown 

origins, and many northern NIS that are native to the southern hemisphere are not in the scope of 

this study and are therefore excluded from the analysis. 

Long-term climate conditions in the northeast Pacific, including San Francisco Bay, 

Puget Sound and Prince William Sound are inferred from monthly average climate time series 

data for the Pacific Ocean and western Americas (Cayan et al. 1991). These data extend over 

approximately 100 years up to 1986. Global records of sea surface temperature and precipitation 

minus evaporation (http://www.cdc.noaa.gov/ 1998) are used for comparisons of temperature 

among ocean regions. The term “western ocean” is used in reference to the Pacific Ocean 

bordering the east Asian coast and the Atlantic Ocean bordering the eastern North American 

coast. The term “eastern ocean” refers to the ocean areas bordering the west coasts of Europe 

and North Africa and the west coast of North America. 

Amphipods in Ballast Water 

Ballast water samples were collected in vertical plankton tows from “dedicated” ballast 

tanks, which are not contaminated by oil. Amphipods retained in the 0.25 mm mesh plankton 

nets preserved in 5% formalin, subsequently transferred to 70% ethanol for final sorting and 

identification. Samples were initially sorted under stereo microscopes at the Smithsonian 

Environmental Research Center or at the field office in Port Valdez and final amphipod 

identifications were performed at HMSC, Oregon State University. 

The origins of the ballast water sampled (Table 9C4.3) were the Los Angeles-Long 

Beach area, the San Francisco Bay area, Puget Sound (Anacortes) and the open ocean. One ship, 

from the San Francisco Bay area, exchanged the ballast water at sea during transit. The potential 

for dispersal of nonindigenous species is assessed from the presence of nonindigenous species in 

samples. 

Results 

North - South Climate 

The maximum, minimum, mean and range of monthly sea surface temperatures of the 

eastern Pacific vary by 5 o C or less between 28 o and 52 o N (Figure 9C4.2). The 11 to 14 o C 

maximum average monthly temperatures of Seward, Kodiak and Sitka, Alaska (between 58 and 

60 o N) overlap the Neah Bay, Washington maximum surface temperatures at 48 o N and are 

similar to the 13 o C average sea surface temperatures adjacent to San Francisco at 38 o N (Figure 

2). Nonindigenous species expected to reach south central Alaska might include those that can 

reproduce within Puget Sound in summer or in average San Francisco temperatures (Figure 

9C4.2).


Figure 9C4.2. Sea surface temperature monthly average minimum, mean, maximum and range of northeast Pacific 

coastal waters estimate over an approximately 100 years up to 1986 (Cayan 1991). 

North - South Biodiversity 

Of the 106 peracaridan crustacean species identified from the surveys of Prince William 

Sound, Puget Sound and San Francisco Bay, 54 are native, 14 are cryptogenic and 38 are 

introduced (Table 9C4.1). Seven peracaridan crustaceans that prominent members of benthic or 

fouling communities that were recovered in the survey are cryptogenic (Table 9C4.1). They are 

the tanaidacean Leptochelia dubia (Kroyer, 1842) a cosmopolitan species (Miller, 1975); the 

cumacean Cumella vulgaris Hart, 1930 which occurs in Asia (Lomakina 1958) as well as the 

eastern Pacific; the amphipod Monocorophium carlottensis Bousfield and Hoover, 1997 is not 

clearly distinguished from the nonindigenous amphipods Monocorophium acherusicum and 

Monocorophium insidiosum (Ruiz and Hines 1997); the amphipod Hyale plumulosa (Stimpson, 

1857) is reported also from the western Atlantic (Bousfield 1973); the amphipod Jassa staudei 

Conlan 1990 is extemely similar to the cosmopolitan Jassa marmorata Holmes, 1903; the 

amphipod Pontogeneia rostrata Gurjanova, 1938 is reported from the eastern and western 

Pacific (Gurjanova 1938, 1951, Barnard 1962, 1964); the caprellid Caprella depranochir Mayer, 

1880 is reported from the eastern and western Pacific (Arimoto 1976, Kozloff and Price1997).


Table 9C4.1. The 106 peracaridan crustaceans identified as nonindigenous, cryptogenic or native, and the records 

per species collected from San Francisco Bay, California, (w/o “*”), south central Alaska and Prince William Sound 

(with “*”), or Puget Sound, Washington (underlined). Gnorimosphaeroma lutea was collected from San Francisco 

Bay and Prince William Sound only. No species were collected only in Puget Sound. 

Nonindigenous Cryptogenic Native 

Records Records Records 

Mysidacea Tanaidacea Mysidacea 

Accanthomysis aspera 1 Leptochelia dubia 3 Mysis littoralis* 1 

Tanaidacea Cumacea Isopoda 

Tanais stanfordi 2 Cumella vulgaris 3 Dynamenella glabra* 2 

Isopoda Gammaridea Gnorimosphaeroma lutea 2 

Asellus sp. 1 Ampithoe lacertosa 3 Gnorimosphaeroma oregonense 3 

Dynoides dentisinus 1 Dulichia sp. 1 Ianiropsis kincaidi* 2 

Euylana arcuata 1 Hyale plumulosa 3 Idotea montereyensis 3 

Ianiropsis serricadus 1 Ischyrocerus sp. 2 Idotea obscura* 1 

Limnoria quadripunctata 1 Jassa staudei 3 Idotea resecata 2 

Limnoria tripunctata 2 Monocorophium carlottensis* 2 Idotea wosnsenskii 3 

Munna ubiquita 1 Pontogeneia rostrata 3 Ligia pallasi* 1 

Paranthura sp. 1 Caprellidea Limnoria lignorum 2 

Sphaeroma quoyanum 1 Caprella depranochir* 2 Cumacea 

Synidotea laevidorsalis 1 Caprella laeviuscula 3 Diastylis alaskensis* 1 

Cumacea Caprella penantus 1 Diastylis sp.* 1 

Nippoleucon hinumensis 2 Caprella verrucosa 2 Lamprops beringi* 1 

Gammaridea Tritella sp. 1 Lamprops quadriplicata* 2 

Ampelisca abdita 1 Gammaridea 

Ampithoe valida 2 Allorchestes angusta 3 

Crangonyx sp. 1 Americorophium brevis* 2 

Eochelidium sp. 2 Americorophium salmonis* 2 

Gammarus daiberi 1 Americorophium spinicorne 2 

Grandidierella japonica 2 Ampithoe dalli* 2 

Hyalella azteca 1 Ampithoe kussakini* 1 

Jassa marmorata 2 Ampithoe sectimanus* 1 

Laticorophium baconi 2 Ampithoe simulans* 2 

Leucothoe alata 1 Anisogammarus pugettensis 3 

Melita nitida 2 Aoroides columbiae 3 

Melita sp. 1 Aoroides intermedius* 2 

Monocorophium 

2 Calliopius carinatus* 2 

acherusicum 

Monocorophium insidiosum 2 Calliopius pacificus* 2 

Monocorophium 

1 Eogammarus confervicolus 3 

oaklandense 

Monocorophium uenoi 1 Eogammarus oclairi* 1 

Trasorchestia enigmatica 1 Gammaridae n. gen. n. sp.* 1 

Paradexamine sp. 1 Gnathopleustes pugettensis 2 

Parapleustes derzhavini 1 Hyale frequens 3 

Senothoe valida 2 Lagunogammarus setosus* 2 

Sinocorophium 

1 Locustogammarus locustoides* 1 

heteroceratum 

Caprellidea Megamoera subtener* 2 

Caprella acanthogaster 2 Microdeutopus schmitti 1 

Caprella bidentata 1 Najna n. sp.* 1 

Caprella californica 2 Paracalliopiella pratti* 2 

Caprella equilibra 2 Parallorchestes ochotensis 3 

Table 9C4.1. Continued


Paramoera bousfieldi* 2 

Paramoera mohri* 1 

Parampithoe humeralis.* 2 

Parampithoe mea* 1 

Photis brevis 2 

Pontogeneia inermis 3 

Pontogeneia ivanovi* 1 

Pontoporeia femorata* 2 

Spinulogammarus subcarinatus* 1 

Trasorchestia traskiana 3 

Caprellidea 

Caprella alaskana* 1 

Caprella gracilior* 1 

Caprella irregularis* 2 

Metacaprella kennerlyi* 2 

Total Species 38 14 54 

Records/Species 1.4 2.1 1.8 

The diversity of species collected is nearly nearly constant among sites (66 to 60 and 59, 

respectively, between San Francisco, Puget Sound and Prince William Sound). Of the 54 native 

species collected in Alaska, 52 (96%) were collected also in San Francisco Bay or Puget Sound 

(Table 9C4.1). The common pool of native species and the similar species diversities collected 

among the three areas both indicate that the habitat selection, collection and sample processing 

methods of the rapid assessment surveys were consistent among the three areas. Little variation 

in NIS diversity among these three sample sets was therefore likely to result from sample biases. 

From San Francisco Bay north to Puget Sound and Prince William Sound, introduced 

peracaridan species declined from 38 to 15 to 0 (Figure 9C4.3, X 2 > 27.01; p < 0.0001; df = 2), 

while the frequencies of cryptogenic peracaridan species were nearly constant at 11, 10 and 8, 

respectively (X 2 = 2.0; p > 0.73; df = 2). In contrast, the frequencies of native species increased 

to the north from 17 to 35 and 52 (Figure 9C4.3, X 2 > 27.01; p < 0.0001; df = 2 ). All 

peracaridan NIS and all but two of the cryptogenic peracaridan species at any site also occurred 

in San Francisco Bay while only 17 of the 54 native species were recovered from San Francisco 

Bay (Table 9C4.1, Figure 9C4.3). The peracaridan NIS that managed to invade northeast Pacific 

estuaries thus have adaptations to lower latitude climates than do the native species. The 

affinities of peracaridan NIS for low latitude climates closely corresponds to a pattern that would 

be expected if the peracaridan NIS are poorly adapted to cold water conditions or other factors of 

climate that vary with latitude.


Native Cryptogenic Nonindigenous 

60 

50 

Species 

40 

30 

20 

10 

0 

SFB 

PS 

PWS 

Figure 9C4.3. Native, cryptogenic, nonindigenous and total species in samples from San Francisco Bay, California, 

Puget Sound, Washington and Prince William Sound, Alaska (SFB, PS and PWS, respectively). 

Some NIS may be among the 8 cryptogenic peracaridan species of Prince William Sound 

(Figure 9C4.3). Six of these cryptogenic Alaskan peracaridan species have broad thermal 

tolerance ranges and occur also in Puget Sound and San Francisco Bay (Table 9C4.1). The 

nearly complete faunal peracaridan overlap between Alaska and the two more southern sites 

indicate that the low diversity of Alaskan NIS peracaridans are not likely to result from unique 

types of NIS peracaridans that were not collected in the survey. The nearly uniform pool of 

native peracaridan species and uniform pool of NIS peracaridans among the areas require few 

qualifications or assumptions to arrive at conclusions of the patterns of diversity. The possibility 

of overlooking populations of nonindigenous peracaridan crustaceans in Alaska that occur at 

similar diversities and abundances as in Puget sound or San Francisco Bay, by these rapid 

assessment methods, is remote. The presence of peracaridan NIS that are not among the 

cryptogenic species of peracardans in Alaskan estuaries and marine waters is not significantly 

different from zero. 

These results are not surprising. The extreme climate of Alaska was previously assumed 

to limit the survival of nearly all NIS peracaridans. However, these data include only a single 

coast and three major areas of reference. NIS pericarican diversity was therefore compared 

between other geographical regions with climates that vary similarly to the variation that occurs 

between San Francisco and south central Alaska as a further test of the climate pattern. 

East -West Climate 

Sea surface temperatures are relatively constant between latitudes of 25 o and 50 o N in 

eastern oceans compared to western ocean areas (Figures 9C4.4 and 9C4.5). Eastern ocean 

species from 32 o to 50 o N evolved in temperature ranges that span only latitudes 40 to 42 o N in


western oceans (Figure 9C4.6). Eastern ocean species thus have not evolved in conditions that 

would create broad temperature tolerance ranges. In contrast, most western ocean organisms 

survive temperature ranges exceeding the total temperature range of broad eastern ocean areas. 

Eastern oceans below 50 o N, are thus broad thermal targets for western ocean species over space 

and time while western ocean coasts are narrow thermal targets for eastern ocean species. 

N.W. Pacific 

N.E. Pacific 

30 

30 

25 

25 

SST (Deg. C) 

20 

15 

10 

5 

‘ 

SST (Deg. C) 

20 

15 

10 

5 

0 

-5 

JF MA MJ JA SO ND 

60 

50 

30 

40 

Lat. N 

0 

-5 


60 

50 

30 

40 

Lat. N 

Months 

Months 

N.W. Pacific 

N.E. Pacific 

5 

5 

4 

4 

Precip. (mm / day) 

3 

2 

1 

0 

-1 

-2 

-3 


30 

60 

50 

40 

Lat. N 


3 

2 

1 

0 

-1 

-2 

-3 


30 

60 

50 

40 

Lat. N 

Months 

Months 

Figure 9C4.4. Northwest and northeast Pacific sea surface temperature in degrees Celsius and bimonthly 

precipitation minus evaporation in mm -d at ten degree latitude intervals for the bimonthly periods of Jan-Feb, Mar- 

Apr, May-Jun, Jul-Aug, Sep-Oct and Nov-Dec. (Note: axes of latitude are reversed between graphs of temperature 

and


N.W. Atlantic 

N.E. Atlantic 

30 

30 

25 

25 

SST (DEG. C) 

20 

15 

10 

SST (DEG. C) 

20 

15 

10 

5 

0 

JF MA MJ 

JA SO ND 

60 

30 

40 

50 

Lat. N. 

5 

0 


60 

30 

40 

50 

Lat. N. 

Months 

Months 

N. W. Atlantic 

N.E. Atlantic 

4 

4 

3 

3 


2 

1 

0 

-1 

-2 

-3 

JF MA MJ 

JA 

Months 

SO 

ND 

30 

60 

50 

40 

Lat. N 

Precip. (mm/day) 

2 

1 

0 

-1 

-2 

-3 

JF 

MA 

MJ 

JA 

Months 

SO 

ND 

30 

60 

50 

40 

Lat. N 

Figure 9C4.5. Northwest and northeast Atlantic sea surface temperature in degrees Celsius and bimonthly 

precipitation minus evaporation in mm -d at ten degree latitude intervals for the bimonthly periods of Jan-Feb, Mar- 

Apr, May-Jun, Jul-Aug, Sep-Oct and Nov-Dec. (Note: axes of latitude are reversed between graphs of temperature 

and precipitation.)


Degrees Celsius 

30 

25 

20 

15 

10 

5 

0 

25 30 35 40 45 50 55 60 

North Latitude 

6 

Precip - Evap / Day 

4 

2 

0 

-2 

-4 

25 30 35 40 45 50 55 60 

North Latitude 

NWA NEA NWP NEP 

Figure 9C4.6. Maximum and minimum monthly sea surface temperature in degrees Celsius (A) and monthly 

precipitation minus evaporation in mm d -1 (B) at ten degree latitude intervals for the northwest (dashed lines) and 

northeast (solid lines) Atlantic (thin lines) and Pacific (thick lines). Northeast Pacific, northwest Pacific, northeast 

Atlantic and northwest Atlantic are NEP, NWP, NEA and NWA, respectively. 

Precipitation, and thus salinity in estuaries, also vary from east to west in patterns that 

resemble the south to north pattern from San Francisco to Alaska. The broadest ranges of 

precipitation occur in the eastern Pacific north of 35 o N. Lat. (Figure 9C4.4). The narrowest 

ranges of precipitation and negative net precipitation (desert conditions) occur in the eastern 

Pacific and Atlantic south of 35 o N. Lat. (Figure 9C4.4). Desert conditions do not occur at low 

latitudes in the northwest Pacific and occur in the northwest Atlantic only below 30 o N (Figure


9C4.4). The latitudinal range and areal extent of low salinity estuaries is therefore less in eastern 

oceans than in western oceans and the climates are more uniform. 

The seasonal patterns of precipitation (Figures 9C4.4 and 9C4.5) also differ consistently 

between eastern and western oceans. More precipitation occurs in western oceans during 

summer when temperatures are maximum while most precipitation occurs in eastern oceans in 

winter when temperature are low (Figures 9C4.4 and 9C4.5). Where snow-melt is not important, 

and in the absence of major water impoundments, the salinity-temperature pattens of eastern and 

western ocean estuaries are out of synchrony. In high latitude regions, such as Alaska, runoff 

varies most with snow-melt and salinity is lower in warm seasons in correspondence with 

western ocean climates. 

East and West Biodiversity 

Peracaridan NIS diversity varied among the four northern hemisphere ocean coasts ( X 2 = 

17.27, p = 0.001 df = 3), with five times as many introductions to eastern ocean coasts (Table 

9C4.2; X 2 = 16.05, p = 0.001 df = 1). Except for the gammaridean amphipods Orchestia 

gammarella (Pallas, 1766) and Corophium volutator (Pallas, 1766) in the tidal mudflats and 

marshes of the Bay of Fundy, peracaridan NIS abundance and diversity in western Atlantic 

estuaries appeared to be low. Similarly, of the 28 peracaridan species included in the east to 

west analysis, only 2 - 3 NIS were in the northwest Pacific, compared to 20-23 in the 

northeastern Pacific and 10-12 in the northeast Atlantic (Table 9C4.2). None of the common 

northeast Pacific peracaridans were introduced to other areas of the world. Only 2 of the 

northeast Atlantic species were clearly introduced to other regions compared to13 from the 

northwest Pacific and 14 from the northwest Atlantic (Table 9C4.2). Remarkably, 5 - 9 of the 28 

NIS (Table 9C4.2) have been reported on two coasts and 4 of these species have been discovered 

on 3 coasts. 

Climate and NIS Diversity 

The ranks of climates from least to most similar based on overall temperature variation 

and seasonal precipitation (Figures 9C4.2, 9C4.3 and 9C4.4) were, northeast Pacific, northeast 

Atlantic, northwest Pacific and northwest Atlantic (NEP, NEA, NWP and NWA, respectively). 

The ranks of NIS invaders of these regions (Imports, Table 1) were from highest to lowest: NEP, 

NEA, NWP and NWA. The ranks of native species that have been introduced to other regions 

(Exports, Table 1) were from lowest to highest: NEP, NEA, and NWP and NWA. The east and 

west variations in NIS imports and exports were thus correlated with climate variation (Kendall 

coefficient of concordance W = 1.0; X 2 = 8.2; p < 0.02; df = 2) (Siegal 1956) in a similar pattern 

to the south to north pattern of NIS between San Francisco Bay and Prince William Sound.


Table 9C4.2. The east and west destinations and sources of common introduced peracaridan crustaceans of the 

Northwest Pacific Northeast Pacific, Northwest Atlantic and Northeast Atlantic (NWP, NEP NWA and NEA, 

respectively), their native, introduced, or probable introduced status (N, I, and I, respectively) and the numbered 

reference sources. 

Common Introduced Nonindigenous Estuarine Peracaridan 

Crustaceans of the Northern Hemisphere 

NWP NEP NWA NEA Sources 

Mysidacea 

Acanthomysis aspera N I 1,2 

Cumacea 

Nippoleucon hinumensis N I 1,3 

Isopoda 

Asellus communis I N I 4 

Caecidotea racovitzai I N 7 

Dynoides dentisinus N I 1 

Ianiropsis serricatus N I 1,8 

Paranthura sp. N I 1 

Synidotea laevidorsalis N I I I 1,5 

Amphipoda 

Ampelisca abdita I N 1,6,10 

Ampithoe valida I N 1,9,10 

Apocorophium lacustre N I 13,14 

Caprella acanthogaster N I I 1,10,11 

Corophium sp. N I 15,16 

Corophium volutator I N 13,14,17 

Crangonyx floridanus I N 7 

Crangonyx pseudogracilis N I 18 

Gammarus daiberi I N 1,7 

Gammarus tigrinus N I 13,14,18,19 

Grandidierella japonica N I I 10,20,21 

Jassa marmorata I I N I 10,22,23 

Leucothoe alata N I 1,24 

Melita nitida I I N 1,6,10 

Microdeutopus gryllotalpa I N I 13,25 

Moncorophium uenoi N I 9,10 

Monocorophium acherusicum I I N I 1,9,10 

Monocorophium insidiosum I I N I 1,9,10 

Orchestia gammarella I N 13,14,19 

Parapleustes derzhavini N I 1,6,10 

Sinocorophium heteroceratum N I 1,12 

Total Natives (Exports) 13 0 14 2 

Total NIS (Imports) 1 - 3 21 - 23 2 - 3 10 - 12 

N = native; I = introduced; = distribution not completely resolved but probable 

1) Cohen and Carlton 1995; 2) Modlin and Orsi 1997; 3) Watling 1991; 4) Williams 1972; 5) Chapman and Carlton 1991, 1994; 6) 

Chapman 1988; 7) Toft et al. 1999; 8) Kussakin 1988; 9) Barnard 1975; 10) Carlton 1979; 11) Platvoet et al. 1995; 10) Carlton 1979; 

12) Chapman and Cole In Prep.; 13) Bousfield 1973; 14) Lincoln 1979; 15) Hirayama 1984, 1986; 16) Janta 1995; 17) Chapman and 

Smith In prep.; 18) Costello 1993; 19) Watling 1979; 20) Chapman and Dorman 1975; 21) Smith et al. 1999; 22) Conlan 1990; 23) 

Mills et al. 1999; 24) Nagata 1965a-d; 25) Chapman and Miller In Prep.


Table 9C4.3. Ballast and Port Valdez amphipos zooplankton samples subdivided into major source areas (Port 

Valdez, Benicia water exchanged at sea, San Francisco Bay area and southern California, respectively).


Amphipods in Ballast Water 

The 125 specimens recovered include one hyperiid species and fourteen species and ten 

families of gammaridean amphipods. Five of the gammaridean amphipods are NIS in the 

northeast Pacific and were present in ballast tanks discharged into waters of Port Valdez. These 

preliminary data indicate that ballast water traffic is a potential mechanism for transporting 

amphipods among harbors and coastal U. S. waters. The amphipod diversity in these samples 

(Table 9C4.4) is high given the small number of specimens involved. Except for the Ocean 

exchanged water and the water from Anacortes, Puget Sound, the heterogeneity among 

zooplankton sources is almost complete with a significant difference among the four species 

represented by more than 8 specimens (O 2 ; p < 0.001, df = 12). Only Ampelisca abdita and 

Gammarus daiberi occurred in more than one zooplankton source. Descriptions of the 

amphipods in these records are given in Appendix Table 94C. 8. The occurrence of amphipod 

species as a function of temperature and salinity of ballast water is shown in Figures 9C4.7 and 

9C4.8, respectively. 

Table 9C4.4. The average salinities, teperatures, subtotals, cryptogenic, introduced and native origins and total 

numbers of amphipod Crustacea collected in Port Valdez waters and from dedicated ballast tanks containing water 

exchanged at sea, or entrained from Puget Sound, San Francisco Bay or southern California. 

Discussion 

Most estuarine peracaridan NIS of the northeast Pacific are from the western sides of the 

Pacific or the Atlantic oceans. Peracaridan crustacean NIS diversity coincided with particular 

climates between 25 and 60 o N. Lat. Annual sea surface temperatures at latitudes below 50 o N 

vary less along northeast Pacific and Atlantic coasts than along western ocean coasts and, also in 

contrast to western ocean coasts, low salinity conditions occur in winter months rather than the 

summer months.


Figure 9C4.7. Native, introduced and cryptogenic amphipod numbers with temperature from 34 ballast water 

samples. 

Figure 9C4.8. Native, introduced and cryptogenic amphipod numbers with salinity from 34 ballast water samples. 

The great diversity of invading species in northeast Pacific estuaries may thus result, in 

part, from the great diversity of climates that invading species are adapted to relative to the 

narrow range of climates in the region. The low diversity of native northeast Pacific species that 

invade other areas may also result, in part, from the relatively broad range of climate variations 

they must endure to survive elsewhere. The decline of northeast Pacific NIS diversity from 

south to north coincides with fewer introductions occurring where greater annual variations in 

temperature occur and where low salinity conditions occur during warm water periods. 

All of the peracaridan NIS known from Puget Sound occurred also in San Francisco Bay. 

The absence of cold water NIS in Southern Alaska is consistent with a pattern of introductions


occurring less in continental climates of high temperature variations and low summer salinities. 

The overall pattern of NIS peracaridan diversity in the northern hemisphere strongly suggests 

that northeast Pacific estuarine peracaridan NIS of San Francisco Bay and north are 

predominantly from lower latitudes. Few of the presently known recognized northeast Pacific 

NIS peracaridans are thus likely to become established in southern Alaskan waters, even though 

Alaskan weather variations resemble western ocean climates from where most NIS originate. 

Aquatic species with nearly any life history and from nearly any taxon can be introduced 

(e.g. Carlton 1985, Cohen and Carlton 1995, Eno et al. 1997, Smith et al. 1999, Hewitt et al. 

1999). The many vectors, directions, distributions, routes of introduction (e.g., Carlton 1979, 

1985, 1987, 1999, Carlton and Geller 1993, Ruiz et al. 1997) and taxa available for introductions 

over the last 500 years (e.g. Carlton 1992, Carlton and Hodder 1995, Ruiz et al. 1997) have 

moved broad diversities of species to many suitable and unsuitable areas. The present 

distributions of NIS are a mosaic of surviving populations composed of a broad diversity of taxa 

and life histories. These surviving nonindigenous populations are surrounded by unsuitable 

areas in which they were also introduced but failed to survive. The patterns of transport 

mechanisms superimposed on this mosaic reveal where introductions fail. Resolution of this 

global pattern can reveal sources, destinations, targets and vectors of NIS and which ecosystems 

are most vulnerable. 

The probability of particular species introductions from one region to another cannot be 

determined from these data. NIS invasions in northern hemisphere estuaries, even confined to 

peracaridan crustaceans, are more complex than Tables 9C4.1 and 9C4.2 indicate. Additionally 

many evolutionary and ecological processes that are likely to contribute to the patterns of NIS 

invasions are not addressed. Western ocean estuaries may be older, with more diverse biotas and 

could be less intensely altered by human activities, for instance, than eastern ocean estuaries. 

When and how climates control NIS distributions are complicated by other processes including 

human and natural disturbances, and the timing, geography and magnitude of transport vectors. 

Also, the complexity of climate variations are drastically simplified in Figure 9C4.2, and 

Figures 9C4.4 - 9C4.6 on the assumption that large populations distributed over broad 

geographic areas are more likely to reflect average conditions over extended periods. However, 

other time intervals for integration could be better than the one and two month averages selected 

here for climate analyses. These simplifications may reduce the fit between biogeographical 

boundaries and climate and thus obscure interactions between NIS distributions and climate. 

These correlations between west ocean to east ocean NIS peracaridan invasions and 

climate nevertheless deserve close inspection. Source and destination climates for peracaridan 

NIS may be easier to identify than particular species that are likely to be introduced by particular 

dispersal vectors. Moreover, peracaridans appear to be a sufficient taxon to sample NIS 

biogeography. Similar east to west patterns of other NIS taxa have also been noted (Cohen and 

Carlton 1995, Leppakoski and Olin 2000, Miller and Chapman 2000). Moreover, the patterns of 

NIS invasions must correspond to climate patterns if climate is an important ecological and 

evolutionary mechanism controlling the geography species.


The correlations between marine and estuarine peracaridan NIS invasions with 

temperature and salinity conditions may reveal where, and perhaps how, climates control NIS 

distributions. NIS peracaridans are unlikely to be as suited to local climates as native species 

that have had more evolutionary time to adapt. From a maximum possible of 3, the average 

occurrence of NIS between San Francisco Bay, Puget Sound and Prince William Sound is 1.4 

compared to 1.8 records per native species (Table 9C4.2). This restriction of NIS peracaridan 

distributions in the northeast Pacific relative to native peracaridan species may result from their 

different adaptations to climate. NIS peracaridan dispersal vectors are sufficient to move NIS 

peracaridans to Prince William Sound. Port Valdez is the third largest ballast water port in the 

U.S. and the associated nonindigenous ballast water species that it receives are diverse and 

abundant (Hines et al. 1999) but none of these peracaridan species have been discovered in 

Prince William Sound. At the same time, the same nonindigenous ballast water species are 

invading San Francisco Bay at an accelerating rate (Cohen and Carlton 1998). The greater 

diversity of NIS in San Francisco Bay (Figure 9C4.1), the complete overlap between Puget 

Sound NIS and San Francisco Bay NIS and the absence of NIS in the Prince William Sound 

collections (Table 9C4.1), indicate that peracaridans have warm water origins and survive poorly 

in the cold-water areas of the northeast Pacific. 

The introductions of Corophium volutator and Orchestia gammarellus (Table 9C4.2) 

from Europe to eastern North America are exceptions to the western ocean to eastern ocean 

pattern of introductions listed here. Corophium volutator is confined in North America to the 

Bay of Fundy. Orchestia gammarellus is confined in North America to the Bay of Fundy and 

the outer coasts north to Newfoundland (Bousfield 1973, Watling 1979). The climates of these 

areas are isolated from the Gulf Current and have narrower temperature ranges than areas either 

to the north or south (Bousfield 1973). The successes of these two species, and other European 

species such as the green crab Carcinus maenus and the European littorine snail Littorina littorea 

in this region, and the Bay of Fundy in particular, may result from the closer match between the 

climate of this region and the climate of northern Europe. 

The occurrence of 5 NIS species of amphipods in segregated ballast water of tankers 

discharging into Port Valdez indicates that this is an active mechanism of transport introduding 

NIS arriving to Prince William Sound, even though none of these species appears to have 

become established yet. 

Conclusions 

These results reveal that climate and evolution interact to prevent estuarine peracaridan 

NIS from invading south central Alaska. Western ocean introductions of peracaridans to eastern 

ocean estuaries are common while few eastern ocean peracaridan species spread in the opposite 

direction. Unless Arctic Ocean passages of ballast water from the north Atlantic become 

possible, or global climate changes significantly, or massive shipping from high latitudes of the 

southern hemisphere occurs, ballast water sources NIS peracaridans from climates similar to 

south central Alaska are too remote to pose a significant risk.


Like a lock and key, the adaptations of successful invaders must be sufficient to survive 

in climates and types of disturbances that occur in the new invaded areas. Thermal adaptations 

of western ocean species may thus fit eastern ocean climates while the thermal adaptations of 

most eastern ocean species may be insufficient for western ocean climates. By the same 

mechanism, seasonal salinity disturbances may confine western ocean NIS peracaridans in 

eastern ocean estuaries to areas where the most stable salinity conditions occur and control which 

taxonomic groups of NIS peracaridans predominate in particular estuaries. This pattern of 

invasion appears to be consistent among many taxa even though the particular interacting 

mechanisms of climate and adaptation creating the pattern remain unresolved and the particular 

species that invade remain unpredictable. However, predicting where species can survive 

provides a means to identify, particular mechanisms of introduction, source regions for NIS of 

greatest concern, the likely secondary dispersal routes of newly introduced species and the 

possible role of climate changes on further invasions. 

Seasonal temperature variations are difficult for shallow water organisms to avoid. The 

extreme high and low temperature ranges of western ocean climates (Figures 9C4.4 and 9C4.5) 

create adaptations that span most eastern ocean temperature ranges. The reverse is less likely 

and must prevent survival of many western ocean introductions of eastern ocean species. The 

entire sea surface temperature range of the northeast Pacific between 35 and 50 o N is overlapped 

by the temperature ranges of the western ocean coasts between 37 and 42 o N (Figure 9C4.6). 

Thermal limits for the NIS peracaridans at 60 o N (Prince William Sound) are apparent 

also for the Pacific oyster, Crassostrea gigas (Thunberg, 1795), native to the western Pacific and 

cultured commercially in Asia as far north as Hokkaido, Japan (Quayle, 1969) at about 44 o N. 

Crassostrea grows but does not spawn in the low temperatures of Prince William Sound or south 

central Alaska (Foster 1991, Hines and Ruiz 1997). The minimum monthly temperatures of 

Prince William Sound match western Atlantic and western Pacific minimum temperatures as far 

south as 44 o N (Figure 9C4.6). Thus, lethal low temperatures might not be encountered in the 

sound by western ocean NIS peracaridans from 44 o N or even slightly farther south. However, 

maximum northeast Pacific sea surface temperatures at 60 o N match western Atlantic and 

western Pacific maximums only as far south as 48 o N (Figure 9C4.6) the inability of C. gigas to 

spawn in Alaska is therefore not surprising. 

The poor match of seasonal precipitation across oceans (Figures 9C4.4 and 9C4.5) may 

also affect the patterns of introduction. The range of coinciding temperature and salinity 

tolerances that peracaridan species require to survive and reproduced in south central Alaska 

may prohibit NIS peracaridans that also survive in Long Beach, San Francisco or Puget Sound. 

The low summer salinities in Prince William Sound (Hines and Ruiz 1997) due to snow melt, 

more closely matches a western ocean climate (Figures 9C4.4 and 9C4.5). Only the largest 

eastern ocean estuaries or estuaries with impounded freshwater sources, such as San Francisco 

Bay, are likely to have stable haloclines in summer that match those of western ocean estuaries. 

The high diversity and predominance of benthic peracaridan NIS in the northeast Pacific 

(Table 9C4.1) may result in part from their superior adaptive responses to large salinity ranges


(Figure 9C4.6). All peracaridan life stages are highly mobile, and their short reproductive cycles 

allow dispersal away from unsuitable conditions and rapid recruitment when conditions improve 

(e.g., Watkin 1941). Peracaridans can avoid rapid changes in salinity (Figures 9C4.4 and 9C4.5) 

by migrating short vertical distances or by swimming into water masses in which transport them 

to higher or lower salinity areas. 

San Francisco Bay may be particularly suited to NIS peracaridans that cannot avoid or 

quickly adapt to changing salinities. The particular predominance and high diversity of large, 

long-lived, suspension feeding NIS in San Francisco Bay (Carlton 1979, Nichols et al. 1990, 

Thompson 1998) may result from human water impoundments that limit major freshwater runoff 

events. San Francisco Bay is the largest estuary of the eastern Pacific. Massive water diversions 

and impoundments in the San Francisco Bay watershed, aided by its large size, create a stable 

salinity structure more typical of western ocean estuaries. Sedentary, long-lived species, 

including molluscs (Nicholls et al. 1990), burrowing decapods (Posey et al. 1991, Grosholz and 

Ruiz 1995, Cohen and Carlton 1997) and sedentariate polychaetes (Pearson and Rosenberg 

1978) predominate in benthic communities in the absence of major salinity disturbances. These 

taxa can control the trophic dynamics in estuaries (Nichols et al. 1986, Kimmerer et al. 1994, 

Barber 1997, Thayer et al. 1997, Thompson 1998) but are slow to repopulate areas following 

disturbances. 

The vulnerability of estuaries to invasion and the potentials of particular taxa and life 

history types to become invasive may be increasing globally and increase the risk of NIS 

invasions of Alaskan waters in the future. Water diversions and impoundments and land use 

practices on western coasts combined with global temperature increases are reducing the 

differences between climates of eastern and western ocean estuaries. The convergence of 

climates will increase the potential for biological exchanges. 

The predominantly western ocean to eastern ocean direction of peracaridan invasions and 

south to north gradient in peracaridan invasions support a lock and key hypothesis in which 

peracaridan NIS cannot be introduced outside of their tolerance ranges. Obvious predictions of 

this hypothesis that should be tested are: 1) sources of eastern ocean invaders are from a narrow 

range of western ocean latitudes; 2) eastern ocean invaders of western oceans have restricted 

geographical ranges; 3) specific physiological tolerances and life history adaptations of most NIS 

exceed the stresses experienced in the climates invaded; 4) southern hemisphere NIS invasions 

superimpose on and are superimposed upon by northern hemisphere NIS when their origins are 

from similar climates; 5) climates affect the life histories and taxonomic composition of invaders 

and; 6) invaders of western to western or eastern to eastern oceans have broader ecological and 

geographical ranges than mixed climate invaders. 

The analysis of climates and the geography of introductions must include more species 

and taxa in more detail than is possible here. However, failure to determine the origins of the 

cryptogenic peracaridan species (Table 9C4.1) are a more critical short-coming of this risk 

analysis. The eight cryptogenic peracaridans are abundant over broad ecological distributions 

within the south central Alaska. Even though none of the cryptogenic peracaridan species


appear to be associated with ballast water as a means of introduction, these species are abundant 

and wide spread. If introduced, they are proof that even a few NIS invaders of Alaskan estuaries 

can increase to ecologically catastrophic densities. Moreover surveys of NIS diversity, such as 

this one, are an insufficient for estimating ecological risks if these species are introduced. Due to 

the potential for a single species to produce massive impacts, conclusions of risk from global 

patterns of diversity of diversity and climate therefore could conflict with conclusions of risk 

based on presence of even a single NIS in a system. The occurrence of 5 NIS species of 

amphipods in segregated ballast water of tankers discharging into Port Valdez indicates that this 

is an active mechanism of transport introduding NIS arriving to Prince William Sound, even 

though none of these species appears to have become established yet. 

Acknowledgments 

James Carlton, Williams College, Maritime Studies Program provided many inspirations 

leading to this project, pointing out problems and providing solutions. Todd Miller read an early 

draft of this manuscript. Robert Malouf and Jan Auyong, NOAA Oregon State University Sea 

Grant College Program; and Leon Cammen, NOAA National Sea Grant College Program and 

the Oregon State legislature for assistance with in kind and direct financial support from Grants 

NA0061RD09 and NA0061RD19; Patrick Clinton and Bryan Coleman, OAO, and Walter 

Nelson, US EPA, Coastal Ecology Branch, Newport, Oregon for technical assistance and 

facilities use. The Prince William Sound survey was greatly assisted by Joel Kopp and Robert 

Benda, Regional Citizens’ Advisory Council of Prince William Sound (RCAC), Nora Foster, 

Howard Feder and Mike Stekoll, University of Alaska; Anson Hines, Greg Ruiz and George 

Smith, Smithsonian Environmental Research Center; Gary Sonnevil, US Fish and Wildlife 

Service; the personnel of the Alyeska Pipeline Service Company and Valdez Marine; Todd 

Miller and Gayle Hansen, Oregon State University, Milos Falta, owner, and Capt. Richard 

Vodica of the “Kristina”, Steve Totemov and Gary Kompkoff, of Tatitlek and Gail Irvine of the 

USGS, Anchorage. The Puget Sound survey was funded by the United States Department of 

Natural Resources, the Washington State Department of Natural Resources through Helen Berry 

and Betty Bookheim and facilities were provided by the University of Washington (UW), 

Western Washington State University (WWU), organized by Claudia Mills, (UW) and Andrew 

Cohen, San Francisco Estuary Institute, participants included Brian Bingham,, James Carlton, 

Leslie Harris, L.A. County Museum, Alan Kohn, UW, Terry Klinger, Charles and Gretchen 

Lambert, Kevin Li, David Secord, WWU, Jason Toft, UW and Marjorie Wonham, UW. The 

San Francisco Bay survey was funded through grants from the USFWS, Oregon Sea Grant and 

PEW travel funds to James Carlton and organized by Andrew Cohen and James Carlton. 

Participants included Claudia Mills, Leslie Harris, John Holliman, Gretchen Lambert, Jan 

Thompson, USGS, Menlo Park, Terry Gosliner, California Academy of Sciences, Jeff Crooks, 

U.C. San Diego. 

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Appendix Table 9C4.5. Site Descriptions 

Chapt 9C4. Peracaridan Crustaceans, page 9C4- 32

Appendix Table 9C4.5. Continued 


Appendix Table 9C4.6. Peracardian Crustacea August 1999 


Appendix Table 9C4.7. UAF unidentified Amphipoda 



APPENDIX TABLE 9C4.8 Descriptions of Amphipod Species Identified in Ballast Water 

Tanks. 

Hyperidae 

Hyperidae 

Hyperia cf. medusarum (Bowman 1973:6-10, figs. 2-6 and references therein; Brusca 

1981:21, fig. 9e; Vinogradov et al. 1996:323-327, fig. 131 and references therein). Hyperiaa 

medusarum is a morphologically variable bipolar pelagic species of cold and moderately cold 

water marine regions of both hemispheres, occurring in the Bering Sea, the Gulf of Alaska and 

the coastal waters of Canada and the western U. S. (Vinogradov et al. 1996). Specimens of this 

study are all less than 4 mm in length, and none are mature. Gnathopods, mandibles and 

pereopods closely resemble H. medusarum but the identifications are tentative. The sutures 

between pereonites 1 and 2 and between coxal plates and pereionites are faint. Their appearance 

only in the Anacortes samples and in the open ocean exchanged ballast water samples (Table 1) 

are consistent with the life history of typical hyperiids and this species is very likely native to the 

region. 

Gammaridea 

Ampeliscidae 

Ampelisca abdita Mills, 1964a; Northeast Pacific records of Ampelisca milleri are: Jones 

1961:253-254; Filice 1959a:183; Filice 1959b:10; Chapman and Dorman 1975:107, 106; 

Chapman 1988:365-368, fig. 2 (and references therein); Dickensen 1982:15-17, fig. 9; (not 

Barnard 1954b:9-11). The range of A. abdita on the east coast of the U.S.extends from Maine to 

the eastern Gulf of Mexico (Mills, 1967, Bousfield 1973, Chapman 1988). 

Ampelisca abdita from San Francisco Bay, Bolinas Lagoon and Tomales Bay was 

confused with Ampelisca milleri Barnard 1954 for 40 years and was probably introduced into 

San Francisco Bay from the eastern U. S. with shipments of the eastern oyster Crassostrea 

virginica(Chapman 1988). The species dominates soft, subtidal sedimentst’s of San Francisco 

Bay at salinities between 10 and 25 PSU and must occasionally occur as zooplankton in massive 

numbers as it seasonally exits and repopulates shallow subtidal an intertidal mudflats (Mills 

1967). 

Arigissidae 

Argissa hamatipes (Norman 1869); Syrrhoe hamatipes Norman 1869:279; Boeck 

1871:125; Argissa stebbingi Bonnier 1896:626-630, pl.36, fig.4; Argissa typica Sars 

1895:141-142, pl.48; Chevreux & Fage 1925:90, figs.81-82; Ruffo 1982:159-161, figs. 106-107; 

Argissa hamatipes Walker 1904:246; Stebbing 1906:277; Shoemaker 1930:37-40, figs.15-16; 

Stephensen 1935:140; Stephensen 1940:41; Stephensen 1944:52; Gurjanova 1951:327-328, 

fig.193; Gurjanova 1962:392-393; J.L. Barnard 1962c:151; J.L. Barnard 1964a:218-219; Nagata 

1965:154-155, fig.7; J.L. Barnard 1966a:61; J.L. Barnard 1967a:14-15, fig.1d-i; J.L. Barnard 

1969b:159, fig. 65; J.L. Barnard 1971b:9; Bousfield 1973:121-122, pl.XX; Griffiths 1975:; 

Lincoln 1979:334, fig.157; Ledoyer 1982:144-146, fig. 50; Hirayama 1983:147-149, figs.38-41; 

Barnard & Barnard 1983:607-608; Thomas & McCann 1997:22, fig.2.1. 

Argissa hamatipes is an entirely marine, nearly cosmopolitan species with an extensive 

bathymetric distribution that ranges throughout coastal north Pacific shelf regions from southern 

California to southern Japan and the north Atlantic from North Carolina Greenland and Iceland,


throughout northern Europe and the western Mediterranean, Madagascar and South Africa. 

Many of the the synonymies are unclear. This extremely dispersed species is likely to be a 

species complex and the identity of the North Pacific population is probably incorrect. 

Nevertheless, the very broad open ocean distribution in the northeast Pacific population strongly 

indicates that it is a native to the region. 

Corophiidae 

Monocorophium acherusicum Podacerus cylindricus Lucas 1842:232; Corophium 

cylindricum Smith 1873:566; Paulmier 1905:167, fig. 37; Holmes 1905:521-522, fig. ; C. 

cylindricus Stebbing 1914:372-373; Kunkel 1918:171-173, fig. 52; Corophium contractum 

Thompson 1881:220-221, fig. 9; Corophium bonnellii K. H. Barnard 1932:244 (in Crawford 

1937); Corophium acherusicum 1853:178; Costa 1857:232, Fig. 1827; Bate 1862:282; Heller 

1867:51-52, pl 4 fig. 14; DeElla Valle 1893:364-367, pl. I, Fig. II, Pl. 8, figs. 17, 18, 20-41; 

Sowinsky 1897:9; Sowinsky 1898:455; Chevreux 1900a:109; Graeffe 1902:20; Holmes 

1905:521-522, fig. ; Stebbing 1906:692-740; Chevreux 1911:271; K. H. Barnard 1916:272-274; 

Stebbing 1917a:448; Ussing and Stephensen 1924:78-79; Chevreux 1925c:271; Chevreux and 

Fage 1925:368, fig. 376; Chevreux 1926:392; Cecchini 1928e:8, pl. 1, fig. 6a; Cecchini 

1928b:309-312, fig. 1; Schellenberg 1928:672;Schijfsma 1931a:22-25; Monod 1931a:499; Fage 

1933:224; Candeias 1934:3; Shoemaker 1934c:24-25; Cecchini-Parenzan 1935:227-229, fig. 52; 

Shoemaker 1935c:250; Crawford 1936:104; Schellenberg 1936c:21; Schijsfma 1936:122-123; 

Crawford 1937:617-620, 650, fig. 2; Monod 1937:13; Miloslavskaya 1939:148-149; K. H. 

Barnard 1940:482; Bassindale 1941:174; Stephensen 1944a:134; Shoemaker 1947:53, figs. 2, 3; 

Shoemaker 1949a:76; Soika 1949:210-211; Gurjanova 1951:977-978, fig. 680; Reid 1951:269; 

Stock and Bloklader 1952:4-5; J.L. Barnard 1954a:36; Hurley 1954e:442-445, figs. 35-39; J. L. 

Barnard 1955a:37; Irie 1957:5-6, fig. 6; Irie 1958c:145; Irie 1959: tab. 4; J.L.Barnard 1959:38; 

Nayar 1959:43-44, pl. XV, figs. 14-20; Irie In Okada and Ochida et al. 1960:122, pl. 61, fig. 12; 

Nagata 1960:177; J. L. Barnard 1961:182; Irie and Nagata 1962:20; Nagata 1964:10; Barnard 

1964a:111, chart 5; Nagata 1965c:317; Nagata 1966:334; Kikuchi 1966:, tab 21; Kikuchi 

1968:179; Reish and J. L. Barnard 1967:16; Ledoyer 1968:214; Fearn-Wannan 1968b:134-135; 

Mordhukai-Boltovskoi 1969:485, pl. 25, fig. 2; Sivaprakasam 1969d:156, fig. 14; Bellan-Santini 

1971:260-261; J. L. Barnard 1971a:59; J. L. Barnard 1972b:48; Bousfield 1973:201, Pl. LXII.2; 

Griffiths 1974:181-182;Griffiths 1974b:228; Griffiths 1974c:281; Griffiths 1975:109; Fox and 

Bynum 1975:225; Hirayama 1984:13, fig. 50; Azuma 1986:77; Sudo et al. 1987:1570; Inaba 

1988:141; Monocorophium acherusicum Bousfield and Hoover 1997:111-114, figs. 26-27. 

