Biological Invasions of Cold-Water Coastal Ecosystems - Aquatic ...
Biological Invasions of Cold-Water Coastal Ecosystems - Aquatic ...
Biological Invasions of Cold-Water Coastal Ecosystems - Aquatic ...
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BIOLOGICAL INVASIONS OF COLD-WATER COASTAL ECOSYSTEMS:<br />
BALLAST-MEDIATED INTRODUCTIONS IN<br />
PORT VALDEZ / PRINCE WILLIAM SOUND, ALASKA<br />
PRESENTED TO:<br />
FINAL PROJECT REPORT<br />
15 March 2000<br />
Regional Citizens’ Advisory Council <strong>of</strong> Prince William Sound<br />
P.O. Box 3089, 154 Fairbanks Drive<br />
Valdez, AK 99686<br />
USA<br />
Telephone: 907-835-5957 Fax: 907-835-5926<br />
Email: www.pwsrcac.org<br />
U.S. Fish and Wildlife Service<br />
PO Box 1670, 43655 Kalifornski Beach Road<br />
Soldatna, AK 99669<br />
National Sea Grant Program<br />
Washington, DC<br />
Alaska Sea Grant Program, University <strong>of</strong> Alaska Fairbanks<br />
Oregon Sea Grant Program, Oregon State University<br />
SeaRiver Maritime, Inc.<br />
ARCO Marine, Inc.<br />
British Petroleum, Inc.<br />
American Petroleum Institute<br />
Alyeska Pipeline Company<br />
PRESENTED BY:<br />
Lead Principal Investigators:<br />
• Dr. Anson H. Hines, Ph.D., Marine Ecologist<br />
• Dr. Gregory M. Ruiz, Ph.D., Marine Ecologist<br />
Smithsonian Environmental Research Center<br />
P.O. Box 28, 647 Contees Wharf Road<br />
Edgewater, MD 21037-0028<br />
USA<br />
Telephone: 301-261-4190 ext. 208 (Hines), 227 (Ruiz) Fax: 301-261-7954<br />
Email: hines@serc.si.edu, ruiz@serc.si.edu
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Co-Investigators:<br />
• Dr. John Chapman, Ph.D., Marine Biologist<br />
• Dr. Gayle I. Hansen, Ph.D., Marine Phycologist<br />
Hatfield Marine Science Center<br />
Oregon State University<br />
Newport, OR 97365-5296<br />
USA<br />
• Dr. James T. Carlton, Ph.D., Director<br />
Maritime Studies Program<br />
Mystic Seaport<br />
Williams College<br />
Mystic, CT 06355-0990<br />
USA<br />
• Ms. Nora Foster, M.S., Curator<br />
University <strong>of</strong> Alaska Museum<br />
University <strong>of</strong> Alaska Fairbanks<br />
Fairbanks, AK 99775<br />
USA<br />
• Dr. Howard M. Feder, Ph.D., Pr<strong>of</strong>essor<br />
Institute <strong>of</strong> Marine Sciences<br />
University <strong>of</strong> Alaska Fairbanks<br />
Fairbanks, AK 99775<br />
USA<br />
Contributing Taxonomic Experts:<br />
• Dr. Jeffrey Cordell. School <strong>of</strong> Fisheries, University <strong>of</strong> Washington<br />
• Dr. Jeffrey Goddard, University <strong>of</strong> California, Santa Barbara<br />
• Dr. Paul W. F<strong>of</strong>on<strong>of</strong>f, Smithsonian Environmental Research Center<br />
• Dr. Jerry Kudenov, University <strong>of</strong> Alaska, Anchorage<br />
• Dr. Gretchen Lambert, Friday Harbor Laboratories, University <strong>of</strong> Washington<br />
• Dr. Claudia Mills, Friday Harbor Laboratories, University <strong>of</strong> Washington<br />
• Ms. Lise Schickel, University <strong>of</strong> California, Santa Barbara<br />
• Dr. Judith Winston, Virginia Museum <strong>of</strong> Natural History<br />
• Ms. Lea Ann Henry, University <strong>of</strong> Toronto<br />
• Ms. Olga Kalata, School <strong>of</strong> Fisheries, University <strong>of</strong> Washington<br />
<strong>Biological</strong> Technicians:<br />
• Melissa Frey, Smithsonian Environmental Research Center<br />
• George Smith, Smithsonian Environmental Research Center<br />
• Max Hoberg, University <strong>of</strong> Alaska Fairbanks<br />
Field Assistants:<br />
• James Wade, Valdez
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• Stephan Benda, Valdez<br />
• Todd Miller, Hatfield Marine Science Center, Oregon State University<br />
Ballast <strong>Water</strong> Exchange Experiments and Plankton Analysis:<br />
• Safra Altman, Smithsonian Environmental Research Center<br />
• Sara Chaves, Smithsonian Environmental Research Center<br />
• Tami Huber, Smithsonian Environmental Research Center<br />
• Dani Lipski, Smithsonian Environmental Research Center<br />
• Kate Murphy, Smithsonian Environmental Research Center<br />
• Kimberly Philips, Smithsonian Environmental Research Center<br />
• Brian Steves, Smithsonian Environmental Research Center
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EXECUTIVE SUMMARY<br />
PROJECT OVERVIEW<br />
This study assesses the risk <strong>of</strong> biological invasion by nonindigenous species (NIS) associated<br />
with oil tanker traffic and ballast water management for Port Valdez / Prince William<br />
Sound (PWS), Alaska. This study included 8 major components:<br />
• Review <strong>of</strong> risk factors for NIS invasions and ship-mediated transfer <strong>of</strong> species relevant to<br />
PWS, a high latitude, cold-water marine ecosystem.<br />
• Analysis <strong>of</strong> ballast water delivery patterns and plankton communities associated with ballast<br />
water on tankers that arrived to PWS.<br />
• Experimental analysis <strong>of</strong> initial survivorship <strong>of</strong> ballast water organisms in temperaturesalinity<br />
combinations typical <strong>of</strong> receiving waters <strong>of</strong> Port Valdez.<br />
• Experimental measurements <strong>of</strong> the effect <strong>of</strong> ballast water exchange and voyage duration on<br />
plankton communities arriving on tankers to PWS.<br />
• Characterization <strong>of</strong> organisms fouling hulls and in sea chests <strong>of</strong> crude oil tankers.<br />
• Characterization <strong>of</strong> organisms in sediments <strong>of</strong> tanker ballast tanks.<br />
• Determination <strong>of</strong> NIS established within Alaska, as detected by field surveys and reviews <strong>of</strong><br />
existing collections and literature conducted by experienced naturalists and taxonomic<br />
experts.<br />
• Analysis <strong>of</strong> the biodiversity <strong>of</strong> PWS.<br />
This study advances our understanding <strong>of</strong> invasion processes in many significant ways.<br />
• Our study provides the most comprehensive analysis worldwide <strong>of</strong> the abundance and<br />
taxonomic composition <strong>of</strong> plankton communities in the segregated ballast water <strong>of</strong> tankers as<br />
well as domestic ballast transfer by any vessel type.<br />
• We have undertaken an ambitious set <strong>of</strong> experimental and quantitative measures to (a)<br />
compare directly, for the first time, the relative efficiency <strong>of</strong> exchange methods (Empty–<br />
Refill and Flow-Through) for any vessel type or taxon, and (b) the effect <strong>of</strong> voyage duration<br />
on plankton survivorship in the ballast water <strong>of</strong> oil tankers.<br />
• We provide the first synthesis <strong>of</strong> NIS known in Alaska, resulting from an extensive literature<br />
review and field-based surveys.<br />
• The large scope <strong>of</strong> this study provides an unusually comprehensive analysis <strong>of</strong> the risks,<br />
mechanisms, and patterns <strong>of</strong> invasion in PWS.<br />
The project represents a cooperative and successful partnership <strong>of</strong> industry, citizen,<br />
agency, and scientific groups. This strong cooperative program addresses critical gaps in our<br />
understanding <strong>of</strong> invasion risks, as well as facilitates information exchange and participation<br />
among a broad spectrum <strong>of</strong> industry, citizen, agency, and scientific groups.<br />
• From a science perspective, this program results in a comprehensive analysis <strong>of</strong> invasion<br />
processes and risks for PWS, representing the first such study in the world for a high-latitude<br />
/ cold-water marine ecosystem.