Monocorophium acherusicum could be the most widely distributed and widely 

introduced estuary invertebrate in the world, occurring in fouling and benthic mud communities 

of shallow and intertidal areas on all continents except Antarctica. C. acherusicum has been 

reported at all latitudes between 60 o North and South. However, many of the records of this 

species at latitudes greater than 60 o , including all records from Alaska are doubtful (Hines et al. 

1999) due likely confusion with other species. M. acherusicum was not found in the present 

surveys of south central Alaska, including Port Valdez and Prince William Sound. 

Parthenogenic populations of an extremely similar species, that closely resembles 

Monocorophium carlottensis Bousfield and Hoover, 1997, occurs in nearly all fouling 

communities of these areas (Chapman 1999, Peracarida and Decapoda). All Alaskan records of


the nonindigenous species, M. insidiosum and M. acherusicum are probably referrable to M. 

carlottensis. 

Sinocorophium heteroceratum (Yu, 1938) Corophium heteroceratum Yu 1938:93-101, 

figs. 7-11; Sinocorophium heteroceratum Bousfield and Hoover 1997:75, 78. 

Sinocorophium heteroceratum was introduced into San Francisco Bay and Los Angeles 

Harbor from Asia in the mid 1980s where it presumed to have been transported with ballast 

water traffic (Cohen and Carlton 1995). This species occurs predominantly in soft sediments in 

high salinity, sub-tidal areas of San Francisco Bay and Los Angeles Harbor (Chapman and Cole, 

MS in preparation). Sinocorophium heteroceratum are also unlikely to have been introduced 

with aquaculture industries, or ship fouling in San Francisco Bay or Los Angeles Harbor because 

its lack of association with fouling communities. 

Grandidierella japonica Stephensen, 1938:179-184; Nagata 1960:179; Nagata 

1965c:320-321; Chapman and Dorman 1975:104-108, 4 figs., Nagata 1984:15, fig. 53-56; Muir 

1997; Smith et al. 1999:8-9, fig. 3. 

Grandidierella japonica occurs in salinities from 5-40 PSU predominantly in warm 

intertidal areas of nearly all estuaries from Puget Sound to San Diego. Dense populations are 

especially common in low-salinity tidepools and seepage areas; mixed sediment. The species has 

been introduced to Hawaii (Muir 1997) England (Smith et al. 1999), and estuaries of the 

northeast Pacific between Puget Sound and San Diego (Chapman and Dorman 1975, Staude 

1997, Bay et al. 1988). Grandidierella was most likely introduced to the northeast Pacific with 

Pacific oysters from Japan between the 1930s and 1950s (Chapman and Dorman 1975). Its 

recent arrival in Europe (Smith et al. 1999) is more likely to be associated with ballast water. It 

was misidentified in Hawaii for 25 years as Neogamphopus cabinae (Muir 1997) where the 

mechanism of its introduction is unclear. The massive ballast water traffic from San Francisco 

Bay to Hawaii is a possible mechanism for its introduction there. 

Cyphocharidae 

Cyphocharis challengeri Stebbing 1888: ;Birstein and Vinogradov 1955:212 (with 

references); J. L. Barnard 1961b:31; Bowman and McCain 1967:1-14, figs. 1-9 (with 

references). 

The most common epipelagic gammaridean amphipod in sub-Arctic offshore and coastal 

waters of the North Pacific is Cyphocharis challengeri (Bowman and McCain 1967). Bousfield 

and McCain (1967) demonstrated that most of the variation in the anterior protrusion of 

pereonite 1 varies with size. The species is endemic to the North Pacific but has been reported 

from a broad range of latitudes in the Atlantic and Indian Ocean. . 

Eusiridae 

Pontogeneia rostrata Gurjanova, 1938; Gurjanova, 1938:330,398, fig.39; Gurjanova, 

1951:719, fig.500; Nagata 1960:171-173, pl.14; J.L.Barnard, 1962b:81; J.L.Barnard, 

1964b:114-116, fig.20; Nagata 1965b:185, fig. 26; Nagata 1965e:563; Nagata 1966:334; 

J.L.Barnard, 1969a:111,112,114; Itoh 1970:29; Mukai 1971:178; Itoh, Honma and Kakimoto 

1972:25; Honma and Kitami 1978:40; Azuma 1980:28; Barnard 1979a:49, figs. 25-27 (part); 

Imada et al. 1981:127; Itoh 1981:24; Itoh, Honma and Kitami 1982:41; Hirayama 1985a:28; 

Azuma et al. 1985:4; Azuma 1986:74; Sudo et al. 1987:1570; Inaba 1988:146; Ariyama 

1988:129, fig. 13f; Barnard and Karaman 1991:334; Ishimaru 1994:45.


A widely reported species in the North Pacific from Bahia de San Quintin, Mexico 

(Barnard 1964) to Alaska and down the Asian coast to southern Japan (Ishimaru, 1994). 

Pontogeneia rostrata was recovered in the nonindigenous species surveys of Prince William 

Sound (Hines 1999). However, P. rostrata can easily be confused with species of Accedemorea 

its extremely broad range in the Northeast Pacific and uncertain taxonomic status prevents clear 

resolution of whether it is endemic or introduced. This species is cryptogenic (Carlton 1996). 

Gammaridae 

Gammarus daiberi Bousfield, 1969; Bousfield 1969:10(1)4-8, figs. 1& 4; Bousfield 

1973:52, pl. IV.2; Toft et al. 1999:36. 

A probable 1980s ballast water introduction into San Francisco Bay from the eastern 

Unites States. Gammarus daiberi is an estuarine species most abundant in the low salinity 

ranges between 1.5 and 15 PSU and is largely pelagic. Except for this report, this species is 

unknown in the northeast Pacific outside of San Francisco Bay. Its range in the eastern U.S. 

extends from Delaware Bay to South Carolina (Bousfield 1973). Gammarus daiberi is a 

probable ballast water introduction into San Francisco Bay that arrived in the 1980s. 

Eogammarus confervicolus Mara confervicola Stimpson 1856:90; Gammarus 

confervicolus Stimpson 1857:520-521; Bate 1862:218, pl. 38, fig. 9; Holmes 1904:239; Melita 

confervicola Stebbing 1906:428; Anisogammarus confervicolus Saunders 1933:248 (in part); J.L. 

Barnard 1954a:9-12, pls.9-10; Shoemaker 1964:423, figs. 14-15; Pamamat 1968:211; Bousfield 

and Hubbard 1968:3; Anisogammarus (Eogammarus) confervicolus Schellenberg 1937a:274; 

Tzvetkova 1975:145-147, fig. 57; Bousfield 1958a:86, fig. 10; Eogammarus confervicolus 

Bousfield 1979:317-319, fig.4. 

Eogammarus confervicolus is the only native species recovered from ballast water that 

was also collected in the Prince William Sound nonindigenous species survey (Chapman 1999). 

Eogammarus confervicolus is extremely euryhaline and is occasionally pelagic. It occurs mainly 

in estuaries and protected coastal shores. Eogammarus confervicolus is the most common and 

widely distributed gammaroidean amphipod of the North American Pacific coast (Bousfield 

1979). 

Megaluropidae 

Gibberosus longimerus Hoek; Megaluropus longimerus; J.L.Barnard 1962b:103, 

figs.20-21; J.L.Barnard, 1964a:224; J.L. Barnard 1966b:19; J.L.Barnard, 1969a:126; J.L. 

Barnard 1971b:15; Gibberosus myersi (McKinney 1980); Megaluropus myersi McKinney 1980; 

Megaluropus longimerus of Cadien et al. NEP (in part); (not Schellenberg 1925:151-153, 

fig.14). 

The genus is poorly studied north of northern California (Cadien et al. 1997). 

Schellenberg (1925) figured only two appendages from his specimens from Lagos, Nigeria and 

the type specimens have not been compared to California material. The possibility of these 

populations comprising a single species seems remote. The common occurrence of this species 

over a broad range of coastal and nearshore waters (Barnard 1962) indicates that it is endemic. 

Melphidipiidae 

Melphisana bola Barnard 1962b:81-82, fig.7; Thomas & McCann 1997:42, fig.2.20. 

This native species is limited to depths no greater than 130 m on the southern California 

coastal shelf (Barnard 1962b, Thomas and McCann 1997). Appendages are usually missing this 

species on recovery from benthic samples (Barnard 1962b, Thomas and McCann 1997) greatly


complicating identifications. These specimens are in excellent condition is in sharp contrast ot 

previous material on which the species is described. 

Oedicerotidae 

Hartmanodes hartmanae Monoculodes hartmanae J.L. Barnard 1962:363, figs. 6-7; 

Barnard & Karaman 1991:560; Hartmanodes hartmannae Bousfield & Chevrier 1996:92-93, fig. 

10. Bousfield and Chevrier (1996) did not find this species in their 200 samples from the shelf 

benthos and coastal waters of Canada and it is not listed in by Staude (1997). This species is 

native to southern California marine waters where it occurs at less than 40 m depths (Barnard 

1962). Hartmanodes hartmannae is the most abundant gammaridean amphipod found in the 

samples. 

Westwoodilla caecula Halimedon caecula Bate 1857:140; Halimedon Molleri Boeck 

1871:169-170; Halimedon Mulleri Sars 1895:327-329, pl.115; Halimedon acutifrons Sars 

1895:329-330, pl.116, fig.1; Westwoodilla caecula Enequist 1950:333-338, figs.40-56; 

Gurjanova 1951:541-543, pl. 357; Mills 1962:5-9, fig.1; J.L. Barnard 1962e:370; J.L. Barnard 

1964a:235; J.L. Barnard 1966a:80 (forma acutifrons); J.L. Barnard 1966b:27; J.L. Barnard 

1971b:51; Lincoln 1979:354, fig.167; Thomas & McCann 1997:58, figs. 2.37, 2.38; Beare, D. J. 

and P. G. Moore 1998. 

This species may not be part of a complex of similar species distributed around the Arctic 

Ocean, the Japan Sea, the north east Pacific from British Columbia to southern California and the 

North Atlantic from Greenland to the Gulf of St. Lawrence and northern Europe. This extremely 

widespread, common, species in offshore marine soft-sediment environments is most likely 

endemic to cold water areas of the northeast Pacific. Its occurrence in Port Valdez zooplankton 

samples is not surprising. 

Synopiidae 

Tiron sp. 

The short dactyls, spineless telson, tiny mandibular palp, smooth dorsal urosome of this 

single specimen do not agree with either of the local species Tiron tropakis J. L. Barnard, 1972 

or Tiron biocellata J. L. Barnard, 1962. Pelagic dispersal of benthic peracaridans usually occurs 

as adults and often is preceded by slight morphological changes that are adaptive for swimming. 

These changes are poorly understood. The low morphological correspondence of this single 

specimen with a known species is therefore not surprising. The specimen is therefore considered 

more likely to be a member of one of the above native species than an introduced species. 

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Vinogradov, M. E., A. F. Volkov and T. N. Semenova, 1996., Hyperiid amphipods (Amphipoda, 

Hyperiidea) of the world oceans,,, Science Publishers, Inc., 10 Water Street, #310, Lebanon, 

NH 03766, xxvii + 632 pp. pp. 

Williams, A. B. and K. H. Bynum 1972. A ten-year study of meroplankton in North Carolina 

estuaries: Amphipods, Chesapeake Sci., 13:175-192 

Williams, R. J., F. B. Griffiths, E. J. Van der Wal and J. Kelly 1988. Cargo vessel ballast water 

as a vector for the transport of nonindigenous marine species, Estuar. Coast. Shelf. Sci., 

26:409-420 

Yu, S. C. 1938. Descriptions of two new amphipod Crustacea from Tankgu, Bulletin of the Fan 

Memorial Institute of Zoology, 8:83-1

Chapt 9C5. Copepod Crustaceans, page 9C5- 1 

Chapter 9C5. Focal Taxonomic Collections: Copepod Crustaceans 

Jeffery R. Cordell, Wetland Ecosystems Team, Fishery Sciences, University of Washington, 

Seattle 

Methods 

Copepods were identified from three types of samples. The first method consisted of 

sweeps that were made through algal and fouling assemblages on the underside of docks, using a 

small hand-held net consisting of 130 µm mesh material attached to a stainless steel hoop of 

approximately 15 cm diameter. An effort was made to disturb algal and bivalve holdfasts in 

order to capture copepods from those microhabitats. The second type of samples taken were 

vertical water column plankton hauls made either off a dock or from a small boat in the harbor 

with a 0.25 m diameter 250 µm mesh plankton net. The net was lowered to the bottom, and after 

waiting approximately 1 minute for disturbance to dissipate, the net was slowly pulled to the 

surface. The third type of samples were taken from settling plates used for collecting fouling 

macro-invertebrates by briefly soaking plates in 5% formaldehyde solution in a plastic tub and 

washing the residue through 130 µm mesh. 

In the laboratory, each sample was examined under a dissecting microscope, and several 

representatives of each species of copepod were removed. Each species was then further 

examined under a compound microscope. Identification was made as far as possible without 

dissection of individuals (with the exception of occasional removal of the abdomen to facilitate 

viewing the fifth leg). 

Results and Discussion 

We identified 68 species of harpacticoid copepods, six calanoids, four cyclopoids, and 

several unidentified poecilostomatoid species (Table 9C5.1). Of these, none is a confirmed 

introduced species. 

For harpacticoids, our results were similar to those of Kask et al. (1982) for the Nanaimo 

estuary in southern British Columbia in both the number of species found (75 in B.C.) and in that 

most of the species that were identified were either northern Pacific, broadly distributed boreal 

species (i.e., Pacific and Atlantic records), or probably undescribed species. Kask et al. (1982) 

speculated that the presence of many of the species that they found might have been the result of 

introductions from ship fouling communities. Harpacticoid copepods may be particularly likely 

to be transported and introduced because as a group they have successfully occupied almost all 

benthic and epibenthic habitats. Also, many species have multiple life history modes (e.g., 

resting and planktonic stages) that may also increase their chance of being transported and 

introduced. However, these same factors may also explain wide natural distributions. The 

paucity of studies of harpacticoid taxonomy in the northeastern Pacific makes it nearly 

impossible to determine whether or not a given species has been introduced without extensive 

distribution or genetic studies. Many of the species described in Lang’s monograph on the 

harpacticoids of central California (Lang, 1965) occur in Puget Sound (J. Cordell, unpublished 

data) and southern British Columbia (Kask et. al., 1982), and Lang’s species that also occur in 

Prince William Sound (Table 9C5.2) probably have continuous distributions. Other species that 

we encountered have Arctic and circumboreal distributions. An example of this is the important


Table 9C5.1. Copepoda 

Order Harpacticoida 

Fam. Ameiridae 

Ameira longipes Boeck, 1865 

Ameira sp. 1 

Ameira sp. 2 

Ameiridae, unid. sp. 1 

Fam. Ancorabolidae 

Arthropsyllus serratus Sars, 1909 

Fam. Canthocamptidae 

Mesochra pygmaea (Claus, 1863) 

Mesochra sp. 1 

Fam. Canthocamptidae, incertae sedis 

Leimia vaga Willey, 1923 

Fam. Cletodidae 

Acrenhydrosoma sp. 

Fam. Danielsseniidae 

Danielssenia typica 

Fam Diosaccidae 

Diosaccus spinatus Lang, 1965 

Diosaccus sp. 1 

Amphiascopsis cinctus 

Amphiascus minutus 

Amphiascus sp. 1 

Amphiascoides cf. debilis 

Amphiascoides sp. 1 

Amonardia perturbata Lang, 1965 

Amonardia normani 

Robertsonia sp. 

Stenhelia peniculata Lang, 1965 

Fam. Ectinosomatidae 

Ectinosoma sp. 

Halectinosoma sp. 1 



Microsetella norvegica 

Fam. Tachidiidae 

Microarthridion littorale (Poppe, 1881)


Table 9C5.1. (continued) Copepoda 

Fam. Harpacticidae 

Harpacticus uniremis Kröyer, 1842 

Harpacticus septentrionalis Klie, 1941 

Harpacticus compressus Frost, 1967 

Harpacticus sp.- uniremis group 1 

Harpacticus sp.- obscurus group 1 


Zaus sp. 

Fam. Laophontidae 

Echinolaophonte sp. 

Heterolaophonte discophora (Willey, 1929) 

Heterolaophonte longisetigera Klie, 1950 

Heterolaophonte variabilis Lang, 1965 

Heterolaophonte sp. 1 

Laophonte elongata Boeck, 1872 

Laophonte applanata 

Laophonte sp. 1 

Pseudonychocamptus spinifer Lang, 1965 

Paralaophonte cf. congenera (Sars, 1908) 

Paralaophonte pacifica Lang, 1965 

Paralaophonte perplexa (T. Scott, 1898) 

Paralaophonte hyperborea (Sars, 1909) 

Paralaophonte sp. 1 

Laophontidae, unid. 

Fam. Longipediidae 

Longipedia sp. 

Fam. Parastenheliidae 

Parastenhelia sp. 1 


Fam. Tegastidae 

Tegastes sp. 1 


Fam. Thalestridae 

Idomene sp. 

Diarthrodes sp. 2 

Parathalestris sp. 1 



Dactylopusia vulgaris Sars, 1905 

Dactylopusia glacialis Sars, 1909 

Dactylopusia cf. glacialis 

Dactylopusia paratisboides Lang, 1965 

Dactylopusia sp. 1 

Paradactylopodia sp. 1


Table 9C5.1. (continued) Copepoda 

Fam. Tisbidae 

Tisbe cf. furcata (Baird, 1837) 

Tisbe spp. 

Scutellidium arthuri Poppe, 1884 

Order Cyclopoida 

Fam. Cyclopinidae 

Fam. Cyclopidae 

Euryte sp. 

Halicyclops sp. 

Fam. Oithonidae 

Oithona similis Claus, 1863 

O. spinirostris Claus, 1863 

Order Poecilostomatoida 

Unidentified spp. 

Order Calanoida 

Fam. Acartiidae 

Acartia cf. clausi Giesbrecht, 1889 

Acartia longiremis (Lilljeborg, 1853) 

Fam. Centropagidae 

Centropages abdominalis Sato, 1913 

Fam. Paracalanidae 

Paracalanus sp. 

Fam. Pseudocalanidae 

Pseudocalanus spp. 

Fam. Temoridae 

Eurytemora herdmani Thompson and Scott, 1897


Table 9C5.2. Copepods Collected at 6 Locations in Southcentral Alaska in August 1999. 

1999 Copepoda Homer Whittier Cordova Valdez Seward Shotgun 

Cove 

Order Harpacticoida 

Fam. Ameiridae 

Ameira longipes 

x 

Ameira sp. 1 x x x x 

Ameira sp. 2 x x 

Family Ancorabolidae 

Arthropsyllus serratus 

x 

Fam. Canthocamptidae 

Mesochra pygmaea x x 

Mesochra sp. 1 

x 

Fam. Canthocamptidae, incertae sedis 

Leimia vaga 

x 

Fam. Danielsseniidae 

Danielssenia typica 

x 

Fam Diosaccidae 

Diosaccus spinatus x x 

Amphiascopsis cinctus x x x 

Amphiascus minutus x x x x 

Amphiascus sp. 1 x x 

Amphiascoides sp. 1 

x 

Amonardia perturbata x x 

Fam. Ectinosomatidae 

Ectinosoma sp. 

Fam. Harpacticidae 

Harpacticus uniremis x x x x x 

H. septentrionalis x 

H. compressus x 

H. unidentified sp. x 

Harpacticus sp. A- uniremis group 

x 

Harpacticus sp.- obscurus group 1 x x x 


x 

Zaus sp. 

Fam. Laophontidae 


x 

Heterolaophonte discophora x x 

Heterolaophonte longisetigera x x x 

Laophonte elongata 

x 

Pseudonychocamptus spinifer x x 

Paralaophonte cf. congenera 

x 

Paralaophonte pacifica 

x 

Paralaophonte perplexa x x x 

Paralaophonte sp. 1 

x 

Family Parastenheliidae 


x 


x


Table 9C5.2. (continued) Copepods Collected at 6 Locations in Southcentral Alaska in 

August 1999. 

1999 Copepoda Homer Whittier Cordova Valdez Seward Shotgun 

Cove 

Fam. Tegastidae 

x 

Tegastes sp. 1 x x 


Fam. Thalestriade 


x 


x 


x 


x 


x 

Dactylopusia vulgaris x x x x 

Dactylopusia cf. glacialis 

x 

Paradactylopodia sp. 

x 

Fam. Tisbidae 

Tisbe cf. furcata x x 

Tisbe sp. x x x x 

Scutellidium arthuri x x 

Order Cyclopoida 

Family Cyclopinidae 

x 

Family Cyclopidae 

Euryte sp. x x x 

Halicyclops sp. x x 

Order Poecilostomatoida 

x 

Total Number of Taxa 28 11 20 1 3 23


juvenile salmon prey species Harpacticus uniremis, which occurs in the arctic and as far south as 

the English Channel in the Atlantic (Kask et al. 1982) and La Jolla, California in the Pacific 

(Gunnill, 1982). 

Settling plates appear to be a good way to sample the diversity of harpacticoid and other 

epibenthic/epiphytic copepods. In each case where we had dock sweep samples to compare with 

settling plate samples, more species were collected from the settling plates. Only one species, 

the algal blade dwelling Scutellidium arthuri was found only in the dock sweep samples. 

Reduced numbers of copepod taxa in the dock sweep samples was probably due to low and/or 

highly fluctuating surface salinities and/or temperatures. This assertion is supported by the fact 

that dock sweep samples taken in harbors with high freshwater input (e.g., Valdez, Seward) had 

extremely low taxa numbers, and those without much freshwater (Homer, Shotgun Cove) had the 

highest number of taxa. 

One species of harpacticoid copepod that we found in dock sweep collections, Leimia 

vaga, may be regarded as “probably introduced”. This species, which was described from Nova 

Scotia, is also abundant in many estuaries in Oregon and Washington, where it is restricted to 

brackish reaches (J. Cordell, unpublished data); however, this species was not reported from the 

Nanaimo estuary by Kask (1982). The fact that L. vaga has restricted habitat requirements and 

apparently disjunct populations on the Pacific coast may indicate that it has been introduced. 

Although a number of Asian planktonic copepods have become established in California, 

Oregon, and Washington estuaries (e.g., Cordell and Morrison, 1996, Orsi and Ohtsuka, 1999), 

we found no introduced species in the vertical haul samples. In fact, overall planktonic copepod 

diversity was quite low, and almost all of the copepod numbers were made up of only three taxa: 

Acartia longiremis, Pseudocalanus spp., and Oithona similis. Also, we did not encounter several 

taxa that have been previously reported from ballast water arriving to Prince William Sound 

(Ruiz and Hines, 1997; Hines et al., 1998; Chapt. 3 Biological Characteristics of Ballast Water). 

As with dock-associated harpacticoids, our shallow sampling depths that were probably subject 

to large fluctuations in salinity and temperature may have decreased diversity of planktonic 

copepods in our samples. 

References 

Gunnill, F.C. 1982. Effects of plant size and distribution on the numbers of invertebrate species 

and individuals inhabiting the brown alga Pelvetia fastigiata. Mar. Biol., 69: 263-280. 

Kask, B.A., J.R. Sibert, and B. Windecker. 1982. A check list of marine and brackish water 

harpacticoid copepods from the Nanaimo estuary, southwestern British Columbia. Syesis, 15: 

25-38. 

Lang, K. 1965. Copepoda Harpacticoidea from the Californian Pacific coast. Kungliga Svenska 

Vetenskapsakademiens Handlingar, (4)10(2), 1-560. 

Cordell, J.R. and S.M. Morrison. 1996. The invasive Asian copepod Pseudodiaptomus inopinus 

in Oregon, Washington, and British Columbia estuaries. Estuaries, 19 (3): 629-638.


Orsi, J.J., and S. Ohtsuka. 1999. Introduction of the Asian copepods Acartiella sinensis, 

Tortanus dextrilobatus (Copepoda: Calanoida), and Limnoithona tetraspina (Copepoda: 

Cyclopoida) to the San Francisco estuary, California, USA. Plank. Biol. Ecol., 46 (2): 128-131.

Chapt 9C6 Decapod Crustaceans, page 9C6- 1 

Chapter 9C6. Focal Taxonomic Collections: Decapod Crustaceans 

Anson Hines, Smithsonian Environmental Research Center 

Lise Schickel, University of California, Santa Barbara 

Nora Foster, University of Alaska Museum 

Results 

We collected a total of 21 species of decapods, including 12 species of brachyuran crabs, 

3 species of lithodid crabs, 3 species of hermit crabs, and 5 species of caridean shrimp (Table 

9C6.1). All of these species were collected by hand or dip net in the intertidal and shallow 

subtidal zones, or on float fouling communities. None of these species is a new record for the 

region (Jensen, 1995). 

We also noted parasitic castrators (rhizocephalan cirripedes, entoniscid isopods) of 

decapods in the collections during August 1999. Several of the samples of the hermit crab 

Pagurus hirsutiusculus exhibited infections by rhizocephalan parasites. The crab 

Lophopanopeus bellus also has a high prevalence of infection by the rhizocephalan 

“Loxothylacus panopaei” at Tatitlek. (Note: The name given to this rhizocephalan in the 

literature by Boschma (1955) for the west coast of North America is undoubtedly incorrect and 

should be renamed, as this is not the same species that is found in xanthid crabs of the eastern 

and gulf coasts.) The population of the shore crab Hemigrapsus oregonensis also had high 

prevalence of the entoniscid isopod Portunion conformis. 

References 

Boschma, H. 1955. The described species of the family Sacculinidae. Zool. Verhandel. 27: 48- 

76. 

Jensen, G.C. 1995. Pacific Coast Crabs and Shrimps. Sea Challengers, Monterey, California. 87 

pp.

Chapt 9C6 Decapod Crustaceans, page 9C6- 2 

Table 9C6.1. Decapod Crustaceans in Field Surveys 

Decapod Crustaceans 

1997; 1998; 1999 

Homer Sew 

- 

ard 

Whittier 

Port Saw- 

Valdez Mill 

Bay 

Rocky 

Pt., 

Gallen 

a Bay 

Busby 

Is. 

Tatitlek 

Cordovcier 

Gla- 

Is. 

Green 

Island 

Consta- 

Tine 

Harbor 

Port 

Chalmers 

Brachura 

Cancer magister X X X X X 

Cancer oregonensis X X X X X X X X X 

Cancer productus X X 

Cancer gracilis 

X 

Chionoecetes bairdi 

X 

(molts) 

Chorilla longipes 

X 

Hemigrapsus oregonensis X X X X X X,9 X,4 X,7 X X 

Lophopanopeus bellus X X X,5 X X 

Oregonia gracilis X X X X 

Pugettia gracilis X X X 

Scyra acutifrons 

X 

Telmessus cheiragonus X X X X X X X X X 

Anomura 

Cryptolithodes typicus 

X 

Hapalogaster grebnitzii X X X X 

Phyllolithodes papillosus 

X 

Pagurus beringanus X X 

Pagurus granosimanus X X X X 

Pagurus hirsutiusculus X,1 X,2 X X,3 X X X X,6 X,8 X X X X 

Caridea 

Eualus b9iunguis 

X 

Hyppolyte clarki X X 

Heptacarpus stimpsoni X X X 

Spirontocaris ochotensis X X X X 

Spirontocaris prionota X X 

Parasite Notes: 

Aug 1999: 

1 = 7/9 with bopyrid isopod 

2 = 1/32 with rhizocephalan Peltogaster paguri; 1/32 with rhizocephalan 

Peltogasterella gracilis 

3 = 0/9 infected 

4 = 30/45 with entoniscid isopod Portunion 

conformis 

5 = 19/44 with rhizocephalan Loxothylacus 

panopei 

6 = 1/6 with bopyrid isopod; 2/6 with rhizocephalan Peltogaster paguri; 1/6 with isopod 

and P. paguri 

7 = 0/13 with Portunion conformis 

8 = 3/30 with Peltogaster paguri; 

1/30 with Peltogasterella gracilis 

June 1998: 9 = 2/93 with 

rhizocephalan

Chapt 9C7. Shelled Molluscs, page 9C7- 1 

Chapter 9C7. Focal Taxonomic Collections: Shelled Molluscs 

Nora Foster, Aquatic Collection, University of Alaska Museum 

Methods 

Molluscs were collected and identified by N. Foster at sites in Prince William Sound 

during June 20-28, 1998 and August 8-14, 1999. In 1999 the field survey was expanded to 

include sites in Homer, Seward, and Hinchinbrook Entrance (see Fig. 9A.2 above). Also, J. 

Goddard and L. Schickel participated in the second year’s field collecting and identification, 

providing expertise in opisthobranch taxonomy (see Chapt 9C8. Opisthobranch Molluscs). 

Site information. Site information was collected by J. Chapman & T. Miller (see Chapt 9B). 

Sampling and processing. Presence and relative abundance (recorded as abundant, common, or 

rare) of the easily identified and abundant species were recorded in the field. Field notes were 

then transferred to spreadsheets maintained by J. Chapman. Voucher specimens of small or rare, 

mollusks were collected and or were preserved in 10% formalin. Identifications were made by 

N. Foster at the University of Alaska Museum Aquatic Collection. 

Identifications and distributions. No comprehensive reference is available for Alaskan marine 

mollusca. Major references used for both identifications and distribution records include Foster 

(1991) , Coan and Scott (unpublished draft ), Baxter (1987), Turgeon (1998), Kozloff (1996), 

and Harbo (1997). Voucher collections form this study will be accessioned into the UAM 

Aquatic Collection as accession 1998-003 and 1999-001. 

Results 

Seventy-nine mollusc species were collected or identified from 27 sites in 1998 (Table 

9C7.1). Sixty- six three species were collected or identified from 16 sites in 1999 (Table 9C7.2). 

Mollusks were collected from three general types of habitats: 

1. Human-made structures, including oyster lantern nets, and associated floats and buoys, and 

docks. Characteristic mollusks from fouling communities are Mytilus trossulus, Lacuna 

vincta, Hiatella arctica. This represented the richest habitat for nudibranchs, including 12 

new records. Small mytilids were examined, to look for Musculista stenhousia. 

2. Mudflats. Upper intertidal areas of mudflats, near Cordova and Valdez and Homer were 

dominated by Mya arenaria. A search was made for Nuttalia obscurata, Batillaria cumingi, 

Ilyanassa obsoleta, and Nassa fratercula, but these potenial NIS were not found at any site of 

our surveys. 

3. Rocky intertidal zones of beaches, sheltered bays with eelgrass. Fauna of rocky intertidal 

areas, both in bays and headlands is fairly well documented. It is the habitat that is richest in 

species, but has fewest NIS candidates or new collecting records. 

In combination for the two years, 115 species of molluscs were sampled (Table 9C7.3). 

Of these species, the soft-shelled clam Mya arenaria is an NIS (Strasser 1999) that was widely 

distributed as a self-sustaining population in intertidal soft sediments of protected bays 

throughout Prince William Sound. Recently, Foster examined archaeological evidence for


Table 9C7.1. Shelled 

Molluscs 1998 Survey 

Harbors 

Valdez 

Cordova 

Whittier 

Mud bays 

Valdez 

Bligh Island 

Valdez Arm 

Growler Island 

Cordova 

Headlands & 

Reefs 

Valdez Arm 

Busby Island 

Green Island 

Rocky 

Bays 

Gastropoda 

x 

Acmaea mitra x x x 

Acteocina harpa 

x 

Aglaja ocelligera 

Alia gauspata x x x x 

Alvania sp. x x x x x 

Archidoris montereyensis x x x x 

Barleeia sp. 

x 

Buccinum baeri x x x 

Cerithiidae 

x 

Cerithiopsis sp. x x 

Crepidula dorsata 

x 

Crepidula perforans 

x 

Crepidula sp. 

x 

Cryptobranchia concentrica 

x 

Cryptonatica affinis x x 

Dendronotus 

Northwest Bay 

Fouling 

Valdez 

Cordova 

Whittier 

Bligh Island 

Eagilik Bay 

Evans Island 

Knight Island 

Lake Bay 

Main bay 

Sheep Bay 

Shotgun Cove 

Squaw Bay 

Port Valdez 

Valdez Arm 

Windy Bay 

Dendronotus albopunctatus 

x 

Dendronotus frondosus x x 

Diplodonta impolita 

x 

Dorid nudibranch x x x 

Doridella steinbergi x x x 

Eubranchus olivaceous x x 

Fusitriton oregonensis 

x 

Granulina margaritula 

x 

Haminoea virescens x x x 

Hermissenda crassicornis x x x x x 

Lacuna sp. x x x x x x x 

Lacuna marmorata x x x x x 

Lacuna vincta x x x x x x x 

Lepeta caeca 

x


Table 9C7.1. 

(Continued) Shelled 


Harbors 

Valdez 

Cordova 

Whittier 

Mud bays 

Valdez 

Bligh Island 

Valdez Arm 


Cordova 

Headlands & 

Reefs 

Valdez Arm 

Busby Island 

Green Island 

Rocky 

Bays 

Northwest Bay 

Valdez 

Cordova 

Whittier 

Bligh Island 

Eagilik Bay 

Evans Island 

Knight Island 

Lake Bay 

Main bay 

Sheep Bay 

Shotgun Cove 

Squaw Bay 

Port Valdez 

Valdez Arm 

Lepetidae 

x 

Lirularia lirulata 

x 

Littorina scutulata x x x x x x x 

Littorina sitkana x x x x x x x x x x x 

Lottia pelta x x x x x x x x x x x x x 

Lottia sp. x x x x x x x x x x 

Margarites berhigensis x x 

Margarites helicinus x x 

Margarites pupilliis x x x x x x x x 

Margarites sp. 

Nassarius mendicus x x 

Neptunea lyrata 

x 

Nucella lamellosa x x x x x 

Nucella lima x x x 

Ocenebra interfossa 

x 

Odostomia spp. 

x 

Olivella baetica x x x 

Onepota spp. x x 

Scabrotrophon maltzani 

x 

Searlesia dira x x x 

Tectura fenestrata x x 

Tectura persona x x x x x x x x 

Tectura scutum x x x x x x x 

Trichotropis cancellata x x x 

Trichotropis insignis x x 

unidentified Turridae 

x 

Velutina rubra 

x 

Bivalvia 

Axinopsida sp. 

x 

Bankia setacea x x 

Chlamys rubida x x x x 

Clinocardium nuttalli x x x x x x 

Crassostrea gigas 

x 

Fouling 

Windy Bay


Table 9C7.1. 



Harbors 

Valdez 

Cordova 

Whittier 

Mud bays 

Valdez 

Bligh Island 

Valdez Arm 

Growler 

Island 

Cordova 

Headlands & 

Reefs 

Valdez Arm 

Busby Island 

Green Island 

Rocky 

Bays 

Northwest 

Bay 

Fouling 

Valdez 

Cordova 

Whittier 

Bligh Island 

Eagilik Bay 

Evans Island 

Knight Island 

Lake Bay 

Main bay 

Sheep Bay 

Shotgun Cove 

Squaw Bay 

Port Valdez 

Valdez Arm 

Gari californica 

x 

Hiatella arctica x x x x x x x x x x x x x x x x x x x 

Macoma balthica x x x x x x x x 

Macoma inquinata x x x x x x x 

Macoma nasuta x x 

Macoma sp. x x 

Modiolus modiolus x x 

Montacutidae x x 

Musculus vernicosus x x x x x 

Mya arenaria x x x x 

Mya pseudoarenaria 

x 

Mya sp. x x x x 

Mya truncata x x x x 

Mysella tumida x x x 

Mytilus trossulus x x x x x x x x x x x x x x x x x x x x x 

Pododesmus macroschisma x x x x x x x 

Protothaca staminea x x x x x x x 

Pseudopythinia compressa x x 

Saxidomus giganteus x x x x x x x x x x 

Serripes groenlandicus x x x x 

Serripes laperousii 

x 

Tresus capax x x 

Turtonia minuta x x 

Yoldia hyperborea 

x 

Polyplacophora 

Cryptochiton stelleri 

x 

Katherina tunicata 

x 

Lepidozona interstinctus 

x 

Mopalia ciliata x x x 

Mopalia lignosa x x x x x 

Mopalia sp. x x 

Schizoplax brandtii x x 

Tonicella insignis x x 

Tonicella lineata x x x 

Windy Bay


Table 9C7.2. Shelled 


Homer 

Seward 

Lowell Pt 

Whittier 

Shot-gun 

Cove 

Fairmont 

Bay 

Valdez 

Cloudman 

Bay 

Busby Is. 

Cordov 

a 

Windy 

Bay 

Constantine 

Harbor 

Tatitlek 

a = abundant 

r = rare 

c = common 

p = present 

floats 

benthic 

homer 

spit 

benthic 

floats 

float 

fouling 

benthic 

grab 

benthic 

benthic 

floats 

floats 

benthic 

Gastropoda 

Alia gauspata 

r 

Archidoris montereyensis 

r 

Boreotrophon 

r 

Buccinum baeri 

c 

Cerithiidae 

r 

Collisella digitalis 

c 

Collisellla strigatellla c c 

Crepipatella dorsata 

r 

Cryptobranchia alba r r r r 

Cryptonatica offinis r r 

dorid nudibranch c c 


Haminoea vesicula c r r c 

Hermissenda crassicornis 

c 

Lacuna vincta c c c c c c 

Littorina sitkana a c c a c r c 

Llttoria scutulata c c 

Lottia pelta a c c c c 

Lottia strigatella 

r 

Margarites pupillis r r 


c 

Nassarius mendicus 

r 


p 

Nucella lamellosa c r 

Nucella lima c c r 

Ocinebrina interfossa 

r 

Odostomia sp. r r r 

Oenopota sp 

r 

Olivella baetica r r 

Onchidoris bilammelata 

c 

Polycera zosterae 

c 

Tectura fenestrata 

c 

Tectura persona c c 

Tectura scutum c c c 

Trichotropis cancellata c r 

Velutina plicatilis 

r 

Polyplacophora 

Mopalia cf M. imporcata r 

Mopallia cf. M.spectabilis 

r 

Moplalia ciliata r r 

Schizoplax brandtii 

r 

Tonicella lineata r r 

Tonicella rubra 

r 

Bivalvia 

Bankia setacea 

p 

Chlamys rubida r r 

Clinocardium californiense r 

Clinocardium nuttalli c c c c 

Clinocardium sp. c p 

Crassicardia crassidens 

r 

Cryptomya californica 

r 

r


Table 9C7.2. 



a = abundant 

r = rare 

c = common 

p = present 

Homer Valdez Cordov 

a 

floats 

benthic 

homer 

spit 

Seward 

Lowell Pt 

benthic 

Whittier 

floats 

Shot-gun 

Cove 

float 

Fairmont 

Bay 

fouling 

benthic 

grab 

Cloudman 

Bay 

benthic 

Busby Is. 

benthic 

floats 

Windy 

Bay 

Constantine 

Harbor 

floats 

benthic 

Tatitlek 

Cyclocardia sp. r r 

Hiatella acrtica c c c c c p c c c 

Humilaria kennerleyi 

r 

Macoma balthica a c a c a c 

Macoma calcarea 

p 

Macoma inquinata c c a c c c 

Macoma lama 

r 

Macoma obliqua r r 

Macoma sp. p p 

Modiolus modiolus r r 

Mya arenaria c a c 

Mya pseudoarenaria c c 

Mya sp. 

p 

Mya truncata c c r 

Mytilus trossulus a a a a c a a a a a a 

Neaeromya compressa 

r 

Pododesmus macroschisma 

r 

Protothaca staminea c c p c 

Saxidomus giganteus c c p c 

Serripes groenlandicus p r 


r 

Siliqua patula p p 

Spisula polynyma c c c 

Tresus capax c c 

Vilasina vernicosa c c c 

Yoldia myalis 

p


Table 9C7.3. All Mollusks from Both 1998 and 1999 

Species Distribution NIS status 

Acmaea mitra 

NE Pac 

Acteocina harpa 

NE Pac 

Aglaja ocelligera NE Pac new record 

Astyris gauspata 

NE Pac 

Alvania sp. 

Archidoris montereyensis NE Pac 

Axinopsida sp. 

Bankia setacea 

NE Pac 

Barleeia sp. 

Boreotrophon 

Buccinum baeri 

NE Pac 

Cerithiidae 

Cerithiopsis 

Chlamys rubida 

NE Pac 

Clinocardium californiense NE Pac 

Clinocardium nuttallii 

NEW Pac 

Clinocardium sp. 

Lottia digitalis 

NE Pac 

Lottia strigatellla 

NE Pac 

Cyclocardia crassidens NEW Pac 

Crassostrea gigas 

introduced 

Crepidula sp. 

Crepipatella dorsata 

NE Pac 

Cryptobranchia alba 

NE Pac 

Cryptobranchia concentrica NEW Pac 

Cryptomya californica 

NEW Pac 

Cryptonatica affinis 

arctic circumboreal 

Cyclocardia sp. 

Dendronotus frondosus northern hemisphere 

Dendronotus sp. 

Diplodonta impolita 

NE Pac 

dorid nudibranch 

Acanthodoris 

Doridella steinbergi 

NE Pac 

Eubranchus olivaceous NE Pac new record 

Fusitriton oregonensis 

NE Pac 

Gari californica 

NE Pac 


NE Pac 

Haminoea vesicula 

NE Pac 

Haminoea virescens 

NE Pac 

Hermissenda crassicornis NE Pac 

Hiatella arctica 

northern hemisphere 

Humilaria kennerleyi 

NE Pac 

Lacuna marmorata 

NE Pac 

Lacuna sp. 

Lacuna vincta 

amphiboreal 

Lepidozona interstinica 

NE Pac 

Lepetidae 

Lirularia lirulata 

NE Pac


Table 9C7.3. (Continued) All Mollusks from Both 1998 and 1999 


Littorina scutulata 

NE Pac 

Littorina sitkana 

NE Pac 

Lottia pelta 

NE Pac 

Lottia sp 

Lottia strigatella 

NE Pac 

Macoma balthica circumboreal poss. cryptogenic 

Macoma calcarea 

ampiboreal 

Macoma inquinata 

NE Pac 

Macoma lama 

NEW pac 

Macoma nasuta 

NE Pac 

Macoma obliqua 

NE Pac 

Macoma sp. 

Margarites beringensis 

AK 

Margarites helicinus 

circumboreal 

Margarites pupillus 

NE Pac 

Margarites sp. 

Melebe leonina 

NE Pac 

Modiolus modiolus 

circumboreal 

Montacutidae 

Mopalia cf M. imporcata NE Pac 

Mopalia ciliata 

NE Pac 

Mopalia lignosa 

NE Pac 

Moplalia cf. M.spectabilis NE Pac 

Mya arenaria amphiboreal introduced 

Mya pseudoarenaria 

NE Pac 

Mya sp. 

Mya truncata 

amphiboreal 

Mysella tumida 

NE Pac 

Mytilus trossulus 

NE Pac 

Nassarius mendicus 

NE Pac 

Neaeromya compressa NE Pac 


NE Pac 

Nucella lamellosa 

NE Pac 

Nucella lima 

NE Pac 

Ocinebrina interfossa 

NE Pac 

Odostomia spp. 

Oenopota sp. 

Olivella baetica 

NE Pac 

Onchidoris bilamellata 

amphiboreal 

Onchidoris sp. 

Pododesmus macroschisma NE Pac 

Polycera zosterae 

NE Pac 

Protothaca staminea 

NE Pac 

Saxidomus giganteus 

NE Pac 

Scabrotrophon maltzani NE Pac 

Schizoplax brandtii 

NEW Pac 

Searlesia dira 

NE Pac 

Serripes groenlandicus circumboreal 


circumboreal 

Siliqua patula 

NE Pac


Table 9C7.3. (Continued) All Mollusks from Both 1998 and 1999 


Spisula polynyma 

amphiboreal 

Tectura fenestrata 

NE Pac 

Tectura persona 

NE Pac 

Tectura scutum 

NE Pac 

Tonicella insignis 

AK 

Tonicella lineata 

NE Pac 

Tonicella rubra 

circumboreal 

Tresus capax 

NE Pac 

Trichotropis cancellata 

NE Pac 

Trichotropis insignis 

NEW Pac 

Turtonia minuta 

amphiboreal 

Velutina plicatilis 

arctic circumboreal 

Velutina rubra 

AK 

Vilasina vernicosa 

NEW Pac 

Yoldia hyperborea 

circumboreal 

Yoldia myalis 

amphiboreal


occurrence of Mya arenaria in Alaskan waters before the 1890s. No M. arenaria were found in 

samples from shell middens from Sitka (1070-470 yr BP) (Foster, unpublished data) and S. 

Hawkins Island (500-200 yr BP and 2150-1380 yr BP) (Yarborough, unpublished data), and 

faunal lists from two other Prince William Sound sites (ca. 400-200 yr BP) (Yarborough, 

unpublished data), and from Resurrection Bay and Aialik Bay (1700s –1800s AD) (Yarborough, 

unpublished data). These negative findings are consistent with M. arenaria being an invasive 

species that was introduced in the late 1880s. In addition, the Asian oyster Crassostrea gigas 

was also widely distributed in protected bays of Prince William Sound and Katchemak Bay, 

where it is cultured abundantly in nets suspended vertically in the water column. However, it is 

sustained as an aquaculture species by importation of laboratory produced spat, and is not a selfsustaining 

population, probably because water temperatures are too cold for gonad development 

and spawning. The common clam Macoma balthica may be viewed as a cryptogenic species that 

may be introduced from the north Atlantic; but because of its circumboreal distribution and 

variation in shell morphology, it is difficult to reconstruct the history of this species’ 

biogeography. 

References: 

Baxter, R. 1987. Mollusks of Alaska. Shells and Sea Life. Bayside, California, 163pp. 

Behrens, D.W. 1991. Pacific coast nudibranchs. Second edition. Monterey California. 207 pp. 

Coan, E.V. and P.V. Scott. 1999. Personal Communication. (unpublished draft of guide to 

Pacific coast bivalves.) 

Feder, H.M. and G.E.M. Matheke. 1980. Distribution, abundance, community structure and 

tropic structure of the infauna of the northeast Gulf of Alaska. Inst. Mar. Sci. Rept. R78-8, Univ. 

of Alaska, Fairbanks. 209 pp. 