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• From an industry and management perspective, this program assesses the effectiveness and<br />
trade-<strong>of</strong>fs involved for various management strategies that are now required in Prince<br />
William Sound, and are being promoted on a national and international scale.<br />
• From a public perspective, this program disseminates findings and serves as a key source <strong>of</strong><br />
information, especially through groups like the Smithsonian Environmental Research Center,<br />
the Regional Citizens’ Advisory Council <strong>of</strong> Prince William Sound, U.S. Fish & Wildlife<br />
Service, and NOAA Sea Grant.<br />
RESULTS<br />
Background<br />
<strong>Biological</strong> invasions <strong>of</strong> marine ecosystems in Alaska are a major environmental concern.<br />
• <strong>Biological</strong> invasions <strong>of</strong> coastal bays and estuaries are common throughout the world and are<br />
having significant ecological and economic impacts.<br />
• High-latitude / cold-water regions are also subject to biological invasions by many species<br />
with potential ecological and economic consequences similar to those reported for more<br />
temperate latitudes.<br />
• Transport <strong>of</strong> coastal planktonic organisms in ballast water <strong>of</strong> commercial ships appears to be<br />
the major source <strong>of</strong> new invasions worldwide in recent years.<br />
• Tankers arriving to Port Valdez release the third largest volume <strong>of</strong> ballast water <strong>of</strong> any U.S.<br />
port.<br />
BW Delivery Patterns and <strong>Biological</strong> Characteristics<br />
A large quantity <strong>of</strong> ballast water arrives to PWS in oil tankers.<br />
• For the past decade, tanker arrivals to Port Valdez have averaged 713 ships per year.<br />
• Tankers arriving to PWS in 1998 carried an estimated average <strong>of</strong> 65,775m 3 <strong>of</strong> total ballast<br />
water, including both segregated (non-oily) and nonsegregated (or oily) ballast water.<br />
• Segregated ballast water comprised an average <strong>of</strong> 54.7% <strong>of</strong> the total ballast water arriving to<br />
PWS in tankers.<br />
• Overall, an estimated 17,000,000 m 3 <strong>of</strong> segregated ballast water (an average <strong>of</strong> 32,715 m 3 per<br />
arrival) was discharged into PWS by oil tankers in 1998.<br />
Most ballast water delivered to PWS by crude oil tankers originates from U.S. domestic<br />
ports.<br />
• Tankers arriving directly from western U.S. ports accounted for 95.8% <strong>of</strong> the total tanker<br />
traffic, and 96% <strong>of</strong> the total segregated ballast water delivered by tankers, to PWS in 1998.<br />
• Arrivals from Puget Sound, San Francisco, and Long Beach comprised approximately 82.7%<br />
<strong>of</strong> all tanker traffic, as well as 86% <strong>of</strong> all segregated ballast water delivered by tankers, to<br />
PWS in 1998.<br />
• Most (69.6%) <strong>of</strong> the tankers arriving to Port Valdez from overseas came directly from Korea<br />
in 1998.<br />
• Tankers arriving from domestic ports transfer ballast water directly from that port to PWS,<br />
whereas foreign arrivals have replaced coastal ballast water with open-ocean exchange prior<br />
to their arrival.