Fisher, W.K. 1930. Asteroidea of the North Pacific and adjacent waters. Part 3. Forcipulata 

(concluded). Smithsonian Institution. U. S. Nat. Mus. Bull 76. 245 pp. 

Foster, N.R. 1981. A synopsis of the marine prosobranch and bivalve mollusks in Alaskan 

waters. Univ. Alaska, Institute of Marine Science Technical Rept IMS 81-3, 479 p. 

Foster, N. R. 1991. Intertidal Bivalves: A Guide to the Common Marine Bivalves of Alaska. 

University of Alaska Press, Fairbanks, Alaska. 152 pp. 

Harbo, R. M. 1997. Shells and Shellfish of the Pacific Northwest A field guide. Harbour 

Publishing, Madiera Park, BC, Canada. 270 pp. 

Kozloff, E. N. 1996. Marine Invertebrates of the Pacific Northwest. University of Washington 

Press. 539 pp. 

Strasser, M. 1999. Mya arenaria- an ancient invader of the North Sea Coast. Helgolander 

Meeresuntersuchungen 52: 309-324.


Turgeon , D.A. (ed.) 1998. Common and Scientific Names of Aquatic Invertebrates from the 

United States and Canada: Mollusks, Second Edition. American Fisheries Society Special 

Publication 26. 526 pp.

Chapt 9C8. Opisthobranch Molluscs, page 9C8- 1 

Chapter 9C8. Focal Taxonomic Collections: Opisthobranch Molluscs 

Jeffrey H. R. Goddard, Marine Science Institute, University of California, Santa Barbara 

Introduction 

The Smithsonian Environmental Research Center, in conjunction with the University of 

Alaska, U.S. Fish and Wildlife Service and Prince William Sound Advisory Council, conducted 

in August 1999 a survey of Prince William Sound for non-indigenous marine species. Owing to 

the abundance of opisthobranch molluscs observed in a similar survey the previous summer, I 

was invited to help identify members of this taxon in the 1999 survey. The following report 

summarizes the 1999 collection of opisthobranch molluscs and attempts to identify nonindigenous 

and cyptogenic members of the fauna. 

Methods 

Specimens were collected by hand from floating docks, intertidal mudflats and rocky 

shores, jetties, and from settling panels deployed earlier in the year. Observations were made of 

living animals, either with a dissecting microscope in the laboratory, or with a hand lens while 

traveling to the next collecting station. Voucher specimens were fixed in either 5 to 10 % 

formalin or 70 % ethanol. Specimens not identified during the expedition were examined, 

dissected and identified in the laboratory at UCSB. 

Results 

Twenty-eight species were found, consisting primarily of dorid nudibranchs. These are 

listed below with notes on their classification, habitats, and prey; an asterisk marks range 

extensions. Distributions and abundance in our field collections are summarized in Table 9C8.1 

following the list. 

Cephalaspidea 

Aglaja ocelligera (Bergh, 1894) 

Numerous specimens were found on the intertidal mudflats west of the Cordova marina. 

Haminoea sp. (either vesicula or virescens) 

Egg masses of this species were abundant on Zostera in the Cordova marina, where Nora Foster 

collected a single specimen. 

Melanochlamys diomedea (Bergh, 1894) 

Adults and egg masses of this species were abundant on mudflats just west of the Cordova 

marina. 

Sacoglossa 

*Alderia modesta (Loven, 1844) 

Adults and their egg masses were abundant on Vaucheria sp. on the high intertidal mudflats 

immediately west of the Cordova marina. Range extension from Vancouver Island, British 

Columbia (Millen, 1980).


*Olea hansineensis Agersborg, 1923 

One specimen of this opisthobranch egg eating sacoglossan was found on the egg masses of 

Melanochlamys diomedea on the mudflat west of the Cordova marina. Range extension from 

Sechelt Inlet, British Columbia (Millen, 1980). 

Nudibranchia, Doridacea 

Acanthodoris nanaimoensis O’Donoghue, 1921 

A specimen about 35 mm long was found in the low intertidal on the jetty at the Cordova marina. 

This species was recently collected from Little Green Island, Prince William Sound (Nora 

Foster, personal communication, 1999). 

Acanthodoris pilosa (Abildgaard, 1789) 

Three specimens, 2.2 to 5 mm long, were found on drift pieces of a large, unidentified, 

fleshy, orange-brown colored ctenostome bryozoan on the mudflats just west of the Cordova 

marina. Note: unidentified species of cheilostome bryozoans were growing on the above 

ctenostome and may have been the prey of these specimens of A. pilosa. Two additional 

specimens, 5.5 and 7 mm long, were found on settling panels in the Cordova marina. 

*Adalaria jannae Millen, 1987 

This species was abundant, along with its ribbon shaped egg masses, on Membranipora sp. 

growing on Laminaria on the docks at Whittier and on the moored buoy at Shotgun Cove. A. 

jannae closely resembles Onchidoris muricata, but lacks the medial radular teeth found in the 

latter; A. jannae also has 4 to 6 small lateral teeth on each half row of the radula, as well as a 

ribbon shaped egg masses (Millen, 1987). The radular formula from an 8 mm long (preserved) 

specimen from Shotgun Cove was 30 (4.1.0.1.4.). Range extension from Sointula, British 

Columbia (Millen, 1987). 

Adalaria proxima (Alder & Hancock, 1854) 

Two specimens, 3 mm long (alive) and 10 mm long (preserved) were found on the unidentified 

ctenostome bryozoan mentioned above for Acanthodoris pilosa and on the jetty, respectively, at 

Cordova (the above note for the prey of Acanthodoris pilosa also applies to this species). The 

radular formula for these two specimens was 30 (7.1.1.1.7) and 36 (9.1.1.1.9), respectively, and 

the changes in shape of the teeth with increasing body size match that described for this species 

by Thompson & Brown (1984). Note: previous Alaskan records of this species can be found 

under the names Adalaria albopapillosa, A. pacifica, and A. virescens (see Lee & Foster, 

1985:444), all three of which Millen (1987) considered junior synonyms of A. proxima. 

*Adalaria sp. 1 of Behrens (1991) and Millen (1987:2701) 

One specimen, 3.3 mm long (preserved) of this distinctive species was found on the low 

intertidal rocky shore at Tatitlek. Range extension from Ketchikan, Alaska (Millen, 1989). 

Adalaria sp. 

Two specimens, 2 and 3 mm long, were found on the same fleshy ctenostome bryozoan that 

Acanthodoris pilosa was found at Cordova (and the same note on prey also applies here). An


additional specimen, 3 mm long, was found on a settlement panel at Valdez. The radular 

formula (24 x 4.1.0.1.4 in a 3 mm specimen), tooth shape, and bipinnate gills inserted in separate 

pits place this onchidoridid in the genus Adalaria, but the presence of long slender dorsal 

papillae lacking spicules does not appear to match any described species. 

*Ancula pacifica MacFarland, 1905 

A single specimen, lacking orange lines on the body, was found on the floating docks in the 

Cordova marina. This species (or the color form of A. pacifica lacking orange lines on the body) 

may be a junior synonym of Ancula gibbosa (Risso, 1818). Range extension from Grant Island, 

Ketchikan, Alaska (Millen, 1989). 

Archidoris montereyensis (Cooper, 1863) 

Two specimens were found eating the sponge Halichondria panicea growing on oysters in 

Fairmont Bay, and two specimens were found in the marina at Cordova. 

Doridella steinbergae (Lance, 1962) 

This species was found on its prey, Membranipora sp., on Laminaria on the floating docks at 

Whittier, and on drift Laminaria on the mudflats at Cordova. The range of this species was 

extended by Foster (1987) northward to Prince William Sound and more recently westward to 

Mink Island, Katmai National Park (Nora Foster, personal communication, 1999). 

*Geitodoris heathi (MacFarland, 1905) 

Four specimens were found on the low intertidal rocky shore at Tatitlek. Range extension from 

Ketchikan, Alaska (Millen, 1989). 

Onchidoris bilamellata (Linnaeus, 1767) 

This circumboreal species feeds on Balanus spp. and was abundant, along with its egg masses, in 

the Homer marina, on the settling panels at Fairmont Bay and Tatitlek, and on rocks in and 

around the Cordova marina. 

Onchidoris muricata (Müller, 1776) 

Atotal of five small specimens were found, two on settling panels at Valdez, and three on settling 

panels at Cordova. Note: previous Alaskan records of this species can be found under the names 

O. hystricina and O. varians (see Lee & Foster, 1985:444), which Millen (1985) synonomized 

with O. muricata. 

*Palio zosterae (O’Donoghue, 1924) 

Adults and egg masses were abundant on the bryozoan Membranipora sp. growing on Laminaria 

on the floating docks at Whittier; a few specimens were also found on the buoy at Shotgun Cove 

and on the docks at Cordova. P. zosterae may be a junior synonym of P. pallida Bergh, 1880; if 

not these specimens represent a small westward range extension from Hawkins Island, Prince 

William Sound (Nora Foster, personal communication, 1999). 

Triopha catalinae (Cooper, 1863) 

Three specimens were found under cobbles in the low intertidal at Tatitlek.


Nudibranchia, Dendronotacea 

Dendronotus frondosus (Ascanius, 1774) 

Specimens were found on Obelia-like hydroids on settling panels at Seward, Fairmont Bay, 

Potato Point in Valdez narrows, and Tatitlek. Adults and egg masses were also common on the 

docks at Cordova. 

Melibe leonina (Gould, 1852) 

Specimens were found on Zostera in Fairmont Bay, on a settling panel at Tatitlek, and on the 

docks at Cordova. 

Nudibranchia, Arminacea 

Dirona albolineata MacFarland in Cockerell & Eliot, 1905 

Four small specimens were found on the floating docks in the Cordova marina, an additional 

juvenile specimen was found on a settling panel at Tatitlek. 

*Janolus fuscus (O’Donoghue, 1924) 

Two specimens, 60 to 70 mm long, were found with their egg masses on Bugula sp. on docks in 

the Homer marina. Range extension from Klu Bay, Revillagigedo Island, Alaska (Robilliard & 

Barr, 1978). 

Nudibranchia, Aeolidacea 

Aeolidia papillosa (Linnaeus, 1761) 

One 70 mm long specimen was found on the docks at Homer. 

*Cuthona albocrusta (MacFarland, 1966) 

A single specimen of this distinctive species was found on the docks at Cordova. Range 

extension from White Rock, southern British Columbia (Millen, 1983). 

*Cuthona pustulata (Alder & Hancock, 1864) 

Four specimens, 4 to 5 mm long, were found feeding on the hydroid Sarsia sp. on a dock in the 

marina at Homer. These specimens resembled Gosliner & Millen’s (1984) description of 

Cuthona pustulata from British Columbia, especially with regard to overall shape of the body, 

cerata, and head tentacles. Our specimens differed however by lacking large white spots on the 

cerata (they did have smaller opaque white flecks) and by having slightly fewer rows of cerata 

with fewer cerata per row. The radula and shape of the radular teeth of our specimens are 

virtually identical to that described by Gosliner & Millen (1984) but differed in having 4 to 5 

lateral denticles, compared to 5 to 9. Range extension from Galiano Island, British Columbia 

(Gosliner & Millen, 1984). 

*Eubranchus olivaceus (O’Donoghue, 1922) 

This distinctive aeolid was found with its egg masses on Obelia-like hydroids on the docks at 

Homer and at Whittier. Range extension from Prince William Sound (Nora Foster, personal 

communication, 1999).


Hermissenda crassicornis (Eschscholtz, 1831) 

Hermissenda was found at Whittier, Shotgun Cove, Fairmont Bay, Tatitlek, and in the Cordova 

marina. Specimens were small at all sites except Cordova, where they were up to 35 mm long. 

Discussion 

Most of the opisthobranchs collected during this survey are cold-temperate species endemic to 

the northern Pacific Ocean, especially the northeastern Pacific. The remainder are circumboreal 

species, a few of which (e.g., Dendronotus frondosus and Onchidoris bilamellata) penetrate the 

Arctic Ocean (see information on distributions in Marcus, 1961; McDonald, 1983; Thompson & 

Brown, 1984; Behrens, 1991). To my knowledge, none of the species collected in our survey are 

non-indigenous in the Prince William Sound region. While native/non-native status can not be 

assigned with certainty to the unidentified species of Adalaria or the tentatively identified 

specimens of Cuthona pustulata, the radiation of both of these genera in boreal waters suggests 

that both of these species are probably also indigenous to Prince William Sound. 

References 

Behrens, D.W. 1991. Pacific coast nudibranchs. Sea Challengers: Monterey, California. 107 pp. 

Foster, N.R. 1987. Range extension for Doridella steinbergae (Lance, 1962) to Prince William 

Sound, Alaska. The Veliger 30: 97-98. 

Gosliner, T.M. and S.V. Millen, 1984. Records of Cuthona pustulata (Alder & Hancock, 1854) 

from the Canadian Pacific. The Veliger 26: 183-187. 

Lee, R.S. and N. R. Foster. 1985. A distributional list with range extensions of the opisthobranch 

gastropods of Alaska. The Veliger 27: 440-448. 

Marcus, E. 1961. Opisthobranch mollusks from California. The Veliger 3(Supplement): 1-85. 

McDonald, G.R. 1983. A review of the nudibranchs of the California coast. Malacologia 24: 

114-276. 

Millen, S.V. 1980. Range extensions, new distribution sites, and notes on the biology of 

sacoglossan opisthobranchs (Mollusca: Gastropoda) in British Columbia. Canadian Journal of 

Zoology 58: 1207-1209. 

Millen, S.V. 1983. Range extensions of opisthobranchs in the northeastern Pacific. The Veliger 

25: 383-386. 

Millen, S.V. 1985. The nudibranch genera Onchidoris and Diaphorodoris (Mollusca, 

Opisthobranchia) in the northeastern Pacific. The Veliger 28: 80-93. 

Millen, S.V. 1987. The nudibranch genus Adalaria, with a description of a new species from the 

northeastern Pacific. Canadian Journal of Zoology 65: 2696-2702.


Millen, S.V. 1989. Opisthobranch range extensions in Alaska with the first records of Cuthona 

viridis (Forbes, 1840) from the Pacific. The Veliger 32: 64-68. 

Robilliard, G.A. and L. Barr. 1978. Range extensions of some nudibranch molluscs in Alaskan 

waters. Canadian Journal of Zoology 56: 152-153. 

Thompson, T.E. and G.H. Brown. 1984. Biology of opisthobranch molluscs, Vol. II. Ray 

Society: London. 229 pp.


Table 9C8. 1. Numbers of opisthobranch molluscs collected at 8 sites on the Kenai Peninsula and in Prince William Sound, Alaska, 

August 1999. A number followed by (S) means those specimens were found on fouling plates only. 

Table 9C8.1 Opisthobranch Homer Seward Whittier Shotgun Fairmont Tatitlek Cordova 

Molluscs 1999 

Species marina mudflat marina Lowell Pt. marina Cove Bay Valdez rocky shore mudflats marina 

Aeolidia papillosa 1 

Eubranchus olivaceus >10 1 

Cuthona pustulata 4 

Janolus fuscus 2 

Onchidoris bilamellata >10 >10(S) 3(S) >10* >10 

Melanochlamys diomedea 1 >100 

Dendronotus frondosus 7(S) >10 2(S) >1(S) >10 

Palio zosterae >100 >1 >1 

Adalaria jannae >100 >10 

Hermissenda crassicornis 5 >10 >10 >1 >10 

Doridella steinbergae 2 >10* 

Archidoris montereyensis 1 2 

Melibe leonina 1 >1(S) 1 

Onchidoris muricata 2(S) 3(S) 

Adalaria sp. 1(S) 2* 

Adalaria sp. 1 of 

Behrens (1991) 1 

Geitodoris heathi 4 

Triopha catalinae 3 

Aglaja ocelligera >10 

Acanthodoris pilosa 3* 2(S) 

Adalaria proxima 1* 1 

Alderia modesta >10 

Olea hansineensis 1 

Haminoea sp. 1 

Acanthodoris nanaimoensis 1 

Dirona albolineta 1(S) 4 

Ancula pacifica 1 

Cuthona albocrusta 1 

Number of species per site 5 1 1 0 5 3 5 3 8 9 14 

* specimens found on cobbles, drift kelp, or drift bryozoans lying on the mudflat.

Chapt 9C9. Echinoderms, page 9C9- 1 

Chapter 9C9. Focal Taxonomic Collections: Echinoderms 



Results 

Echinoderm collections over the two years of field sampling included 4 species of 

holothuroid, 1 species of ophiuroid, 6 species of asteroid, and 1 species ophiurioid (Table 

9C9.1). No crinoid species were collected. All of these species are reported as native to Alaska. 

However, Asterias amurensis, a northwest Pacific seastar, was found at Homer spit of Kachemak 

Bay, off Cook Inlet. This is a range extension form the Shumagin Islands (Fisher 1930) and 

from the Kodiak Shelf and Izhut Bay, Kodiak Island, and Bering and Chukchi Seas, where it was 

found by Jewett and Feder (1981). Specimens of A. amurenis are not represented in the 

University of Alaska Museum’s collections for trawl surveys in either the Gulf of Alaska (Foster, 

1999 pers. comm.) or Cook Inlet (Feder and Paul 1981, Feder et al. 1981). The survey of Cook 

Inlet also included scuba surveys. It would have been very unusual for such collections to miss 

a large conspicuous sea star (30 cm from ray tip to ray tip). Prior experienced naturalists 

working in Kachemak Bay reported the sudden recent appearance of this species. This species is 

an invasive species in Tasmania, where it was probablyintroduced by ballast water (it has a longlived 

planktotrophic larva). In Tasmania it is having adverse impacts on native bivalves and is 

considered to be a serious pest. We consider this species to be cryptogenic in Cook Inlet, with 

characteristics that are very suspicious of an NIS. Because it is a voracious predator, it could 

have a major impact on benthic communities. 

References 

Feder, H.M. and A.J. Paul. 1981. Distribution and abundance of some epibenthic invertebrates 

of Cook Inlet, Alaska. Univ. Alaska Fairbanks, Institute of Marine Science Tech. Rept. R80-3, 

154 p. 

Feder, H.M., A.J. Paul, M. Hoberg, and S. Jewett. 1981. Distribution, abundance, community 

structure and trophic relationships of the nearshore benthos of Cook Inlet. In: Environmental 

assessment of the Alaskan Continental Shelf. Final Reports, Biological Studies 14:45-676. 

Fisher, W.K. 1930. Asteroidea of the North Pacific and Adjacent Waters. Part 3. Forcipulata 

(concluded). Smithsonian Institution. U. S. Nat. Mus. Bull 76. 245 pp. 

Jewett, S.C. and H. M Feder. 1981. Epifaunal invertebrates of the continental shelf of the Bering 

and Chukchi Seas. Pp. 1131-1153 in Hood, D.W. and J. Clader (ed.), The Eastern Bering Sea 

Shelf: Oceanography and Resources. Vol. II, U.S. Dept of Commerce. Distributed by the 

University of Washington Press, Seattle. 1339 p.

Chapt 9C9. Echinoderms, page 9C9- 2 

Table 9C9.1. Echinoderms 1998, 1999 

Homer Seward Whittier Growler Sawmill 

Island Bay 

Port 

Valdez 

Rockey 

Point 

Busby 

Island 

Tatitlek Cloudman 

Bay 

Cordova Constantine 

Harbor 

Port 

Chalmers 

Green 

Island 

Chenega Northwest 

Bay 

Main 

Bay 

Eupentacta X 

pseudoquesemita 

Cucumaria 

X 

frondosa japonica 

Psolus chitonoides X X 

Cucumaria vegae 

X 

Ophiopholis 

X X X 

aculeata 

Pycnopodia X X X X X X X X X X X X X 

helianthoides 

Evasterias 

X X X X X X 

troschelii 

Pisaster ochraceus X X X 

Asterias amurensis X 

Dermasterias 

X X X X 

imbricata 

Leptasterias sp. X X X 

Strongylocentrotus 

droebachiensis 

X X X

Chapt 9C10. Ascidians, page 9C10- 1 

Chapter 9C10. Focal Taxonomic Collections: Ascidians 

Gretchen Lambert, Friday Harbor Laboratories, University of Washington 

Results 

A total of 12 species of ascidians were collected during this expedition. The highest 

number of species and of individuals was at Homer, Tatitlek and Cordova, the three locations 

with the highest salinity. No ascidians were recorded from Seward or Whittier, or the floats at 

Valdez, where the salinity was only 10-12 parts per thousand, although fouling panels at Valdez, 

submerged at about 7m, did have some ascidians. Ascidians require at least 25-27 parts per 

thousand and prefer a salinity of 30 or higher. Three of the 12 species are colonial, the other 9 

are solitary forms. Although there were not a large number of species collected, they do 

represent a good diversity. Both orders and all three suborders of the class Ascidiacea are 

represented and include a total of 6 families. Seven of the 12 species are known to brood their 

embryos and release swimming tadpoles; all 7 of these did harbor mature embryos. At the end of 

this report is a list of the ascidian voucher specimens collected in 1998; there were no additional 

species to the 1999 collections. 

The collection includes an undescribed species of Distaplia, a colonial form in the 

suborder Aplousobranchia. It is surprising that this species is very abundant at both Homer and 

Cordova marinas, yet has gone unrecognized for nearly a century. It is possible that if Ritter 

(1901) or Huntsman (1912) collected it, they may have considered it merely a variant of 

Distaplia occidentalis. On the other hand, this species may have been much rarer during Ritter’s 

and Huntsman’s time. Its preferred habitat is apparently sheltered surfaces, shallow but never 

exposed at low tide and away from very much light. It is thus “pre-adapted” for the huge surface 

area and specialized environment provided by marina floats and the many submerged ropes up to 

3m. in length that are often suspended from floats. Before the building of large marinas, this 

particular type of habitat was not abundant. The marina environment is also somewhat unstable, 

subject to changing seasonal conditions. Many ascidians are well adapted to take advantage of 

this instability. Distaplia spp., like most aplousobranchs, are very fast growing, reach sexual 

maturity in a few weeks, reproduce and then die back to a dedifferentiated basal portion than can 

survive until environmental conditions are again suitable for rapid growth. This new species is 

somewhat similar to a cold-water form from the Kamchatka Peninsula of Russia, and an analysis 

and description of it is being prepared in collaboration with Dr. Karen Sanamyan of the 

Kamchatka Institute of Science. 

Two species of the genus Ascidia were collected and present a taxonomic problem. One 

corresponds to Ascidia adhaerens Ritter, 1901, the other to Ascidiopsis columbiana Huntsman, 

1912. (The genus Ascidiopsis is no longer valid; it was synonymized under the genus Ascidia.) 

Van Name (1945) synonymized these two NE Pacific species under Ascidia callosa, probably 

incorrectly; A. callosa was described in 1852 by Stimpson from U.S. east coast specimens from 

Massachusetts. The recent genetic analysis of many Atlantic and Pacific species of various taxa 

that closely resemble each morphologically and were originally lumped into the same species has 

resulted in many cases in their separation into different species (R. Strathmann pers. comm.). 

However, a recent re-examination of Stimpson’s syntypes of Ascidia callosa and Ritter’s 

paratypes of Ascidia adhaerens, borrowed from the Smithsonian National Museum of Natural


History, has confirmed that these are the same species. Thus, Ascidia adhaerens remains a 

synonym under A. callosa. The type specimen of Huntsman’s Ascidiopsis columbiana was 

borrowed from the Royal Ontario Museum in Toronto. I was able to confirm that this species is 

indeed distinct from Ascidia callosa and thus should be resurrected as a valid species, which will 

be done in a publication now in preparation. 

Botrylloides violaceus is not native to Alaska, or to any part of the NE Pacific. It is a 

Japanese species that appeared on the U.S. Pacific coast about 20 years ago (J. Carlton pers. 

comm.). Since that time it has spread and become extremely abundant from southern California 

to British Columbia (Lambert and Lambert, 1998). This is the first report of its presence in any 

part of Alaska, however. Although large colonies were not observed, the fouling panels at 

Tatitlek contained numerous newly settled zooids most of which appeared healthy. Thus 

somewhere close to the panels there had to be mature colonies which were supplying the shortlived 

tadpoles that settled on the panels. B. violaceus incubates its embryos; the tadpoles that are 

released are huge, complex, and usually swim for just a very brief period, perhaps only a few 

minutes, before settling. The unusual tadpole morphology allows for easy recognition of this 

invasive species (Saito et al., 1981). 

Molgula retortiformis and Ascidia callosa are apparently the only species collected on 

this expedition that occur in both the North Pacific and North Atlantic (Van Name, 1945). 

Chelyosoma productum, Corella inflata and C. willmeriana, Distaplia occidentalis, Pyura 

haustor and Styela truncata have been recorded only from the NE Pacific. Halocynthia 

hilgendorfi is known from both the NW and NE Pacific. 

Sixty-five species of ascidians are known to occur in Alaskan waters. Most of these were 

obtained by dredging many decades ago and may be restricted to the Bering Sea or the Arctic 

Ocean. The ascidian fauna of Alaska is still mostly unexplored, and probably there exist a 

number of undescribed species. 

Publication of the present work, including a description of the new Distaplia species and 

redescription of the two Ascidia species, is in preparation with Dr. Karen Sanamyan of the 

Kamchatka Institute of Ecology. 

References 

Huntsman, A.G. 1912. Ascidians from the coasts of Canada. Trans. Canad. Inst. 9: 111-148. 

Kott, P. 1985. The Australian Ascidiacea. Mem. Qd. Mus. 23: 1-440. 

Lambert, C.C. & G. Lambert 1998. Non-indigenous ascidians in southern California harbors and 

marinas. Mar. Biol. 130: 675-688. 

Lambert, G., C.C. Lambert & D.P. Abbott 1981. Corella species in the American Pacific 

Northwest: distinction of C. inflata Huntsman, 1912 from C. willmeriana Herdman, 1898 

(Ascidiacea, Phlebobranchia). Can. J. Zool. 59: 1493-1504. 

Nishikawa, T. 1991. The ascidians of the Japan Sea. II. Publ. Seto Mar. Biol. Lab. 35: 25-170.


O’Clair, R.M. & C.E. O’Clair 1998. Southeast Alaska’s Rocky Shores. . Plant Press, Auke Bay, 

AK, 564 pp. 

Ritter, W.E. 1901. Papers from the Harriman Alaska Expedition. XXIII. The ascidians. Proc. 

Wash. Acad. Sci. 3: 225-266. 

Ritter, W.E. 1913. The simple ascidians from the northeastern Pacific in the collection of the 

United States National Museum. Proc. U.S. Natl. Mus. 45: 427-505. 

Saito, Y., H. Mukai & H. Watanabe 1981. Studies on Japanese compound styelid ascidians II. A 

new species of the genus Botrylloides and redescription of B. violaceus Oka. Publ. Seto Mar. 

Biol. Lab. 26: 357-368. 

Sanamyan, K. 1993. Ascidians from the north-western Pacific region. 2. Molgulidae. Ophelia 

38: 127-135. 

Sanamyan, K. 1996. Ascidians from the north-western Pacific region. 3. Pyuridae. Ophelia 45: 

199-210. 

Van Name, W.G. 1945. The North and South American Ascidians. Bull. Amer. Mus. Nat. Hist. 

84: 1-476.


Table 9C10.1 Sites visited: 

1. Aug. 8 Homer Marina, Kachemak Bay, Cook Inlet 

Salinity 27 % o 1 ft. depth, 30 % o at 2m depth; Temp. 10 o C 

Most of the following records are from suspended ropes. 

Ascidia callosa--common esp. about halfway between the inner and outer ends of the marina. 

Some contain brooded larvae in atrial chamber. Two small specimens from fouling panel: 

one from E1 and one from OH--2-P2. 

Corella inflata--reported by John Chapman but not personally seen in spite of extensive 

sampling of floats and ropes. Samples lost; record not verified. 

Distaplia new sp. very abundant at most locations in the marina, esp. on ropes 1-2 m. deep. A 

few small colonies on fouling panel. 

Molgula retortiformis: 2 large and 8 small specimens on ropes. Two small specimens on fouling 

plate E1. The large animals contain brooded embryos in the atrial chamber. 

Styela truncata - 1 small individual. 

2. Aug. 9: Seward Marina. 

Salinity 11 % o about a foot down. 

Temperature 11.5 o C. 

No ascidians either at the marina or Lowell Pt. rocky intertidal (11 % o , 11.5 o C.) 

3. Aug. 10, 1999 Whittier marina. 

No sal. or temp. readings taken. Sal. very low; no ascidians on floats. 

4. Aug. 10, 1999 Fairmont Bay oyster farm, Prince Wm. Sound. 


Temperature 14 o C. 

Ascidia columbiana--numerous small specimens on concrete blocks of the fouling panels and 

one large one on bottom of submerged 4 ft long oyster bag. 

5. Aug. 11, 1999 Valdez Marina 

No salinity or temp. readings taken 

No ascidians observed on marina floats or suspended ropes. 

6. Aug. 11-12, 1999 Tatitlek 


Temperature 17 o C. surface 

a) Fouling panels and frames, 3m depth: 

Ascidia columbiana -- a few small specimens on the concrete blocks for the fouling panels and 

one on panel PL3-l. 

Botrylloides violaceus -- numerous very small zooids on panels. Largest one with 2 zooids in 

colony. Youngest appeared to be only hours post metamorphosis. 

Corella inflata -- common on both the panels and frames about 3m depth, with brooded larvae in 

the atrial chamber.


Table 9C10.1 (Continued) Sites visited: 

b) Intertidal rocks, low tide, morning Aug. 12 

Ascidia columbiana -- one large specimen coll. by J. Goddard. 

Chelyosoma productum -- about 12 seen, 2 collected. 

Halocynthia igaboja -- one very small immature specimen coll. by J. Goddard. 

Pyura haustor -- 4 seen, one incomplete specimen collected. Difficult to collect; wedged tightly 

into rock crevices. 

c) oyster cages from across bay: one small Corella inflata. 

7. Aug. 13, 1999 Cordova Marina. Tide just beginning to come in. 


Temperature 13 o C. surface 

Ascidia callosa-- numerous on floats and especially on suspended ropes. Some contain brooded 

larvae in atrial chamber. 

Corella inflata -- large, common, full of brooded larvae in atrial chamber. 

Distaplia new sp. --abundant colonies, with many mature larvae. 

Distaplia occidentalis -- common; several color morphs, large heads, with mature larvae. 

Styela truncata -- numerous. Some contain brooded larvae in atrial chamber. 

b) Fouling panels 

Ascidia callosa -- 12 small specimens from I dock, Pl. 3, 3 in a second vial. 

Corella inflata -- one small specimen from I dock, Pl. 3; two in a second vial. 

Corella willmeriana-- (specimens identified on site, not saved apparently.) 

Distaplia new sp. -- numerous small colonies from I dock Pl. 3, one in a second vial. 

Styela truncata -- 8 on fouling panels in voucher collection in one vial, 3 in another. 

8. Aug. 14, 1999 Valdez fouling panels, Berth 5 

Corella willmeriana -- 3 individuals, 1 large and 2 small. Specimens identified on site; not saved 

apparently. 

Ascidia columbiana -- one small specimen from fouling panel (or perhaps the concrete block 

anchoring the panels) at 20 ft. (sampled 8/14).


Table 9C10.2 Ascidian vouchers from fouling panels 1998, identified Sept. 1999 

Data taken from jar labels. 

Homer 3/9/98 -- Styela truncata (1) 

Homer: 9/3/98 -- Ascidia callosa (1) 

Homer Oct. 1998 -- Distaplia new sp. collected by G. Sonnevil. Colonies without gonads or 

larvae. 

Homer 5/11/99 -- Distaplia new sp. collected by G. Sonnevil. Colonies small, immature. 

Chenega 7/9/98 -- Corella inflata (2) Port San Juan, Chenega sm. boat hbr 

Chenega 9/7/98 -- Ascidia callosa (1) 

Chenega no date -- Corella inflata (7) panel MI - 05 - (can’t read label) 

Valdez 9/8/98 -- Corella inflata (2) at Alyeska berths 

No location given -- Ascidia sp. (1) too small to ID to species. Panel WH-04-P3.


Table 9C10.3 Systematics 

Class Ascidiacea 

OrderAplousobranchia 

Family Holozoidae 

Distaplia occidentalis Bancroft, 1899 

Distaplia new species 

Order Phlebobranchia 

Family Corellidae 

Chelyosoma productum Stimpson, 1864 

Corella inflata Huntsman, 1912 

Corella willmeriana Herdman, 1898 

Family Ascidiidae 

Ascidia callosa Stimpson, 1852 

Ascidia columbiana (Huntsman, 1912) 

Order Stolidobranchia 

Family Styelidae 

Botrylloides violaceus Oka, 1927 

Styela truncata Ritter, 1901 

Family Pyuridae 

Halocynthia igaboja Oka, 1906 

Pyura haustor (Stimpson, 1864) 

Family Molgulidae 

Molgula retortiformis Verrill, 1871 

Table 9C10. 4 Ascidian References by Depth and Habitat 

Depth(m) 

Selected References 

intertidal, floats;to 10m Ritter ’01, ’13; VName 45; PWS Expedition 1999 

us. 0-60m 

Ritter’01,Huntsman’12,VName45,Abbott66,O’Clair98;PWS Exped.’99 

0-few m. Saito et al 81,Lambert & Lambert 98, PWS Exped. 1999 

intertidal - 50m Huntsman ’12,VName 45,O’Clair98; PWS Exped. ’99 

0-50m Huntsman ’12,VName 45; PWS Exped. ’99 

0-few m. Huntsman ’12,VName 45; PWS Exped. ’99 

float PWS Expedition 1999 

floats; intertidal-few m. VName 45, O’Clair 98, PWS Expedition 1999 

to 57m VName 45, Nishikawa 91,Sanamyan 96,O’Clair98, PWS Exped. ’99 

4-75m; floats Ritter 01,VName 45,Abbott66,Sanamyan 93, PWS Exped. ’99 

intertidal - 114m VName 45, Sanamyan 96,O’Clair98; PWS Exped. ’99 

ropes on floats;0-20m Ritter ’01, Van Name 45, PWS Exped. ’99


Table 9C10.5 Ascidan Species Distribution in Alaska 

Genus Species Author/date Family Distribution in Alaska 

Ascidia callosa Stimpson, 1852 Ascidiidae Homer;PWS:Fairmont Bay 

oyster 

farm,Tatitlek,Cordova,Valdez, 

Chenega 

Ascidia columbiana (Huntsman, 

1912) 

Ascidiidae Arctic coast to Alask. Penin. & 

SE Alaska 

Botrylloides violaceus Oka, 1927 Styelidae PWS: Tatitlek 

Chelyosom 

a 

productum Stimpson, 1864 Corellidae PWS: Tatitlek; Sitka 

Sound:Passage Is. 

Corella willmeriana Herdman, 1898 Corellidae PWS: Cordova, Valdez 

Corella inflata Huntsman, 1912 Corellidae PWS: Tatitlek, Cordova, 

Valdez, Chenega 

Distaplia new sp. Holozoidae Cook Inlet: Homer Marina; 

PWS: Cordova Marina 

Distaplia occidentalis Bancroft, 1899 Holozoidae PWS: Cordova Marina; 

Chichagof Is. 

Halocynthia igaboja Oka, 1906 Pyuridae Gulf of Alaska: Kodiak 

Is.;Chichagof Is.; PWS: 

Tatitlek; Prince Rupert BC 

Molgula retortiformis Verrill, 1871 Molgulidae SE Bering Sea to Sitka; Canoe 

Bay; SE Chukchi Sea; Homer 

Marina 

Pyura haustor (Stimpson, 1864) Pyuridae Alaska Gulf:Sanak Is., 

Shumagin Is.; Sitka 

Sound:Alice Is.; PWS: Tatitlek 

Styela truncata Ritter, 1901 Styelidae Yakutat Bay; Cook Inlet: 

Homer Marina; PWS: Cordova 

Marina

Chapt 9D. Fouling Community Surveys, page 9D- 1 

9D. Fouling Community Surveys 



9D1. Purpose 

Fouling communities in many bays, harbors and estuaries are frequently invaded by NIS 

associated with shipping traffic (Cohen & Carlton 1995, Hewitt 1993, Coles et al. 1999). 

Fouling communities have major impact on ships and floating structures, ranging from buoys 

and floats to aquaculture pens and nets, as well as surfaces of oysters and mussels themselves. 

Consequently, fouling communities are well-studied in many parts of the world, but they have 

received little study in Alaskan waters. We conducted surveys of fouling communities in Prince 

William Sound, Seward and Homer using experimental fouling plates. The use of fouling plates 

provides a replicated standardized assay for NIS in a community that is prone to invasions, but 

which has received little prior ecological analysis in Alaska. We also surveyed fouling 

communities on floats, pilings, and buoys to compare these substrates with our experimental 

plates. 

9D2. Methods 

We conducted two surveys that focused on fouling communities that settled on natural 

surfaces and experimental plates, which we deployed in earlier spring months at an array of sites 

in Homer, Seward and Prince William Sound. We sampled these fouling plates in 7-17 

September 1998 and in 8-16 August 1999. 

The team for the fouling community surveys consisted of: 

• Greg Ruiz (SERC), molluscs, parasites, fouling communities; 

• Anson Hines (SERC), barnacles and decapod crustaceans; 

• James Carlton (Mystic Seaport), marine/estuarine invertebrates and global NIS 

• Melissa Frey (SERC); technical and field assistance 

• George Smith (SERC); technical and field assistance 

• Lea Ann Henry (University of Ontario), hydroid identifications; 

• Judy Winston (Virginia Museum of Natural History), bryozoan identifications. 

We deployed arrays of fouling plates at 12 locations, including 2 on the Kenai Peninsula 

(Homer, Seward) and 10 in Prince William Sound (Table 9D.1). At each site we deployed 5 

arrays suspended at depths of either 1 meter (3 arrays) or 3 meters (2 arrays) below mean low 

water level. Each array consisted of settling plates attached to a frame made of 2 crossed pieces 

of pvc pipe (20 mm diameter, 50 cm long), which was suspended in a horizontal position by a 

line attached to a dock or float and weighted by a concrete block. Settling plates made of 14 cm 

x 14 cm x 7 mm thick pvc (3 plates) or plywood (1 plate) were attached to each of the 4 ends of 

the cross in a horizontal position. The horizontal orientation assured that sediment would not 

accumulate on the underside, providing a clean surface addition to several irregular surfaces of 

the frame and top surfaces for settlement. To maximize the chance of detecting possible NIS 

within each site, the 5 arrays were dispersed as widely as possible to sample the range of 

microhabitats present. We deployed the plates during April-May of each year and retrieved them 

in August-September, providing a “soak time” of about 4 months.


Table 9D.1. Sampling Locations for Fouling Community Surveys. 

Site 1998 1999 

Kenai Sites 

Homer X X 

Seward X X 

Prince William Sound 

Port Valdez 

Marine Terminal X X 

Valdez Area X X 


X 

Whittier 

X 

Chenega 

X 

Port Chalmers 

X 

Fairmont Bay 

X 

Tatitlek 

Oyster Culture 

X 

Docks 

X 

Cordova 

X 

At the time of retrieving the arrays, the fouling plates were removed from their frame and 

placed individually into ziplock plastic bags for transport to a laboratory. Each plate was 

examined under a dissecting microscope and sessile and motile species were scored as present or 

absent. Voucher specimens were preserved and sent to taxonomic experts for authoritative 

identification. At the present stage of this project, we have used the fouling plates to develop an 

inventory of species. However, using this technique, the occurrence and abundance of species 

can be quantified and compared statistically for frequency of plates with species present. This 

type of comparison will proceed in new work during 2000 and 2001 as we sample other locations 

in Alaska and the major ports of western North America along the lower 48 states. 

9D3. Results 

During the soak time of ca. 4 months, the fouling plates accumulated dense community 

assemblages with large biomasses of mussels, barnacles, ascidians, hydroids, and bryozoans 

(Table 9.D.2). The fouling communities were rich in species, particularly for bryozoans, 

hydroids, nudibranchs, and ascidians Combining the 2 years of study across the12 sampling 

locations, we recorded more than 107 taxa/species on the fouling plates (Table 9D.2), including: 

8 families of polychaete worms, 13 species of acidians, 34 species of bryozoans, 1 species of sea 

anemone, 3 species of barnacle and 13 taxa of motile crustacea, 2 species of echinoderms, 12 

species of hydroids, 29+ taxa of molluscs, and 1 species of protozoan. Diversity of hydroids was 

particularly high at Homer (11 species). Byozoans were most diverse at Chenega (10 species), 

Homer (13 species), Port Valdez (14 species), and Tatilek (13 species). Acidians were most 

diverse at Homer (6 species) and Cordova (6 species).


Table 9D.2. Taxa Recovered on Fouling Plates. Asterisk denotes NIS. x denotes presence in 1998 and or 1999. 

Site Key: AL=Alyeska terminal; CH=Chenega Bay; GR=Growler Isl.; HO=Homer, MI=Montague Isl.; 

SE=Seward;VA=Port Valdez; WH=Whittier; CORD=Cordova; FRMNT=Fairmount Bay; PTOPT=Potato Pt. 

TAT=Tatitlek 

Site 

AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT 

Protozoan Foliculina x x x x x x x x x 


Cnidaria Metridium senile x 

Metridium sp. x x x 


Hydroids Calycella syringa x 

Clytia hemispherca 

x 

Clytia kincaidi 

x 

Companulina rugosa 

x 

* Garveia franciscana x 

Gonoththyraea clarki x x x x 

Obelia longissima x x x x x x x x 

Obelia sp. x x x x x x x x 

Opercularella lacerata 

x 

Sarsia eximia 

x 

Sarsia tubulosa 

x 

Sertularia robusta 

x 


Annelida Capitellidae x x x x 

Cirratulidae x x 

Nereidae x x x x x x x x x 

Polynoidae x x x x x x x x x 

Sabellidae x x x 

Serpulidae x x x x x x x x x x 

Spirorbidae x x x x x x x x x x x 

Syllidae x x x x x x 

Terebellidae 

x 


Mollusca Acanthodoris sp. x 

Acmaeidae 

x 

Aelolididae 

x 

Alvania sp. x x x x x 

Chlamys sp. x x x 

Clinocardium sp. 

x 

Dendronotus frondosus x x 

Dendronotus sp x x 

Fusitron oregonensis 

x 

Haminoea sp. x x x 

Hermissenda sp. x x x x x x x x x x 

Hiatella arcitca x x x x x x x x x x x 

Hinnites sp. 

x 

Lacuna sp. x x x x 

Lacuna vincta 

x 

Limacina helicina 

x


Table 9D.2. Continued 


Mollusca Lottia sp. x 

cont. Margarites helinicus x 

Margarites pupillis x x 

Mytilus trossulus x x 

Mytilus sp. x x x x x x x x x x x x 

Odostomia sp. x x x x 

Onchidoris sp. x x x x 

Onchidoris murricata x x 

Onchidoris sp. x x x x x x 

Pododesma macroshisma 

x 

Pododesmus sp. x x 

Sacoglossan sp. x x 

Tritonia sp. 

x 

Vilisina vernicosa 

x 


Crustacea Ampithoe sp. x 

Balanus cariosus x x x x x x x 

Balanus crenatus x x x x x x x x x x x x 

Semibalanus carriosus x x x 

Cancer magister x x 

Caprellidae x x x x x x 

Corophium sp. x x x x 

Eogammarus sp. 

x 

Gnorimosphaeroma oregonense x x x x x 

Harpacticus uniramus 

x 

Jassa sp. x x 

Munna sp. 

x 

Oregonia gracilis 

x 

Scyra acutifems 

x 

Spirontocaris sp. 

x 


Bryozoa Alcyonidium sp. x x x x x x 

Alcyonidium hirsutum x x 

Bugula californica x x 

Bugula pacifica x x x 

Bugula sp. x x x x x x x x 

Calloporella craticula x x x 

Calloporella "lineata" 

x 

Cauloramphus pseudospinifer 

x 

Celleporella hyalina x x x x x x 

Cribrilina sp. x x 

Cribrilina corbicula x x x x x x 

Cribrilina sp. x x x x x x x 

Crisia serrulata 

x 

Cryptosula okadai 

x 

Electra sp. x x x 

Fenestruloides eopacifica x x 

Filicrisia franciscana x x x 

Harmeria scutulata 

x 

Hippothoa sp. x x x 

Lichenopora sp. 

x


Table 9D.2. continued 


Bryozoa Membranipora menbrenacea x 

cont. Membranipora sp. x x 

Microporella germana 

x 

Parasmittina trispinosa 

x 

Phidolopora pacifica x x 

Porella major x x 

Rhamphostomella gigantea 

x 

Rhamphostomella hincksi 

x 

* Schizoporella "unicornis" x 

Tegella armifera 

x 

Tegella aquilirostris 

x 

Tubulipora tuba x x x 

Tubulipora sp. 

x 


Echino- Psolus sp. x 

dermata Strongylocentrotus sp. x x 


Ascidia Ascidia sp. x 

Ascidia callosa x x x 

Ascidia columbiana x x x 

* Botrylloides violaceus x 

Corella sp. x x x 

Corella inflata x x x x 

Corella willmariana x x 

Distaplia sp. (new) x x 

Distaplia alaskensis x x 

Molgula retortiformis 

x 

Molgula sp. x x 

Ritterella sp. x x x 

Styela truncata 

x 

Among these species, there were 3 NIS: the ascidean Botrylloides violaceus, the bryozoan 

Schizoporella unicornis, and the hydroid Garvieia franciscana. These are the first records for 

two of these species (B. violaceus and G. franciscana) in Alaska. Botrylloides violaceus 

(=Botryllus aurantius is a colonial tunicate that is native to the northwest Pacific (Japan), and 

may have been first found on the West Coast in 1973, in San Francisco Bay (Cohen and Carlton 

1995). It is now widespread, from southern California to British Columbia (Cohen et al. 1998; 

Lambert and Lambert 1998). B. violaceus was abundant on fouling plates in Prince William 

Sound in 1999 (G. Lambert 1999 pers. comm.). Schizoporella unicornis is a bryozoan found in 

the northwest Pacific but was first collected in the Eastern Pacific in 1927, in Puget Sound 

(Carlton 1979; Cohen and Carlton 1995). Its first Alaska collection was made between 1944 and 

1949, in Kodiak (U. S. Navy 1951; Carlton, pers. comm.). Schizoporella unicornis may have 

been introduced in ship fouling or with plantings of Pacific Oysters (Cohen and Carlton 1995). 

In 1999, it was found in Tatilek. This form, while definitely introduced to the Pacific coast, may 

actually be a complex of several species (Winston 1999 pers. comm.). Garveia franciscana 

(Rope Grass Hydroid) has been found in many estuaries around the world, but its origin is 

uncertain. The Indo-Pacific and the Black--Caspian Sea basin have been suggested as possible 

native regions (Cohen and Carlton 1995; Calder 1997 pers. comm.) It was first described from 

San Francisco Bay in 1902, which was its only known location on the west coast of North 

America (Cohen and Carlton 1995), until we found it near Homer in 1999 (Lea-Anne Henry


pers. comm. 1999). In other regions of the world, this hydroid has been an economically 

important fouling organism, adversely affecting ships, power plants and fishing gear (Simkina 

1963; Andrews 1973; McLean 1972). In addition, fouling plates at Homer and Cordova had 

large biomasses of the new/undescribed ascidian, Distaplia sp. nov, which has many suspicious 

characteristics of NIS. Diastaplia sp. nov. is a new, undescribed species of tunicate, which is 

very abundant in fouling communities on floats and man-made substrates in marinas at Homer 

and Cordova. It was first collected in 1998 in Homer and was found in both Homer and Cordova 

in 1999. It was not found at other sites within Prince William Sound where other, native species 

of tunicate were common in fouling communities but lack similar shipping/boating traffic (e.g., 

Tatitlek, Chenega, Port Chalmers). Its appearance is also suspicious, because it was not found in 

1901 when tunicates were collected in the region at nearby sites. This tunicate could be a 

formerly rare native species that has taken advantage of the newly created marina habitat, or a 

recent introduction (G. Lambert 1999 pers. comm.). 

Fouling communities on the plates suspended at 1m depth were greatly diminished at 

Seward, Whittier and Port Valdez during the summer, when snow and glacial melt produced 

markedly low salinities in the surface waters and also thick sediment deposits that covered the 

plates. Fouling plate arrays at 3 m depth in these locations experienced higher salinities, but still 

suffered from heavy sediment deposits. It was evident, however, that these plates had developed 

rich fouling communities prior to the summer season of greatest freshwater runoff and siltation. 

Species composition of fouling communities on the experimental plates were generally 

quite similar to the composition on surrounding surfaces (floats, pilings, lines, oysters, etc) at 

comparable tidal levels. Since the arrays were suspended below mean low water, the main 

groups of specie s that the plates did not sample adequately were species on pilings in the 

intertidal zone, such as Balanus glandula, Semibalanus balanoides, and various species of 

acmaeid limpets.

Chapt 9E. Museum, Reference and Voucher Specimens, page 9E- 1 

Chapter 9E. Re-Examination of Museum, Reference, and Voucher Specimens 

Nora Foster, University of Alaska Museum, Fairbanks 

9E1. Purpose 

As a result of environmental assessment work in Port Valdez during the 1970’s for 

construction of the Valdez Marine Terminal, and the Exxon Valdez oil spill (EVOS), a large 

ecological database exists for Prince William Sound. Through a careful scrutiny of both the 

species lists and specimens archived in museum, reference and voucher collections for the past 

work, this part of our study was intended to test the idea that this extensive prior work in Prince 

William Sound should have detected NIS. However, in spite of the amount of sampling done in 

Prince William Sound, some species lists were developed without input from taxonomic experts, 

so that the biota is still poorly known. Few prior surveys have focused on biogeography or 

taxonomy, and until this study, none has specifically looked for NIS. 

9E2. Methods 

Literature sources for Prince William Sound invertebrates include: 

• Reports on marine fauna of the northeastern Gulf of Alaska, compiled as a result of the Outer 

Continental Shelf Environmental Assessment Project (OCSEAP);(Feder and Matheke, 1980; 

Feder and Jewett, 1988); 

• Thesis by M. Hoberg (1986); 

• Project reports on the Port Valdez environment (Cooney and Coyle, 1988; Feder et al., 1976; 

Feder and Bryson-Schwafel, 1988; Feder and Keiser, 1980; Jewett and Feder, 1977); 

• Eyerdam’s (1924) species lists; 

• Haven (1971); and 

• Several taxonomic references (Butler, 1980; Hart, 1982; Lambert, 1981; 1991; Behrens, 1991 

and others) for additional distribution records in Alaskan waters. 

A vast array of preserved biological samples from Prince William Sound and adjacent 

Gulf of Alaska are housed in the Aquatic Collection of the University of Alaska Museum 

(UAM). The major part of these are collections made as part of the OCSEAP projects in the mid- 

1970’s, environmental monitoring of Port Valdez, and Max Hoberg’s thesis material from three 

bays in the outer part of the Sound. Intertidal and benthic biota from the soft bottom 

communities of Prince William Sound are well represented in the collection. Species lists for 

this material were generated using the museum accession records. 

EVOS Specimen Archives 

The specimens collected as part environmental surveys after the 1989 Exxon Valdez oil 

spill supplement the UAM collection. Subtidal invertebrates in 1,154 lots have been transferred 

to the UAM from warehouse storage for examination. These represent sampling conducted 

during 1990, 1991, 1993, and 1995, from 15 selected unoiled sites. The collections provide 

samples of shallow subtidal fauna from silled fjords, eelgrass beds, and habitats dominated by 

Laminaria. With limited resources, each specimen from this large number of samples cannot be 

examined individually, so it has been necessary to select subset of specimens for careful reexamination.


The subset of reference and voucher specimens examined as part of this study was 

selected by first compiling a list of 88 invertebrate taxa by comparing the lists of known taxa 

from Prince William Sound (discussed above) with lists of known NIS from west coast source 

ports: Puget Sound, San Francisco Bay, southern California, and other localities (see Ruiz & 

Hines, 1997). This compilation of 88 species includes: 

• species that may be confused with known NIS in Alaska; 

• NIS known from western North America (especially those extending to higher latitudes); 

• taxonomically difficult species and species complexes; 

• biogeographic outliers; and 

• range extensions. 

For example, polychaete specimens of most Boccardia and Polydora, were removed for reexamination 

because for their resemblance to Polydora ligni. Similarly, capitellid and many 

syllid polychaetes were selected to check for the presence of Barontolla americana, Capitella 

capitata, and Decamastus spp. The gastropod slipper shell Crepidula was partitioned from the 

collections to check for Crepidula fornicata and C. plana. Specimens of the amphipod 

Corophuim were selected to check for several NIS Corophium reported from Puget Sound. This 

original list was considered a working document and has been refined as the project has 

progressed (Table 9.E.1). 

Table 9.E.1. Museum and Reference Samples Selected for Re-Examination 

Family Genus Species Specimen Source 

Amphitoidae Amphitoe sp. Jewett samples 

Amphitoidae Amphitoe simulans Jewett samples 

Corophidae Corophium sp. Jewett samples 

Corophidae Corophium brevis Jewett samples 

Gammaridae Melita sp. UAM uncataloged 

Gammaridae Gammarus sp. Jewett samples 

Gammaridae Jassa sp. Jewett samples 

Gammaridae Melita dentata UAM uncataloged 

Cardiidae Clinocardium fucanum Jewett samples 

Glycymeridae Glycymeris septentrionalis Jewett samples 

Kelliidae Pseudopythina* compressa Jewett samples 

Kelliidae Pseudopythina* rugifera* Jewett samples 

Mytilidae Dacrydium vitreum* Jewett samples 

Mytilidae Dacrydium pacificum* UAM cataloged 

Ungulinidae Diplodonta orbella Port Valdez uncataloged 

Ungulinidae Diplodonta impolita UAM cataloged 

Adontorhina sp. Jewett samples 

Cryptosula okaidai Jewett samples 

Atyidae Haminoea virescens Little Green Island 

Atyidae Haminoea vesicula PWS 98 samples 

Atyidae Haminoea sp. Jewett samples 

Calyptraeidae Crepidula dorsata* Jewett samples 

Calyptraeidae Crepidula sp. Jewett samples 

Calyptraeidae Crepidula grandis Jewett samples 

Calyptraeidae Crepidula sp. Jewett samples 

Diaphanidae Diaphana minuta* Jewett samples


Table 9.E.1. Continued 

Diaphanidae Diaphana brunnea Jewett samples 

Diaphanidae Diaphana sp. UAM cataloged 

Fissurellidae Puncturella cooperi* Jewett samples 

Lacunidae Lacuna sp. UAM cataloged 

Lacunidae Lacuna vincta* Jewett samples 

Lacunidae Lacuna variegata* UAM cataloged 

Lacunidae Lacuna marmorata* UAM cataloged 

Lamellariidae Velutina sp. UAM cataloged 

Rissoidae Barleeia acuta* Jewett samples 

Scaphandridae Cylichna alba UAM cataloged 

Scaphandridae Cylichna occulta UAM cataloged 

Scaphandridae Cylichna sp. UAM cataloged 

Scaphandridae Cylichna attonsa Jewett samples 

Scaphandridae Cylichnella culcitella Jewett samples 

Scaphandridae Cylichnella harpa Jewett samples 

Scaphandridae Cylichnella sp. Jewett samples 

Turridae Kurtziella plumbea Jewett samples 

Turridae Taranis strongi Jewett samples 

Limnoriidae Limnoria lignorum Jewett samples 

Limnoriidae Limnoria Jewett samples 

Capitellidae Barantolla americana Jewett samples 

Capitellidae Barantolla sp. Jewett samples 

Capitellidae Capitella sp. Jewett samples 

Capitellidae Capitella capitata Jewett samples 

Capitellidae Decamastus sp. Jewett samples 

Capitellidae Decamastus sp. Jewett samples 

Capitellidae Heteromastus sp. Jewett samples 

Capitellidae Heteromastus filliformis Jewett samples 

Capitellidae Mediomastus sp. Jewett samples 

Capitellidae Notomastus sp. Jewett samples 

Nereidae Platynereis bicanaliculata Jewett samples 

Polynoidae Harmothoe imbricata Jewett samples 

Spionidae Malacoceros sp. Jewett samples 

Spionidae Nerine cirratulus Jewett samples 

Spionidae Polydora sp. Jewett samples 

Spionidae Polydora cf P.bracycephalata Jewett samples 

Spionidae Polydora socialis Jewett samples 

Spionidae Prionospio cirrifera Jewett samples 

Spionidae Prionospio sp. Jewett samples 

Spionidae Prionospio steenstrupi Jewett samples 

Spionidae Prionospoi malmgreni Jewett samples 

Spionidae Pygospio sp. Jewett samples 

Spionidae Pygospio elegans Jewett samples 

Spionidae Rhynchospio glutaeus Jewett samples 

Spionidae Scolelepis sp. Jewett samples 

Spionidae Scolelepis squamata Jewett samples 

Spionidae Spio cirrifera Jewett samples 

Spionidae Spio fillicornis Jewett samples 

Spionidae Spio sp. Jewett samples 

Spionidae Spiophanes berkeleyorum Jewett samples 

Spionidae Spiophanes bombyx Jewett samples 

Spionidae Spiophanes sp. Jewett samples


Table 9.E.1. Continued 

Spionidae Sternapsis scutata Jewett samples 

Syllidae Typosyllis armillaris UAM uncataloged 

Syllidae Typosyllis fasciata Jewett samples 

Syllidae Typosyllis harti UAM uncataoged 

Syllidae Typosyllis sp. UAM uncataoged 

Jewett samples 

Biological Technician Max Hoberg (UAF) used the suspect list to remove 1,154 specimens for 

screening: 154 crustaceans, 1081 polychaetes, 307 molluscs, 30 bryozoans. From these, 

specimens in the best condition were removed for further study. Thirty-five small crustacea 

were loaned to focal taxonomic expert J. Chapman (OSU), and similarly polychaete specimens 

were loaned to focal taxonomic expert J. Kudenov (UA Anchorage). Analyses of these 

specimens (presently on-going) will be reported in the 2000 final report for the University of 

Alaska Sea Grant. 

9E3. Results 

Mollusc Taxa, including One Known NIS 

Most east Pacific records of the soft shell clam, Mya arenaria, are the result of its 

accidental introduction along with Atlantic oysters, starting in San Francisco Bay in 1869, and 

Puget Sound in 1888 or 1889. We found that the clam was abundant in mud sediments in 

Cordova. We also found it in Port Valdez, Tatitlek, Constantine Harbor, Homer and Seward. 

Mya arenaria is also documented in Alaska from Nunivak Island, Norton Sound, and Kodiak 

Island (UAM collection records) as well as southeastern Alaska. 

Nora Foster examined mollusc specimens from both the EVOS specimens and cataloged 

and uncataloged specimens the UA Museum, selecting the following taxa for re-examination: 

• small venerids and Turtonia, were examined as possibly misidentified specimens of potential 

NIS Protamcorbicula amurensis, Venerupsis philipinarium, or Nuttalia obscurata; 

• small individuals of Musuculus spp. were reexamined as possibly misidentified specimens of 

the potential NIS Musculista stenhousei; 

• Cerithiidae were re-examined were reexamined as possibly misidentified specimens of the 

potential NIS Battilaria; 

• Ocenibrina were reexamined as possibly misidentified specimens of the potential NIS 

Urosalpinx cineria and Ocenebra inornata, two introduced oyster drills; and 

• Crepidula specimens were reexamined as possibly misidentified specimens of the potential 

NIS Crepidula fornicata. 

None of these potential NIS taxa were found. However, to date it has been possible to examine 

carefully only a small subsample of the collections, and we are faced with the situation of there 

being too many samples and too little time. 

University of Alaska Sea Grant funding for this analysis of existing museum and 

reference collections continues until June 30, 2000. The goals for this remaining time frame are 

to: 

• Incorporate information from additional taxonomic experts and ecologists.


• Check database against additional grey literature sources (e.g., O’Clair & Zimerman,1987). 

• Add to the database a biogeographical descriptor, indicating whether the PWS record 

represents a range extension, and a reference source for the animal’s distribution. 

• Publish with J. Goddard an analysis of our 1999 collections of opisthobranch molluscs of 

Prince William Sound (see also focal taxonomic subsection 9C8 Opisthobranch Molluscs, 

above). 

• Incorporate of additional specimens collected as part of this assessment of EVOS collections 

into the UAM Aquatic Collection. 

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Chapt 10. Biodiversity, page 10- 1 

Chapter 10. Biodiversity of Prince William Sound 


Howard M. Feder, Institute of Marine Sciences, University of Alaska Fairbanks 

10A. Summary 

Basic taxonomic, biogeographic, and habitat information for 1876 species of marine 

plants and animals of Prince William Sound were compiled into structured data sets. This species 

inventory is intended as a baseline from which biodiversity responses to future environmental 

changes can be assessed measured. The data sets include 39 possibly undescribed species, 89 

range extensions, and 17 nonindigenous species. Two hundred thirty-one marine plant species, 

mostly algae, are present. Twenty invertebrate phyla are represented, and Annelida, Mollusca, 

and Crustacea account for over 60% of the invertebrates. Vertebrates include 175 fish, 14 

mammal, and 113 bird species. The fauna and flora of Prince William Sound is a mixture of 

species with biogeographic affinities that overlap the northeastern and northwestern Pacific as 

well as the Arctic and Atlantic regions. 

10B. Purpose 

A single inventory of Prince William Sound biodiversity is currently lacking. This gap in 

taxonomic and biogeographic knowledge is an impediment to efforts to monitor environmental 

change, to understand the diversity of plant and animal life and to clarify biogeographic 

relationships among the living organisms in the rich waters of Prince William Sound and the 

adjacent shelf of the northern Gulf of Alaska. (Hines et al. 2000) The purpose of this report is to 

compile a database for the plant and animal species of Prince William Sound. The project is one 

component of a research project on potential introductions of nonindigenous species into Prince 

William Sound, especially through the discharge of ballast water from oil tankers traveling into 

Port Valdez. Annotated species lists were developed for the project to help taxonomic experts 

establish a current baseline for the status of nonindigenous species in Prince William (Hines et 

al. 2000). 

Working versions of the data sets were compiled by M. Fry and expanded by N. Foster 

with assistance by M. Hoberg, and review by K. Coyle (zooplankton), J. Goddard 

(opisthobranch gastropods), J. Kudenov (polychaete annelids), G. Lambert (urochordates), P. 

Lambert (echinoderms), C. Mecklenberg (fishes), C. Mills (Cnidaria and Ctenophora), J. 

Norenburg (nemerteans) and others. Additional polychaete identifications were accomplished for 

this report by J. Kudenov. The draft versions of the data sets were included in the final project 

report (Hines et al. 2000). 

10C. Scope 

Taxonomic 

The data sets list free-living macrophytes and animals . Protists and most endoparasites 

are outside the scope of the project. 

Geographic 

Prince William Sound waters are well-defined We have used literature and collecting 

records from Port Bainbridge east to Orca Inlet, and south and east to the outer coasts of


Montague and Hinchinbrook islands. However, for some taxa, (e.g. Bryozoa), the fauna of 

adjacent areas (northeastern Gulf of Alaska and Kodiak Island waters) are better documented and 

some records for those areas are included. Migratory birds and marine mammals that inhabit 

Prince William Sound waters seasonally are included in the data set. Planktonic Cnidaria and 

Crustacea from the northern Gulf of Alaska listed in the data sets occur in Prince William Sound 

rarely, with their presence dependent on oceanographic conditions. 

10D. Methods 

Literature sources 

A large number of species included in the data sets are based on their listing in the 

environmental surveys of the Gulf of Alaska, Prince William Sound, and Port Valdez in the late 

1970’s and early 80’s. We compiled species lists from Cooney et al. (1973) and Cooney and 

Coyle (1988):plankton, Feder et al. (1976):sediments, Feder and Bryson-Schwafel (1988):the 

intertidal zone, Feder and Jewett(1987):subtidal benthos, Feder and Keiser (1980):intertidal 

biology, Feder and Matheke (1980):benthic infauna, Feder et al. (1979):benthos of Resurrection 

and Aialik bays, Feder and Paul (1973):the intertidal zone, Feder and Paul (1980):Cook Inlet, 

Hoberg (1986):benthic infauna, Jewett and Feder (1977):Port Valdez, and Rogers et al. 

(1986):nearshore fishes. 

Species names from field guides and taxonomic revisions of characteristic northeastern 

Pacific taxa have been added, for example Banse and Hobson (1968) (benthic errantiate 

polychaetes), Barnard (1969) (gammaridean amphipods), Butler (1980) (shrimp), Coan et al. 

(2000) (bivalve mollusks), Dick and Ross (1988) (Bryozoa), Lambert (1981, 1997) (sea stars and 

sea cucumbers), and Schultz (1969) (isopods). For detailed information on the geographical 

ranges, we have relied on the same taxonomic revisions and regional publications. However, this 

information in a consistent format has been difficult to obtain for many species. When specific 

reference to a species’ occurrence in Prince William Sound was not available, we used Austin 

(1985). We have also deleted inaccurate records that have crept into the literature, and have 

made changes in nomenclature based on reviews and taxonomic revisions accomplished after 

the environmental surveys mentioned above. 

Biogeographic analyses presented here use conventions based on a global system 

developed by the World Conservation Union and described in Kelleher et al. (1995) (Figures 

10.1 and 10.2). The biogeographic summaries are based on numbers of species for which we 

have confident identifications, leaving out unidentified or undescribed species. Some of the 

analyses exclude birds, because the migratory habits of some species fit poorly into the 

bioregional classification used for algae, invertebrates and fishes.


Figure 10.1. Biogeographic areas of the North Pacific. 

Figure 10.2. Biogeographic areas of the North Atlantic.


Specimens and observations 

The second major source for names of Prince William Sound species is the collections of 

the University of Alaska Museum (UAM). Names of algae, mollusks, bryozoans, and fishes were 

entered from specimen catalogues. Voucher specimens for many of the species listed in the 

environmental studies are in the UAM Aquatic Collection, but may not be catalogued. A large 

number of specimens was also collected in 1990-1995 by Dr. Stephen Jewett as part of damage 

assessment after the 1989 Exxon Valdez oil spill. While not accessioned into the UAM, these 

specimens were available for this study . Each species entry should be traceable to a specimen 

identified to the species level by an expert. Citing specimens from other museum collections, 

however, goes beyond the scope of the project. 

Rapid community assessments, focal taxonomic collections, and fouling plate surveys in 

Prince William Sound in 1998 and 1999 were intended to detect well-established nonindigenous 

species, especially in areas at risk for invasion (Hines and Ruiz 2000). Taxonomic experts who 

participated in the field work in Prince William Sound and the Kenai Peninsula contributed 

names and distribution information for marine plants (G. Hansen), Cnidaria and Ctenophora (C. 

Mills), opisthobranch gastropods (J. Goddard), polychaete annelids (J. Kudenov), Crustacea (J. 

Chapman and J. Cordell), Bryozoa (J. Winston), and urochordates (G. and C. Lambert). 

Results 

The data sets 

The data sets comprising the bulk of this report are in the form of separate Excel 

worksheets for algae and vascular plants, Cnidaria and Ctenophora, Annelida, Mollusca, 

Arthropoda , Bryozoa, Echinodermata, miscellaneous invertebrate taxa, fishes, birds, mammals 

(Tables 10.1 – 10.11). Data are presented in tabular form, one species per row. 

The following data fields, as columns, are common to all data sets: 

• Family, Genus, Species 

An attempt has been made to use the currently accepted name 

• Other Name 

The column contains family, genus or species names under which the taxon has 

appeared in the cited literature, museum catalogs, or specimens labels, which may not 

reflect revisions made in the past 20 years. 

• A Source for the name 

UAM specimen available in the University of Alaska Museum Aquatic Collection 

EVOS specimen archived as part of Exxon Valdez oil spill damage assessment studies 

Other sources are in Literature Cited section of this report. 

• Habitat 

For most invertebrates the following conventions are used: 

I intertidal Inf infauna P plankton 

ST subtidal Epi epifauna


Abbreviations used in the fish, bird and mammal sections will be given with the 

Summary Information for those taxa. 

• Bioregion 

An “x” marks the presence of a species in the large scale bioregions: Northwestern 

Pacific (NWP), Arctic (AR), and Northeastern Atlantic (NWA) (Figures 1, 2). 

Occurrence of each species with in the northeastern Pacific bioregion is further 

designated by Roman numerals denoting a small scale bioregion (Figure 3). 

• Origin 

For species that represent a new collecting record or nonindigenous species, “origin” 

indicates the area closest to Prince William Sound from which the species is likely to 

have spread. 

• NIS status 

nr 

NR 

C 

definite 

possible 

range extension within Alaska to Prince William Sound 

new record for Alaska 

cryptogenic 

definite nonindigenous species 

likely a nonindigenous species 

• Reference to Distribution 

Literature source for distributional information. 

Summary information for major taxa 

Marine Plants 

Red, brown, and green algae account for over 90% of the 231 marine plants listed in the 

data set (Table 10.1). Ten taxa could not be determined to the species level. The non-vascular 

plant species lists is based on G. Hansen’s (2000) report and cataloged UAM specimens. Records 

of Chrysophyta and Ascomycota are based on Feder and Keiser 1980, and Feder and Bryson- 

Schwafel 1988. Additional distribution information is derived from Scagel et al. 1986 and 

Lindstrom 1977. Ten vascular plants species are included. The surfgrasses and eelgrass are 

obvious vascular plants, but a few other species are included because of their presence in the 

upper intertidal and splash zones. Sources used are Scheel et al. 1997; Hulten 1968, and Hansen 

2000. Hansen (2000) found 17 new distributional records for marine plants in Prince William 

Sound. There is one undescribed species, a Coilodesme and six species are possibly introduced.


Table 10.1. Marine plants. 

BIOREGION 

SPECIMEN or 

SOURCE 

COMMUNI 

TY NEP NWP AR NWA ORIGIN 

NIS 

STATUS 

REFERENCE to 

DISTRIBUTION 

PHYLUM FAMILY GENUS SPECIES OTHER NAMES 

Feder and Bryson- 

Ascomycota Verrucaricaea Verrucaria mucosa 

Schwafel 1988 I II III IV x x C Hansen 2000 

Ascomycota Verrucaricaea Verrucaria maura Hansen 2000 I II III IV x x C Hansen 2000 

Chlorophyta Acrosiphoniaceae Acrosiphonia saxitilis Spongomorpha Hansen 2000 I II III IV x Hansen 2000 

Chlorophyta Acrosiphoniaceae Acrosiphonia arcta Hansen 2000 I II III IV x x C Hansen 2000 

Chlorophyta Acrosiphoniaceae Acrosiphonia coalita Hansen 2000 I II III IV Hansen 2000 

Chlorophyta Acrosiphoniaceae Urospora penicilliformis Hansen 2000 I x x x x C Hansen 2000 

Chlorophyta Capsosiphonaceae Capsosiphon fulvescens Hansen 2000 I II III IV x x BC NR, C Hansen 2000 

Chlorophyta Chlorocystidaceae Halochlorococcum moorei Hansen 2000 I II III IV x x BC NR, C Hansen 2000 

Chlorophyta Cladophoraceae Chaetomorpha capillaris/ cannabina Hansen 2000 I II III IV x x Hansen 2000 

Chlorophyta Cladophoraceae Chaetomorpha recurva Hansen 2000 I II III IV WA NR Hansen 2000 

Chlorophyta Cladophoraceae Cladophora sericea C. gracilis Hansen 2000 I II III IV x C Hansen 2000 

Chlorophyta Cladophoraceae Cladophora albida Hansen 2000 I II III IV x x C Hansen 2000 

Chlorophyta Cladophoraceae Cladophora hutchinsiae Hansen 2000 I II III IV x x C Hansen 2000 

Chlorophyta Cladophoraceae Cladophora stimpsonii Hansen 2000 I II III IV Hansen 2000 

Chlorophyta Cladophoraceae Rhizoclonium implexum R. implexum Hansen 2000 I x x x x C Hansen 2000 

Chlorophyta Cladophoraceae Rhizoclonium riparium Hansen 2000 I x x x x C Hansen 2000 

Chlorophyta Cladophoraceae Rhizoclonium tortuosum Hansen 2000 I x x x x C Hansen 2000 

Chlorophyta Codiaceae Codium fragile subsp. fragile Hansen 2000 I II III IV V VI SE Alaska nr Hansen 2000 

Chlorophyta Codiaceae Codium 

fragile subsp. 

tomentosoides fragile Hansen 2000 I x x x x WA probable Hansen 2000 

Chlorophyta Collinsiellaceae Collinsiella tuberculata 


Schwafel 1988 I II III IV Scagel et al. 1986 

Chlorophyta Gayraliaceae Gayralia oxyspermum Hansen 2000 I II III IV x x C Hansen 2000 

Chlorophyta Monostromataceae Monostroma fractum Hansen 2000 I II III IV WA NR Hansen 2000 

Chlorophyta Monostromataceae Monostroma grevillei/arcticum M. arcticum Hansen 2000 I II III IV x x C Hansen 2000 

Chlorophyta Ulotrichaceae Ulothrix flacca U. pseudoflacca 


Schwafel 1988 I II III IV x x C Scagel et al. 1986 

Chlorophyta Ulotrichaceae Ulothrix implexa ( non flacca) Hansen 2000 I II III IV x x C Scagel et al. 1986 

Chlorophyta Ulvaceae Blidingia chadefauldi Hansen 2000 I x x x C Hansen 2000 

Chlorophyta Ulvaceae Blidingia marginata Hansen 2000 I x x x x BC NR, C Hansen 2000 

Chlorophyta Ulvaceae Blidingia minima Hansen 2000 I x x x x C Hansen 2000 

Chlorophyta Ulvaceae Blidingia subsalsa Hansen 2000 I II III IV x x C Hansen 2000 

Chlorophyta Ulvaceae Enteromorpha compressa 


Schwafel 1988 I x x x x C Scagel et al. 1986 

Chlorophyta Ulvaceae Enteromorpha clathrata E. crinita Hansen 2000 I x x x x C Scagel et al. 1986 

Chlorophyta Ulvaceae Enteromorpha intestinalis Hansen 2000 I x x x x C Hansen 2000 

Chlorophyta Ulvaceae Enteromorpha linza Hansen 2000 I x x x x C Hansen 2000 

Chlorophyta Ulvaceae Enteromorpha prolifera/torta Hansen 2000 I x x x x C Hansen 2000 

Chlorophyta Ulvaceae Kornmannia leptoderma Hansen 2000 I/epilithic II III IV x x North Atlantic NR, C Hansen 2000


Table 10.1. Continued. 

BIOREGION 


SPECIMEN or 

SOURCE 

COMMUNI 


NIS 

STATUS 

REFERENCE to 

DISTRIBUTION 

Chlorophyta Ulvaceae Kornmannia zostericola Hansen 2000 I/epiphytic II III IV x x C Hansen 2000 


Chlorophyta Ulvaceae Ulva fenestrata U. lactuca Schwafel 1988 I II III IV x Scagel et al. 1986 

fenestrata/expansa/ 

Chlorophyta Ulvaceae Ulva 

lactuca Hansen 2000 I x x x x C Hansen 2000 

Chlorophyta Ulvaceae Ulvaria obscura 

Monostroma 

fuscum Hansen 2000 I II III IV x x C Hansen 2000 

Chlorophyta Ulvellaceae Ulvella setchelli Hansen 2000 

I/epiphytic/e 

ndophytic II III IV V VI x C Hansen 2000 

Chlorophyta Cladophoraceae Percursaria percursa Hansen 2000 brack/I x x x x C Hansen 2000 

Cyanophyta Calothryx crustacea Hansen 2000 I x x x x C Hansen 2000 

Cyanophyta Rivularia atra Hansen 2000 I x x x x C Hansen 2000 

Phaeophyta Alariaceae Alaria prelonga/marginata Hansen 2000 I/ST II III IV V x Hansen 2000 

Phaeophyta Alariaceae Alaria 

taeniata/augusta/ 

crispa Hansen 2000 I/ST II III IV x Hansen 2000 

Phaeophyta Alariaceae Alaria 

tenuifolia/pylaii/ 

membranacea Hansen 2000 I/ST II III IV x Hansen 2000 

Phaeophyta Chordaceae Chorda filum Hansen 2000 I/ST II III IV x x Hansen 2000 

Phaeophyta Chordariacea Chordaria flagelliformis Hansen 2000 I/ST II III IV x x x C Hansen 2000 

Phaeophyta Chordariacea Chordaria gracilis Hansen 2000 I/ST II III IV x C Hansen 2000 

Phaeophyta Chordariaceae Eudesme virescens Hansen 2000 I II III IV x x Hansen 2000 

Phaeophyta Chordariaceae Saundersella simplex UAM I/epiphytic II III IV x Scagel et al. 1986 

Phaeophyta Coilodesmaceae Coilodesme bulligera C. polygnampta Hansen 2000 I II III IV x x C Hansen 2000 

Phaeophyta Coilodesmaceae Coilodesme californica Hansen 2000 

I/ST/ 

epiphytic II III IV V VI Hansen 2000 

Phaeophyta Coilodesmaceae Coilodesme undescribed Hansen 2000 I II III x Hansen 2000 

Phaeophyta Cystoseiraceae Cystoceria germinata Hansen 2000 I/ST II III IV x Hansen 2000 

Phaeophyta Desmarestiaceae Desmarestia lingulata UAM I/ST II III IV V VI Scagel et al. 1986 

Phaeophyta Desmarestiaceae Desmarestia viridis D. media UAM I/ST II III IV V VI x x C Scagel et al. 1986 

Phaeophyta Desmarestiaceae Desmarestia viridis Hansen 2000 I/ST II III IV V VI x x Hansen 2000 

Phaeophyta Desmarestiaceae Desmarestia aculeata D. intermedia UAM I/ST II III IV x x C Hansen 2000 

Phaeophyta Dictyosiphonaceae Dictyosiphon foeniculaceus Hansen 2000 I II III IV x x C Hansen 2000 

Phaeophyta Dictyosiphonaceae Dictyosiphon sinicola UAM I II III IV Scagel et al. 1986 

Phaeophyta Ectocarpaceae Ectocarpus parvus Hansen 2000 I II III IV V VI Hansen 2000 

Phaeophyta Ectocarpaceae Ectocarpus siliculosus Hansen 2000 I x x x x C Hansen 2000 

Phaeophyta Ectocarpaceae Ectocarpus acutus Hansen 2000 I II III IV V BC nr Hansen 2000 

Phaeophyta Ectocarpaceae Ectocarpus dimorpha E. dimorphus Hansen 2000 I II III IV V BC nr Hansen 2000 

Phaeophyta Ectocarpaceae Ectocarpus sp. (Acinetospora) Hansen 2000 I II III IV V Hansen 2000 

Phaeophyta Ectocarpaceae Pilayella littoralis Hansen 2000 I x x x x Hansen 2000



BIOREGION 

SPECIMEN or 

SOURCE 

COMMUNI 


NIS 

STATUS 

REFERENCE to 

DISTRIBUTION 


littoralis/ 

Phaeophyta Ectocarpaceae Pilayella 

washingtoniensis Hansen 2000 I x x x x C Hansen 2000 

Phaeophyta Ectocarpaceae Pilayella unidentified Hansen 2000 I/epiphytic II Hansen 2000 

Phaeophyta Ectocarpiaceae Spongonema tomentosum Hansen 2000 I/epiphytic II III IV V VI x x x C Hansen 2000 

Phaeophyta Elachistaceae Elachista fucicola Hansen 2000 I/epiphytic II III IV V x x Hansen 2000 

Phaeophyta Elachistaceae Elachista lubrica Hansen 2000 I/epiphytic II III x x Hansen 2000 

gardneri/distichus/ 

Phaeophyta Fucaceae Fucus 

evanensis Hansen 2000 I II III IV V C Hansen 2000 

Phaeophyta Fucaceae Fucus cottoni Hansen 2000 I II III IV x x North Atlantic probable Hansen 2000 

Phaeophyta Fucaceae Fucus spiralis UAM I II III IV C Hansen 2000 

Phaeophyta Heterochordariaceae Analipus japonicus UAM I II III IV V x Scagel et al. 1986 

Phaeophyta Laminariaceae Agarum clathratum (cribrosum) Hansen 2000 ST II III IV x x C Hansen 2000 

Phaeophyta Laminariaceae Costaria costata Hansen 2000 I II III IV x Hansen 2000 

Phaeophyta Laminariaceae Cymathere triplicata Hansen 2000 ST II III IV x Hansen 2000 

Phaeophyta Laminariaceae Hedophyllum sessile UAM I II III IV V Scagel et al. 1986 

Phaeophyta Laminariaceae Laminaria groenlandica 


Schwafel 1988 I/ST II III IV Scagel et al. 1986 

"groenlandica"/ 

Phaeophyta Laminariaceae Laminaria 

bongardiana Hansen 2000 ST II III IV x x Hansen 2000 

Phaeophyta Laminariaceae Laminaria dentigera UAM ST II III IV x Scagel et al. 1986 

Phaeophyta Laminariaceae Laminaria longipes UAM I/ST II III IV x Scagel et al. 1986 

Phaeophyta Laminariaceae Laminaria yezoensis Hansen 2000 I II III IV x Hansen 2000 

Phaeophyta Laminariaceae Laminaria saccharina Hansen 2000 I II III IV x x C Hansen 2000 

Phaeophyta Laminariaceae Pleurophycus gardneri UAM ST II III IV V Scagel et al. 1986 

Phaeophyta Leathesiaceae Leathesia difformis 


Schwafel 1988 I x x x x C Scagel et al. 1986 

Phaeophyta Leathesiaceae Leathesia nana Hansen 2000 I II III IV V VI Hansen 2000 

II III IV V VI 

Phaeophyta Lessoniaceae Macrocystis integrifolia UAM ST VII VIII IX SE Alaska definate Hansen 2000 

Phaeophyta Lessoniaceae Nereocystis leutkeana UAM ST II III IV V Scagel et al. 1986 

Phaeophyta Punctariaceae Delamarea attenuata Hansen 2000 I II III IV x x 

Commander 

Islands NR, C Hansen 2000 


Phaeophyta Punctariaceae Myelophycus intestinalis Melanosiphon Schwafel 1988 I II III IV x x C Scagel et al. 1986 

latifolia 

Phaeophyta Punctariaceae Punctaria 

(Desmotrichium) Hansen 2000 ST II III IV x x SE Alaska nr Hansen 2000 

Phaeophyta Punctariaceae Punctaria lobata Hansen 2000 

E/ST/ 

epiphytic II III IV Hansen 2000 

Phaeophyta Punctariaceae Punctaria plantagenea Hansen 2000 I II III IV x x Japan NR, C Hansen 2000



BIOREGION 


SPECIMEN or 

SOURCE 

COMMUNI 


NIS 

STATUS 

REFERENCE to 

DISTRIBUTION 

Phaeophyta Punctariaceae Punctaria tenuimissima Hansen 2000 I II III IV x x C Hansen 2000 

Phaeophyta Punctariaceae Soranthera ulvoidea Hansen 2000 I II III IV x Hansen 2000 

Phaeophyta Punctariaceae Soranthera ulvoidea f. difformis Hansen 2000 I/epiphytic II III IV x Hansen 2000 

Phaeophyta Ralfsiaceae Microspongium globosum Hansen 2000 I/epiphytic II III IV x x Japan probable Hansen 2000 

Phaeophyta Ralfsiaceae Ralfsia fungiformis Hansen 2000 I II III IV x x C Hansen 2000 

Phaeophyta Scytosiphonaceae Colpomenia peregrina Hansen 2000 I x x x x C Hansen 2000 

Phaeophyta Scytosiphonaceae Colpomenia bullosa Hansen 2000 I/epiphytic II III IV x Hansen 2000 

Phaeophyta Scytosiphonaceae Petalonia fascia Hansen 2000 I/ST x x x x Hansen 2000 

Phaeophyta Scytosiphonaceae Scytosiphon lomentaria 


Schwafel 1988 I x x x x Scagel et al. 1986 

Phaeophyta Scytosiphonaceae Scytosiphon simplicissima Hansen 2000 I x x x x C Hansen 2000 

Phaeophyta Sphacelariaceae Sphacelaria racemosa Hansen 2000 I/ST II III IV x x C Hansen 2000 

Phaeophyta Sphacelariaceae Sphacelaria rigidula Hansen 2000 I x x x x C Hansen 2000 

Rhodophyta Acrochaetiaceae Acrochaetium unidentified UAM I C 

Rhodophyta Acrochaetiaceae Audouienlla membranaceum 

Rhodochorton 

membranaceum UAM I x x x x C Scagel et al. 1986 

Rhodophyta Acrochaetiaceae Audouinella purpurea Hansen 2000 I x x x x C Hansen 2000 

Rhodophyta Bangiaceae Bangia atropurpurea Hansen 2000 I x x x x C Hansen 2000 

Rhodophyta Bangiaceae Porphyra cuneiformis Hansen 2000 I/ST II III IV Hansen 2000 

Rhodophyta Bangiaceae Porphyra miniata Hansen 2000 I/ST II III IV x x 

Commander 

Islands NR Hansen 2000 

Rhodophyta Bangiaceae Porphyra mumfordii Hansen 2000 I/ST II III IV Hansen 2000 

Rhodophyta Bangiaceae Porphyra nereocystis Hansen 2000 

ST/ 


Rhodophyta Bangiaceae Porphyra perforata Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Bangiaceae Porphyra purpureo-violacea Hansen 2000 I/ST II III IV x x North Atlantic NR, C Hansen 2000 

Rhodophyta Bangiaceae Porphyra rediviva Hansen 2000 I II III IV WA NR Hansen 2000 

Rhodophyta Bangiaceae Porphyra cf. P. thuretti 



Rhodophyta Bangiaceae Porphyra torta/abbottae Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Ceramiaceae Antithamnion dendroideum UAM ST 


VII VIII IX Scagel et al. 1986 

Rhodophyta Ceramiaceae Antithamnionella pacifica Hansen 2000 

ST/ 


Rhodophyta Ceramiaceae Antithamnionella spiriographidis Hansen 2000 I II III IV x x x C Scagel et al. 1986 

Rhodophyta Ceramiaceae Callithamnion acutum Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Ceramiaceae Callithamnion pikeanum v. laxum Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Ceramiaceae Callithamnion pikeanum v. pikeaum Hansen 2000 I II III IV Hansen 2000



BIOREGION 

SPECIMEN or 

SOURCE 

COMMUNI 


NIS 

STATUS 

REFERENCE to 

DISTRIBUTION 



Rhodophyta Ceramiaceae Callophyllis unidentified 

Schwafel 1988 I II III IV 

Rhodophyta Ceramiaceae Ceramium gardneri Hansen 2000 I II III IV Hansen 2000 

pacificum/ 

Rhodophyta Ceramiaceae Ceramium 

washingtoniensis Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Ceramiaceae Ceramium rubrum/kondoi Hansen 2000 I x x x x Hansen 2000 

Rhodophyta Ceramiaceae Ceramium cimbricum Hansen 2000 I II III IV x x Hansen 2000 

Rhodophyta Ceramiaceae Ceramium sinicola Hansen 2000 I II III IV CA probable Hansen 2000 

Rhodophyta Ceramiaceae Ceramium rubrum UAM I x x x x Scagel et al. 1986 

Rhodophyta Ceramiaceae Hollenbergia unidentified UAM I/ST Scagel et al. 1986 

Rhodophyta Ceramiaceae Hollenbergia subulata UAM I/ST II III IV V Scagel et al. 1986 

Rhodophyta Ceramiaceae Microcladia borealis Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Ceramiaceae Neoptilota asplenioides Hansen 2000 I/ST II III IV x Hansen 2000 

Rhodophyta Ceramiaceae Platythamnion pectinatum Hansen 2000 ST II III IV Hansen 2000 

Rhodophyta Ceramiaceae Plenosporium cf. P. vancouverianum Hansen 2000 ST II III IV x Hansen 2000 

Rhodophyta Ceramiaceae Ptilota serrata ( incl. pectinata) Hansen 2000 I/ST II III IV x x C Hansen 2000 

Rhodophyta Ceramiaceae Ptilota californica UAM I/ST II III IV x C Scagel et al. 1986 

Abbott and Hollenberg 

Rhodophyta Ceramiaceae Ptilota plumosa P. p v. filicina UAM I/ST II III IV V VI x C 

1976 

Rhodophyta Ceramiaceae Ptilota filicina P. tenuis Hansen 2000 I/ST II III IV Hansen 2000 

Rhodophyta Ceramiaceae Scagiella americana Hansen 2000 I II III IV x x C Hansen 2000 

Rhodophyta Choreocolacaceae Leachiella pacifica Hansen 2000 I II III IV V VI x Hansen 2000 

Rhodophyta Cladophoraceae Chroodactylon ramosus Hansen 2000 I II III IV x x CA probable Hansen 2000 

Rhodophyta Corallinaceae Bossiella chiloensis UAM I/ST II III IV Scagel et al. 1986 

Rhodophyta Corallinaceae Bossiella cretacea UAM I/ST II III IV x Scagel et al. 1986 

Rhodophyta Corallinaceae Bossiella plumosa UAM I II III IV Scagel et al. 1986 

Rhodophyta Corallinaceae Calliarthron unidentified UAM I Scagel et al. 1986 

Rhodophyta Corallinaceae Clathromorphum reclinatum I/epiphytic II III IV V VI x Scagel et al. 1986 

Rhodophyta Corallinaceae Corallina frondescens Hansen 2000 I II III IV Hansen 2000 


Rhodophyta Corallinaceae Corallina officianalis v. chilensis Hansen 2000 I/ST VII VII VIII IX x Hansen 2000 

Rhodophyta Corallinaceae Corallina vancouverensis UAM I II III IV Scagel et al. 1986 

Rhodophyta Corallinaceae Lithophyllum dispar Hansen 2000 II III IV Hansen 2000 

Rhodophyta Corallinaceae Lithothamnion unidentified I Scagel et al. 1986 

Rhodophyta Corallinaceae Mesophyllum lamellatum Scagel 1986 I II III IV V VI Scagel et al. 1986 

Rhodophyta Delesseriacaea Delesseria decipiens Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Delesseriaceae Phycodrys riggii Hansen 2000 I/ST II III IV x Hansen 2000 

Rhodophyta Delesseriaceae Tokidadendron kurilensis T. bullata Hansen 2000 I II III IV x Hansen 2000



BIOREGION 


SPECIMEN or 

SOURCE 

COMMUNI 


NIS 

STATUS 

REFERENCE to 

DISTRIBUTION 

Rhodophyta Dumontiacaea Cryptosiphonia woodii Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Dumontiacaea Weeksia coccinea Hansen 2000 II III IV Hansen 2000 

Rhodophyta Dumontiaceae Constantinea subulifera Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Dumontiaceae Constantinea simplex 


Schwafel 1988 I II III IV V Scagel et al. 1986 

Rhodophyta Dumontiaceae Dumontia simplex Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Dumontiaceae Dumontia contorta D. incrassata Hansen 2000 I II III IV x x C Hansen 2000 

Rhodophyta Dumontiaceae Farlowia mollis UAM I/ST II III IV V VI x Scagel et al. 1986 

Rhodophyta Dumontiaceae Neodilsea unidentified UAM I Hansen 2000 

Rhodophyta Endocladiaceae Endocladia muricata Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Endocladiaceae Gloiopeltis furcata Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Erythropeltidaceae Erythrotrichia carnea Hansen 2000 I x x x x Hansen 2000 

ST/ 

Rhodophyta Erythropeltidiaceae Smithoria naiadum Hansen 2000 epiphytic II III IV C Hansen 2000 

Rhodophyta Gigartinaceae Gigartina unidentified UAM I/ST Hansen 2000 

Rhodophyta Gigartinaceae Iridaea cordata UAM I/ST II III IV x Scagel et al. 1986 

Rhodophyta Gigartinaceae Iridaea cornucopiae 


Schwafel 1988 I II III IV x Scagel et al. 1986 

Rhodophyta Gigartinaceae Rhodoglossum latissimum UAM I II III IV Scagel et al. 1986 

Rhodophyta Halemeniaceae Cryptonemia obovata Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Halemeniaceae Cryptonemia borealis Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Helminthocladiaceae Nemalion helminthoides Hansen 2000 I x x x x C Hansen 2000 

Rhodophyta Hildenbrandiaceae Hildenbrandia rubra H. prototypus Hansen 2000 I x x x x C Scagel et al. 1986 

Rhodophyta Nemastomataceae Neodilsea borealis 

N. americana/ 

Schizymenia UAM I II III IV Scagel et al. 1986 

Rhodophyta Palmariaceae Devaleraea ramentacea Hansen 2000 I II III IV x x C Hansen 2000 

Rhodophyta Palmariaceae Halosaccion glandiforme Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Palmariaceae Halosaccion firmum Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Palmeriaceae Palmaria mollis/palmata Hansen 2000 I II III IV x x C Hansen 2000 

Rhodophyta Palmeriaceae Palmaria 

calophylloides/ 

stenogona Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Palmeriaceae Palmaria hecatensis Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Petrocelidaceae Mastocarpus cf. M. pacificus Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Petrocelidaceae Mastocarpus papillatus complex Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Petrocelidaceae Mastocarpus papillatus Gigartina cristata UAM I 


VII VII VIII IX Scagel et al. 1986 

Rhodophyta Phyllophoraceae Ahnfeltia plicata UAM I II III IV x x Scagel et al. 1986 

Rhodophyta Phyllophoraceae Ahnfeltia fastigata Hansen 2000 I x x x x C Hansen 2000 

Rhodophyta Phyllophoraceae Ahnfeltiopsis gigartinoides Arnfeltia Hansen 2000 I II III IV Hansen 2000



BIOREGION 


SPECIMEN or 

SOURCE 

COMMUNI 


NIS 

STATUS 

REFERENCE to 

DISTRIBUTION 

Rhodophyta Rhodomelaceae Polysiphonia brodiaei Hansen 2000 I x x x x C Hansen 2000 

Rhodophyta Rhodomelaceae Polysiphonia eastwoodae Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Rhodomelaceae Polysiphonia 

hendryi v. 

deliquescens Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Rhodomelaceae Polysiphonia hendryi v. hendryi Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Rhodomelaceae Polysiphonia hendryi v. luxurians Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Rhodomelaceae Polysiphonia hendryi v. pacifica Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Rhodomelaceae Polysiphonia 

pacifica v. 

determinata Hansen 2000 I/ST II III IV Hansen 2000 

Rhodophyta Rhodomelaceae Polysiphonia pacifica v. pacifica Hansen 2000 I/ST II III IV Hansen 2000 

Rhodophyta Rhodomelaceae Polysiphonia pungens Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Rhodomelaceae Polysiphonia hendryi P. collinsii UAM I II III IV x Scagel et al. 1986 

Rhodophyta Rhodomelaceae Polysiphonia pacifica 



Rhodophyta Rhodomelaceae Polysiphonia senticulosa Hansen 2000 I II III IV SE Alaska nr Hansen 2000 

Rhodophyta Rhodomelaceae Polysiphonia stricta (ureceolata) Hansen 2000 I II III IV x x Hansen 2000 

Rhodophyta Rhodomelaceae Polysiphonia ureceolata Hansen 2000 I x x x x Hansen 2000 

Rhodophyta Rhodomelaceae Pterosiphonia bipinnata Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Rhodomelaceae Rhodomela lycopodioides Hansen 2000 I II III IV x x C Hansen 2000 

Rhodophyta Rodomelaceae Neorhodomela larix Rhodomela Hansen 2000 I/ST II III IV x Hansen 2000 

Rhodophyta Rodomelaceae Neorhodomela aculaeta Hansen 2000 I/ST II III IV x Hansen 2000 

Rhodophyta Rodomelaceae Neorhodomela oregona Hansen 2000 I/ST II III IV x Hansen 2000 

Rhodophyta Rodomelaceae Odonthalia floccosa Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Rodomelaceae Odonthalia kamtschatica Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Rodomelaceae Odonthalia setacea Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Rhodymeniaceae Rhodymenia pertusa Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Solieriaceae Opuntiella californica Hansen 2000 ST II III IV Hansen 2000 

Rhodophyta Gigartinaceae Mazzaella heterocarpa Hansen 2000 I II III IV Hansen 2000 

Rhodophyta Gigartinaceae Mazzaella phyllocarpa Hansen 2000 I II III IV x Hansen 2000 

Rhodophyta Gigartinaceae Mazzaella splendescens Hansen 2000 I II III IV Hansen 2000 

Xanthophyta Vaucheria longicaulis () Hansen 2000 I II III IV x x BC NR Hansen 2000 

vascular plant Caryophyllaceae Honchneya peploides Scheel et al. 1997 I II III IV x Hulten 1968 

vascular plant Cyperaceae Carex unidentified Feder and Bryson-SchwI Hulten 1968 

vascular plant Gramineae Puccinellia nutkaensis Scheel et al. 1997 I II III IV Hulten 1968 

vascular plant Gramineae Puccinellia plumila Hulten 1968 I II III IV Hulten 1968 

vascular plant Primulaceae Glaux maritima Scheel et al. 1997 I II III IV x x Hulten 1968 

vascular plant Rosaceae Potentilla unidentified Feder and Bryson-SchwI Hulten 1968 

vascular plant Sparganiaceae Phyllospadix scouleri Hansen 2000 I/ST II III IV Hulten 1968 

vascular plant Sparganiaceae Phyllospadix serrulatus Hansen 2000 I/ST II III IV Hulten 1968 

vascular plant Sparganiaceae Zostera marina Hansen 2000 I/ST II III IV x x Hulten 1968


Cnidaria and Ctenophora 

One hundred two species of Cnidaria and Ctenophora are listed in this report (Table 

10.2). Twenty-seven of the taxa have been identified to the generic level, and not to species. 