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The voyage duration <strong>of</strong> tankers arriving to Port Valdez is relatively short compared to<br />
traffic arriving at other commercial ports, where invasions are common.<br />
• Ballast water spent an average <strong>of</strong> 6.6 days in the ballast tanks <strong>of</strong> oil tankers before arrival to<br />
Port Valdez, ranging between 4.8 to 10.2 days.<br />
A large quantity <strong>of</strong> planktonic organisms is released into PWS with segregated ballast<br />
water from oil tankers.<br />
• An average <strong>of</strong> 12,637 total organisms per m 3 (excluding chain-forming diatoms) was<br />
measured in our ballast water samples from 169 tanker arrivals, including those from both<br />
domestic and foreign source ports.<br />
• Overall, we estimate that roughly 264 billion organisms were delivered to PWS in the<br />
segregated ballast water <strong>of</strong> oil tankers during 1998.<br />
• Importantly, these estimates include only the largest plankton and miss many small<br />
planktonic organisms (e.g., bacteria, viruses, and other microorganisms), that would likely<br />
increase overall densities many fold.<br />
The abundance <strong>of</strong> planktonic organisms was greater in segregated ballast water from<br />
domestic source ports compared to that from foreign source ports.<br />
• Total density (across all taxonomic groups) <strong>of</strong> organisms was greatest on average in<br />
segregated ballast water from domestic arrivals compared to foreign arrivals.<br />
• Average densities <strong>of</strong> most taxonomic groups were 10- to 100-fold greater in segregated<br />
ballast water from domestic versus foreign sources.<br />
• The magnitude <strong>of</strong> density differences between domestic and foreign sources was much less<br />
for copepods and solitary diatoms.<br />
• Din<strong>of</strong>lagellates were a notable exception to the general pattern, as average density was<br />
greatest in ballast water <strong>of</strong> the foreign arrivals.<br />
Significant variation existed in abundance <strong>of</strong> taxonomic groups in the segregated ballast<br />
water arriving from the major source ports.<br />
• Total density <strong>of</strong> organisms was lowest on average in ballast water from foreign arrivals<br />
compared to arrivals from each <strong>of</strong> the three major domestic ports (Puget Sound, San<br />
Francisco, and Long Beach)<br />
• Total density on average declined among the four major ports with increasing voyage<br />
duration.<br />
• In contrast, the greatest average densities for individual taxonomic groups (e.g., protozoans,<br />
brachyuran crabs, and bryozoans) did not always correspond to the shortest voyage duration.<br />
The abundance <strong>of</strong> plankton arriving in segregated ballast water from the major domestic<br />
ports varied both spatially and temporally.<br />
• The greatest densities occurred for all taxonomic groups, individually and combined, during<br />
the spring and summer months.<br />
• However, the timing <strong>of</strong> peak densities differed among taxonomic groups and among source<br />
ports.
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• The magnitude <strong>of</strong> seasonal variation in plankton densities also differed among source ports,<br />
being greatest for Puget Sound and San Francisco Bay compared to Long Beach.<br />
• Furthermore, significant annual variation also existed in the densities <strong>of</strong> plankton arriving to<br />
PWS from each <strong>of</strong> the major domestic source ports.<br />
NIS are present in the segregated ballast water released by oil tankers in PWS.<br />
• We identified 14 different nonindigenous species (13 crustaceans and 1 fish) arriving to Port<br />
Valdez in the ballast water <strong>of</strong> oil tankers.<br />
• To date, all <strong>of</strong> these identified NIS have been in ballast water from San Francisco Bay or<br />
Long Beach.<br />
• Importantly, these numbers are clearly underestimates, since only a subset <strong>of</strong> the plankton<br />
can be identified to species and only the largest fraction <strong>of</strong> planktonic organisms were<br />
included in our analyses.<br />
Organisms discharged in tanker ballast water, including known NIS, have high potential <strong>of</strong><br />
initial survival in the salinity-temperature conditions <strong>of</strong> Port Valdez and PWS.<br />
• Seasonal cycles <strong>of</strong> salinities and temperatures in Port Valdez waters encompass the range <strong>of</strong><br />
salinities and temperatures <strong>of</strong> arriving ballast waster, providing a good match between source<br />
ports and receiving waters.<br />
• Laboratory experiments indicate that a wide range <strong>of</strong> ballast water species (including some<br />
NIS) can survive the salinity and temperature conditions <strong>of</strong> Port Valdez upon initial<br />
discharge from tankers.<br />
Other Mechanisms <strong>of</strong> NIS Transport by Tankers<br />
Tankers also transfer organisms that are not in ballast water and that may become<br />
established in PWS.<br />
• Tanker hulls and sea chests sampled in dry dock sometimes carried a diverse array <strong>of</strong> fouling<br />
and nektonic organisms, including several NIS.<br />
• Sediment taken into ballast tanks during ballasting in shallow ports sometimes carried<br />
diverse and abundant bottom-dwelling organisms, including reproductive adult individuals.<br />
Ballast <strong>Water</strong> Exchange Experiments<br />
Preliminary experimental results suggest that ballast water exchange is as effective for oil<br />
tankers as for other vessel types.<br />
• Initial analyses suggest roughly 80-99% <strong>of</strong> the resident water is replaced per ballast water<br />
exchange event.<br />
• The efficacy <strong>of</strong> exchange appears to differ between exchange method, with Empty–Refill<br />
Exchange replacing the greatest proportion <strong>of</strong> water.<br />
• The efficacy <strong>of</strong> exchange also appears to differ among taxa.<br />
• Importantly, all analyses for the exchange experiments are still underway, and final results /<br />
conclusions are therefore pending project completion (anticipated in June 2000).