Anthozoa are probably under-counted, because they are difficult to identify based on preserved 

specimens. The hydrozoans and scyphozoans include 34 species, four of which, the hydrozoans 

Tiaropsis multicirrata, Eperetmus typus, Gonionemus vertens and Proboscidactyla flavicirrata, 

are distributional range extensions (Mills 2000). Thirty species records are derived from T. 

Cooney‘s (1987) compilation of Gulf of Alaska plankton. 

Table 10.2. Cnidaria and Ctenophora. 

PHYLUM/ 

SPECIMEN or 

CLASS FAMILY GENUS SPECIES OTHER NAMES SOURCE HABITAT NEP NWP AR NWA ORIGIN 

Cnidaria: 

Anthozoa Actiniidae Anthopleura artemesia 

Cnidaria: 

Anthozoa Actiniidae Cribirnopsis unidentified Rosenthal 1977 ST/Epi 

Cnidaria: 

Anthozoa Actiniidae Urticina crassicornis 

NIS 

STATUS 

REFERENCE to 

DISTRIBUTION 


Schwafel 1988 I II III IV Austin 1985 

Feder and Jewett 

1987 I/ST/Epi II III IV x Austin 1985 

Cnidaria: 

Anthozoa Actinostolidae Stomphia unidentified Scheel et al. 1997 ST/Epi 

Cnidaria: 

Anthozoa Carophllidae Carophyllia alaskensis UAM ST II III IV Austin 1985 

Cnidaria: 

Anthozoa Cerianthidae Pachycerianthus unidentified UAM ST 

Cnidaria: 

Anthozoa Epizoanthidae Epizoanthus unidentified UAM ST 

Cnidaria: 

Anthozoa Halcalvidae Peachia unidentified 

Cnidaria: 

Anthozoa Metridiidae Metridium senile 

Feder and Matheke 

1980 ST 


1987 I/ST/Epi II x x x Austin 1985 

Cnidaria: 

Anthozoa Metridiidae Metridium unidentified NRF observation ST 

Cnidaria: 

Anthozoa Pennatulidae Ptilosarcus gurneyi Feder et al. 1979 ST II III IV Austin 1985 

Cnidaria: 

Anthozoa Primnoidae Callogorgia unidentified UAM ST 

Cnidaria: 

Anthozoa Primnoidae Primnoa unidentified UAM ST 

Cnidaria: 

Anthozoa Virgularidae Acanthoptilum ptile 


1980 ST II III 

Cnidaria: 

Anthozoa Virgularidae Stylatula elongata Rosenthal 1977 ST II III IV V 

Cnidaria: 

British 

columbia NR Austin 1985 

Hydrozoa Aeginidae Aegina citrea A. rosea Cooney 1987 P II III IV V VI x x Austin 1985 

Cnidaria: 

Hydrozoa Aequoreidae Aequorea 

aequorea v. 

albida Mills 2000 ST II III IV 

Wrobel and Mills 

1998 

Cnidaria: 

Hydrozoa Aequoreidae Aequorea unidentified Cooney:SEA ST 

Cnidaria: 

Hydrozoa Aequoreidae Aequorea 

victoria/ A. 

aequorea v. 

aequorea Mills 2000 ST II III IV 


1998 

Cnidaria: 

Hydrozoa Agalmidae Agalma elegans Cooney 1987 P II III x Austin 1985 

Cnidaria: 

Hydrozoa Agalmidae Nanomia unidentified Cooney 1987 P 

Cnidaria: 

Hydrozoa Bougainvilliidae Bougainvillia superciliaris Mills 2000 ST II III IV x Austin 1985 

Cnidaria: 

Hydrozoa Bougainvilliidae Garveia franciscana Mills 2000 ST x x x x uncertain definate 

Cnidaria: 

Hydrozoa Calycopsidae Calycopsis simulans 

Cnidaria: 

Hydrozoa Campanulariidae Clytia gregaria 

Cnidaria: 

Hydrozoa Campanulariidae Clytia hemispherca 


1998 P x 

Phialidium 

gregarium Mills 2000 I II III IV 

BIOREGION 

Hines and Ruiz 

2000 


1998 


1998 

Hines and Ruiz 

2000 ST/Epi II III IV V VI x Austin 1985 

Cnidaria: 

Hydrozoa Campanulariidae Clytia kincaidi Mills 2000 ST x x x x Austin 1985 

Cnidaria: 

Hydrozoa Campanulariidae Gonothyraea clarki Mills 2000 I II III IV x Austin 1985 

Cnidaria: 

Hydrozoa Campanulariidae Obelia longissima Mills 2000 I II x x Austin 1985 

Cnidaria: 

Hydrozoa Campanulariidae Obelia borealis O. dichotoma Cooney 1987 P II III IV V x Austin 1985 

Cnidaria: 

Hydrozoa Campanulariidae Obelia unidentified Mills 2000 I 

Cnidaria: 

Hydrozoa Campanulariidae Obelia spp. (hydriods) Mills 2000 I 

Cnidaria: 

Hydrozoa Campanulariidae Phialidium gregarium P. languidum Cooney 1987 P II III IV V Austin 1985


Table 10.2 continued. 

PHYLUM/ 

SPECIMEN or 


Cnidaria: 

Hydrozoa Campanulariidae Verticillina verticillata 


1980 ST II III IV V x x 

NIS 

STATUS 

REFERENCE to 

DISTRIBUTION 

Cnidaria: 

Hydrozoa Clausophyidae Chuniphyes multidentata Cooney 1987 P II III IV V VI Austin 1985 

Cnidaria: 

Hydrozoa Corymorphidae Euphysa japonica Cooney 1987 P II III IV x x x 


1998 

Cnidaria: 

Hydrozoa Corymorphidae Euphysa unidentified Mills 2000 I/ST/Epi 

Cnidaria: 

Hydrozoa Corynidae Sarsia spp. (medusae) Mills 2000 I/ST/Epi 

Cnidaria: 

Hydrozoa Corynidae Sarsia tubulosa Mills 2000 I/ST/Epi x x x x Austin 1985 

Cnidaria: 

Hydrozoa Corynidae Sarsia eximia Mills 2000 I/ST/Epi II x x Austin 1985 

Cnidaria: 

Hydrozoa Corynidae Sarsia princeps Cooney 1987 P II III IV x Austin 1985 

Cnidaria: 

Hydrozoa Corynidae Sarsia rosaria Cooney 1987 P II III IV x Austin 1985 

Cnidaria: 

Hydrozoa Corynidae Sarsia/Coryne sp. (hydroids) Mills 2000 I/ST/Epi 

Cnidaria: 

Hydrozoa Cuninidae Cunina globosa Cooney 1987 P II III IV Cooney 1987 

Cnidaria: 

Hydrozoa Diphyidae Dimophyes arctica Cooney 1987 P 


VII VIII x x x Austin 1985 

Cnidaria: 

Hydrozoa Diphyidae Lensia concoidea Cooney 1987 P II III IV V VI x x Austin 1985 

Cnidaria: 

Hydrozoa Diphyidae Muggiaea atlantica Cooney 1987 P II III x x Austin 1985 

Cnidaria: 

Hydrozoa Eirenidae Eutonina indicans 


1998 P II III IV x 


1998 

Cnidaria: 

Hydrozoa Eutimidae Eutonima indicans Mills 2000 ST II III IV Austin 1985 

Cnidaria: 

Hydrozoa Haleciidae Grammaria immersa Rosenthal 1977 ST II III x Austin 1985 

Cnidaria: 

Hydrozoa Halicreatidae Halicreas unidentified Cooney:SEA P 

Cnidaria: 

Hydrozoa Halimedusidae Halimedusa typus Cooney 1987 P II III IV Austin 1985 

Cnidaria: 

Hydrozoa Hippopodiidae Vogtia serrata V. pentacantha Cooney 1987 P II III x x Austin 1985 

Cnidaria: 

Hydrozoa Laodicidae Cuspidella unidentified Cooney:SEA P 

Cnidaria: 

Hydrozoa Laodicidae Staurophora mertensii Mills 2000 ST II x x Austin 1985 

Cnidaria: 

Hydrozoa Laodicidae Stegopoma plicatile 

BIOREGION 


1980 ST II x x Austin 1985 

Cnidaria: 

Hydrozoa Lovenellidae Calycella springe Mills 2000 I II x x Austin 1985 

Cnidaria: 

Hydrozoa Melicertidae Melicertum campanula Cooney:SEA ST II III IV x x Austin 1985 

Cnidaria: 

Hydrozoa Melicertidae Melicertum octocostatum Mills 2000 ST II x x 

Cnidaria: 

Hydrozoa Mitrocomidae Mitrocoma cellularia Halistaura Mills 2000 ST II III IV V 

Cnidaria: 

Hydrozoa Mitrocomidae Tiaropsidium unidentified Cooney 1987 P 

Cnidaria: 

Hydrozoa Mitrocomidae Tiaropsis multicirrata Mills 2000 I/ST/Epi II III IV x x SE Alaska nr 

Cnidaria: 

Hydrozoa Olindiasidae Eperetmus typus Mills 2000 ST II III IV SE Alaska nr 

Cnidaria: 

Hydrozoa Olindiasidae Gonionemus unidentified Cooney:SEA ST 

Cnidaria: 

Hydrozoa Olindiasidae Gonionemus vertens Mills 2000 ST II III IV SE Alaska nr 


1998 


1998 


1998 


1998 


1998



PHYLUM/ 

SPECIMEN or 


NIS 

STATUS 

REFERENCE to 

DISTRIBUTION 

Cnidaria: 

Hydrozoa Pandeidae Catablema multicirrata Mills 2000 ST/Epi II III IV x Austin 1985 

Cnidaria: 

Hydrozoa Pandeidae Halitholus unidentified Mills 2000 ST 

Cnidaria: 

Hydrozoa Pandeidae Leuckartiara breviconis Neoturris Cooney 1987 P II III IV Austin 1985 

Cnidaria: 

Hydrozoa Pandeidae Leuckartiara nobilis 

Catablema 

nodulosa Cooney 1987 P II III IV Austin 1985 

Cnidaria: 

Hydrozoa Pandeidae Leuckartiara octona Cooney 1987 P II III IV Austin 1985 

Cnidaria: 

Hydrozoa Pandeidae Leuckartiara unidentified Mills 2000 I/ST/Epi 

Cnidaria: 

Hydrozoa Pandeidae Neoturris breviconis 


1998 P II II III IV V 

Cnidaria: 

Hydrozoa Pandeidae Perigonimus unidentified Cooney:SEA P 

Cnidaria: 

Hydrozoa Pandeidae Stomotoka atra 


1998 P II III 

Cnidaria: 

Hydrozoa Periphyllidae Periphylla periphylla P. hyacinthina Cooney 1987 P II III IV V VI x 


1998 


1998 


1998 

Cnidaria: 

Hydrozoa Phialellidae Campanulina rugosa Mills 2000 ST II III IV Austin 1985 

Cnidaria: 

Hydrozoa Phialellidae Opercularella lacerata Mills 2000 ST II x x Austin 1985 

Cnidaria: 

Hydrozoa Polyorchidae Polyorchis penicillatus 


1998 P II III IV V VI 


1998 

Cnidaria: 

Hydrozoa Prayidae Nectopyramis diomedeae Cooney 1987 P II III IV V VI x Austin 1985 

Cnidaria: 

Hydrozoa Prayidae Praya reticulata Nectodroma Cooney 1987 P 


VII VIII Austin 1985 

Cnidaria: 

Hydrozoa Proboscidactylidae Proboscidactyla flavicirrata Mills 2000 I/ST/Epi II x x SE Alaska nr 

Cnidaria: 

Hydrozoa Protohydridae Protohydra unidentified 


Schwafel 1988 


1998 

Cnidaria: 

Hydrozoa Rathkeidae Rathkea jaschnowi Cooney:SEA P II x Bering Sea nr Naumov 1960 

Cnidaria: 

Hydrozoa Rathkeidae Rathkea octopunctata R. blumenbachii Cooney 1987 P II III IV x x 

Cnidaria: 

Hydrozoa Rathkeidae Rathkea unidentified Cooney:SEA P 

Cnidaria: 

Hydrozoa Rhopalonematidae Aglantha digitale 

Cooney and Coyle 

1988 P/ST II III IV x x 


1998 


1998 

Cnidaria: 

Hydrozoa Rhopalonematidae Pantachogon haekeli Cooney 1987 P II III IV x Austin 1985 

Cnidaria: 

Hydrozoa Sertulariidae Sertularia robusta Mills 2000 ST II x x Austin 1985 

Cnidaria: 

Hydrozoa Tubularidae Hybocodon prolifer Cooney 1987 P II III x Austin 1985 

Cnidaria: 

Hydrozoa Tubularidae Plotocnide borealis Cooney:SEA P II III IV x x Austin 1985 

Cnidaria: 

Hydrozoa Tubularidae Tubularia prolifera Cooney:SEA P II x x x Austin 1985 

Cnidaria: 

Scyphozoa Cyaneidae Cyanea capillata Mills 2000 ST II x x 

Cnidaria: 

Scyphozoa Pelagiidae Chrysaora melangaster Cooney 1987 P II III IV V VI x 

Cnidaria: 

Scyphozoa Ulmariidae Aurelia aurita Mills 2000 ST II x x 

Cnidaria: 

Scyphozoa Ulmariidae Aurelia labiata Mills 2000 ST II III IV x 

Cnidaria: 

Scyphozoa Ulmariidae Aurelia unidentified Mills 2000 P 

Cnidaria: 

Scyphozoa Ulmariidae Phacellophora camtschatica Catablema Cooney:SEA P II III IV V VI x x 

Ctenophora Beroidae Beroe unidentified Cooney 1987 P 


1998 


1998 


1998 


1998 


1998 

Ctenophora Bolinopsidae Bolinopsis infundibulum Mills 2000 ST II x x 


1998 

Ctenophora Mertensiidae Mertensia sp. (ovum) 


1988 P II x x Austin 1985 

Ctenophora Pleurobranchiidae Hormiphora cucumis 


1998 P II III x 

Ctenophora Pleurobranchiidae Pleurobrachia bachei Mills 2000 ST II III IV 

BIOREGION 


1998 


1998


Annelida 

The region’s polychaete annelid fauna is particularly rich, with 233 species in 46 

families. Kudenov (2000) has noted three range extensions, for Phyllodoce medipapillata, 

Chaetozone senticosa, and Rhynchospio glutaea. Some species of Eumida, Scoloplos, Exogone, 

Nephtys, Glycera, and the archaeannelid Polygordius are likely to be undescribed. Seven taxa, 

including members of the Spirorbidae have not been identified below the generic level. Nineteen 

new records from the UAM collection and Exxon Valdez oil spill specimens represent range 

extensions. 

Hirudinea and Oligochaeta are not included in the data sets (Table 10.3), because we lack 

reliable identifications for specimens found in Prince William Sound. At least one species of 

marine leech is present in UAM samples from the adjacent Gulf of Alaska. Kozloff (1987) lists 

14 leech species with distributions from Oregon north. Oligochaetes have been collected, but no 

effort has been made to identify them to family or lower taxon. Austin (1985) lists eight 

oligochaete species whose ranges include Alaska or southeast Alaska. 

Information on distribution, habitat, and nomenclatural changes have been drawn from 

Austin (1985), Banse and Hobson (1968), Berkeley and Berkeley (1948, 1952), Hobson and 

Banse (1981), Kudenov (2000) and Ushakov (1955). 

Table 10.3. Annelida. 

GENUS SPECIES OTHER NAMES 

SPECIMEN or 

SOURCE HABITAT NEP NWP AR NWA ORIGIN 

REFERENCE to 

NIS STATUS DISTRIBUTION 

Ampharete acutifrons EVOS ST/Inf II III IV Austin 1985 

Ampharete finmarchia A. arctica 

Feder et al. 

1979 ST/Inf II III IV x x x Austin 1985 

Amphicteis 

gunneri 

Feder and 

Matheke 1980 ST/Inf II III IV x Ushakov 1955 

Amphicteis 

scaphobranchiata 

Feder et al. 

1979 ST/Inf II III IV Austin 1985 

Anobothrus 

gracilis 

Feder et al. 

1979 ST/Inf II III IV x Austin 1985 

Lysippe 

labiata 

Feder and 

Matheke 1980 ST/Inf II III IV Austin 1985 

Melinna 

cf. M. cristata 

Feder et al. 


Melinna 

cristata 

Feder and 

Matheke 1980 ST/Inf II III IV x Austin 1985 

Melinna 

elisabethae 

Feder and 


Pseudosabellides lineata Asabellides EVOS ST/Inf II III IV x Austin 1985 

Pseudosabellides sibirica Asabellides littoralis UAM ST/Inf II III IV Austin 1985 

Sosanella unidentified UAM 

Aphrodita japonica UAM ST/Inf II III IV x Austin 1985 

Drilonereis 

falcata minor 

Feder and 

Matheke 1980 ST/Inf II III IV 

BIOREGION 

Drilonereis longa UAM ST/Inf II x x 

British 

Columbia NR Austin 1985 

British 


Abarenicola 

pacifica 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf II III IV x Austin 1985 

Barantolla americana EVOS I/ST/Inf II III IV C Austin 1985 

Capitella 

capitata 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf II x x x C Austin 1985 

Heteromastus filiformis EVOS ST/Inf II III IV V x x possible Austin 1985 

Mediomastus unidentified EVOS I/ST/Inf 

Mesochaetopterus taylori EVOS ST/Inf II III IV 

British 


Feder and 

Spiochaetopterus costarum 

Matheke 1980 ST/Inf x x x x Austin 1985



BIOREGION 


SPECIMEN or 


REFERENCE to 


Chrysopetalium occidentale Kudenov 2000 II III IV V VI x Kudenov 2000 

Chaetozone senticosa Kudenov 2000 ST/Inf II IV V California NR Kudenov 2000 

Feder and 

Chaetozone 

setosa 

Matheke 1980 ST/Inf x x x x Austin 1985 

Cirratulus cingulatus Kudenov 2000 ST/Inf II III IV Kudenov 2000 

Cirratulus cirratus EVOS ST/Inf x x x x 

British 


Tharyx 

monilaris 

Feder and 

Bryson- 

Schwafel 1988 ST/Inf II III IV Austin 1985 

Tharyx 

multifilis 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf II III IV Austin 1985 

Tharyx 

parvus 

Feder and 


Tharyx secundus EVOS ST/Inf II III IV 

British 


Cossura 

longocirrata 

Feder and 


Dorvillea rudolphi EVOS ST/Inf II III IV 

Dorvillea 

pseudorubrovittata 

Feder and 


Protodorvillea gracilis EVOS ST/Inf II III IV Austin 1985 

Schistomeringos unidentified UAM ST 

Eunice valens E. kobiensis UAM ST/Inf II III IV x Austin 1985 

Brada 

granulata 

Feder and 


Flabelligera affinis EVOS ST/Inf II III IV x Austin 1985 

Pherusa plumosa EVOS I/ST/Inf II III IV x Austin 1985 

Pherusa 

papillata 

Feder and 


Glycera sp. undescribed Kudenov 2000 Kudenov 2000 

Glycera cf. G. nana Kudenov 2000 

Glycera nana G. capitata 

Feder and 

Bryson- 

Schwafel 1988 ST/Inf II III IV x Austin 1985 

Hemipodus borealis EVOS I/ST/Inf II III IV Austin 1985 

Glycinde armigera EVOS ST/Inf II III IV x Austin 1985 

Glycinde 

Goniada 

Goniada 

picta 

annulata 

maculata 

Feder and 

Bryson- 


Feder and 

Jewett 1988 ST/Inf II III IV Austin 1985 

Feder and 


Microphthalmus sczelkowii 

Feder and 

Bryson- 

Schwafel 1988 ST/Inf II x x California NR Austin 1985 

Micropodarke dubia EVOS ST/Inf II III IV Austin 1985 

Feder and 

Podarkeopsis glabrus Gyptis brevipalpa 


Pelagobia longicirrata Cooney 1987 P II III IV V VI x x Austin 1985 

Feder et al. 


Lumbrineris 

latreilli 

1979 ST/Inf VII VIII IX x Austin 1985 

Lumbrineris limicola UAM ST/Inf II III IV SE Alaska nr 

Lumbrineris 

Lumbrineris 

luti 

similabris 

Feder and 

Jewett 1988 ST/Inf II III IV 

British 


Feder and 


Lumbrineris 

zonata 

Feder and 

Bryson- 


Ninoe 

gemmea 

Feder and 


Ninoe 

simpla 

Feder et al. 

1979 ST/Inf II III IV Austin 1985 

Magelona berkeleyi EVOS ST/Inf II III IV 

British 


Magelona hobsonae M. pitekai 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf II III IV 

British 

Columbia NR Austin 1985




SPECIMEN or 


REFERENCE to 


Metasychis disparidentata Asychis 

Feder et al. 


Nicomache personata UAM ST/Inf II III IV Austin 1985 

Notoproctus 

pacificus 

Feder and 


Petaloproctus borealis P. tenuis EVOS ST/Inf II III IV x Austin 1985 

Praxillella praetermissa EVOS ST/Inf II III IV x Austin 1985 

Praxillella 

gracilis 

Feder and 

Jewett 1988 ST/Inf II III IV Austin 1985 

Praxillella pacifica P. affinis 

Feder and 


Rhodine 

bitorquata 

Feder and 


Aglophamus 

rubella anops 

Feder and 


Nephtys cornuta cornuta UAM ST/Inf II III IV Austin 1985 

Nephtys ferruginea EVOS ST/Inf II III IV Austin 1985 

Nephtys 

punctata 

Feder and 

Jewett 1988 ST/Inf II III IV x Austin 1985 

Nephtys 

caeca 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf II III IV V x x x Austin 1985 

Nephtys ciliata EVOS I/ST/Inf II x x Austin 1985 

Nephtys 

cornuta 

Feder and 

Matheke 1980 I/ST/Inf II III IV Austin 1985 

Nephtys 

cornuta franciscana 

Feder and 


Nephtys sp. undescribed Kudenov 2000 

Chelonereis cyclurus Kudenov 2000 ST/Inf II III IV V x Austin 1985 

Nereis grubei UAM I/ST/Inf II III IV x Austin 1985 

Nereis pelagica UAM I/ST/Inf II III IV Austin 1985 

Nereis 

procera 

Feder and 

Matheke 1980 ST/Inf II III IV V VI Austin 1985 

Nereis 

vexillosa 

Feder and 

Bryson- 

Schwafel 1988 I/inf II III IV V VI Austin 1985 

Nereis 

zonata 

Feder and 

Matheke 1980 ST/Inf II III IV V VI x Austin 1985 

Platynereis bicanaliculata P. agassizi EVOS I/ST/Inf II III IV Austin 1985 

Onuphis iridescens UAM ST/Inf II III IV x Austin 1985 

Onuphis 

conchylega 

Feder and 

Matheke 1980 ST/Inf II III IV x 

BIOREGION 

Banse and Hobson 

1968 

Onuphis 

geophiliformis 

Feder and 


Onuphis 

parva 

Feder and 

Matheke 1980 ST/Inf II III IV x 


1968 

Onuphis stigmatis ST/Inf II III IV Austin 1985 

Armandia brevis EVOS ST/Inf II III IV Austin 1985 

Ophelia 

limacina 

Ophelina acuminata Amnotrypane aulogaster 

Travisia 

forbesii 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf x x x x Austin 1985 

Feder and 

Matheke 1980 ST/Inf II III IV V VI x Austin 1985 

Feder et al. 

1979 ST/Inf II x x x Austin 1985 

Leitoscoloplos 

panamensis 

Feder and 

Bryson- 


Leitoscoloplos pugettensis L. elongatus 

Feder and 


Naineris dendritica Kudenov 2000 ST/Inf II III IV V Austin 1985 

Naineris drenritica N. laevigata UAM ST/Inf II III IV Austin 1985 

Naineris uncinata EVOS ST/Inf II III IV Austin 1985 

Naineris 

unidentified 

Feder and 


Scoloplos acmeceps EVOS ST/Inf II III IV Austin 1985 

Scoloplos armiger EVOS ST/Inf II III IV x x Austin 1985 

Scoloplos sp. undescribed Kudenov 2000 

Feder and 

Myriochele oculata M. heeri 


Owenia 

fusiformis 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf II III IV x x Austin 1985 

Aricidea ramosa EVOS ST/Inf II III IV x Austin 1985



BIOREGION 


SPECIMEN or 


REFERENCE to 


Aricidea 

lopezi 

Feder and 


Aricidea 

neosuecica 

Feder and 


Aricidea nolani A. suecica UAM ST/Inf II III IV Austin 1985 

Cirrophoras lyra EVOS ST/Inf II III IV Austin 1985 

Cirrophoras branchiatus UAM ST/Inf II III IV Austin 1985 

Paraonis 

gracilis 

Feder and 

Matheke 1980 ST/Inf II III IV x x Austin 1985 

Amphictene moorei A. auricoma 

Feder and 

Matheke 1980 I/ST/Inf II III IV x Austin 1985 

Cistenides granulata C. brevicoma 

Feder and 

Bryson- 


Pectinaria californiensis EVOS ST/Inf II III IV Austin 1985 

Pholoides asperus Polyodontidae:Peisidice 

Feder and 


Anaitides groenlandica Phyllodoce UAM ST/Inf II III IV x x Austin 1985 

Anaitides 

mucosa 

Feder and 


Anaitides 

maculata 

Feder and 


Eteone 

longa 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf II III IV x C Austin 1985 

Eulalia viridis UAM ST/Inf II III IV x Austin 1985 

Eumida sp. undescribed Kudenov 2000 ST/Inf 

Hypoeulalia bilineata Eulalia UAM ST/Inf II III IV x x Austin 1985 

Mysta barbata UAM ST/Inf II x x Chukchi Sea nr Ushakov 1955 

Nereiphylla castanea Phyllodoce, Genetylus UAM ST/Inf II III IV x Austin 1985 

Notophyllum tectum UAM ST/Inf II III IV Austin 1985 

Phyllodoce medipapillata Kudenov 2000 ST/Inf II III IV V California NR Kudenov 2000 

Ancistrosyllis 

hamata 

Feder and 


Polygordius sp. undescribed UAM ST/Inf Kudenov 2000 

Feder and 

Antinoella 

sarsi 

Matheke 1980 ST/Inf II III IV x x Ushakov 1955 

Arcteobia 

spinelytris 

Feder and 


Feder and 

Byglides macrolepidus Antinoella 


Eunoe depressa UAM ST/Inf II III IV x Austin 1985 

Eunoe oerstedi EVOS ST/Inf II III IV Austin 1985 

Eunoe senta UAM ST/Inf II III IV Austin 1985 

Gattyana 

ciliata 

Feder and 


Gattyana 

cirrosa 

Feder and 

Matheke 1980 ST/Inf II III IV x x Austin 1985 

Gattyana 

treadwelli 

Feder and 


Halosydna brevisetosa UAM ST/Inf II III IV Austin 1985 

Harmothoe extenuata UAM ST/Inf II III IV x x Austin 1985 

Harmothoe imbricata EVOS I/ST/Inf II III IV x x C Austin 1985 

Hesperonoe 

complanata 

Feder and 


Lepidonotus helotypus UAM ST/Inf II III IV Ushakov 1955 

Feder and 

Lepidonotus 

squamatus 


Nemida tamarae UAM ST/Inf II Bering Sea nr Ushakov 1955 

Feder et al. 

Polynoe 

canadensis 


Idanthyrsus 

armatus 

Feder and 

Matheke 1980 ST/Inf 


VII VIII IX Austin 1985 

Idanthyrsus ornamentatus EVOS ST/Inf II III IV Austin 1985 

Chone magna EVOS ST/Inf II III IV Austin 1985 

Chone cincta UAM ST/Inf II III IV x Austin 1985 

Euchone hancocki EVOS ST/Inf II III IV Austin 1985 

Euchone longifissuirata UAM ST/Inf II III IV x x Ushakov 1955 

Euchone 

analis 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf II III IV x x Ushakov 1955 

Fabricia 

sabella 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf II x x Austin 1985



BIOREGION 


SPECIMEN or 

REFERENCE to 

SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS DISTRIBUTION 

Laonome 

kroyeri 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf II x x 

Berkeley and 

Berkeley 1952 

Myxicola 

infundibulum 

Feder and 


Potentilla ocellata UAM ST/Inf II III IV Austin 1985 

Schizobranchia insignis Kudenov 2000 II III IV V Austin 1985 

Scalibregma 

inflatum 

Feder and 


Crucigera 

irregularis 

Feder and 

Matheke 1980 ST/Epi II III IV Austin 1985 

Crucigera zygophora UAM ST/Epi II III IV x Austin 1985 

Pseudochitinopoma occidentalis EVOS ST/Epi II III IV Austin 1985 

Serpula vermicularis Kudenov 2000 ST/Epi II III IV x Kudenov 2000 

Pholoe 

sp. "minuta" 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf II III IV x x C Austin 1985 

Sphaerodoropsis minuta EVOS ST/Inf II III IV x Austin 1985 

Sphaerodoropsis sphaerulifer EVOS ST/Inf II III IV x Austin 1985 

Sphaerodorum papillifer EVOS ST/Inf II x x x Austin 1985 

Dipolydora cf. D. bidentata Kudenov 2000 Kudenov 2000 

Dipolydora cf. D. giardi Kudenov 2000 Kudenov 2000 

Dipolydora cf. D. socialis Kudenov 2000 Kudenov 2000 

Laonice cirrata EVOS ST/Inf x x x x Austin 1985 

Berkeley and 

Rhynchospio glutaea Malacoceros EVOS ST/Inf II x x nr 

Berkeley 1952 

Polydora limicola Kudenov 2000 ST/Inf II III IV V Austin 1985 

Polydora cf. P.brachycephalata UAM ST/Inf II III IV Austin 1985 

Polydora ciliata P. pygidialis 

Feder and 

Bryson- 

Schwafel 1988 I/ST/Inf x x x x Austin 1985 

Dipolydora quadrilobata EVOS I/ST/Inf II III IV C Austin 1985 


Polydora socialis Dipolydora UAM ST/Inf VII VIII IX Austin 1985 

Prionospio cirrifera EVOS ST/Inf II III IV x Austin 1985 

Prionospio steenstrupi EVOS I/ST/Inf x x x x Austin 1985 

Pygospio elegans EVOS I/ST/Inf II x x Austin 1985 

Scolelepis squamata Nerine cirratulus EVOS ST/Inf II III IV Austin 1985 

Spio filicornis EVOS ST/Inf x x x x Austin 1985 

Spiophanes berkeleyorum EVOS ST/Inf II III IV x Austin 1985 

Spiophanes bombyx EVOS ST/Inf x x x x Austin 1985 

Feder et al. 

Spiophanes cirrata S. kroyeri 

1979 ST/Inf II x x Austin 1985 

Circeis 

spirillum 

Feder and 

Bryson- 

Schwafel 1988 I/Epi II III IV Austin 1985 

Dexiospira 

unidentified 

Feder and 

Bryson- 


Spirorbis 

unidentified 

Feder and 

Matheke 1980 ST/Epi II III IV x x Austin 1985 

Sternaspis 

scutata 

Feder and 


Autolytus alexandri A. verrilli UAM ST/Inf II III IV Austin 1985 

Autolytus cornatus A. prismaticus UAM ST/Inf II x x x Austin 1985 

Eusyllis 

bloomstrandi 

Feder and 

Matheke 1980 ST/Inf II x x Austin 1985 

Exogone sp. undescribed Kudenov 2000 

Exogone cf. E. dwisula Kudenov 2000 Kudenov 2000 

Exogone 

lourei 

Feder and 

Bryson- 

Schwafel 1988 ST/Inf II III IV Austin 1985 

Exogone naidina E. gemmifera 

Feder and 

Bryson- 


Feder and 

Exogone 

verugera 


Sphaerosyllis cf. C. californiensis Kudenov 2000 Kudenov 2000



BIOREGION 


SPECIMEN or 

REFERENCE to 

SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS DISTRIBUTION 

Feder and 

Bryson- 

Schwafel 1988 ST/Inf II III IV x 


1968 

Sphaerosyllis 

erinaceous 

Feder and 

Syllis armillaris Typosyllis 

Matheke 1980 ST/Inf II III IV V VI x x x Austin 1985 

Syllis fasciata Typosyllis EVOS ST/Inf II III IV Austin 1985 

Syllis harti Typosyllis EVOS ST/Inf II III IV 

Feder and 

Syllis harti armillaris Typosyllis 

Matheke 1980 ST/Inf II III IV 

British 


Syllis harti unguicula Typosyllis UAM ST/Inf II III IV 

British 

Columbia NR 

Feder and 

Syllis heterochaeta Langerhansia cornuta Matheke 1980 ST/Inf II III IV x Austin 1985 

Syllis hyalina Typosyllis Kudenov 2000 I/ST/Inf x x x x Austin 1985 

Trypanosyllis gemmipara Kudenov 2000 II III IV V Austin 1985 

Typosyllis stuarti Kudenov 2000 II III Austin 1985 

Typosyllis alternata Kudenov 2000 I/ST/Inf II III IV x Austin 1985 

Amphitrite cirrata Kudenov 2000 II III IV V x C Austin 1985 

Artacama conifera UAM ST/Inf II III IV Austin 1985 

Artacama 

proboscidea 

Feder et al. 


Lanassa venusta EVOS ST/Inf II III IV x x Austin 1985 

Laphania boecki EVOS ST/Inf II x x Austin 1985 

Leana abranchiata UAM ST/Inf II III IV x x Austin 1985 

Feder and 

Lysilla 

loveni 


Pista 

cristata 

Feder and 


Feder and 

Pista vinogradovi P. pacifica 


Polycirrus medusa UAM ST/Inf II x x Austin 1985 

Proclea graffi UAM ST/Inf II III IV Austin 1985 

Terbellides 

stroemi 

Feder and 

Jewett 1988 ST/Inf x x x x Austin 1985 

Tomopteris septentrionalis Cooney 1987 P x x x Austin 1985 

Tomopteris pacificus Cooney 1987 P II III IV V x Austin 1985 

Tomopteris 

unidentified 

Cooney and 

Coyle 1988 P 

Trichobranchus 

glacialis 

Feder and 

Matheke 1980 ST/Inf II x x x Austin 1985 

Disoma carica Disomidae:Trichochaeta UAM ST/Inf II III IV x Austin 1985 

Disoma multisetosum Disomidae:Trichochaeta UAM ST/Inf II III IV x Austin 1985 

Typhloscolex mulleri Cooney: SEA ST II III IV x x NE Pacific NR Austin 1985 

Mollusca 

Three hundred fifteen mollusk species, representing all classes except 

monoplacophorans, are included in the data set (Table 10.4). There is one aplacophoran, and 108 

bivalves, 179 gastropods, 17 polyplacophorans, three scaphopods, and eight cephalopods. 

Mollusks in the UAM comprise one of the chief sources for the data set. Information on 

nomenclature and distribution derives form Austin (1985), Baxter (1987), Behrens (1991), Coan 

et al. (2000), and Turgeon et al. (1998). 

The shelled fauna is quite well known, but opisthobranchs, because they require special 

techniques to collect and preserve, have not been adequately documented in Alaska. It is not 

surprising to find potentially undescribed species and geographical range extensions within the 

Prince William Sound fauna. A paper describing range extensions for 11 species of 

opisthobranchs is in preparation by J. Goddard and N. Foster.


Table 10.4. Mollusca. 

BIOREGION 


SPECIMEN or 

SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS 

REFERENCE to 

DISTRIBUTION 

Chaetoderma robustum UAM ST/Inf II III IV Baxter 1987 

Pododesmus macroschisma UAM ST/Epi II III IV Coan et al. 2000 

Astarte borealis UAM ST/Inf II x x x Coan et al. 2000 

Astarte compacta A. polaris UAM ST/Inf II III IV Coan et al. 2000 

Astarte elliptica A. alaskensis UAM ST/Inf II x x Coan et al. 2000 

Astarte esquimaulti UAM ST/Inf II III x Coan et al. 2000 

Astarte ovata A. borealis UAM ST/Inf II x x x Coan et al. 2000 

Clinocardium blandum C. fucanum Coan et al. 2000 I/ST/Inf II III IV Coan et al. 2000 

Clinocardium californiense UAM I/ST/Inf II III IV x Coan et al. 2000 

Clinocardium ciliatum UAM I/ST/Inf II x x x Coan et al. 2000 

Clinocardium nuttallii UAM I/ST/Inf II III IV x Coan et al. 2000 

Serripes groenlandicus UAM I/ST/Inf II x x x Coan et al. 2000 

Serripes laperousii UAM I/ST/Inf II x x x Coan et al. 2000 

Serripes notabilis UAM ST/Inf II III IV x Coan et al. 2000 

Cyclocardia crebricosta UAM ST/Inf II III IV x Coan et al. 2000 

Cyclocardia ventricosa UAM ST/Inf II III IV Coan et al. 2000 

Miodontiscus prolongata Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000 

Cardiomya behringensis Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000 

Cardiomya pectinata UAM ST/Inf II III IV Coan et al. 2000 

Cardiomya planetica UAM ST/Inf II III IV Coan et al. 2000 

Glycymeris septentrionalis EVOS ST/Inf II III IV Coan et al. 2000 

Hiatella arctica UAM I/ST/Epi/Inf x x x x Coan et al. 2000 

Panomya ampla Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000 

Panomya norvegica P. arctica UAM ST/Inf II x x x Coan et al. 2000 

Kellia suborbicularis Kelliidae Coan et al. 2000 ST/Inf II III IV Coan et al. 2000 

Mysella planata Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000 

Neaeromya compressa Pseudopythina UAM I/ST/Inf II III IV Coan et al. 2000 

Rochefordia tumida Mysella UAM ST/Inf II III IV Coan et al. 2000 

Limatula attenuata L. subauriculata Coan et al. 2000 ST/Epi II III IV x Coan et al. 2000 

Lucina tenuisculpta Parvilucina UAM ST/Inf II III IV Coan et al. 2000 

Lucinoma annulatum UAM ST/Inf II III IV Coan et al. 2000 

Entodesma navicula E. saxicola UAM I/Epi/Inf II III IV x Coan et al. 2000 

Lyonsia bracteata UAM ST/Inf II III IV Coan et al. 2000 

Mactromeris polynyma UAM I/ST/Inf II x Coan et al. 2000 

Tresus capax Coan et al. 2000 I/ST/Inf II III IV Coan et al. 2000 

Tresus nuttallii UAM I/ST/Inf II III IV Coan et al. 2000 

Malletia pacifica M. cuneata UAM ST/Inf II III IV Coan et al. 2000 

Cryptomya californica Coan et al. 2000 I/ST/Inf II III IV x Coan et al. 2000 

Mya arenaria UAM I/Inf II III IV x N Atlantic definate Coan et al. 2000 

Mya truncata UAM I/ST/Inf II x x x Coan et al. 2000 

Crenella decussata UAM I/ST/Epi II x x x Coan et al. 2000 

Dacrydium vitreum UAM ST/Inf II x x x Coan et al. 2000 

Modiolus modiolus UAM ST/Inf II x x Coan et al. 2000 

Musculus discors UAM ST/Inf II x x Coan et al. 2000 

Musculus glacialis M. corrugatus UAM ST/Inf II x x x Coan et al. 2000 

Musculus niger UAM ST/Inf II x x x Coan et al. 2000 

Mytilus trossulus M. edulis UAM I/ST/Epi II III IV Coan et al. 2000 

Solamen columbianun Megacrenella Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000 

Vilasina seminuda Musculus Coan et al. 2000 ST/Epi II III IV x Coan et al. 2000 

Vilasina vernicosa Musculus UAM ST/Epi II III IV x Coan et al. 2000 

Nuculana conceptionis Perrisonota UAM ST/Inf II III IV Coan et al. 2000 

Nuculana minuta UAM ST/Inf II x x x Coan et al. 2000 

Nuculana navisa Coan et al. 2000 ST/Inf II III IV Coan et al. 2000 

Nuculana pernula N. fossa UAM ST/Inf II x x x Coan et al. 2000 

Eunucula tenuis Nucula UAM ST/Inf II x x Coan et al. 2000 

Crassostrea gigas UAM I/ST/Epi II III IV x NW Pacific definate Coan et al. 2000 

Pandora wardiana P. grandis UAM ST/Inf II III IV x Coan et al. 2000 

Pandora bilirata Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000 

Pandora filosa Coan et al. 2000 ST/Inf II III IV Coan et al. 2000 

Pandora glacialis UAM ST/Inf II x x x Coan et al. 2000 

Chlamys rubida UAM ST/Epi II III IV x Coan et al. 2000 

Crassdoma gigantea Hinnites Coan et al. 2000 ST/Epi II III IV Coan et al. 2000 

Delectopecten vancouverensis D. randolphi UAM ST/Epi II III IV x Coan et al. 2000 

Parvamussium alaskense UAM ST/Epi II III IV Coan et al. 2000 

Patinopecten caurinus UAM ST/Epi II III IV Coan et al. 2000 

Periploma aleuticum UAM ST/Inf II x x x Coan et al. 2000 

Siliqua alta Solenidae: S. sloati UAM I/ST/Inf II III IV x Coan et al. 2000 

Siliqua patula Solenidae Coan et al. 2000 I/Inf II III IV Coan et al. 2000 

Philibrya setosa Coan et al. 2000 ST/Inf II III IV Coan et al. 2000 

Penitella penita Coan et al. 2000 I/ST/Inf II III IV Coan et al. 2000



BIOREGION 


SPECIMEN or 


REFERENCE to 

DISTRIBUTION 

Gari californica UAM I/Inf II III IV x Coan et al. 2000 

Macoma dexioptera UAM ST/Inf II Coan et al. 2000 

Macoma elimata UAM ST/Inf II III IV Coan et al. 2000 

Macoma moesta UAM ST/Inf II III IV x x Coan et al. 2000 

Macoma balthica UAM I/Inf x x x x C Coan et al. 2000 

Macoma brota UAM ST/Inf II III IV Coan et al. 2000 

Macoma calcarea UAM ST/Inf II x x x Coan et al. 2000 

Macoma carlottensis UAM ST/Inf II III IV Coan et al. 2000 

Macoma expansa UAM I/ST/Inf II III IV Coan et al. 2000 

Macoma golikovi M. obliqua UAM I/ST/Inf II III IV Coan et al. 2000 

Macoma inquinata UAM I/ST/Inf II III IV Coan et al. 2000 

Macoma lipara Coan et al. 2000 ST/Inf II III IV Coan et al. 2000 

Macoma nasuta UAM I/ST/Inf II III IV Coan et al. 2000 

Tellina modesta UAM ST/Inf II III IV Coan et al. 2000 

Bankia setacea UAM ST/Inf II III IV x Coan et al. 2000 

Thracia challisiana Coan et al. 2000 ST/Inf II III IV Coan et al. 2000 

Thracia condoni Coan et al. 2000 ST/Inf II III IV Coan et al. 2000 

Thracia myopsis UAM ST/Inf II x x x Coan et al. 2000 

Thracia trapeziodes UAM ST/Inf II III IV Coan et al. 2000 

Adontorhina cyclia UAM I/ST/Inf II III IV x Coan et al. 2000 

Axinopsida serricata UAM I/ST/Inf II x x x Coan et al. 2000 

Conchocoele bisecta Thyasira Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000 

Mendicula ferruginosa Odontogena borealis UAM ST/Inf II x x x Coan et al. 2000 

Thyasira flexuosa UAM ST/Inf II x x x Coan et al. 2000 

Turtonia minuta UAM I/Inf II x Coan et al. 2000 

Diplodonta impolita UAM I/Inf II III IV Coan et al. 2000 

Compsomyax subdiaphana UAM ST/Inf II III IV Coan et al. 2000 

Humilaria kennerleyi UAM I/ST/Inf II III IV Coan et al. 2000 

Liocyma fluctuosa UAM ST/Inf II x x x Coan et al. 2000 

Nutricola lordi Psephidia ovalis UAM I/Inf II III IV Coan et al. 2000 

Nutricola tantilla Transenella UAM I/ST/Inf II III IV Coan et al. 2000 

Protothaca staminea UAM I/ST/Inf II III IV x Coan et al. 2000 

Saxidomus gigantea UAM I/Inf II III IV Coan et al. 2000 

Yoldia hyperborea Y. amygdalea UAM ST/Inf II x x x Coan et al. 2000 

Yoldia 

montereyensis 

Megayoldia martyria/ 

M. secunda/ M. 

beringiana UAM ST/Inf II III IV Coan et al. 2000 

Yoldia myalis UAM ST/Inf II x Coan et al. 2000 

Yoldia seminuda Y. scissurata UAM ST/Inf II III IV x Coan et al. 2000 

Yoldia thraciaeformis Megayoldia UAM ST/Inf II x x x Coan et al. 2000 

Gonatus 

unidentified 


1988 P 

Benthoctopus leioderma 

Scheel- possible 

for PWS ST/Epi II III IV Austin 1985 

Octopus dofleini Enteroctopus 

Scheel et al. 

1997 ST/Epi II III IV Austin 1985 

Octopus 

rubescens 

Scheelobservation 

ST/Epi II III IV Austin 1985 

Rossia pacifica UAM ST/Epi II III IV x Austin 1985 

II III IV V 

Chiroteuthis veranayi C. calyx Cooney 1987 P 

VI Austin 1985 

Gliteuthis armata G. phyllura Cooney 1987 P 

II III IV V 

VI x Austin 1985 

Acanthodoris 

hudsoni 

Lee and Foster 

1985 I/ST/Epi II III IV 


1985 


1985 

Acanthodoris nanaimoenisis Goddard 2000 I/ST/Epi II III IV 


Acanthodoris pilosa 

1985 I/ST/Epi II x x Behrens 1991 

Acmaea mitra UAM I/Epi II III IV Austin 1985 

Aeolidia 

papillosa 


1985 I/ST/Epi x x x x Behrens 1991 


Aglaja ocelligera Melanochlamys Goddard 2000 I/ST/Epi II III IV 

1985 

Melanochlamys diomedea Aglaja 


Schwafel 1988 I/ST/Epi II III IV SE Alaska nr 


1985



BIOREGION 


SPECIMEN or 


REFERENCE to 

DISTRIBUTION 

Archidoris montereyensis I/ST/Epi II III IV 


1985 

Archidoris 

Armina 

odhneri 

californica 


1985 I/ST/Epi II III IV 


1985 ST/Epi II III IV 

Haminoea vesicula Foster 2000 I/ST/Epi II III IV 


1985 


1985 


1985 


1985 

Haminoea virescens Baxter 1987 I/ST/Epi II III IV 

Barleeia subtenuis UAM I/Inf II III IV Baxter 1987 

Barleeia haliotiphila Baxter 1987 I/Inf II III IV Baxter 1987 

Pseudodiala acuta Barleeia UAM I/Inf II III IV 

British 

Columbia NR Baxter 1987 

Ancistrolepis eucosmuis UAM ST/Epi II III IV Baxter 1987 

Beringius kennicotti UAM ST/Epi II III IV Austin 1985 

Buccinum baerii UAM I/Epi II III IV Austin 1985 

Buccinum plectrum UAM ST/Epi II III IV x x Austin 1985 

Colus aphelus C. hypolispus UAM ST/Epi II III IV x x 

Turgeon et al. 

1998 

Colus halli UAM ST/Epi II III IV Austin 1985 

Lirabuccinum dira Searlesia UAM I/ST/Epi II III IV Baxter 1987 

Lussivolutopsius filosa Volutopsius UAM I/ST/Epi II III IV Baxter 1987 

Neptunea lyrata UAM ST/Epi II III IV x x Austin 1985 

Neptunea pribiloffensis UAM ST/Epi II III IV x Austin 1985 

Pyrulofusus harpa Volutopsius UAM I/ST/Epi II III IV Austin 1985 

Volutopsius undescribed sp. Baxter 1987 I/ST/Epi II III IV 


1998 

Caecum crebricinctum Micranellum UAM I/ST/Inf II III IV Baxter 1987 

Caecum occidentale Fartulum UAM I/Inf II III IV Baxter 1987 

Calliostoma ligatum Trochidae UAM I/ST/Epi II III IV Baxter 1987 

Calyptraea fastigiata UAM I/ST/Epi II III IV Baxter 1987 

Crepidula perforans Baxter 1987 I/ST/Epi II III IV Austin 1985 

Crepipatella lingulata 

Calyptraea fastigiata/ 

Crepiptella dorsata UAM I/ST/Epi II III IV Baxter 1987 

Admete viridula A. couthouyi UAM ST/Inf II x x x Baxter 1987 

Neadmete modesta UAM ST/Inf II III IV Baxter 1987 

Trichotropis borealis Trichotropidae UAM I/ST/Epi II x Baxter 1987 

Trichotropis cancellata Trichotropidae UAM I/ST/Epi II III IV Baxter 1987 

Trichotropis insignis Trichotropidae UAM I/ST/Epi II III IV x Baxter 1987 

Stylidium eschrichti Bittium Baxter 1987 I/Inf II III IV Austin 1985 

II III IV V 

Cadlina luteomarginata Rosenthal 1977 ST/Epi VI SE Alaska nr Behrens 1991 

Cooney and 


Clione 

limacina 

Coyle 1988 P x x x x 

1998 

Cocculina baxteri UAM ST/Epi II III Baxter 1987 

Alia carinata Mitrella I/ST/Epi II III IV Baxter 1987 

Amphissa columbiana UAM I/ST/Inf II III IV Baxter 1987 

Amphissa reticulata UAM ST/Epi II III IV Baxter 1987 

Astrys gauspata Alia UAM I/ST/Inf II III IV Baxter 1987 

Grantotoma incisula Turridae: Oenopota UAM ST/Inf II x Baxter 1987 

Kurtziella plumbea Turridae EVOS I/ST/Epi II III IV Austin 1985 

Oenopota bicarinata Turridae UAM ST/Inf II x Austin 1985 

Oenopota excurvata Turridae UAM ST/Inf II III IV Austin 1985 

Oenopota krausei Turridae UAM ST/Inf II III IV Austin 1985 

Oenopota rugulata Turridae UAM ST/Inf II x Austin 1985 

Oenopota turricula Turridae UAM ST/Inf II x Austin 1985 

Propebela arctica 

Turridae: Oenopota 

viridula UAM ST/Inf II x Baxter 1987 

Pseudotaranis strongi Turridae: Tarais EVOS ST/Epi II III IV Austin 1985 

Doridella steinbergi Suhinia EVOS ST/Epi II III IV Behrens 1991 

Flabellina trophina F. fusca 




1985 ST/Epi II x x x 


1985 


1985 

Flabellina salmonacea 

Euclio pyamidata Clio Cooney 1987 P x x x x Austin 1985 

Granulina margaritula Marginellidae UAM I/ST/Epi II III IV Baxter 1987




Dendronotus 

Dendronotus 

Dendronotus 

Diaphana 

albus 

frondosus 

iris 

minuta 

BIOREGION 

SPECIMEN or 


REFERENCE to 

DISTRIBUTION 


1985 ST/Epi II III IV Behrens 1991 


1985 ST/Epi II x Behrens 1991 




1985 ST/Inf II x 

Diaphana brunnea UAM II III IV 


1998 


1985 

Dirona 

albolineata 



Dirona 

aurantia 



Anisodoris lentigenosa 



Anisodoris nobilis 



Dialula 

sandiegensis 



Geitodoris heathi Goddard 2000 ST/Epi II III IV 

British 

Columbia NR Behrens 1991 

Eubranchus olivaceous Goddard 2000 ST/Epi II III IV SE Alaska nr Behrens 1991 

Melanella columbiana Vitreolina UAM II III IV Baxter 1987 

Melanella micrans Balcis UAM ST/Epi II III IV Baxter 1987 

Hermissenda crassicornis Goddard 2000 I/ST/Epi II III IV Behrens 1991 

Diadora aspera Baxter 1987 I/ST/Epi II III IV Austin 1985 

Puncturella cooperi UAM ST/Epi II III IV Austin 1985 

Puncturella galeata UAM ST/Epi II III IV Austin 1985 

Puncturella noachina UAM ST/Epi II III IV x x Austin 1985 


Gastropteron pacificum 

Schwafel 1988 I/Epi II III IV Behrens 1991 

Ancula pacifica Goddard 2000 ST/Epi II III IV SE Alaska nr Behrens 1991 

Alderia modesta Goddard 2000 ST/Epi II x 

British 


Cryptobranchia alba UAM I/ST/Epi II III IV x Baxter 1987 

Cryptobranchia concentrica UAM I/ST/Epi II III IV x Baxter 1987 

Lepeta caeca UAM ST/Epi II III IV x Baxter 1987 

Limacina 

helicina 

Cooney and 

Coyle 1988 P x x x x 


1998 

Lacuna carinata Lacunidae UAM I/ST/Epi II III IV Austin 1985 

Lacuna marmorata Lacunidae Baxter 1987 I/ST/Epi II III IV Baxter 1987 

Lacuna variegata Lacunidae Baxter 1987 I/ST/Epi II III IV Baxter 1987 

Lacuna vincta Lacunidae UAM I/ST/Epi II x Austin 1985 

Littorina scutulata UAM I/Epi II III IV Austin 1985 

Littorina sitkana UAM I/Epi II III IV Austin 1985 

Lottia borealis UAM I/Epi II III IV Austin 1985 

Lottia instabilis Baxter 1987 I/ST/Epi II III IV Austin 1985 

Lottia ochracea UAM I/ST/Epi II III IV Baxter 1987 

Lottia pelta UAM I/ST/Epi II III IV Austin 1985 

Lottia triangularis Baxter 1987 I/ST/Epi II III IV Austin 1985 

Lottia digitalis UAM I/Epi II III IV Baxter 1987 

Tectura fenestrata UAM I/Epi II III IV Austin 1985 

Tectura persona UAM I/Epi II III IV Austin 1985 

Tectura scutum UAM I/ST/Epi II III IV Austin 1985 

Tectura testudinalis Baxter 1987 I/ST/Epi II x Austin 1985 

Boreotrophon clathratus UAM I/ST/Epi II x x x Baxter 1987 

Boreotrophon multicostata UAM I/ST/Epi II III IV 


1998 

Boreotrophon truncatus B. pacificus/ B. beringi UAM I/ST/Epi II x x x 


1998 

Nucella canaliculata UAM I/Epi II III IV Baxter 1987 

Nucella lamellosa UAM I/Epi II III IV Austin 1985 

Nucella lima UAM I/Epi II III IV x Baxter 1987 

Ocenebrina interfossa Ocenebra UAM I/Epi II III IV Baxter 1987 

Ocenebrina lurida Ocenebra UAM I/Epi II III IV Austin 1985 

Scabrotrophon maltzan Trophonopsis lasius UAM I/Epi II III IV Baxter 1987



BIOREGION 


SPECIMEN or 


REFERENCE to 

DISTRIBUTION 

Nassarius mendicus UAM I/ST/Inf II III IV Austin 1985 

Cryptonatica affinis 

Natica russa/ Natica 

aleutica/ Natica clausa UAM I/ST/Inf II III IV x x x Austin 1985 

Euspira pallida Polinices pallidus UAM I/ST/Inf II x x x Austin 1985 

Olea hansinensis Goddard 2000 ST/Epi II III IV 

British 


Olivella baetica UAM I/ST/Inf II III IV Baxter 1987 

Onchidella 

borealis 


1985 I/Epi II III IV 


1985 

British 



1985 

Adalaria jannae Goddard 2000 ST/Epi II III IV 


Adalaria 

proxima 

1985 ST/Epi II x x 

Adalaria sp. 1 of Behrens Goddard 2000 ST/Epi II III IV SE Alaska nr Behrens 1991 

Onchidoris 

bilamellata 


1985 ST/Epi II x x 


1985 


1985 

Onchidoris muricata Goddard 2000 ST/Epi II x x 

Policera zosterae Goddard 2000 ST/Epi II III IV SE Alaska nr Behrens 1991 



Triopha catalinae Triophidae 

1985 ST/Epi II III IV x 

1985 

Cyclostremella concordia Cyclostremellidae UAM II III IV Baxter 1987 

Odostomia angularis UAM I/ST/Inf II III IV Austin 1985 

Odostomia arctica UAM I/ST/Inf II x x Arctic nr 


1998 

Odostomia iliuliukensis UAM I/ST/Inf II III IV Austin 1985 

Odostomia cassandra 

Feder and 


Odostomia pesa Baxter 1987 I/ST/Inf II III IV Baxter 1987 

Turbonilla canadensis Baxter 1987 I/ST/Inf II III IV Austin 1985 

Turbonilla eyerdami Baxter 1987 I/ST/Inf II III IV Austin 1985 

Turbonilla lordii Baxter 1987 I/ST/Inf II III IV Austin 1985 

Turbonilla lyalli Baxter 1987 I/ST/Inf II III IV Austin 1985 

Turbonilla middendorffi Baxter 1987 I/ST/Inf II III IV Austin 1985 

Turbonilla rinella Baxter 1987 I/ST/Inf II III IV Austin 1985 

Fusitriton oregonensis Cyamatiidae UAM I/ST/Epi II III IV Baxter 1987 

Retusa obtusa R. perteuis/ R. semen UAM I/ST/Inf II x Bering Sea nr 


1985 

"Rissoella" alaskensis 

Feder and 

Bryson-Schwafel 

1988 I/ST/Inf II III IV Austin 1985 

Alvania compacta UAM I/ST/Inf II III IV Baxter 1987 

Alvania moerchi Baxter 1987 I/ST/Inf II x x x Baxter 1987 

Alvania rosana UAM I/ST/Inf II III IV Baxter 1987 


Boreocingula martyni Cingula katherenae Schwafel 1988 I/ST/Inf II III IV Austin 1985 

Falsicingula aleutica Falsicingulidae UAM I/ST/Inf II III IV Baxter 1987 

Onoba dalli Alvania Baxter 1987 I/ST/Inf II III IV Baxter 1987 

Onoba cerinella Baxter 1987 I/ST/Inf II III IV Austin 1985 

Onoba kyskensis Baxter 1987 I/ST/Inf II III IV Austin 1985 

Onoba montereyensis Alvania carpenteri Baxter 1987 I/ST/Inf II III IV Austin 1985 

Acteocina culcitella Cylichnella EVOS I/ST/Inf II III IV Austin 1985 

Acteocina harpa A. oldroydi 


1985 I/ST/Inf II III IV Behrens 1991 

Cylichna attonsa UAM I/ST/Inf II III IV Austin 1985 


Cylichna occulta UAM I/ST/Inf II x 

1985 

Cylichna alba UAM I/ST/Inf II x x x Austin 1985 

Siphonaria thirsites UAM I/Epi II III IV Baxter 1987 

Hermaea vancouverensis Hermaeidae Foster 1987 ST/Epi II III IV Behrens 1991 

Cuthona albocrusta Goddard 2000 ST/Epi II III IV Washington NR Goddard 2000 

Cuthona pustulata Goddard 2000 ST/Epi II III IV 

British 

Columbia NR Goddard 2000 

Melibe 

leonina 


1985 P II III IV 


1985




Tritonia 

Tritonia 

diomedea 

festiva 

BIOREGION 

SPECIMEN or 


REFERENCE to 

DISTRIBUTION 








1985 


1985 


1985 

Trochuina tetraquetra 

Cidarina cidaris UAM ST/Epi II III IV Baxter 1987 

Lirularia succincta UAM I/Inf II III IV Austin 1985 

Margarites beringensis UAM I/Inf II III IV Baxter 1987 

Margarites helicinus UAM ST/Inf II x x x Austin 1985 

Margarites pupillus UAM ST/Inf II III IV Austin 1985 

Solariella varicosa UAM ST/Inf II x Baxter 1987 

Solariella obscura UAM ST/Inf II x Austin 1985 

Spiromoellaria kachemakens UAM I/ST/Epi II III IV Baxter 1987 

Aforia circinata UAM ST/Inf II III IV Baxter 1987 

Tachyrhynchus erosus Baxter 1987 ST/Inf II x Austin 1985 

Tachyrhynchus reticulatus Baxter 1987 ST/Inf II III IV Austin 1985 

Velutina plicatilis Baxter 1987 ST/Epi II x Austin 1985 

Velutina prolongata Baxter 1987 ST/Epi II III IV Austin 1985 

Velutina rubra Foster 2000 I/Epi II III IV Austin 1985 

Velutina undata Baxter 1987 ST/Epi II x x x Austin 1985 

Volumitra alaskana UAM ST/Epi II III IV Baxter 1987 

Arctomelon stearnsii UAM ST/Epi II III IV Austin 1985 

Janolus fuscus Goddard 2000 ST/Epi II III IV SE Alaska nr Goddard 2000 


Clio pyramidata Cooney 1987 P x x x 

1998 

Cryptochiton stelleri UAM I/ST/Epi II III IV Baxter 1987 

Lepidozona interstincta Ischnochiton UAM ST/Epi II III IV Baxter 1987 

Stenonemus albus Ischnochiton UAM ST/Epi II x Baxter 1987 

Leptochiton rugatus UAM I/ST/Epi II III IV x Austin 1985 

Katherina tunicata UAM I/Epi II III IV Baxter 1987 

Mopalia laevior UAM I/ST/Epi II III IV Austin 1985 

Mopalia lignosa UAM I/ST/Epi II III IV Baxter 1987 

Mopalia ciliata UAM I/ST/Epi II III IV Baxter 1987 

Mopalia cirrata UAM I/ST/Epi II III IV Baxter 1987 

Mopalia mucosa UAM I/ST/Epi II III IV Baxter 1987 

Mopalia spectabilis Baxter 1987 I/ST/Epi II III IV Baxter 1987 

Mopalia swanii UAM I/ST/Epi II III IV Austin 1985 

Placiphorella rufa UAM I/ST/Epi II III IV Austin 1985 

Schizoplax brandtii UAM I/Epi II III IV x Austin 1985 

Tonicella insignis Ischnochitonidae Foster 2000 I/ST/Epi II III IV Baxter 1987 

Tonicella lineata Ischnochitonidae UAM I/ST/Epi II III IV Baxter 1987 

Tonicella rubra Ischnochitonidae UAM I/ST/Epi II x Baxter 1987 

Rhabdus rectius Dentalium dalli UAM ST/Inf II III IV Baxter 1987 

Pulsellum salishorum UAM ST/Inf II III IV Baxter 1987 

Polyschides tolmei Cadulus stearnsi UAM ST/Inf II III IV Baxter 1987


Arthropoda 

Chelicerata and Pycnogonida are relatively scarce in Prince William Sound compared to 

Crustacea. Seven of the ten pycnogonid species are included in the data set based on UAM 

specimens, One Acarina and one pseudoscorpion are frequently observed in the high intertidal 

zone of southeastern and south-central Alaska. 

Crustacea make up about 28% of the animal species listed in the data set (Table 10.5). 

We list 172 copepods, (including planktonic species from the Gulf of Alaska which may be 

present occasionally in Prince William Sound), 105 decapods, 105 amphipods, 21 cumacean, 19 

isopods, and smaller numbers of others. 

Several taxa (e.g. Ostracoda) are probably very much under-counted. Six copepods, one 

amphipod, and two ostracods are identified to the family level, and no further. A large number 

of small crustacea, mostly harpacticoid copepods and amphipods have not been identified to 

species. Two species, the amphipod Jassa sp./marmorata and the copepod Leimia vaga are 

possible NIS, but this number probably represents an underestimate. Cordell (2000) pointed out 

that “Harpacticoid copepods may be particularly likely to be transported and introduced because 

as a group they have successfully occupied almost all benthic and epibenthic habitats. … The 

paucity of studies of harpacticoid taxonomy in the northeastern Pacific makes it nearly 

impossible to determine whether or not a given species has been introduced without extensive 

distribution or genetic studies.” We have not included an inventory of parasitic cirripedia or 

isopods, but several species of these were recorded by Hines et al. (2000). 

Sources for information of Crustacea include Barnard 1969 (amphipods), Butler 1980 

(shrimp), Hart 1982 (crabs), and Schultz 1969 and Squires and Figueria 1974 (isopods). J. 

Chapman (2000) contributed to the information on Pericarida, and J. Cordell (2000) to our 

knowledge of the harpacticoid Copepoda. Records for the planktonic crustacea derive from 

unpublished data compiled from T. Cooney’s Sound Ecosystem Analysis project of 1994-1998. 

We found eleven new records for the occurrence of Crustacea. 

The third major group of Arthropoda, marine and brackish water insects, have not been 

surveyed in any detail in the area. For insects, Austin (1985) lists eight Coleoptera, and 12 

Diptera with ranges that include Alaska.


Table 10.5. Arthropoda. 

BIOREGION 

HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES 

SPECIMEN or 


REFERENCE to 


Arachnida: Acarina Nemolgus littoralis 

Foster/Hobergobservation 

I/Epi 

Arachnida: 

Pseudoscorpionida unidentified unidentified UAM I/epi 

Crustacea: Amphipoda Acanthonotozomatidae Odius carinatus EVOS ST/Epi II x Austin 1985 

Crustacea: Amphipoda Ampeliscidae Ampelisca birulai 

Feder and 

Matheke 1980 I/ST/Inf II III IV x x 

Coyle and Highsmith 

1994 

Crustacea: Amphipoda Ampeliscidae Ampelisca brevisimulata EVOS ST II III IV Austin 1985 

Crustacea: Amphipoda Ampeliscidae Ampelisca caryei EVOS ST II III IV Austin 1985 

Crustacea: Amphipoda Ampeliscidae Ampelisca pugetica EVOS ST II III IV Austin 1985 

Crustacea: Amphipoda Ampeliscidae Ampelisca eschrichti 

Feder and 


Feder and 

Crustacea: Amphipoda Ampeliscidae Ampelisca macrocephala A. careyi 


Feder and 

Crustacea: Amphipoda Ampeliscidae Haploops tubicola 


Crustacea: Amphipoda Amphitoidae Ampithoe kussakini Chapman 2000 I/Inf II III IV x Chapman 2000 

Crustacea: Amphipoda Amphitoidae Ampithoe sectimanus Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Amphitoidae Ampithoe simulans 


Schwafel 1988 I/ST x x x x Austin 1985 

Crustacea: Amphipoda Amphitoidae Peramphithoe eoa Ampithoe mea Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Amphitoidae Peramphithoe humeralis Ampithoe Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Anisogammaridae Anisogammarus pugettensis EVOS ST/Epi II III IV x Chapman 2000 

Crustacea: Amphipoda Anisogammaridae Eogammarus confervicolus Anisogammarus Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Anisogammaridae Eogammarus oclairi Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Anisogammaridae Eogammarus unidentified Chapman 2000 I/ST 

Crustacea: Amphipoda Anisogammaridae Locustogammarus locustoides Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Anisogammaridae Spinulogammarus subcarinatus Chapman 2000 I II III IV Chapman 2000 

Crustacea: Amphipoda Aoridae Aoroides columbiae EVOS ST II III IV x Austin 1985 

Crustacea: Amphipoda Atylidae Atylus collingi EVOS ST II III IV x Austin 1985 

Crustacea: Amphipoda Calliopiidae Calliopiella unidentified Paracalliopiella Chapman 2000 I/Inf




BIOREGION 

SPECIMEN or 


REFERENCE to 


Crustacea: Amphipoda Calliopiidae Calliopius behringi Cooney 1987 I/ST II III x x x Gur’yanova 1951, 1962 

Crustacea: Amphipoda Calliopiidae Calliopius carinatus Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Calliopiidae Calliopius laeviuscula Cooney 1987 P III IV V VI x x Austin 1985 

Crustacea: Amphipoda Caprellidae Caprella drepanochir Chapman 2000 I/Inf II III IV x C Chapman 2000 

Crustacea: Amphipoda Caprellidae Caprella laeviuscula Chapman 2000 I/Inf II III IV x Chapman 2000 

Crustacea: Amphipoda Caprellidae Caprella unidentified Chapman 2000 I/Inf 

Crustacea: Amphipoda Caprellidae Metacaprella kennerlyi Chapman 2000 I/Inf II III IV Chapman 2000 

Feder and 

Crustacea: Amphipoda Corophidae Neohela unidentified 

Matheke 1980 ST/Inf 

Crustacea: Amphipoda Corophidae Americorophium brevis Corophium Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Corophidae Americorophium salmonis Corophium Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Corophidae Americorophium spinicore Corophium Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Corophidae Corophium unidentified EVOS ST 

Crustacea: Amphipoda Corophidae Monocorophium carlottensis Chapman 2000 I/Inf II III IV C Chapman 2000 

Crustacea: Amphipoda Dexamidae Guernea unidentified EVOS ST 

Crustacea: Amphipoda Eusiridae Eusiriella multicalceola Gracilipes Cooney 1987 P II III Austin 1985 

Crustacea: Amphipoda Eusiridae Pontogeneia unidentified EVOS ST 

Crustacea: Amphipoda Eusiridae Rhachotropis natator Cooney 1987 P II III IV V x Austin 1985 

Crustacea: Amphipoda Gammaridae Lagunogammarus setosus Chapman 2000 I/Inf II III IV x Chapman 2000 

Feder and 

Crustacea: Amphipoda Haustoriidae Ericthonius hunteri E. rubricornis 

Matheke 1980 I/ST/Inf II x x Austin 1985 

Crustacea: Amphipoda Hyalellidae Najna unidentified Najnidae EVOS I/St 

Crustacea: Amphipoda Hyalidae Allorchestes angusta Chapman 2000 I/Inf II III IV x Chapman 2000 

Crustacea: Amphipoda Hyalidae Hyale frequena Chapman 2000 I/Inf II III IV Chapman 2000




BIOREGION 

SPECIMEN or 


REFERENCE to 


Crustacea: Amphipoda Hyalidae Hyale plumulosa Chapman 2000 I/Inf II x x C Chapman 2000 

Crustacea: Amphipoda Hyperiidae Euprimno macropa Primno abyssalis Cooney: SEA P 

II III IV V 

VI VII VIII 

IX x x Austin 1985 

Crustacea: Amphipoda Hyperiidae Euprimno unidentified Primno Cooney 1987 P 

Crustacea: Amphipoda Hyperiidae Hyperia medusarum hystryx Cooney 1987 P 

Crustacea: Amphipoda Hyperiidae Hyperia unidentified Cooney 1987 P 

Crustacea: Amphipoda Hyperiidae Hyperoche medusarum Cooney 1987 P 

II III IV V 


II III IV V 


Crustacea: Amphipoda Hyperiidae Hyperoche unidentified Cooney 1987 P 

Crustacea: Amphipoda Hyperiidae Parathemisto gracilipes Thermisto Cooney 1987 P II III x x Austin 1985 

Crustacea: Amphipoda Hyperiidae Parathemisto libellula Thermisto Cooney 1987 P II III IV V x Austin 1985 

Crustacea: Amphipoda Hyperiidae Parathemisto pacifica Thermisto Cooney 1987 P II III IV x Austin 1985 

Crustacea: Amphipoda Isaeidae Photis unidentified EVOS ST/Inf 

Crustacea: Amphipoda Ischyroceridae Ischyrocerus unidentified EVOS I/ST/Inf 

Crustacea: Amphipoda Ischyroceridae Jassa sp./J. marmorata Chapman 2000 I/Inf II possible Austin 1985 

Crustacea: Amphipoda Ischyroceridae Jassa staudel Chapman 2000 I/Inf II III IV C Chapman 2000 

Feder and 

Crustacea: Amphipoda Ischyroceridae Protomedeia unidentified 

Matheke 1980 I/ST/Inf 

Crustacea: Amphipoda Lanceolidae Lanceola pacifica Cooney 1987 P II III x x Gur’yanova 1951, 1962 

Crustacea: Amphipoda Lycaeidae Tryphana malmii Cooney 1987 P II III IV V Austin 1985 

Crustacea: Amphipoda Lysianassidae Andaniexis subabyssis Cooney 1987 P II III IV x Gur’yanova 1951, 1962 

Crustacea: Amphipoda Lysianassidae Anonyx unidentified 

Feder and 


Feder and 

Crustacea: Amphipoda Lysianassidae Crybelocephalus unidentified 

Matheke 1980 P x x x x 

Crustacea: Amphipoda Lysianassidae Cyphocaris anonyx C. micronyx Cooney 1987 P 

II III IV V 

VI VII VIII x Austin 1985




Crustacea: Amphipoda Lysianassidae Cyphocaris challengeri 

BIOREGION 

SPECIMEN or 


REFERENCE to 



1988 S/P x x x x Austin 1985 

Crustacea: Amphipoda Lysianassidae Hippomedon kurilicus H. abyssi UAM ST II III IV x Gur’yanova 1951, 1962 

Crustacea: Amphipoda Lysianassidae Koroga megalops Cooney 1987 P II III x x x Austin 1985 

Crustacea: Amphipoda Lysianassidae Lepidepecerium unidentified EVOS ST 

Crustacea: Amphipoda Lysianassidae Onismius unidentified EVOS ST 

Crustacea: Amphipoda Lysianassidae Orchomene obtusa Cooney: SEA ST 

II III IV V 

VI VII Gur’yanova 1951, 1962 

Crustacea: Amphipoda Lysianassidae Orchomene pacifica EVOS ST II III IV x Austin 1985 

Crustacea: Amphipoda Lysianassidae Orchomene unidentified EVOS ST 

Crustacea: Amphipoda Lysianassidae Paracallisoma alberti Cooney 1987 P II III x Gur’yanova 1951, 1962 

Feder and 

Crustacea: Amphipoda Melitidae Maera loveni M.danae 


Crustacea: Amphipoda Melitidae Melita dentata EVOS ST II III IV x Austin 1985 

Crustacea: Amphipoda Melphidippidae unidentified unidentified Cooney: SEA ST II III x NE Pac NR Austin 1985 

Feder and 

Crustacea: Amphipoda Oedicerotidae Aceroides latipes 


Crustacea: Amphipoda Oedicerotidae Bathymedon unidentified EVOS ST/Inf 

Feder and 

Crustacea: Amphipoda Oedicerotidae Monoculodes diamesus 

Matheke 1980 I/ST/Inf II III IV Gur’yanova 1951, 1962 

Crustacea: Amphipoda Oedicerotidae Synchelidium rectipalmatum EVOS ST/Inf II III IV Austin 1985 

Crustacea: Amphipoda Oedicerotidae Westwoodillia rectangulata W. caecula Cooney 1987 P II x Gur’yanova 1951, 1962 

Crustacea: Amphipoda Oxycephalidae Streesia unidentified Cooney 1987 P 

Crustacea: Amphipoda Paraphronimidae Paraphronima crassipes Cooney 1987 P 

II III IV V 

VI VII x Austin 1985 

Feder and 

Crustacea: Amphipoda Pardaliscidae Nicippe tumida 


Feder and 

Crustacea: Amphipoda Phoxocephalidae Harpiniopsis sanpedroensis Harpinia 


Crustacea: Amphipoda Phoxocephalidae Paraphoxus homilis EVOS ST/Inf II III IV Austin 1985 

Crustacea: Amphipoda Phoxocephalidae Paraphoxus similis EVOS ST/Inf II III IV Austin 1985 

II III IV V 

Crustacea: Amphipoda Phronimidae Phronima sedentaria Cooney 1987 P 

VI x Austin 1985



BIOREGION 


SPECIMEN or 


REFERENCE to 


II III IV V 

Crustacea: Amphipoda Phrosinidae Primno macropa P. abyssalis Cooney 1987 P 

VI x x Austin 1985 

Crustacea: Amphipoda Pleustidae Pleustes cataphractus EVOS I/Inf x x x Austin 1985 

Crustacea: Amphipoda Pontogeneiidae Megamorea subiener Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Pontogeneiidae Paramoera bousfieldi Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Pontogeneiidae Paramoera columbiana 


Schwafel 1988 I/Inf II III IV Austin 1985 

Crustacea: Amphipoda Pontogeneiidae Paramoera mohri Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Pontogeneiidae Pontogeneia ivanovae P. ivanovi Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Pontogeneiidae Pontogeneia rostrata Chapman 2000 I/Inf II III IV C Chapman 2000 

Crustacea: Amphipoda Pontoporeiidae Pontoporeia femorata Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Amphipoda Proscinidae Proscina birsteini Cooney 1987 P II x Gur’yanova 1951, 1962 

II III IV V 

Crustacea: Amphipoda Scinidae Scina borealis UAM I/ST/Inf VI x x Austin 1985 

Crustacea: Amphipoda Scinidae Scina rattayi Cooney 1987 P II III IV x Austin 1985 

Crustacea: Amphipoda Scinidae Scina unidentified Cooney: SEA P 

Crustacea: Amphipoda Scinidae Scina stebbingi Cooney 1987 P II x Gur’yanova 1951, 1962 

Feder and 

Crustacea: Amphipoda Stenothoidae Metopa unidentified 


Crustacea: Amphipoda Stenothoidae Metopelloides erythrophthalmus EVOS II III IV Austin 1985 

Feder and 

Crustacea: Amphipoda Synopiidae Syrrhoe crenulata 

Matheke 1980 I/ST/Inf II x x Austin 1985 

Crustacea: Amphipoda Synopiidae Tiron biocellata EVOS II III IV Austin 1985 

Crustacea: Amphipoda Talitridae Talitrus unidentified 

Crustacea: Amphipoda Urothoidae Urothoe denticulata 


Schwafel 1988 I/Inf 

Feder and 

Matheke 1980 I/ST/Inf II III IV Gur’yanova 1951, 1962 

Crustacea: Amphipoda Vibiliidae Viblia australis Cooney 1987 P II III IV x Austin 1985 

II III IV V 

Crustacea: Cladocera Podonidae Evadne nordmanni Cooney 1987 P 

VI x x x Austin 1985 

Crustacea: Cladocera Podonidae Evadne unidentified Cooney 1987 P



BIOREGION 


SPECIMEN or 


REFERENCE to 


Crustacea: Cladocera Podonidae Evadne tergistina Pseudevadne Cooney 1987 P 

II III IV V 


Crustacea: Cladocera Podonidae Podon leuckarti Cooney 1987 P II III IV x x Austin 1985 

II III IV V 

Crustacea: Cladocera Podonidae Podon polyphmoies Cooney 1987 P 

VI x x x Austin 1985 

Crustacea: Cladocera Podonidae Podon unidentified Cooney 1987 P 

Crustacea: Copepoda: 

Calanoida Acartiidae Acartia cf. A. clausi Cordell 2000 P II III IV Cordell 2000 


Calanoida Acartiidae Acartia longiremis 


Calanoida Acartiidae Acartia unidentified Cooney: SEA P 


Calanoida Acartiidae Acartia tumida 


Calanoida Aetideidae Aetideus divergens A. armatus Cooney 1987 P 




1988 P II III IV x Austin 1985 

II III IV V 



Calanoida Aetideidae Aetideus pacificus Cooney 1987 P II III IV x Austin 1985 


Calanoida Aetideidae Aetideus unidentified Cooney 1987 P 


Calanoida Aetideidae Bradyidius saanichi Cooney 1987 P II III IV Austin 1985 


Calanoida Aetideidae Chiridiella unidentified 


1988 P 


Calanoida Aetideidae Chiridius gracilis Cooney 1987 P II III x x Austin 1985 


Calanoida Aetideidae Chiridius poppei Cooney 1987 P II x Brodsky 1950 


Calanoida Aetideidae Chiridius unidentified 


1988 P 


Calanoida Aetideidae Gaetanus intermedius Gaidius Cooney 1987 P II III IV Austin 1985 


Calanoida Aetideidae Gaidius tenuispinus G. minutus Cooney 1987 P II III IV Austin 1985 


Calanoida Aetideidae Gaidius variabilis Cooney 1987 P II III IV x Austin 1985 


Calanoida Aetideidae Pseudochirella sarsi Euchaetidae: Euchaeta Cooney 1987 P II III IV Austin 1985 


Calanoida Aetideidae Pseudochirella unidentified Cooney 1987 P 


Calanoida Aetideidae unidentified unidentified Cooney: SEA P 


Calanoida Augaptilidae Haloptilus pseudooxycephalus Cooney 1987 P II III IV x Austin 1985 


Calanoida Augaptilidae Pachyptilis pacificus Cooney 1987 P II III x Austin 1985 


Calanoida Calanidae Calanus glacialis E. marshallae Cooney 1987 P II III IV x Austin 1985





Calanoida Calanidae Calanus marshallae C. glacialis 

BIOREGION 

SPECIMEN or 


REFERENCE to 





Calanoida Calanidae Calanus pacificus Cooney 1987 P II III IV Austin 1985 


Calanoida Calanidae Neocalanus cristatus Cooney 1987 P II III IV V x x Austin 1985 


Calanoida Calanidae Neocalanus flemingeri Cooney: SEA P II III IV Miller 1988 


Calanoida Calanidae Neocalanus plumchrus 




Calanoida Calanidae Neocalanus unidentified Cooney: SEA P 


Calanoida Candaciidae Candacia bipinnata Cooney: SEA P II III IV V x x Austin 1985 


Calanoida Candaciidae Candacia columbiae 


Calanoida Centropagidae Centropages abdominalis 


1988 P II III IV x Cooney 1987 


1988 P II III IV x Cordell 2000 


Calanoida Clausocalanidae Clausocalanus arcuicornis Cooney 1987 P II III IV Austin 1985 


Calanoida Clausocalanidae Clausocalanus unidentified Cooney: SEA P 


Calanoida Clausocalanidae Mesocalanus tenuicornis Cooney 1987 P II x x Austin 1985 


Calanoida Clausocalanidae Microcalanus unidentified Cooney 1987 P 


Calanoida Clausocalanidae Pseudocalanus unidentified Cooney: SEA P 


Calanoida Corycaeidae Corycaeus anglicus Cooney 1987 P II III IV V x Austin 1985 


Calanoida Corycaeidae Corycaeus unidentified Cooney: SEA P 


Calanoida Ectinosomatidae Microsetella rosea Cooney 1987 P 


Calanoida Ectinosomatidae Microsetella unidentified Cooney: SEA P 


Calanoida Eucalanidae Eucalanus bungii 


Calanoida Euchaetidae Euchaete elongata 

II III VI V 



1988 P II III IV Austin 1985 




Calanoida Heterhabdidae Heterorhabdus compactus Cooney 1987 P II x x Brodsky 1950 


Calanoida Heterhabdidae Heterorhabdus robustoides Cooney 1987 P II III x 

Gardner and Szabo 

1982 


Calanoida Heterhabdidae Heterorhabdus tanneri Cooney 1987 P II III IV x x Austin 1985 


Calanoida Heterhabdidae Heterostylites longicornis Cooney: SEA P III VI V VI x Austin 1985 


Calanoida Heterhabdidae Heterostylites major Cooney 1987 P II III IV x Austin 1985 


Calanoida Lucicutiidae Lucicutia flavicornis Cooney 1987 P 

II III IV V 

VI x Austin 1985



BIOREGION 


SPECIMEN or 


REFERENCE to 



Calanoida Lucicutiidae Lucicutia ovalis Cooney 1987 P II III IV x Austin 1985 


Calanoida Lucicutiidae Lucicutia unidentified Cooney: SEA P 


Calanoida Metridiidae Metridia curticauda 


Calanoida Metridiidae Metridia okhotensis M. longa 


Calanoida Metridiidae Metridia pacifica 


Calanoida Metridiidae Metridia princeps 


Calanoida Metridiidae Metridia unidentified 








1988 P 


1988 P 

II III IV V 



Calanoida Metridiidae Pleuromamma robusta Cooney 1987 P II III IV x Austin 1985 


Calanoida Metridiidae Pleuromamma scutullata Cooney 1987 P II III IV x Austin 1985 


Calanoida Monstrillidae Cymbasoma rigidum Cooney 1987 P II III x Austin 1985 


Calanoida Monstrillidae Monstrilla canadensis Cooney 1987 P II III IV x 


Calanoida Monstrillidae Monstrilla helgolandica Cooney 1987 P II III IV x 


Calanoida Monstrillidae Monstrilla longiremus Cooney 1987 P II III IV 


Calanoida Monstrillidae Monstrilla unidentified Cooney: SEA P 


Calanoida Monstrillidae Monstrilla wandlii Cooney 1987 P II III IV x 


Calanoida Paracalanidae Paracalanus parvus Cooney 1987 P 


Calanoida Paracalanidae Paracalanus unidentified Cordell 2000 P 


Calanoida Phaennidae Gaetanus unidentified 


1988 P 


1982 


1982 


1982 


1982 

II III IV V 



Calanoida Pontellidae Epilabidocera longipedata E. amphitrites Cooney 1987 P II III IV x Austin 1985 


Calanoida Pseudocalanidae Pseudocalanus unidentified Cordell 2000 P 


Calanoida Scolecitrichidae Lophothrix frontalis Cooney 1987 P 

II III IV V 



Calanoida Scolecitrichidae Racovitzanus antarcticus Cooney 1987 P III IV x x Austin 1985 


Calanoida Scolecitrichidae Scaphocalanus brevicornis Cooney 1987 P II III IV x x Austin 1985 


Calanoida Scolecitrichidae Scaphocalanus magnus Cooney 1987 P II III IV x x Austin 1985



BIOREGION 


SPECIMEN or 


REFERENCE to 



Calanoida Scolecitrichidae Scolecithricella minor Cooney 1987 P II x x Austin 1985 


Calanoida Scolecitrichidae Scolecithricella ovata S. subdentata Cooney: SEA P 

II III IV V 



Calanoida Spinocalanidae Spinocalanus brevicaudatus Cooney 1987 P II III x x Austin 1985 


Calanoida Temoridae Eurytemora americana Cooney 1987 P II III x Austin 1985 


Calanoida Temoridae Eurytemora herdmani E. pacifica Cordell 2000 P II III IV x Austin 1985 


Calanoida Temoridae Eurytemora unidentified Cooney 1987 P 


Calanoida Tharydidae Undinella unidentified Cooney 1987 P 


Calanoida Tortanidae Tortanus discaudata 


Calanoida Amallothryx inornata P 


Cyclopoida Cyclopidae Euryte unidentified Cordell 2000 P 




Cyclopoida Cyclopidae Halicyclops unidentified Cordell 2000 P 


Cyclopoida Cyclopinidae Cyclopina unidentified Cooney 1987 P 


Cyclopoida Cyclopinidae unidentified unidentified Cordell 2000 P 


Cyclopoida Oithonidae Oithona similis Cordell 2000 P x x x x Cordell 2000 


Cyclopoida Oithonidae Oithona unidentified Cooney: SEA P 


Cyclopoida Oithonidae Oithona spinirostris Cordell 2000 P x x x x Cordell 2000 


Harpacticoida Ameiridae Ameira longipes Cordell 2000 I/Inf II x x Cordell 2000 


Harpacticoida Ameiridae Ameira sp. 1 Cordell 2000 I/Inf 


Harpacticoida Ameiridae Ameira sp. 2 Cordell 2000 I/Inf 


Harpacticoida Ameiridae unidentified sp. 1 Cordell 2000 I/Inf




BIOREGION 

SPECIMEN or 


REFERENCE to 



Harpacticoida Ancorabilidae Arthropsyllus serratus Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Canthocamptidae Mesochra pygmaea 


Schwafel 1988 I/Inf II x x x Cordell 2000 


Harpacticoida Canthocamptidae Mesochra sp. 1 Cordell 2000 I/Inf 


Harpacticoida Canthocamptidae unidentified incertae sedis Cordell 2000 II III IV Cordell 2000 


Harpacticoida Cletodidae Acrenhydrosoma unidentified Cordell 2000 I/Inf 


Harpacticoida Cletodidae Leimia vaga Cordell 2000 I/Inf II x x possible Cordell 2000 


Harpacticoida Cletodidae Nannopus palustris 


Harpacticoida Cletodidae Rhizothrix unidentified 


Harpacticoida Danielsseniidae Danielsennia typica 


Schwafel 1988 I/ST/Inf II x x Austin 1985 


Schwafel 1988 

I/ST/Inf 


Schwafel 1988 I/Inf II x x Cordell 2000 


Harpacticoida Diosaccidae Amonardia normani Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Diosaccidae Amonardia perturbata Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Diosaccidae Amphiascoides cf. D. debilis Cordell 2000 I/Inf II x x Cordell 2000 


Harpacticoida Diosaccidae Amphiascoides sp. 1 Cordell 2000 I/Inf 


Harpacticoida Diosaccidae Amphiascopis cinctus Cordell 2000 I/Inf II x x Cordell 2000 


Harpacticoida Diosaccidae Amphiascus minutus Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Diosaccidae Amphiascus sp. 1 Cordell 2000 I/Inf




BIOREGION 

SPECIMEN or 


REFERENCE to 



Harpacticoida Diosaccidae Diosaccus sp. 1 Cordell 2000 I/Inf 


Harpacticoida Diosaccidae Diosaccus spinatus Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Diosaccidae Robertsonia unidentified Cordell 2000 I/Inf 


Harpacticoida Diosaccidae Stenhelia peniculata Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Diosaccidae Stenhelia unidentified 