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Summary <strong>of</strong> NIS in Prince William Sound and Alaska<br />
A diverse array <strong>of</strong> NIS have been introduced into PWS and Alaska.<br />
• There are 24 species <strong>of</strong> NIS plants and animals in marine and estuarine ecosystems in<br />
Alaska, including 15 species recorded in PWS.<br />
• These NIS are taxonomically diverse and occupy a wide range <strong>of</strong> ecological niches and<br />
habitats, although there appear to be more NIS associated with boat harbors and with<br />
aquaculture activities.<br />
• Our focal taxonomic collections provided the first records <strong>of</strong> 7 NIS in Alaska, including<br />
some species that appear to be very recent introductions.<br />
• Many <strong>of</strong> the Alaskan NIS have larval stages which could be transported in ballast water.<br />
• None <strong>of</strong> the Alaskan NIS is clearly associated with ballast water <strong>of</strong> oil tankers as a primary<br />
mechanism <strong>of</strong> introduction, even though many NIS are frequently found in ballast water<br />
arriving to Port Valdez.<br />
• Instead, the transfer <strong>of</strong> NIS may have resulted from any one <strong>of</strong> multiple transfer mechanisms,<br />
including ballast water, ship fouling communities, and aquaculture.<br />
• Finally, it is important to note that many additional Alaskan marine species are cryptogenic<br />
(possibly introduced), as the historical baseline <strong>of</strong> biogeographic and taxonomic information<br />
is very limited for this biota. For example, we identified at least 29 cryptogenic species in<br />
Alaska (including 24 in PWS), exhibiting either wide global distributions <strong>of</strong>ten associated<br />
with spread by early shipping traffic or a variety <strong>of</strong> characteristics common to NIS.<br />
Biodiversity <strong>of</strong> Prince William Sound<br />
Taxonomy and biogeography <strong>of</strong> species in Alaskan marine ecosystems have received poor<br />
levels <strong>of</strong> study and understanding.<br />
• We discovered 10 new, previously undescribed species, as well as recorded range extensions<br />
for 74 other species from a diverse array <strong>of</strong> taxonomic groups.<br />
• It is now apparent that a large portion <strong>of</strong> many major groups remain undocumented, as well<br />
as cryptogenic in origin, due to limited surveys and historical analysis <strong>of</strong> the Alaskan biota.<br />
We have now initiated a biodiversity data base for marine species in PWS.<br />
• We have established a comprehensive data base for marine invertebrates in PWS.<br />
• The scope <strong>of</strong> this data base will be expanded to include algae, fish, mammals and birds in the<br />
next year.<br />
CONCLUSIONS<br />
Multiple risk factors exist that favor the establishment <strong>of</strong> NIS in PWS.<br />
• Approximately 550 tankers currently arrive per year to PWS and release an estimated<br />
17,000,000 metric tons <strong>of</strong> segregated ballast water.<br />
• Tankers repeatedly deliver ballast water from the same, limited source ports, providing<br />
repeated inoculations <strong>of</strong> the same species.