Schwafel 1988 

I/ST/Inf 


Harpacticoida Ectinosomatidae Ectinosoma unidentified Cordell 2000 I/Inf 


Harpacticoida Ectinosomatidae Halectinosoma finmarchium 


Harpacticoida Ectinosomatidae Halectinosoma gothiceps 






Harpacticoida Ectinosomatidae Halectinosoma sp. 1 Cordell 2000 I/Inf 






Harpacticoida Ectinosomatidae Microarthridion norvegica Cordell 2000 I/Inf x x x x Cordell 2000 


Harpacticoida Harpacticidae Harpacticus compressus Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Harpacticidae Harpacticus septentrionalis Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Harpacticidae Harpacticus 

sp.- obscurus group 

1 Cordell 2000 I/Inf II III IV Cordell 2000




BIOREGION 

SPECIMEN or 


REFERENCE to 






sp.- obscurus group 

2 Cordell 2000 I/Inf II III IV Cordell 2000 

sp.- uniremis group 

1 Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Harpacticidae Harpacticus superflexus 


Harpacticoida Harpacticidae Harpacticus uniremis 


Schwafel 1988 I/ST/Inf II III Cooney 1987 


Schwafel 1988 I/ST/Inf II x x Cordell 2000 


Harpacticoida Harpacticidae Zaus unidentified Cordell 2000 I/Inf 


Harpacticoida Laophontidae Echinolaophonte unidentified Cordell 2000 I/Inf 


Harpacticoida Laophontidae Heterolaophonte discophora Cordell 2000 I/Inf II x x Cordell 2000 


Harpacticoida Laophontidae Heterolaophonte longisetigera Cordell 2000 I/Inf II x x Cordell 2000 


Harpacticoida Laophontidae Heterolaophonte sp. 1 Cordell 2000 I/Inf 


Harpacticoida Laophontidae Heterolaophonte variabilis Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Laophontidae Laophonte applanata Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Laophontidae Laophonte elongata Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Laophontidae Laophonte sp. 1 Cordell 2000 I/Inf 


Harpacticoida Laophontidae Paralaophonte cf. L. congenera Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Laophontidae Paralaophonte hyperborea Cordell 2000 I/Inf II x x Cordell 2000 


Harpacticoida Laophontidae Paralaophonte pacifica Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Laophontidae Paralaophonte perplexa 


Schwafel 1988 I/ST/Inf II III IV x Cordell 2000




BIOREGION 

SPECIMEN or 


REFERENCE to 



Harpacticoida Laophontidae Paralaophonte sp. 1 Cordell 2000 I/Inf 


Harpacticoida Laophontidae Pseudonychocamptus spinifer Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Longipediidae Longipedia unidentified Cordell 2000 I/Inf 


Harpacticoida Parastenheliidae Parastenhelia sp. 1 Cordell 2000 I/Inf 


Harpacticoida Parastenheliidae Parastenhelia sp. 2 Cordell 2000 I/Inf 


Harpacticoida Tachidiidae Microarthridion littorale 


Schwafel 1988 I/Inf II x x Cordell 2000 


Harpacticoida Tegastidae Tegestes sp. 1 Cordell 2000 I/Inf 


Harpacticoida Tegastidae Tegestes sp. 2 Cordell 2000 I/Inf 


Harpacticoida Thalestridae Dactylopausia cf. D. glacialis Dactylopodia Cordell 2000 I/Inf II x x Cordell 2000 


Harpacticoida Thalestridae Dactylopausia glacialis Dactylopodia Cordell 2000 I/Inf II x x Cordell 2000 


Harpacticoida Thalestridae Dactylopausia paratisboides Dactylopodia Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Thalestridae Dactylopausia sp. 1 Dactylopodia Cordell 2000 I/Inf 


Harpacticoida Thalestridae Dactylopausia vulgaris Dactylopodia Cordell 2000 I/Inf II x x Cordell 2000 


Harpacticoida Thalestridae Diarthrodes sp. 2 Cordell 2000 I/Inf 


Harpacticoida Thalestridae Idomene unidentified Cordell 2000 I/Inf 


Harpacticoida Thalestridae Paradactylopodia latipes 


Schwafel 1988 I/ST/Inf II III IV Austin 1985





Harpacticoida Thalestridae Paradactylopodia latipes 

BIOREGION 

SPECIMEN or 


REFERENCE to 





Harpacticoida Thalestridae Paradactylopodia sp. 1 Cordell 2000 I/Inf 


Harpacticoida Thalestridae Parathalestris sp. 1 Cordell 2000 I/Inf 






Harpacticoida Tisbidae Scutellidium arthuri Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Tisbidae Tisbe cf. T. furcata Cordell 2000 I/Inf II III IV Cordell 2000 


Harpacticoida Tisbidae Tisbe gracilis Cooney 1987 I/Inf Cooney 1987 


Harpacticoida Tisbidae Tisbe inflata 




Harpacticoida Tisbidae Tisbe unidentified Cordell 2000 I/Inf 


Poecilostomatoida Oncaeidae Lubbockia wilsonae Cooney 1987 P II III x Austin 1985 


Poecilostomatoida Oncaeidae Oncaea borealis Copepoda: Cyclopoida Cooney 1987 P 

II III IV V 



Poecilostomatoida Oncaeidae Oncaea conifera Cooney 1987 P II III IV x x x Austin 1985 


Poecilostomatoida Oncaeidae Oncaea prolata O. notopus Cooney 1987 P II III IV Austin 1985 


Poecilostomatoida Oncaeidae Oncaea unidentified Cooney: SEA P 


Poecilostomatoida Oncaeidae Pseudolubbokia dilatata Cooney 1987 P II III IV x Austin 1985 


Poecilostomatoida unidentified unidentified Copepoda: Cyclopoida Cordell 2000 

Crustacea: Cumacea Bodoptriidae Vaunthompsonia pacifica EVOS ST 

II III IV V 

VI Austin 1985



BIOREGION 


SPECIMEN or 


REFERENCE to 


Crustacea: Cumacea Diastylidae Brachydiastylis unidentified 

Feder and 


Crustacea: Cumacea Diastylidae Diastylis alaskensis EVOS ST II III IV x Austin 1985 

Crustacea: Cumacea Diastylidae Diastylis bidentata EVOS ST II III IV x Austin 1985 

Crustacea: Cumacea Diastylidae Diastylis koreana EVOS ST II III IV x Austin 1985 

Crustacea: Cumacea Diastylidae Diastylis paraspinulosa Hoberg 1986 ST 

II III IV V 


Crustacea: Cumacea Diastylidae Diastylopsis dawsoni Hoberg 1986 ST 

II III IV V 


Crustacea: Cumacea Diastylidae Leptostylis unidentified EVOS ST II III IV x x Austin 1985 

Crustacea: Cumacea Lampropidae Lamprops beringi Chapman 2000 I/Inf II III IV Chapman 2000 

Crustacea: Cumacea Lampropidae Lamprops quadriplicata EVOS ST II x x Chapman 2000 

Crustacea: Cumacea Lampropidae Lamprops sarsi EVOS ST II III IV x Austin 1985 

Crustacea: Cumacea Leuconiidae Eudorella emarginata 


1988 ST/Inf II x x Austin 1985 

Crustacea: Cumacea Leuconiidae Eudorellopsis biplicata EVOS ST/Inf II x x Austin 1985 

Crustacea: Cumacea Leuconiidae Eudorellopsis deformis EVOS ST/Inf II III IV Austin 1985 

Crustacea: Cumacea Leuconiidae Eudorellopsis dezhavini EVOS ST/Inf II III IV x Austin 1985 

Crustacea: Cumacea Leuconiidae Eudorellopsis integra 

Feder and 


Crustacea: Cumacea Leuconiidae Leptocuma unidentified 


Schwafel 1988 I/ST 

Crustacea: Cumacea Leuconiidae Leucon nasica 

Feder and 

Matheke 1980 ST/Inf II x x x Austin 1985 

Crustacea: Cumacea Leuconiidae Leucon nasica orientalis UAM ST II x x x Austin 1985 

Crustacea: Cumacea Nannastacidae Campylaspis rubicunda UAM II x x x Schultz 1969 

Crustacea: Cumacea Nannastacidae Cumella vulgaris 


Schwafel 1988 I/ST/Epi II III IV C Chapman 2000 

Crustacea: Decapoda Atelecyclidae Telmessus cheiragonus Hart 1968 I/ST II III IV Hart 1968 

Crustacea: Decapoda Callianassidae Callinassa unidentified Hart 1968 I/ST 

Crustacea: Decapoda Cancridae Cancer branneri Hart 1968 I/ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Cancridae Cancer gracilis Hart 1968 I/ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Cancridae Cancer magister Hart 1968 I/ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Cancridae Cancer oregonensis Hart 1968 I/ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Cancridae Cancer productus Hart 1968 I/ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Crangonidae Argis alaskensis UAM ST/Epi II III IV Butler 1980 

Crustacea: Decapoda Crangonidae Argis lar EVOS ST/Epi II III IV x Butler 1980



BIOREGION 


SPECIMEN or 


REFERENCE to 


Crustacea: Decapoda Crangonidae Argis dentata 

Squires and 

Figueira 1974 I/ST II III IV x Butler 1980 

Crustacea: Decapoda Crangonidae Argis levior 

Squires and 

Figueira 1974 ST/Epi II III IV Butler 1980 

Crustacea: Decapoda Crangonidae Crangon alaskensis Cooney 1987 ST/Epi 

II III IV V 


Crustacea: Decapoda Crangonidae Crangon communis 

Squires and 

Figueira 1974 ST/Epi II III IV x Butler 1980 

Crustacea: Decapoda Crangonidae Crangon dalli 

Squires and 


Crustacea: Decapoda Crangonidae Crangon franciscorum 

Squires and 

Figueira 1974 I/ST II III IV Butler 1980 

Crustacea: Decapoda Crangonidae Crangon nigricauda 

Squires and 


Crustacea: Decapoda Crangonidae Lissocrangon stylirostris Crangon 

Squires and 


Crustacea: Decapoda Crangonidae Mesocrangon minuitella EVOS ST/Epi II III IV Butler 1980 

Crustacea: Decapoda Crangonidae Neocrangon alaskensis Crangon 

Squires and 


Crustacea: Decapoda Crangonidae Paracrangon echinata 

Squires and 


Crustacea: Decapoda Crangonidae Rhynchocramgon alata EVOS ST/Epi II III IV x Butler 1980 

Squires and 

Crustacea: Decapoda Crangonidae Sclerocrangon boreas 


Crustacea: Decapoda Galatheidae Munida quadrispina UAM ST/Epi II III IV Butler 1980 

Crustacea: Decapoda Grapsidae Hemigrapsus nudus Hart 1968 I/ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Grapsidae Hemigrapsus oregonensis 

Crustacea: Decapoda Hippolytidae Eualus fabricii 


Schwafel 1988 I/ST/Epi II III IV Hart 1968 

Squires and 


Crustacea: Decapoda Hippolytidae Eualus macropthalmalmus UAM ST/Epi II III IV Butler 1980 

Crustacea: Decapoda Hippolytidae Eualus suckleyi UAM ST/Epi II III IV x Butler 1980 

Crustacea: Decapoda Hippolytidae Eualus avinus UAM ST/Epi II III IV Butler 1980 

Squires and 

Crustacea: Decapoda Hippolytidae Eualus gaimardii 

Figueira 1974 ST/Epi II III IV x Austin 1985




Crustacea: Decapoda Hippolytidae Eualus pusiolus 

Crustacea: Decapoda Hippolytidae Eualus townsendi 

Crustacea: Decapoda Hippolytidae Heptacarpus decorus 

Crustacea: Decapoda Hippolytidae Heptacarpus brevirostris 

Crustacea: Decapoda Hippolytidae Heptacarpus camtschaticus 

Crustacea: Decapoda Hippolytidae Heptacarpus carinatus 

Crustacea: Decapoda Hippolytidae Heptacarpus paludicola 

Crustacea: Decapoda Hippolytidae Heptacarpus pictus 

Crustacea: Decapoda Hippolytidae Heptacarpus sitchensis 

Crustacea: Decapoda Hippolytidae Heptacarpus stimpsoni 

Crustacea: Decapoda Hippolytidae Heptacarpus stylus 

Crustacea: Decapoda Hippolytidae Heptacarpus tridens 

Crustacea: Decapoda Hippolytidae Hippolyte clarki 

Crustacea: Decapoda Hippolytidae Lebbeus groenlandicus 

BIOREGION 

SPECIMEN or 


REFERENCE to 


Squires and 

Figueira 1974 ST/Epi II x x Butler 1980 

Squires and 


Squires and 


Squires and 


Squires and 


Squires and 


Squires and 


Squires and 

Figueira 1974 I/ST II III IV Austin 1985 

Squires and 


Squires and 


Squires and 


Squires and 


Squires and 


Squires and 


Crustacea: Decapoda Hippolytidae Spirontocaris arcuata UAM I/ST II III IV x Butler 1980 

Crustacea: Decapoda Hippolytidae Spirontocaris lamellicornis UAM I/ST II III IV Butler 1980 

Crustacea: Decapoda Hippolytidae Spirontocaris ochotensis 

Squires and 


Crustacea: Decapoda Hippolytidae Spirontocaris phippsi 

Squires and 

Figueira 1974 I/ST II x x Austin 1985 

Crustacea: Decapoda Lithodidae Acantholithodes hispidus Hart 1968 I/ST II III IV Hart 1968 

Crustacea: Decapoda Lithodidae Cryptolithodes sitchensis Hart 1968 I/ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Lithodidae Cryptolithodes typicus Hart 1968 I/ST II III IV Hart 1968 

Crustacea: Decapoda Lithodidae Hapalogaster cavicauda Austin 1985 I/ST/Epi II III IV Austin 1985 

Crustacea: Decapoda Lithodidae Hapalogaster grebnitskii Hart 1968 I/ST II III IV x Hart 1968 

Crustacea: Decapoda Lithodidae Hapalogaster mertensii Hart 1968 I/ST/Epi II III IV Hart 1968




BIOREGION 

SPECIMEN or 


REFERENCE to 


Crustacea: Decapoda Lithodidae Lithodes aequispina UAM ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Lithodidae Lithodes couesi UAM ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Lithodidae Lopholithodes foraminatus UAM ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Lithodidae Oedignathus inermis Hart 1968 I/ST/Epi II III IV x Hart 1968 

Crustacea: Decapoda Lithodidae Paralithodes camtschatica UAM ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Lithodidae Phyllolithodes papillosus UAM I/ST II III IV Hart 1968 

Crustacea: Decapoda Lithodidae Placetron wosnessenskii UAM ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Lithodidae Rhinolithodes wosnessenskii UAM ST/Epi II III IV Hart 1968 


Crustacea: Decapoda Majidae Chionecetes bairdi 

1988 ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Majidae Chorilia longipes Hart 1968 ST/Epi II III IV x Hart 1968 

Crustacea: Decapoda Majidae Hyas lyratus Hart 1968 I/ST II III IV Hart 1968 

Crustacea: Decapoda Majidae Oregonia gracilis Hart 1968 I/ST II III IV Hart 1968 

Crustacea: Decapoda Majidae Pugettia gracilis Hart 1968 I/ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Majidae Pugettia producta Hart 1968 I/ST II III IV Hart 1968 

Crustacea: Decapoda Majidae Pugettia richii Hart 1968 I/ST II III IV Hart 1968 

Crustacea: Decapoda Majidae Scyra acutifrons Hines et al. 2000 I/ST II III IV x Hart 1968 

II III IV V 

Crustacea: Decapoda Oplophoridae Hymenadora frontalis Cooney 1987 P 


Crustacea: Decapoda Paguridae Discorsopagurus schmitti UAM I/ST II III IV Hart 1968 

Crustacea: Decapoda Paguridae Elassochirus cavimanus UAM ST/Epi II III IV x Hart 1968 

Crustacea: Decapoda Paguridae Elassochirus gilli UAM I/ST II III IV Hart 1968 

Crustacea: Decapoda Paguridae Elassochirus tenimanius UAM I/ST II III IV Hart 1968 

Crustacea: Decapoda Paguridae Labidochirus splendescens UAM I/ST II III IV x x Hart 1968 

Crustacea: Decapoda Paguridae Pagurus aleuticus UAM I/ST II III IV Hart 1968 

Crustacea: Decapoda Paguridae Pagurus beringanus UAM I/ST II III IV Hart 1968 

Crustacea: Decapoda Paguridae Pagurus capillatus UAM I/ST II III IV Hart 1968




BIOREGION 

SPECIMEN or 


REFERENCE to 


Crustacea: Decapoda Paguridae Pagurus cornutus UAM ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Paguridae Pagurus granosimanus UAM I/ST II III IV Hart 1968 

Crustacea: Decapoda Paguridae Pagurus 

hirsutiusculus 

hirsutiusculus 


Schwafel 1988 I/ST II III IV Hart 1968 

Crustacea: Decapoda Paguridae Pagurus kennerleyi UAM I/ST II III IV Hart 1968 

Crustacea: Decapoda Paguridae Pagurus samuelis UAM I/ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Paguridae Pagurus caurinus Hart 1968 I/ST II III IV V Hart 1968 

Crustacea: Decapoda Paguridae Pagurus confragosus Feder et al. 1979 ST/Epi II III IV Hart 1968 

Crustacea: Decapoda Paguridae Pagurus ochotensis 

Crustacea: Decapoda Pandalidae Pandalopsis dispar 

Crustacea: Decapoda Pandalidae Pandalus danae 


Schwafel 1988 I/ST II III IV Hart 1968 


1988 ST/Epi II III IV Butler 1980 

Squires and 


Crustacea: Decapoda Pandalidae Pandalus jordani UAM ST/Epi II III IV Butler 1980 

Crustacea: Decapoda Pandalidae Pandalus unidentified Cooney: SEA ST/Epi 

Crustacea: Decapoda Pandalidae Pandalus borealis 


1988 ST/Epi II x x Butler 1980 

Crustacea: Decapoda Pandalidae Pandalus goniurus 

Squires and 


Crustacea: Decapoda Pandalidae Pandalus hypsinotus 

Squires and 


Crustacea: Decapoda Pandalidae Pandalus montagui tridens 

Squires and 


Crustacea: Decapoda Pandalidae Pandalus platyceros 

Squires and 


Crustacea: Decapoda Pandalidae Pandalus stenolepis 

Squires and 


Crustacea: Decapoda Pasiphaeidae Pasiphaea pacifica UAM P II III IV x Butler 1980 

Crustacea: Decapoda Pinnotheridae Pinnixa littoralis Hart 1968 I/ST II III IV Hart 1968 

Crustacea: Decapoda Pinnotheridae Pinnixa occidentalis Hart 1968 ST/Epi II III IV Hart 1968 

Feder and 

Crustacea: Decapoda Pinnotheridae Pinnixa schmitti 

Matheke 1980 ST/Epi II III IV Hart 1968




BIOREGION 

SPECIMEN or 


REFERENCE to 


Crustacea: Decapoda Porcellanidae Petrolisthes eriomerus Hart 1968 I/ST II III IV Hart 1968 

Crustacea: Decapoda Sergestidae Sergestes similis Eusergestes Cooney 1987 P 

II III IV V 


Crustacea: Decapoda Upogebiidae Upogebia pugettensis 

Squires and 

Figueira 1974 I/Inf II III IV Hart 1968 

Crustacea: Decapoda Xanthidae Lophopanopeus bellus bellus Hart 1968 I/ST II III IV Hart 1968 

Crustacea: 

Euphausiacea Euphausiidae Euphausia pacifica Cooney 1987 P 

II III IV V 


Crustacea: 

Euphausiacea Euphausiidae Stylocherion unidentified Cooney 1987 P 

Crustacea: 

Euphausiacea Euphausiidae Tessarabranchion oculata Cooney 1987 P II III IV V x Austin 1985 

Crustacea: 

Euphausiacea Euphausiidae Thysanoessa inermis Cooney 1987 P 

II III IV V 


Crustacea: 

Euphausiacea Euphausiidae Thysanoessa longipes Cooney 1987 P III IV x Austin 1985 

Crustacea: 

Euphausiacea Euphausiidae Thysanoessa spinifera Cooney 1987 P 

II III IV V 


Crustacea: 

Euphausiacea Euphausiidae Thysanoessa inspinata Cooney 1987 P II III IV x Austin 1985 

Crustacea: 

Euphausiacea Euphausiidae Thysanoessa raschii 



Crustacea: Isopoda Aegidae Rocinela angustana UAM ST II III IV Schultz 1969 

Crustacea: Isopoda Dajidae Holophryxus alascensis UAM ST II III IV Austin 1985 

Crustacea: Isopoda Gnathiidae Gnathia tridens EVOS ST II III IV California NR Schultz 1969 

Crustacea: Isopoda Gnathiidae Gnathia unidentified 

Feder and 

Matheke 1980 ST/Inf Schultz 1969 

Crustacea: Isopoda Idoteidae Idotea fewkesi EVOS I/ST 

II III IV V 

VI Schultz 1969 

Crustacea: Isopoda Idoteidae Idotea obscura Chapman 2000 I/Epi II III IV Chapman 2000 

Crustacea: Isopoda Idoteidae Pentidotea wosnessenskii Idotea 


Schwafel 1988 I/Epi II III IV Schultz 1969 

Crustacea: Isopoda Idoteidae Synidotea ritteri UAM ST/Epi II III IV V California NR Schultz 1969 

Crustacea: Isopoda Janiridae Janiropsis kincaldi J. pugettensis Chapman 2000 I II III IV Chapman 2000 

Crustacea: Isopoda Ligiidae Ligia pallasii Chapman 2000 I/Epi II III IV Chapman 2000 

Crustacea: Isopoda Limnoriidae Limnoria lignorum EVOS I/ST II III IV Schultz 1969 

cf. M. 

Crustacea: Isopoda Munnidae Munna 

chromocephala UAM ST II III IV Washington NR Schultz 1969 

Crustacea: Isopoda Munnidae Munna ubiquita EVOS ST II III IV Washington NR Schultz 1969 

Crustacea: Isopoda Munnidae Pleurogonium unidentified I/Epi Schultz 1969 

Crustacea: Isopoda Sphaeromatidae Dynamenella sheari UAM I/Epi II III IV Austin 1985 

Crustacea: Isopoda Sphaeromatidae Exosphaeroma amplicauda UAM I/Epi II III IV Schultz 1969 

Crustacea: Isopoda Sphaeromatidae Exosphaeroma unidentified UAM I/Epi Schultz 1969 

Crustacea: Isopoda Sphaeromatidae Gnorimosphaeroma lutea G. insulare Chapman 2000 I/Epi II III IV Chapman 2000




Crustacea: Isopoda Sphaeromatidae Gnorimosphaeroma oregonensis 

BIOREGION 

SPECIMEN or 


REFERENCE to 



Schwafel 1988 I/ST II III IV Schultz 1969 

Crustacea: Leptostraca Nebaliidae Nebalia unidentified EVOS P 

Crustacea: Mysidacea Lophogastridae Gnatahophausia gigas Cooney 1987 P 

Crustacea: Mysidacea Mysidae Acanthomysis nephropthalma Cooney 1987 P 

II III IV V 

VI VII VIII 

IX x x Austin 1985 

II III IV V 


Crustacea: Mysidacea Mysidae Acanthomysis pseudomacraspis Cooney 1987 P II III IV x Austin 1985 

II III IV V 

Crustacea: Mysidacea Mysidae Holmsiella anomala Cooney 1987 P 


Crustacea: Mysidacea Mysidae Meterythrops robusta Cooney 1987 P II III IV x Austin 1985 

Crustacea: Mysidacea Mysidae Mysis littoralis Chapman 2000 P II x x x Chapman 2000 

Crustacea: Mysidacea Mysidae Mysis oculata M. littoralis Cooney 1987 P II III IV x x Austin 1985 

Crustacea: Mysidacea Mysidae Neomysis kadiakensis Cooney 1987 P II III IV Austin 1985 

Crustacea: Mysidacea Mysidae Neomysis rayi Cooney 1987 P II III IV V x Austin 1985 

II III IV V 

Crustacea: Mysidacea Mysidae Pseudomma truncatus Cooney 1987 P 


Crustacea: Ostracoda Halocyprididae Conchoecia alata minor Alacia 


1988 ST/P II III 

Crustacea: Ostracoda Halocyprididae Conchoecia elegans Paraconchoecia UAM ST/P II III 


Crustacea: Ostracoda Halocyprididae Conchoecia unidentified 

1988 ST/P 

British 


British 


Crustacea: Ostracoda Parodoxostomatinae unidentified unidentified UAM ST/P 

Crustacea: Ostracoda Philomedidae Philomedes dentata UAM ST/Inf II III 

Crustacea: Ostracoda Philomedidae Scleroconcha trituberculata UAM ST/Inf II III 

British 


British 


Crustacea: Ostracoda Podocopidae unidentified unidentified UAM 

Crustacea: Tanaidacea Paratanaidae Leptochelia savignyi Chapman 2000 I/Epi II x x C Chapman 2000 

Crustacea: Thoracica Balanamorpha Balanus crenatus 

Crustacea: Thoracica Balanamorpha Balanus glandula 

Crustacea: Thoracica Balanamorpha Chirona evermani 


Schwafel 1988 I/ST II III IV x Austin 1985 



Feder and 

Matheke 1980 ST/Inf II III IV Austin 1985




Crustacea: Thoracica Balanamorpha Semibalanus balanoides Balanus 

Crustacea: Thoracica Balanamorpha Semibalanus cariosus Balanus 

BIOREGION 

SPECIMEN or 


REFERENCE to 





Schwafel 1988 I/Epi II III IV x Austin 1985 

Crustacea: Thoracica Chthamalidae Chthamalus dalli EVOS I/Epi II III IV x Austin 1985 

Crustacea: Thoracica Scalpellidae Scaphellum pacificum 

Feder and 

Matheke 1980 ST/Epi II III IV x Austin 1985 

Pycnogonida Ammotheidae Achelia alaskensis Child 1995 ST/Epi II III IV x Child 1995 

Pycnogonida Ammotheidae Achelia borealis UAM ST/Epi II III IV x x Child 1995 

Pycnogonida Ammotheidae Achelia chaelata UAM ST/Epi II III IV Washington NR Austin 1985 

Pycnogonida Ammotheidae Achelia gracilipes Ammothea UAM ST/Epi II III IV 

British 


Pycnogonida Ammotheidae Achelia latifrons Child 1995 ST/Epi II III IV x Child 1995 

Pycnogonida Ammotheidae Achelia megova Child 1995 ST/Epi II III IV Child 1995 

Pycnogonida Ammotheidae Achelia pribilofensis Ammothea UAM ST/Epi II III IV Bering Sea nr Child 1995 

Pycnogonida Phoxochilidiidae Phoxichilidium femoratum UAM ST/Epi II x x Austin 1985 

Pycnogonida Phoxochilidiidae Phoxichilidium quadrodentatum UAM ST/Epi II III IV Austin 1985 

Pycnogonida Tanystylidae Tanystylum anthostomasti UAM ST/Epi II III IV x Austin 1985


Echinodermata 

Ninety-nine echinoderm species, representing all five classes are listed (Table 10.6). 

Principal sources for taxonomic and distributional information are D‘yakonov (1954) 

(ophiuroids), Lambert (1981, 1997) (Asteroidea and Holothuroidea). No range extensions or 

possibly undescribed species were noted for Prince William Sound. 

Table 10.6. Echinodermata. 

CLASS FAMILY GENUS SPECIES 

BIOREGION 

OTHER 

NAMES SPECIMEN or SOURCE HABITAT NEP NWP AR NWA 

REFERENCE to 

DISTRIBUTION 

Asteroidea Asteriidae Evasterias troschelii 


Schwafel 1988 I/ST/Epi II III IV Lambert 1981 

Asteroidea Asteriidae Leptasterias hexactis UAM I/ST/Epi II III IV Lambert 1981 

Asteroidea Asteriidae Orthasterias koehleri UAM ST II III IV Lambert 1981 

Asteroidea Asteriidae Pisaster ochraceus UAM I/ST/Epi II III IV Lambert 1981 

Asteroidea Asteriidae Pycnopodia helianthoides 


Schwafel 1988 I/ST/Epi II III IV Lambert 1981 

Asteroidea Asteriidae Rathbunaster californianus UAM ST II III IV Lambert 1981 

Asteroidea Asteropseidae Dermasterias imbricata Poraniidae UAM I/ST/Epi II III IV Lambert 1981 

Asteroidea Astropectinidae Dipsacaster borealis UAM ST II III IV Austin 1985 

Asteroidea Astropectinidae Leptychaster anomalus Feder and Matheke 1980 ST II III IV Lambert 1981 

Asteroidea Astropectinidae Leptychaster arcticus UAM ST II III IV x x x Lambert 1981 

Asteroidea Astropectinidae Leptychaster pacificus Lambert 1981 ST II III IV Lambert 1981 

Asteroidea Benthopectinidae Luidiaster dawsoni UAM ST II III IV Lambert 1981 

Asteroidea Benthopectinidae Nearchaster aciculosus UAM ST II III IV Austin 1985 

Asteroidea Echinasteridae Henricia aspera UAM ST II III IV Lambert 1981 

Asteroidea Echinasteridae Henricia asthenactis Lambert 1981 ST II III IV Lambert 1981 

Asteroidea Echinasteridae Henricia leviuscula UAM ST II III IV Lambert 1981 

Asteroidea Echinasteridae Henricia longispina Lambert 1981 ST II III IV x Lambert 1981 

Asteroidea Echinasteridae Henricia sanguinolenta UAM ST II III x x Lambert 1981 

Asteroidea Echinasteridae Poraniopsis inflata UAM ST II III IV x Lambert 1981 

Asteroidea Goniasteridae Ceramaster arcticus UAM ST II III IV Lambert 1981 

Asteroidea Goniasteridae Ceramaster patagonicus Lambert 1981 ST 


VII VIII IX Lambert 1981 

Asteroidea Goniasteridae Gephyreaster swifti Radiasterridae Feder and Jewett 1987 ST II III IV Lambert 1981 

Asteroidea Goniasteridae Hippasteria spinosa UAM ST II III IV Lambert 1981 

Asteroidea Goniasteridae Mediaster aequalis UAM ST II III IV Lambert 1981 

Asteroidea Goniasteridae Pseudarchaster parelii UAM ST II x x Lambert 1981 

Asteroidea Goniopectinidae Ctenodiscus crispatus Ctenodiscidae UAM ST 

I II III IV V 

VI VII IX x x x Lambert 1981 

Asteroidea Luidiidae Luidia foliolata UAM ST II III IV Lambert 1981 

Asteroidea Pedicellasteridae Pedicellaster magister Heliasteridae UAM ST II III IV x Austin 1985 

Asteroidea Pterasteridae Diplopteraster multipes UAM ST II III IV V x x x Lambert 1981 

Asteroidea Pterasteridae Pteraster militaris UAM ST II III IV x Lambert 1981 

Asteroidea Pterasteridae Pteraster tesselatus UAM ST II III IV Lambert 1981 

Asteroidea Solasteridae Crossaster papposus UAM ST II III IV x x x Lambert 1981 

Asteroidea Solasteridae Lophaster furcilliger UAM ST II III IV Lambert 1981 

Asteroidea Solasteridae Solaster dawsoni UAM ST II III IV x Lambert 1981 

Asteroidea Solasteridae Solaster endeca UAM ST II III IV V x x Lambert 1981 

Asteroidea Solasteridae Solaster paxillatus UAM ST II III IV x Lambert 1981 

Asteroidea Solasteridae Solaster stimpsoni Lambert 1981 ST II III IV x Lambert 1981 

Criniodea Antedonidae Florometra serratissima Austin 1985 ST II III IV V Austin 1985 

Criniodea Antedonidae Florometra asperrima 

Helimetra 

gracilis UAM ST II III IV V x Austin 1985 

Criniodea Antedonidae Psathyrometra fragilis Austin 1985 ST II III IV V x Austin 1985 

Criniodea Antedonidae Retiometra alascana Austin 1985 ST II III IV Austin 1985 

Echinoidea Echinarachniidae Echinarachnius parma Foster and Hines 2000 ST II III IV x Pavloskii 1955 

Echinoidea Schizasteridae Brisaster townsendi Feder and Matheke 1980 ST II III IV x Pavloskii 1955 

Echinoidea Strongylocentrotidae Allocentrotus fragilis Feder and Jewett 1987 ST II III IV Austin 1985 

Echinoidea Strongylocentrotidae Strongylocentrotus droebachiensis 


Schwafel 1988 I/ST/Epi II x x Austin 1985 

Echinoidea Strongylocentrotidae Strongylocentrotus franciscanus Rosenthal 1977 ST/Epi II III IV V Austin 1985 

Holothuroidea Cucumariidae Cucumaria frondosa japonica Lambert 1997 ST II III IV Lambert 1997 

Holothuroidea Cucumariidae Cucumaria miniata Lambert 1997 ST II III IV Lambert 1997 

Holothuroidea Cucumariidae Cucumaria vegae Lambert 1997 ST II III IV x Lambert 1997 

Holothuroidea Cucumariidae Ekmania diomedeae Lambert 1997 ST II III IV x Lambert 1997 

Holothuroidea Cucumariidae Pentamera calcigera UAM ST II x x Austin 1985 

Holothuroidea Cucumariidae Thyonidium kurilensis Lambert 1997 ST II III IV x Lambert 1997 

Holothuroidea Molpadiidae Molpadia intermedia UAM ST II III IV Lambert 1997 

Holothuroidea Phyllophoridae Pentamera populifera Lambert 1997 ST II III IV Lambert 1997 

Holothuroidea Phyllophoridae Thyone cf. T. benti UAM ST II III IV Lambert 1997 

Holothuroidea Psolidae Psolus chitonoides Lambert 1997 ST II III IV Lambert 1997 

Holothuroidea Psolidae Psolus squamatus Lambert 1997 ST II x x Lambert 1997 

Holothuroidea Sclerodactylidae Eupentacta pseudoquinquesemita Lambert 1997 ST II III IV Lambert 1997 

Holothuroidea Stichopodidae Parastichopus californicus Lambert 1997 ST II III IV Lambert 1997 

Holothuroidea Synallactidae Pseudostichopus mollis Lambert 1997 ST x x x x Lambert 1997 

Holothuroidea Synallactidae Synallactes challengeri Lambert 1997 ST x x x x Lambert 1997 

Holothuroidea Synaptidae Leptosynapta clarki UAM ST II III IV x Austin 1985 

Ophiuroidea Amphiuridae Amphipolis squamata Feder and Jewett 1987 ST x x x x Austin 1985 

Ophiuroidea Amphiuridae Diamphiodia craterodmeta Amphiodia Feder and Matheke 1980 ST II III IV x D’Yakanov 1954 

Ophiuroidea Amphiuridae Unioplis macraspis Amphioplus Feder and Matheke 1980 ST II III IV x Austin 1985 

Ophiuroidea Ophiactidae Ophiopholis aculeatata O. caryi Rosenthal 1977 ST II II IV x Austin 1985 

Ophiuroidea Ophiuridae Ophiura sarsi Feder and Matheke 1980 ST II x x x Austin 1985 

Ophiuroidea Ophiuridae Ophiura quadrispina Feder and Matheke 1980 ST II III IV D’Yakanov 1954


Bryozoa 

Bryozoa are abundant in Alaskan waters and prevalent in the fouling community. The 

data set consists of 74 species (Table 10.7), which is probably an underestimate. Nineteen 

identifications are confident only to the generic level. Sources for distribution data include Dick 

and Ross (1988), Hines and Ruiz (2000), Kluge (1962), as well as cataloged specimens in the 

UAM collection. J. Winston (2000) contributed the identifications and distributional information 

for 22 species. Schizoporella unicornis is regarded as a nonindigenous species, native to the 

northwestern Pacific. Ten species are possible new records for Alaska, range extensions form 

either the Arctic or Pacific region. 

Table 10.7. Bryozoa. 

ORDER FAMILY GENUS SPECIES 

OTHER 

NAMES 

Cheilostomata: 

Anasca Beaniidae Beania mirabilis UAM ST 

BIOREGION 

SPECIMEN or 


REFERENCE to 



VII VIII IX x x 

British 

Columbia nr Austin 1985 


Anasca Bugulidae Bugula californica Hines and Ruiz 2000 I II III IV Austin 1985 


Anasca Bugulidae Bugula pacifica Hines and Ruiz 2000 I II III IV Austin 1985 


Anasca Bugulidae Bugula unidentified Hines and Ruiz 2000 I 


Anasca Bugulidae Dendrobeania curvirostrata Hines and Ruiz 2000 I II III IV V VI 

British 



Anasca Bugulidae Dendrobeania lichenoides UAM I II III IV Dick and Ross 1988 


Anasca Bugulidae Dendrobeania murrayana UAM ST II x x x Kluge 1962 


Anasca Calloporidae Calloporella craticula Callopora Hines and Ruiz 2000 I/ST II x x x Kluge 1962 


Anasca Calloporidae Calloporella lineata Callopora UAM I/ST II x x x C Kluge 1962 


Anasca Calloporidae Calloporella "lineata" Hines and Ruiz 2000 I/ST II x x Austin 1985 


Anasca Calloporidae Tegella aquilirostris Hines and Ruiz 2000 I/ST II III IV x Austin 1985 


Anasca Calloporidae Tegella armifera Hines and Ruiz 2000 I/ST II III IV x Austin 1985 


Anasca Calloporidae Tegella robertsonae UAM ST II III IV V VI x Osburn 1950 


Anasca Cryptosulidae Cryptosula okadai Hines and Ruiz 2000 I II III IV x Dick and Ross 1988 


Anasca Cryptosulidae Harmeria scutulata Hippothoidae UAM I/ST II x x x Arctic nr Dick and Ross 1988 


Anasca Electridae Cauloramphus pseudospinifer Hines and Ruiz 2000 I II III IV x Dick and Ross 1988 


Anasca Electridae Electra unidentified UAM I 


Anasca Flustridae Terminoflustra membranaceotruncata UAM I/ST II x x x Dick and Ross 1988 


Anasca Hincksinidae Cauloramphus "variegata" C. spiniferum UAM I/ST II III IV V x x Austin 1985 


Anasca Membraniporidae Conopeum sp. (chesapeakensis) Hines and Ruiz 2000 I/ST II III IV V x x x Austin 1985 


Anasca Membraniporidae Membranipora membranipora Hines and Ruiz 2000 I/ST II III x x x Kluge 1962 


Anasca Membraniporidae Membranipora serrilamella UAM I/ST x x x x N Pacific nr Austin 1985 


Anasca Microporidae Micropora unidentified UAM I/ST 


Anasca Microporinidae Microporina articulata UAM I II x x x Dick and Ross 1988 


Anasca Scrupariidae Brettia unidentified 

Bicellariidae/C 

orynoprella UAM I/ST 


Anasca Scrupocellariidae Scrupocellaria unidentified UAM I/ST 


Anasca Scrupocellariidae Tricellaria gracilis UAM I/ST II x x x Kluge 1962 


Anasca Scrupocellariidae Tricellaria occidentalis UAM ST II III IV V VI 

British 



Anasca Scrupocellariidae Tricellaria ternata UAM I II x x x Dick and Ross 1988 


Ascophora Cribrilinidae Cribrilina annulata UAM I/ST II x x x Kluge 1962 


Ascophora Cribrilinidae Cribrilina corbicula Hines and Ruiz 2000 I II III IV C Austin 1985 


Ascophora Cribrilinidae Cribrilina unidentified Hines and Ruiz 2000 I/ST 


Ascophora Hippoporinidae Lepralia unidentified Hippothoa UAM I/ST 


Ascophora Hippothoidae Cellepora craticula Costazia Hines and Ruiz 2000 I/ST x C Kluge 1962



ORDER FAMILY GENUS SPECIES 

OTHER 

NAMES 

BIOREGION 

SPECIMEN or 


REFERENCE to 



Ascophora Hippothoidae Celleporella hyalina Hippothoa UAM I/ST x x x x C Dick and Ross 1988 


Ascophora Hippothoidae Hippothoa unidentified UAM I 


Ascophora Microporellidae Fenestruloides eopacifica Hines and Ruiz 2000 I II III IV V VI California NR Soule et al. 1996 


Ascophora Microporellidae Microporella germana Hines and Ruiz 2000 I II III IV Dick and Ross 1988 


Ascophora Mucronellidae Cystisella bicornia Osburn 1952 ST II III IV x Osburn 1952 


Ascophora Mucronellidae Cystisella saccata Osburn 1952 ST II III x Arctic nr Osburn 1952 


Ascophora Mucronellidae Parasmittina trispinosa Smittinidae Hines and Ruiz 2000 I x x x x C Dick and Ross 1988 


Ascophora Mucronellidae Porella acutirostris Smittinidae UAM I/ST II III IV V VI x x x Dick and Ross 1988 


Ascophora Mucronellidae Porella acutirostris 


Ascophora Mucronellidae Rhamphostomella bilaminata 


Ascophora Mucronellidae Rhamphostomella gigantea 


Ascophora Mucronellidae Rhamphostomella hincksi 

P. major 

Smittinidae Hines and Ruiz 2000 ST II III IV V VI x x x Osburn 1952 

Rhamphostom 

ellidae UAM I/ST x x Kluge 1962 

Rhamphostom 

ellidae Hines and Ruiz 2000 ST x x Arctic nr Osburn 1952 

Rhamphostom 

ellidae Hines and Ruiz 2000 ST II x x Kluge 1962 


Ascophora Mucronellidae Smittina unidentified Smittinidae UAM I/ST 


Ascophora Reteporidae Phidolopora pacifica P. labiata Hines and Ruiz 2000 I II III IV Austin 1985 


Ascophora Schizoporellidae Hippodiplosia unidentified UAM I/ST II x x x Kluge 1962 


Ascophora Schizoporellidae Hippoporina vulgaris Hines and Ruiz 2000 I/ST II 


Ascophora Schizoporellidae Schizomavella porifera Hines and Ruiz 2000 I/ST II x x x Arctic nr Osburn 1952 


Ascophora Schizoporellidae Schizoporella "unicornis" Hines and Ruiz 2000 I x x x x NW Pacific Definate Hines and Ruiz 2000 


Ascophora Umbonulidae Umbonula unidentified UAM I/ST 

Ctenostomata Alcyonidiidae Alcyonidium hirsutum Hines and Ruiz 2000 I/ST II x x x NE Atlantic NR Kluge 1962 

Ctenostomata Alcyonidiidae Alcyonidium polynoum A. mytili Hines and Ruiz 2000 I/ST II III IV V x x x C Austin 1985 

Ctenostomata Alcyonidiidae Alcyonidium unidentified Hines and Ruiz 2000 I/ST 

III IV V VI 

Cyclostomata Crisiidae Crisia occidentalis UAM I/ST VII VIII IX x x Washington NR Austin 1985 

Cyclostomata Crisiidae Crisia serrulata Hines and Ruiz 2000 I II III IV Austin 1985 

Cyclostomata Crisiidae Filicrisia fanciscana Hines and Ruiz 2000 I II III IV x Austin 1985 

Cyclostomata Crisiidae Filicrisia geniculata UAM I/ST II x x Kluge 1962 

Cyclostomata Crisiidae Filicrisia smitti Hines and Ruiz 2000 ST II III x x x Kluge 1962 

Cyclostomata Diaperoeciidae Diaperoecia unidentified UAM ST 

Cyclostomata Diastoporidae Disporella unidentified UAM I/ST 

Cyclostomata Heteroporidae Heteropora magna UAM ST II III IV 

British 


Cyclostomata Heteroporidae Heteropora pacifica UAM ST II III IV Austin 1985 

Cyclostomata Heteroporidae Heteropora unidentified UAM I/ST 

Cyclostomata Lichenoporidae Lichenopora unidentified UAM I/ST 

Cyclostomata Oncousoeciidae Oncousiecia unidentified UAM I/ST 

Cyclostomata Oncousoeciidae Stomatopora granulata UAM I/ST II III IV x Kluge 1962 

Cyclostomata Tubuliporidae Idmonea unidentified Idmoneidae UAM I/ST 

Cyclostomata Tubuliporidae Platonea unidentified UAM I/ST 

Cyclostomata Tubuliporidae Tubulipora flabellaris UAM I/ST II III IV x Kluge 1962 

Cyclostomata Tubuliporidae Tubulipora sp. (tuba) Hines and Ruiz 2000 I/ST II III IV 

Cyclostomata Tubuliporidae Tubulipora tuba 

T. 

occidentalis, 

T. fasciculifera Hines and Ruiz 2000 I/ST II III IV V 

British 


Miscellaneous Invertebrates 

Ninety-five taxa, representing twelve different phyla make up this data set (Table 10.8). 

Nemerteans, chaetognaths, brachiopods, and urochordates , and are fairly well-known or have 

received the attention of the project’s taxonomists. (Lambert 2000; Mills 2000). However, it is 

clear that sponges, flatworms, and nematodes are undercounted. The nonindigenous sponge, 

Cliona thosina was recorded by the fouling community survey (Hines and Ruiz 200). 

Phylum 

taxa reported in this study 

Porifera 4 

Platyhelminthes none identified to species 

Nemertea 54 

Sipunculida 3 

Priapulida 1 

Echiuridae 1


Phoronidae none identified to species 

Brachiopoda 3 

Chaetognatha 5 

Tardigrada 1 

Urochordata 28 

Gnathostomulida, Gastrotrichida, Kinorhyncha have not been documented. 

Table 10.8. Miscellaneous invertebrates. 

Bioregion 

PHYLUM/ CLASS FAMILY GENUS SPECIES OTHER NAMES 

SPECIMEN or 


REFERENCE to 

DISTRIBUTION 

Brachiopoda: 

Articulata Cancellothyrididae Terebratulina unguicula UAM I/ST/Epi II III IV Austin 1985 

Brachiopoda: 

Articulata Hemithyridae Hemithiris psittacea UAM ST II III IV x x Austin 1985 

Brachiopoda: 

Articulata Laqueidae Terebratalia transversa UAM ST II III IV SE Alaska nr Austin 1985 

Brachiopoda: 

Articulata Laqueidae Laqueus californianus UAM ST II III IV x Austin 1985 

Chaetagnatha Eukrohniidae Eukrohnia fowleri Cooney 1987 P II III IV x x x Austin 1985 

Chaetagnatha Eukrohniidae Eukrohnia bathypelagica Cooney 1987 P II III IV x Austin 1985 

Chaetagnatha Eukrohniidae Eukrohnia hamata 


1988 P x x x x 

Smith and Johnson 

1996 

Chaetagnatha Sagittidae Sagitta elegans 


1988 P II III IV 

Smith and Johnson 

1996 

Chaetagnatha Sagittidae Sagitta scrippsae Cooney 1987 P 


VII VIII x Austin 1985 

Echiurida Echiuridae Echiuris 

echiuris 

alaskensis 


Schwafel 1988 I II III IV x Austin 1985 

Nematoda unidentified unidentified 

Feder and 

Matheke 1980 Inf 

Nemertea Amphiporidae Amphiporus angulatus Coe 1910 I II III IV V VI x Coe 1904 

Nemertea Amphiporidae Amphiporus bimaculatus Coe 1910 I 


VII VIII Coe 1904 

Nemertea Amphiporidae Amphiporus exilis A. formidabilis Coe 1910 I II III IV V VI Coe 1904 

Nemertea Amphiporidae Amphiporus gelatinosus Austin 1985 I II III x Austin 1985 

Nemertea Amphiporidae Amphiporus imparispinosus Austin 1985 I 


VII VIII Austin 1985 

Nemertea Amphiporidae Amphiporus lactifloreus I I 

Nemertea Amphiporidae Amphiporus leuciodes Coe 1910 I III Coe 1904 

Nemertea Amphiporidae Amphiporus nebulosus Coe 1910 I II III Coe 1904 

Nemertea Amphiporidae Amphiporus tigrinus Coe 1910 I III IV Coe 1904 

Nemertea Amphiporidae Zygonemertes albida Coe 1910 I III IV V VI Coe 1904 

Nemertea Amphiporidae Zygonemertes thalassina Coe 1910 I II III Coe 1904 

Nemertea Amphiporidae Zygonemertes virescens Austin 1985 I 

I II III IV V VI 

VII VIII IX x Austin 1985 

III IV V VI 

Nemertea Carinomidae Carinoma griffini C. mutabilis Coe 1910 I 


Nemertea Carinomidae Carinomella lactea Tubulanidae Austin 1985 I I II III IV V VI Austin 1985 

Nemertea Cephalothricidae Cephalothrix linearis Procephalothrix spiralis Coe 1910 I II III IV V VI Coe 1904 

Nemertea Cerebratulidae Cerebratulus albifrons Coe 1910 I III IV V VI Coe 1904 

Nemertea Cerebratulidae Cerebratulus herculeus Coe 1910 I III IV V VI Coe 1904 

Nemertea Cerebratulidae Cerebratulus longiceps Coe 1910 I II III IV V Coe 1904 

Nemertea Cerebratulidae Cerebratulus marginatus Coe 1910 I III Coe 1904 

Nemertea Cerebratulidae Cerebratulus montgomeryi Coe 1910 I II III IV V Coe 1904 

Nemertea Cerebratulidae Cerebratulus occidentalis Coe 1910 I II III Coe 1904 

Nemertea Cerebratulidae Cerebratulus unidentified 


1980 ST 

Nemertea Emplectonematidae Emplectonema buergeri Coe 1910 I III Coe 1904 

Nemertea Emplectonematidae Emplectonema gracile Coe 1910 I II III IV V Coe 1904 

Nemertea Emplectonematidae Paranemertes carnea Coe 1910 I II III IV Coe 1904 

Nemertea Emplectonematidae Paranemertes pallida Coe 1910 I II III Coe 1904 


Nemertea Emplectonematidae Paranemertes peregrina 

Schwafel 1988 I II III IV x Austin 1985 

Nemertea Lineidae Lineus torquatus Coe 1910 I II III IV V VI Coe 1904 

Nemertea Lineidae Micrura alaskensis Coe 1910 I 

III IV V VI 


Nemertea Lineidae Micrura verrilli Coe 1910 I II III IV V VI Coe 1904 

Nemertea Lineidae Myoisophagus sanguineus Lineus socialis/vegetus Austin 1985 I II III IV V VI x x Austin 1985 

Nemertea Tetrastemmatidae Tetrastemma aberrans Coe 1910 I II III Coe 1904 

Nemertea Tetrastemmatidae Tetrastemma bicolor Coe 1910 I II III Coe 1904 

Nemertea Tetrastemmatidae Tetrastemma caecum Amphinemertes Coe 1910 I/ST II III Coe 1904 

Nemertea Tetrastemmatidae Tetrastemma candidum Austin 1985 I 


VII VIII x x Austin 1985 

Nemertea Tetrastemmatidae Tetrastemma nigrifrons Austin 1985 I 

I II III IV V VI 

VII VIII IX Austin 1985 

Nemertea Tubulanidae Carinella albocinctus Tubulanus Austin 1985 ST II III IV V VI Austin 1985 

Nemertea Tubulanidae Carinella annulatus Tubulanus Austin 1985 I II III x Austin 1985 

Nemertea Tubulanidae Carinella capistrata Tubulanus capistratus Coe 1910 I II Coe 1904 

Nemertea Tubulanidae Carinella cingulatus Tubulanus Austin 1985 I II III IV V VI Austin 1985 

Nemertea Tubulanidae Carinella dinema Tubulanus sexlineatus Coe 1910 I II III Coe 1904 

Nemertea Tubulanidae Carinella frenatus Tubulanus Austin 1985 I II III IV V VI Austin 1985 

Nemertea Tubulanidae Carinella nothus Tubulanus Austin 1985 I II III x Austin 1985 

Nemertea Tubulanidae Carinella speciosa Tubulanus polymorphus Coe 1910 I II III Coe 1904 

Nemertea Valenciniidae Taeniosoma princeps Baseodiscus Coe 1910 I II III IV Coe 1904 

Phoronida unidentified unidentified 


1980 ST



Bioregion 

PHYLUM/ CLASS FAMILY GENUS SPECIES OTHER NAMES 

SPECIMEN or 


REFERENCE to 

DISTRIBUTION 

Platyhelminthes unidentified unidentified 


Schwafel 1988 Inf 

Porifera Myxillidae Myxilla incrustans UAM ST II III IV x Austin 1985 

Porifera Subertidae Suberites ficus 

Scheel, Foster, and 

Hough, 1997 I/ST/Epi II III IV Austin 1985 

Porifera Spirastrellidae Cliona thosina Hines et al. 2000 boring II unkown definate 

Porifera Spirastrellidae Cliona celata Rosenthal 1977 Epi II III IV V Austin 1985 

Priapulida Priapulidae Priapulus caudatus 


1980 ST II x x x Austin 1985 

Sipunculida Golfingiidae Golfingia margaritacea 


1980 I/ST/Inf II x x Austin 1985 

Sipunculida Golfingiidae Golfingia vulgaris 


1980 ST II x x Austin 1985 

Sipunculida Golfingiidae Phascolion strombi 


1980 ST x x x x Austin 1985 

Sipunculida Phascolosomatidae Phascolosoma agassizii Fisher 1952 I/ST/Inf 


VIII IX Austin 1985 

Tardigrada Hypsibiidae Hypsibius appelloefi 


Schwafel 1988 ST x x x x 

Schuster and 

Grigarick 1965 

Urochordata: 

Ascidiacea Ascidiidae Ascidia adhaerens Lambert 2000 I II x x Lambert 2000 

Urochordata: 

Ascidiacea Ascidiidae Ascidia columbiana A. callosa Lambert 2000 I II III IV x Lambert 2000 

Urochordata: 

Ascidiacea Ascidiidae Ascidia unidentified Lambert 2000 I/ST/Epi 

Urochordata: 

Ascidiacea Clavelinidae Distaplia alaskensis Lambert 2000 ST/Epi 

Urochordata: 

Ascidiacea Clavelinidae Distaplia sp. undescribed Lambert 2000 ST/Epi possible Lambert 2000 

Urochordata: 

Ascidiacea Corellidae Chelysoma columbiana Austin 1985 ST/Epi II III IV Austin 1985 

Urochordata: 

Ascidiacea Corellidae Chelysoma productum Lambert 2000 ST/Epi II III IV V VI Austin 1985 

Urochordata: 

Ascidiacea Corellidae Corella inflata Lambert 2000 I II III IV Austin 1985 

Urochordata: 

Ascidiacea Corellidae Corella unidentified Lambert 2000 I 

Urochordata: 

Ascidiacea Corellidae Corella willmeriana Lambert 2000 I II III IV Austin 1985 

Urochordata: 

Ascidiacea Molgulidae Molgula retortiformis Lambert 2000 I II x x Austin 1985 

Urochordata: 

Ascidiacea Polyclinidae Ritterella unidentified Lambert 2000 I 

Urochordata: 

Ascidiacea Pyruridae Boltenia echinata EVOS ST/Epi II III IV x Austin 1985 

Urochordata: 

Ascidiacea Pyruridae Boltenia ovifera UAM ST/Epi II III IV x Pavloskii 1955 

Urochordata: 

Ascidiacea Pyruridae Halocynthia igoboja UAM ST/Epi II III IV x Austin 1985 

Urochordata: 

Ascidiacea Pyruridae Halocynthia auranticum Tethyum 

Scheel, Foster, and 

Hough, 1997 ST/Epi II III IV x Austin 1985 

Urochordata: 

Ascidiacea Pyruridae Pyura haustor Lambert 2000 ST/Epi II III IV V VI Austin 1985 

Urochordata: 

Ascidiacea Styelidae Botrylloides violaceus Lambert 2000 ST/Epi x Japan definate 

Urochordata: 

Ascidiacea Styelidae Cnemidocarpa finmarkiensis Stylea Rosenthal 1977 ST/Epi II III IV V x x Austin 1985 

Urochordata: 

Ascidiacea Styelidae Metandrocarpa taylori Rosenthal 1977 ST/Epi II III IV V Austin 1985 

Urochordata: 

Ascidiacea Styelidae Styela truncata Lambert 2000 I II III IV Austin 1985 

Urochordata: 

Ascidiacea Clavelinidae Distaplia occidentalis Lambert 2000 ST/Epi III IV V VI 

British 


Urochordata: 

Larvacea Fritillariidae Fritillaria borealis Cooney 1987 P II III IV V VI x x x Austin 1985 

Urochordata: 

Larvacea Fritillariidae Fritillaria unidentified Cooney: SEA P 

Urochordata: 

Larvacea Oikopleuridae Oikopleura labradorensis Cooney 1987 P II III IV V x x x 

Urochordata: 

Larvacea Oikopleuridae Oikopleura unidentified 


1988 P 

Urochordata: 

Larvacea Oikopleuridae Oikopleura vanhoeffeni Cooney 1987 P II III x x x 

Urochordata Oikopleuridae Oikopleura dioica Cooney 1987 P II III IV V 


Urochordata Salpida Salpa fusiformis Cooney 1987 P 



Urochordata Salpida Salpa maxima Cooney 1987 P 



1998 


1998 


1998 


1998 


1998 

Fishes 

The 175 species we list represent 33 families (Table 10.9). The fish data set uses records 

for fish specimens catalogued in the University of Alaska Museum, as well as published and 

manuscript sources. (Baxter unpublished manuscript, Hart 1973). Name changes derive from and 

Humann (1996). Two species, the American shad, Alosa sapidissima, and the Atlantic Salmon, 

Samo salar are nonindigenous. 

Mecklenberg, (letter of 10/30/00) recognizes an undescribed cottid, possibly a 

Malacocottus. It has been identified incorrectly as Thecopterus aleuticus.


The following habitat descriptors used in the data sets are based on usage in Eschmeyer 

et al (1983)and Eschmeyer (1990) and Hart (1973). 

Table 10.9. Fishes. 

ANA anadromous BW brackish D dermesal 

E eelgrass Epip epipelagic Est estuaries 

FW freshwater I intertidal KB kelp bed 

NS nearshore Off offshore Pel pelagic 

PB pebble bottom RI rocky intertidal 

RST rocky subtidal SAB sand bottom 

SB softbottom ST subtidal 

SW saltwater 

BIOREGION 

FAMILY GENUS SPECIES OTHER NAMES 

SPECIMEN or 


REFERENCE to 

DISTRIBUTION 

Acipenseridae Acipenser medirostris Hart 1973 BW, SW, NS IV x x Hart 1973 

Acipenseridae Acipenser transmontanus Hart 1973 ANA IV Hart 1973 

Agonidae Anoplagonus inermis UAM D IV x Hart 1973 

Agonidae Aspidophoroides bartoni UAM SB, SA IV x Baxter 1987ms 

Agonidae Aspidophoroides olrikii UAM SB, SA IV x Baxter 1987ms 

Agonidae Bathyagonus infraspinata Asterotheca UAM D, SB IV Hart 1973 

Agonidae Bathyagonus nigripinnis UAM D, SB IV x x Hart 1973 

Agonidae Bathyagonus pentacanthus Asterotheca Hart 1973 D, SB IV Hart 1973 

Agonidae Bathyagonus alascanus 

Asterotheca 

alascana Hart 1973 RST, D 

II III 

IV Hart 1973 

Agonidae Bothragonus swani Hart 1973 I, ST IV Hart 1973 

Agonidae Hypsagonus quadricornis Hart 1973 D, SB IV x x Hart 1973 

Agonidae Occella verrucosa Hart 1973 D, SB IV Hart 1973 

Agonidae Odontopyxis trispinosa UAM D, SB IV SE Alaska nr Hart 1973 

Agonidae Podothecus acipenserinus Agonus Hart 1973 D IV x Hart 1973 

Ammodytidae Ammodytes hexapterus UAM I, ST, NS, P IV x x Hart 1973 

Anarhichadidae Anarrhichthys ocellatus Hart 1973 RST, D IV x Hart 1973 

Anoplopomatidae Anoplopoma fimbria Feder et al. 1979 ST, D IV x x Hart 1973 

1, ST, SAB, KB, II III 

Aulorhynchidae Aulorhynchus flavidus UAM 

E 

IV Hart 1973 

Bathylagidae Leuroglossus callorhinus L. schmidti UAM P IV x Hart 1973 

Bathymasteridae Bathylagus milleri Hart 1973 P IV x Hart 1973 

Bathymasteridae Bathylagus ochotensis Hart 1973 P IV x x Hart 1973 

Bathymasteridae Bathymaster caeruleofasciatus Rogers et al. 1986 RST 

II III 

IV 

II III 

IV 

Eschmeyer et al. 

1983 


1983 

Bathymasteridae Bathymaster leurolepis Rogers et al. 1986 I/ST 

x 

Bathymasteridae Bathymaster signatus UAM D, SB IV x x Hart 1973 

Bathymasteridae Ronquilus jordani Hart 1973 RST, D IV Hart 1973 

Bothidae Citharichthys sordidus Hart 1973 D, SAB IV Hart 1973 

Carcharhinidae Prionace glauca UAM Epip x x x x Hart 1973 

Clupeidae Alosa sapidissima UAM ANA III IV x Atlantic definate Hart 1973 

Clupeidae Clupea pallasii UAM P, Est II x x Hart 1973 

Cottidae Artedius fenestralis Hart 1973 I, ST IV Hart 1973 

Cottidae Artedius harringtoni Hart 1973 I, RST IV Hart 1973 

Cottidae Artedius lateralis UAM I, ST IV Hart 1973 

Cottidae Blepsias bilobus UAM I, ST IV x x Hart 1973 

Cottidae Blepsias cirrhosus Hart 1973 I, ST IV x Hart 1973 

RI, ST, SAB, E, II III 

Cottidae Clinocottus acuticeps UAM 

KB 

IV Hart 1973 

Cottidae Clinocottus embryum UAM RI IV Hart 1973 

Cottidae Clinocottus globiceps Hart 1973 RI IV Hart 1973 

Cottidae Dasycottus setiger UAM SB IV x Hart 1973 

Cottidae Enophrys bison Hart 1973 RST, SB IV Hart 1973 

Cottidae Gymnocanthus galeatus UAM SB, NS IV Hart 1973 

Cottidae Hemilepidotus hemilepidotus UAM I, RST, NS IV x Hart 1973 

Cottidae Hemilepidotus jordani UAM ST IV x Hart 1973 

Cottidae Hemilepidotus spinosus Hart 1973 I, ST IV SE Alaska nr Hart 1973 

Cottidae Icelinus borealis UAM SB IV Hart 1973 

Cottidae Icelus spiniger Hart 1973 SB IV x x Hart 1973 

Cottidae Leptocottus armatus UAM I, ST IV Hart 1973 

Cottidae Malacocottus kincaidi UAM SB IV x Hart 1973 

Cottidae Malacocottus zonurus UAM SB IV x Hart 1973 

Cottidae Myoxocephalus polyacanthocephalus Hart 1973 I, NS, SB, SAB IV x Hart 1973 

Cottidae Myoxocephalus scorpius 

M. 

groenlandicus Rogers et al. 1986 ST 

II III 

IV x x Baxter 1987ms



BIOREGION 


SPECIMEN or 


REFERENCE to 

DISTRIBUTION 

Cottidae Myoxocephalus verrucosus UAM ST IV x Baxter 1987ms 

Cottidae Nautichthys oculofasciatus Hart 1973 RST IV x x Hart 1973 

Cottidae Oligocottus maculosus Hart 1973 I, RST IV x x Hart 1973 

Cottidae Oligocottus rimensis UAM I, RST IV Hart 1973 

Cottidae Psychrolutes paradoxus Hart 1973 NS, RST, SB IV x x Hart 1973 

Cottidae Psychrolutes sigalutes Gilbertidia Hart 1973 RST, NS, SB IV Hart 1973 

Cottidae Radulinus asprellus UAM NS IV Hart 1973 

Cottidae Rhamphocottus richardsoni UAM I, RST, SAB IV Hart 1973 

Cottidae Malacocottus sp. 

Thecopterus 

aleuticus UAM RST II Bering Sea nr Baxter 1987ms 

Cottidae Triglops macellus Hart 1973 D IV Hart 1973 

Cottidae Triglops pingelii UAM D II x x x Hart 1973 

Cryptacanthodidae Delolepis giganteus Hart 1973 SB IV Hart 1973 

Cryptacanthodidae Lyconectes aleutensis UAM ST, NS, SAB IV Hart 1973 

Cyclopteridae Aptocyclus ventricosus Hart 1973 I, ST IV x x Hart 1973 

Cyclopteridae Careproctus colletti UAM SB, SA II x Hart 1973 

Cyclopteridae Careproctus gilberti Hart 1973 SB, SA IV Hart 1973 

Cyclopteridae Careproctus melanurus UAM ST, SB, SAB IV Hart 1973 

Cyclopteridae Eumicrotremus orbis UAM I, RST IV x Hart 1973 

Cyclopteridae Liparis callyodon Hart 1973 I IV Hart 1973 

Cyclopteridae Liparis cyclopus Hart 1973 NS IV Hart 1973 

Cyclopteridae Liparis dennyi UAM SB, SA IV Hart 1973 

Cyclopteridae Liparis florae UAM I IV Hart 1973 

Cyclopteridae Liparis pulchellus Hart 1973 SB IV x Hart 1973 

Cyclopteridae Liparis rutteri Hart 1973 I, ST IV Hart 1973 

Cyclopteridae Nectoliparis pelagicus UAM P IV x Hart 1973 

Cyclopteridae Paraliparis deani UAM D IV x x SE Alaska nr Hart 1973 

Gadidae Eleginus gracilis UAM P, D IV x Hart 1973 

Gadidae Gadus macrocephalus UAM P, D IV x x Hart 1973 

Gadidae Merluccius productus Hart 1973 P, D IV x Hart 1973 

Gadidae Microgadus proximus UAM D IV Hart 1973 

Gadidae Theragra chalcogramma UAM Off, D IV x x Hart 1973 

Gasterosteidae Gasterosteus aculeatus Hart 1973 FW/SW, NS II x Hart 1973 

Hexagrammidae Hexagrammos decagrammus Hart 1973 RST, KB, SAB IV Hart 1973 

Hexagrammidae Hexagrammos lagocephalus Hart 1973 RST IV x Hart 1973 

Hexagrammidae Hexagrammos octogrammus UAM RST IV x x Hart 1973 

Hexagrammidae Hexagrammos stelleri UAM RST, NS, E IV x x Hart 1973 

Hexagrammidae Ophiodon elongatus Hart 1973 

NS, RST, SB, 

SAB 

II III 

IV Hart 1973 

Lamnidae Cetorhinus maximus Hart 1973 Epip, P, NS x x x x Hart 1973 

Lamnidae Lamna ditropus Hart 1973 Off, Epip, NS IV x x Hart 1973 

Osmeridae Hypomesus pretiosus pretiosus Hart 1973 NS IV x Hart 1973 

Osmeridae Mallotus villosus Hart 1973 Oceanic II x x x Hart 1973 

Osmeridae Spirinchus thaleichthys Hart 1973 ANA IV Hart 1973 

Osmeridae Thaleichthys pacificus UAM ANA IV x Hart 1973 

Petromyzontidae Lampetra japonica Rogers et al. 1986 ANA IV Baxter 1987ms 

Petromyzontidae Lampetra tridentata Hart 1973 ANA IV x x Hart 1973 

Pholidae Apodichthys flavidus Hart 1973 I/RT IV Hart 1973 

Pholidae Pholis gilli UAM I/RT IV Hart 1973 

Pholidae Pholis laeta UAM I, ST IV Hart 1973 

Pleuronectidae Atheresthes stomias UAM SB, Off IV Hart 1973 

Pleuronectidae Eopsetta jordani Hart 1973 SAB IV Hart 1973 

Pleuronectidae Glyptocephalus zachirus UAM SB, SAB IV Hart 1973 

Pleuronectidae Hippoglossoides elassodon UAM SB IV x x Hart 1973 

Pleuronectidae Hippoglossus stenolepis Rogers et al. 1986 ST, NS 

II III 

IV x 


1983 

Pleuronectidae Iopsetta ischyra Hart 1973 NS, Off IV Hart 1973 

Pleuronectidae Limanda aspera UAM NS IV x x Hart 1973 

Pleuronectidae Limanda proboscidea UAM SB IV Hart 1973 

Pleuronectidae Microstomus pacificus UAM SB IV Hart 1973 

Pleuronectidae Platichthys stellatus UAM NS, Est IV x x Hart 1973 

Pleuronectidae Pleuronectes bilineata Lepidosetta UAM NS, Off, PB IV x x Hart 1973 

Pleuronectidae Pleuronectes vetulus Parophrys UAM I, ST IV Hart 1973 

Pleuronectidae Pleuronichthys decurrens Feder et al. 1979 SB IV Hart 1973 

Pleuronectidae Psettichthys melanostictus Hart 1973 NS IV Hart 1973 

Pleuronectidae Reinhardtius hippoglossoides Hart 1973 SAB, PB II x x x Hart 1973 

Rajidae Bathyraja kincaidi Raja UAM D IV Hart 1973 

Rajidae Bathyraja aleutica Wilimovsky 1958 D II III x Wilimovsky 1958 

Rajidae Bathyraja interrupta Escheyer et al. 1983 

Rajidae Raja binoculata Escheyer et al. 1983 D 

II III 

IV 

II III 

IV 

Escheyer et al. 

1983 


1983 


1983 

Rajidae Raja parmifera Escheyer et al. 1983 D II III x 

Rajidae Raja rhina Bathyraja Feder et al. 1979 D IV Hart 1973 

Rajidae Bathyraja stellulata Raja Hart 1973 D IV Hart 1973 

Salmonidae Oncorhynchus clarki Salmo clarki Hart 1973 ANA, FW IV Hart 1973 

Salmonidae Oncorhynchus gorbuscha UAM ANA IV x Hart 1973 

Salmonidae Oncorhynchus keta Hart 1973 ANA IV x x Hart 1973



BIOREGION 


SPECIMEN or 


REFERENCE to 

DISTRIBUTION 

Salmonidae Oncorhynchus kisutch Hart 1973 ANA IV x Hart 1973 

Salmonidae Oncorhynchus mykiss Salmo gardnerii Hart 1973 ANA IV x Hart 1973 

Salmonidae Oncorhynchus nerka Hart 1973 ANA IV x x Hart 1973 

Salmonidae Oncorhynchus tshawytscha Hart 1973 ANA IV x x Hart 1973 

Salmonidae Salmo salar Hart 1973 ANA III IV x 

British 

Columbia definate Hines et al. 2000 

Salmonidae Salvelinus malma Hart 1973 ANA IV x x Hart 1973 

Scombridae Sarda chiliensis Hart 1973 NS IV x Hart 1973 

Scorpaenidae Sebastes aleutianus UAM D IV Hart 1973 

Scorpaenidae Sebastes alutus UAM Off IV x Hart 1973 

II III 


Scorpaenidae Sebastes auriculatus Rogers et al. 1986 ST, NS IV SE Alaska nr 

1983 

Scorpaenidae Sebastes borealis UAM SB IV x Hart 1973 

Scorpaenidae Sebastes brevispinis Hart 1973 D IV Hart 1973 

Scorpaenidae Sebastes caurinus Hart 1973 RST, SAB, NS IV Hart 1973 

Scorpaenidae Sebastes ciliatus Hart 1973 D, NS IV x Hart 1973 

Scorpaenidae Sebastes crameri Hart 1973 SB IV Hart 1973 

Scorpaenidae Sebastes diploproa Hart 1973 Off, SB IV Hart 1973 

Scorpaenidae Sebastes elongatus Hart 1973 RST, SB IV Hart 1973 

Scorpaenidae Sebastes emphaeus Rogers et al. 1986 ST, NS IV Baxter 1987ms 

Scorpaenidae Sebastes flavidus Hart 1973 P IV Hart 1973 

Scorpaenidae Sebastes maliger Hart 1973 RST, NS IV Hart 1973 

Scorpaenidae Sebastes melanops Hart 1973 D, NS, RR IV Hart 1973 

Scorpaenidae Sebastes mystinus Hart 1973 D, RR, KB IV Hart 1973 

Scorpaenidae Sebastes nebulosus Rogers et al. 1986 NS, RST, RR IV SE Alaska nr Baxter 1987ms 

Scorpaenidae Sebastes nigrocinctus Rogers et al. 1986 RST IV SE Alaska nr Baxter 1987ms 

Scorpaenidae Sebastes paucispinis Hart 1973 D, RR IV Hart 1973 

Scorpaenidae Sebastes proriger Hart 1973 D IV Hart 1973 

Scorpaenidae Sebastes ruberrimus Feder et al. 1979 RR IV Hart 1973 

Scorpaenidae Sebastes variegatus UAM D IV Hart 1973 

Scorpaenidae Sebsatsolobus alascanus Hart 1973 SB IV x Hart 1973 

Scorpaenidae Sebsatsolobus altivelis Hart 1973 Off, SB IV Hart 1973 

Scytalinidae Scytalina cerdale Hart 1973 I, NS SAB, PB IV Hart 1973 

Sphyraenidae Sphyraena argentea Hart 1973 NS IV Hart 1973 

Squalidae Somniosus pacificus Hart 1973 I, P, D IV x x Hart 1973 

Squalidae Squalus acanthias Hart 1973 P, D II x x Hart 1973 

Stichaeidae Anoplarchus insignis Hart 1973 RST IV Hart 1973 

Stichaeidae Anoplarchus purpurescens UAM I, RST IV Hart 1973 

Stichaeidae Chirolophis decoratus Hart 1973 RST, KB IV Hart 1973 

Stichaeidae Chirolophis nugator Rogers et al. 1986 RST 

II III 

IV 


1983 

Stichaeidae Lumpenella longirostris UAM Off IV Hart 1973 

Stichaeidae Lumpenus maculatus UAM SAB IV x Hart 1973 

Stichaeidae Lumpenus sagitta UAM Off, NS IV x Hart 1973 

Stichaeidae Phytichthys chirus Hart 1973 I, RST IV Hart 1973 

Stichaeidae Poroclinus rothrocki Hart 1973 SB, SA IV Hart 1973 

Stichaeidae Stichaeus punctatus UAM RST, SAB II x x x Hart 1973 

Stichaeidae Xiphister mucosus UAM I, RST IV Hart 1973 

Trichodontidae Trichodon trichodon UAM NS, SB SAB IV x x Hart 1973 

Zaproridae Zaprora silenus Hart 1973 D IV x x Hart 1973 

Zoarcidae Bothrocara molle UAM SB, SA IV x Hart 1973 

Zoarcidae Lycodapus mandibularis UAM P IV Hart 1973 

Zoarcidae Lycodes brevipes UAM SB, SAB IV x x Hart 1973 

Zoarcidae Lycodes diapterus UAM SB IV x x Hart 1973 

Zoarcidae Lycodes palearis UAM SB- SAB IV x Hart 1973 

Zoarcidae Lycodopsis pacifica Feder et al. 1979 P IV Hart 1973 

Stichaeidae Xiphister atropurpureus Hart 1973 I, RST IV Hart 1973 

Birds 

The rich bird fauna of Prince William Sound, the northern Gulf of Alaska coast and 

Copper River Delta has been described in detail by Isleb and Kessel (1973, 1989). From their 

checklist, we have selected 114 species with habitats designated “ beaches and tidal flats, rocky 

shores, and reefs, inshore waters, and offshore waters” (Table 10.10). 

The American Ornithologists’ Union’s (1983) Check-list of North American Birds, was 

used to update common and scientific names. Usage of terms that define status: migrant, visitor, 

breeder and resident is derived from Isleb and Kessel (1973). 

Nsh nearshore m migrant 

Pel pelagic v visitor 

Int intertidal r rare 

b breeder


Table 10.10. Birds. 

FAMILY GENUS SPECIES COMMON NAME 

OTHER 

NAMES SOURCE HABITAT STATUS 

Gaviidae Gavia immer Common Loon Isleib and Kessel 1973 nsh r 

Gaviidae Gavia adamsii Yellow-billed Loon Isleib and Kessel 1973 nsh r 

Gaviidae Gavia arctica Arctic Loon Isleib and Kessel 1973 nsh r 

Gaviidae Gavia stellata Red-throated Loon Isleib and Kessel 1973 nsh r 

Podicipedidae Podiceps grisegena Red-necked Grebe Isleib and Kessel 1973 nsh r 

Podicipedidae Podiceps auritis Horned Grebe Isleib and Kessel 1973 nsh r 

Porcellariidae Fulmarus glacialis Fulmar Isleib and Kessel 1973 pel v 

Porcellariidae Puffinus creatopus Pink-footed Shearwater Isleib and Kessel 1973 pel v 

Porcellariidae Puffinus carneipes Pale-footed Shearwater Isleib and Kessel 1973 pel v 

Porcellariidae Puffinus griseus Sooty Shearwater Isleib and Kessel 1973 pel v 

Porcellariidae Puffinus tenuirostris Slender-billed Shearwater Isleib and Kessel 1973 pel v 

Porcellariidae Oceandroma furcata Fork-tailed Storm Petrel Isleib and Kessel 1973 pel r 

Phalacrocoracidae Phalacrocorax auritius Double-crested Cormorant Isleib and Kessel 1973 nsh/int r 

Phalacrocoracidae Phalacrocorax penicillatus Brandt’s Cormorant Isleib and Kessel 1973 nsh/int b 

Phalacrocoracidae Phalacrocorax peligicus Pelagic Cormorant Isleib and Kessel 1973 nsh/int r 

Phalacrocoracidae Phalacrocorax urile Red-faced Cormorant Isleib and Kessel 1973 nsh/int r 

Ardeidae Ardea herodias Great blue Heron Isleib and Kessel 1973 int r 

Anatidae Cygnus columbianus Whistling Swan Olor Isleib and Kessel 1973 int m 

Anatidae Cygnus buccinator Trumpeter Swam Olor Isleib and Kessel 1973 int r 

Anatidae Branta canadensis Canada Goose Isleib and Kessel 1973 int r 

Anatidae Branta bernicula Brant B. nigricans Isleib and Kessel 1973 int m 

Anatidae Chen canagica Emperor Goose Philacate Isleib and Kessel 1973 int m 

Anatidae Anser albifrons Greater White-fronted Goose Isleib and Kessel 1973 int m 

Anatidae Chen caerulescens Snow Goose 

C. 

hyperborea Isleib and Kessel 1973 int m 

Anatidae Anas platyrhynchos Mallard Isleib and Kessel 1973 int r 

Anatidae Anas strepera Gadwall Isleib and Kessel 1973 int r 

Anatidae Anas acuta Northern Pintail Isleib and Kessel 1973 int r 

Anatidae Anas crecca Green-winged Teal 

A. 

carolinensis Isleib and Kessel 1973 int r 

Anatidae Anas discors Blue-winged Teal Isleib and Kessel 1973 int m 

Anatidae Anas penelope Europeam Widgeon Mareca Isleib and Kessel 1973 int m 

Anatidae Anas americana American Widgeon Mareca Isleib and Kessel 1973 int r 

Anatidae Anas clypeata Northern Shoveler Spatula Isleib and Kessel 1973 int m/b 

Anatidae Aythya valisineria Canvasback Isleib and Kessel 1973 int m/b 

Anatidae Aythya marila Greater Scaup Isleib and Kessel 1973 int r 

Anatidae Bucephala clangula Common Goldeneye Isleib and Kessel 1973 int/nsh r 

Anatidae Bucephala islandica Barrow’s Goldeneye Isleib and Kessel 1973 int/nsh r 

Anatidae Bucephala albeola Bufflehead Isleib and Kessel 1973 int/nsh r 

Anatidae Clangula hymenalis Oldsquaw Isleib and Kessel 1973 int/nsh r 

Anatidae Histrionicus histrionicus Harlequin Duck Isleib and Kessel 1973 int/nsh r 

Anatidae Polysticta stelleri Steller’s Eider Isleib and Kessel 1973 int/nsh v 

Anatidae Somateria mollissima Common Eider Isleib and Kessel 1973 int/nsh v 

Anatidae Somateria spectabilis King Eider Isleib and Kessel 1973 int/nsh v 

Anatidae Melanitta fusca White-winged Scoter M. deglandi Isleib and Kessel 1973 int/nsh r 

Anatidae Melanitta perspicillata Surf Scoter Isleib and Kessel 1973 int/nsh r 

Anatidae Melanitta nigra Black Scoter Oidema Isleib and Kessel 1973 int/nsh r 

Anatidae Lophodytes cucullatus Hooded Merganser Isleib and Kessel 1973 int/nsh r 

Anatidae Mergus merganser Common Merganser Isleib and Kessel 1973 int/nsh 

Anatidae Mergus serrator Red-breated Merganser Isleib and Kessel 1973 int/nsh r 

Accipitridae Haliaeetus leucocephalus Bald Eagle Isleib and Kessel 1973 int r 

Accipitridae Circus cyaneus Marsh Hawk Isleib and Kessel 1973 int r 

Accipitridae Pandion haliaetus Osprey P. rusticolus Isleib and Kessel 1973 int r 

Accipitridae Falco peregrinus Peregrine Falcon Isleib and Kessel 1973 int r 

Accipitridae Falco columbarius Merlin Isleib and Kessel 1973 int r 

Gruidae Grus canadensis Sandhill Crane Isleib and Kessel 1973 int r 

Hematopodidae Haematopus bachmani Black Oystercatcher Isleib and Kessel 1973 int r 

Caradriidae Charadrius semipalmatus Semipalmated Plover Isleib and Kessel 1973 int m/b 

Caradriidae Charadrius vociferus Killldeer Isleib and Kessel 1973 int v 

Caradriidae Pluvialis dominica American Golder Plover Isleib and Kessel 1973 int m/b 

Caradriidae Pluvialis squatarola Black-bellied Plover Squatarola Isleib and Kessel 1973 int m 

Caradriidae Aphriza virgata Surfbird Isleib and Kessel 1973 int r 

Caradriidae Arinaria interpres Ruddy Turnstone Isleib and Kessel 1973 int m




OTHER 

NAMES SOURCE HABITAT STATUS 

Caradriidae Arinaria melanocephala Black Turnstone Isleib and Kessel 1973 int r 

Caradriidae Galligano galligano Common Snipe Capella Isleib and Kessel 1973 int r 

Caradriidae Numenius phaeopus Whimbrel Isleib and Kessel 1973 int m 

Caradriidae Actitus macularia Spotted Sandpiper Isleib and Kessel 1973 int m 

Caradriidae Trigna solitaria Solitary Sandpiper Isleib and Kessel 1973 int m 

Caradriidae Heteroscelus incanus Wandering Tattler Isleib and Kessel 1973 int m 

Caradriidae Tringa melanoleuca Greater Yellowlegs Tatonus Isleib and Kessel 1973 int m 

Caradriidae Tringa flavipes Lesser Yellowlegs Tatonus Isleib and Kessel 1973 int m 

Caradriidae Calidris canutus Red Knot Isleib and Kessel 1973 int m 

Caradriidae Calidris ptilocnemis Rock Sandpiper Erolia Isleib and Kessel 1973 int m 

Caradriidae Calidris acuminata Sharp-tailed Sandpiper Erolia Isleib and Kessel 1973 int m 

Caradriidae Calidris melanotos Pectoral Sandpiper Erolia Isleib and Kessel 1973 int m 

Caradriidae Calidris baridii Baird’s Sandpiper Erolia Isleib and Kessel 1973 int m 

Caradriidae Calidris minutilla Least Sandpiper Erolia Isleib and Kessel 1973 int m/b 

Caradriidae Calidris alpina Dunlin Erolia Isleib and Kessel 1973 int r 

Caradriidae Limnodromus griseus Short-billed Dowitcher Isleib and Kessel 1973 int m/b 

Caradriidae Limnodromus scolopaceus Long-billed Dowitcher Isleib and Kessel 1973 int m 

Caradriidae Calidris pusilla Semipaliated Sandpiper Ereunetes Isleib and Kessel 1973 int m 

Caradriidae Calidris mauri Western Sandpiper Ereunetes Isleib and Kessel 1973 int m 

Caradriidae Limosa lapponica Bar-tailed Godwit Isleib and Kessel 1973 int m 

Caradriidae Limosa haemastica Hudsonian Godwit Isleib and Kessel 1973 int 

Caradriidae Calidris alba Sanderling Crocethia Isleib and Kessel 1973 int m 

Caradriidae Phalaropus fulicarius Red Phalarope Isleib and Kessel 1973 int/nsh/pel m 

Caradriidae Lobipes lobatus Northern Phalarope Isleib and Kessel 1973 int/nsh/pel m/b 

Stercorariidae Stercorarius pomarinus Pomarine Jaeger Isleib and Kessel 1973 pel m 

Stercorariidae Stercorarius parasiticus Parasitic Jaeger Isleib and Kessel 1973 pel m/b 

Stercorariidae Stercorarius longicaudus Long-tailed Jaeger Isleib and Kessel 1973 pel m 

Stercorariidae Catharacta skua Greater Skua Isleib and Kessel 1973 pel v 

Laridae Larus hyperboreus Glaucus Gull Isleib and Kessel 1973 int/nsh/pel r 

Laridae Larus glaucesens Glaucous-winged Gull Isleib and Kessel 1973 int/nsh/pel r 

Laridae Larus argentatus Herring Gull Isleib and Kessel 1973 int/nsh/pel r 

Laridae Larus canus Mew Gull Isleib and Kessel 1973 int/nsh/pel r 

Laridae Larus philadelphia Bonaparte’s Gulls Isleib and Kessel 1973 int/nsh/pel m/b 

Laridae Rissa tridactyla Black-legged Kittywake Isleib and Kessel 1973 int/nsh/pel r 

Laridae Xema sabini Sabine’s Gull Isleib and Kessel 1973 int/nsh/pel m 

Laridae Sterna paradisaea Arctic Tern Isleib and Kessel 1973 int/nsh/pel m/b 

Laridae Sterna aleutica Aleutian Tern Isleib and Kessel 1973 int/nsh/pel b 

Alcidae Uria aalge Common Murre Isleib and Kessel 1973 nsh/pel r 

Alcidae Uria lomvia Thick-billed Murre Isleib and Kessel 1973 nsh/pel r 

Alcidae Cepphus columba Pigeon Guillemont Isleib and Kessel 1973 nsh/pel r 

Alcidae Brachyramphus marmoratus Marbled Murrelet Isleib and Kessel 1973 nsh/pel r 

Alcidae Brachyramphus brevirostis Kitlitz’s Murrelet Isleib and Kessel 1973 nsh/pel r 

Alcidae Synthliboramphus antiquum Ancient Murrelet Isleib and Kessel 1973 nsh/pel r 

Alcidae Aethia psittacula Parakeet Auklet 

Synthlibora 

mphus Isleib and Kessel 1973 nsh/pel b 

Alcidae Aethia cristatella Crested Auklet Isleib and Kessel 1973 nsh/pel v 

Alcidae Cerorhica monocrata Rhinoceros Auklet Isleib and Kessel 1973 nsh v/b 

Alcidae Fratercula corniculata Horned Puffin Isleib and Kessel 1973 nsh/pel r 

Alcidae Lunda cirrhata Tufted Puffin Isleib and Kessel 1973 nsh/pel r 

Alcedinidae Megaceryle alcyon Belted Kingfisher Isleib and Kessel 1973 int r 

Corvidae Cyanocitta stelleri Steller’s Jay Isleib and Kessel 1973 int r 

Corvidae Corvus corax Common Raven Isleib and Kessel 1973 int r 

Corvidae Corvus caurinus Northwestern Crow Isleib and Kessel 1973 int r 

Mammals 

Eleven marine mammal species are well known from Prince William Sound. We also 

include the river otter, and black and brown bear because of their use of marine resources 

(salmon, intertidal animals) (Table 10.11). Wynne (1992) and UAM mammal collection records 

were used as a source for species names. Distribution, community, and status are inferred from 

range maps in Wynne (1992).


Table 10.11. Mammals. 


BIOREGION 

SPECIMEN or 

SOURE HABITAT STATUS NEP NWP AR NWA Source 

Balaenopteridae Balaenoptera acutorostrata Minke Whale Wynne 1992 nsh/pel m x x x x Wynne 1992 

Balaenopteridae Balaenoptera physalus Fin Whale Wynne 1992 nsh/pel m x x x x Wynne 1992 

Balaenopteridae Megaptera novaeangliae Humpback Whale Wynne 1992 nsh m x x x x Wynne 1992 

Delphinidae Orcinus orca Killer Whale Wynne 1992 nsh/pel r/m x x x x Wynne 1992 

Eschrichtiidae Eschrichtius robustus Grey Whale Wynne 1992 pel m II III IV Wynne 1992 

Mustelidae Enhydra lutris Sea Otter Wynne 1992 int/nsh r II III IV x Hall 1981 

Mustelidae Lontra canadensis River Otter UAM Mammals int r x x x x Hall 1981 

Otariidae Callorhinus ursinus Northern Fur Seal UAM Mammals pel m II III IV x Hall 1981 

Otariidae Eumetopias jubatus Steller’s Sea Lion UAM Mammals nsh r II III IV x Hall 1981 

Phocidae Phoca vitulina Harbor Seal UAM Mammals nsh r II III IV Hall 1981 

Phoecoenidae Phocoena phocoena Harbor Porpoise Wynne 1992 nsh m/r II x x Wynne 1992 

Phoecoenidae Phocoenoides dalli Dall’s Porpoise Wynne 1992 nsh/pel m/r II III IV x Wynne 1992 

Ursidae Ursus americanus Black Bear UAM Mammals int r II III Hall 1981 

Ursidae Ursus arctos Brown Bear UAM Mammals int r II III Hall 1981 

10E. Discussion 

The data sets contain entries for 1878 taxa, of which all but 180 are species-level 

identifications. Thirty-nine species are recognized as possibly undescribed. Eighty-nine species 

had not previously been reported in Prince William Sound, and represent distributional range 

extension. (Table 10.12.) These species were probably overlooked in previous environmental 

surveys, because of their small size, cryptic appearance, similarity to more well-known species, 

or lack of taxonomic experts to work with them. Eighteen species are likely to represent 

nonindigenous species. At least 97 species are uncertain in origin (cryptogenic). Over 70% of the 

animals (1343) are invertebrates. Arthropods 24.3%, mollusks 16.8%, and polychaete annelids 

12.4% make up the largest number of total species. (Table 10.13. Figure 10.4). 

Plants 

Cnidaria/ 

Ctenophora 

Nemertea 

Polychaeta 

Mollusca 

Arthropoda 

probable NIS 5 1 2 1 9 

definate NIS 1 1 2 1 1 1 2 9 

range extension 

within Alaska 4 9 1 8 1 8 31 

range extension to 

Alaska 17 2 19 7 10 2 1 58 

cryptogenic 68 7 1 6 82 

potential new or 

undescribed species 1 6 2 1 1 11 

not identified to 

species 10 27 1 7 1 109 19 5 3 182 

total spcies-level 

taxa 231 102 45 233 315 455 68 74 30 23 178 113 14 1881 

Echinodermata 

Bryozoa 

Urochordata 

Other 

invertebrates 

Fishes 

Birds 

Mammals 

TOTALS 

Table 10.12. Summary of Prince William Sound marine biota by NIS status and identification confidence.


Plants 

Cnidaria/ Ctenophor 

Mollusca 

numer of species 231 102 315 233 455 66 74 98 178 113 14 

percent of total fauna 6.2% 19.1% 14.2% 27.7% 4.0% 4.5% 6.0% 10.6% 6.9% 0.9% 

percent of total biota 12.3% 5.4% 16.8% 12.4% 24.3% 3.5% 3.9% 5.2% 9.3% 6.0% 0.7% 

total invertebrates 1343 

total vertebrates 302 

total plants 233 

total biota considered 1876 

Polychaeta 

Arthropoda 

Echinodermata 

Bryozoa 

Other invertebrates 

Fishes 

Birds 

Mammals 

Table 10.13. Summary of the plants and animals considered. 

Marine Plants 

Ascomycota 

Cyanophyta 

Xanthophyta 

Chlorophyta 

Rhodophyta 

Phaeophyta 

vascular plants 

Marine Biota 

total invertebrates 

total vertebrates 

total plants 

Marine Invertebrates 

Cnidaria/Ctenophora 

Polychaeta 

Mollusca 

Arthropoda 

Echinodermata 

Bryozoa 

other invertebrate phyla 

Vertebrates 

fishes 

birds 

mammals 

Figure 10.4. Species composition of Prince William Sound marine biota by major taxon.


The marine biota of Prince William Sound is a mix of species with ranges that overlap 

several biogeographic provinces. Forty-nine percent of the biota for which reliable data are 

available are northeast Pacific species, not found north or west of Bering Strait. Over one third 

of the species (43.4 %) have geographic ranges that extend into the northwestern Pacific. Fewer 

species have distributions that overlap the north Atlantic 19.6.5% and/or Arctic 16.9%. The 

biogeographic affinities vary greatly among the phyla. 

We caution users of these data sets that in spite of our best efforts the information is only 

as good as our sources. We were unable to obtain data, or found few reliable records, for five 

major groups of animals: sponges, Anthozoa, Nematoda, Oligochaeta , Hirudinea, Ostracoda and 

insects. Two factors account for this, lack of taxonomic experts, and the need for specialized 

collection and preservation techniques. Further, even among several well-known taxonomic 

groups, we found few sources of authentic information and so we have had to rely on 

unpublished reports and manuscripts. We hope that using these data sets will not perpetuate 

erroneous records. In critical situations, users are advised to rely on museum records and 

consultation with other experts. 

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