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• The voyage duration <strong>of</strong> these tankers is usually short (3-7 days), favoring high survivorship<br />
<strong>of</strong> transported plankton and resulting in the dense inoculation <strong>of</strong> competent organisms into<br />
PWS.<br />
• Environmental conditions <strong>of</strong> source ports match those in PWS for some portions <strong>of</strong> the year,<br />
and many organisms arriving in ballast water can tolerate conditions in receiving waters.<br />
• Most (95.6%) <strong>of</strong> arriving tankers do not undergo ballast water exchange, a process which can<br />
limit the transfer rate <strong>of</strong> NIS.<br />
• A large number (tens-to-hundreds) <strong>of</strong> NIS are known from the domestic ports that are the<br />
source <strong>of</strong> unexchanged ballast water arriving to PWS in oil tankers.<br />
• NIS are present in this domestic ballast water arriving to PWS in oil tankers.<br />
Ballast water exchange appears effective at reducing resident plankton on tankers,<br />
although a risk <strong>of</strong> invasion still exists.<br />
• Ballast exchange experiments suggest that tankers arriving to Port Valdez from foreign ports<br />
have reduced resident coastal organisms by > 90% through the current exchange practices.<br />
• Abundance <strong>of</strong> coastal organisms was 10-100 fold lower for oil tankers that were foreign<br />
arrivals (that underwent ballast water exchange) compared to domestic arrivals (that do not<br />
undergo exchange).<br />
• Although roughly equivalent to efficacy <strong>of</strong> exchange estimated for other vessel types, both<br />
data sets suggest that tens to hundreds <strong>of</strong> thousands <strong>of</strong> organisms/ship still arrive with<br />
exchanged ballast water.<br />
Alaskan waters, and those <strong>of</strong> PWS, are susceptible to invasion by NIS.<br />
• It is now evident that a diverse array <strong>of</strong> taxa have become established in Alaska and PWS.<br />
• These NIS occupy a broad range <strong>of</strong> marine and estuarine habitats.<br />
The number <strong>of</strong> marine NIS in Alaska appears to be significantly lower than other marine<br />
ecosystems at lower latitude.<br />
• Our surveys <strong>of</strong> PWS and Alaska were intensive and failed to detect many NIS known from<br />
the domestic source ports <strong>of</strong> oil tankers.<br />
• However, the limited historical record and scope <strong>of</strong> past surveys limits direct comparisons<br />
with low latitude marine ecosystems, for which extensive surveys and knowledge have been<br />
developed over decades to centuries <strong>of</strong> biological research.<br />
In general, the poor resolution <strong>of</strong> taxonomic and biogeographic data in Alaskan marine<br />
ecosystems is a substantial impediment for analysis <strong>of</strong> environmental impacts.<br />
• To date, we have been able to provide only a minimum estimate <strong>of</strong> NIS, as many species<br />
remain undescribed or cryptogenic until further analysis.<br />
• Assessment <strong>of</strong> other environmental impacts, such as oil spills, may also be limited without<br />
adequate baseline data on species composition and abundance.<br />
• We recommend a program <strong>of</strong> standardized surveys across multiple sites in PWS and Alaska<br />
to both improve the existing knowledge <strong>of</strong> NIS and provide a regional baseline <strong>of</strong> data.
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Acknowledgments<br />
Funding for this study was provided by Regional Citizens’ Advisory Council <strong>of</strong> Prince<br />
William Sound (RCAC), US Fish & Wildlife Service (USF&WS), and the NOAA National Sea<br />
Grant Program through Oregon State University (OSU) and University <strong>of</strong> Alaska Fairbanks<br />
(UAF). In-kind support and participation were provided by the Smithsonian Environmental<br />
Research Center (SERC), Maritime Studies Program/Mystic Seaport with Williams College,<br />
Alyeska Pipeline Service Company, and the oil shippers, particularly SeaRiver Maritime, ARCO<br />
Marine and British Petroleum (BP). Additional support for expanded ballast water exchange<br />
experiments was provided by the American Petroleum Institute, SeaRiver and ARCO,<br />
USF&WS, and U.S. Coast Guard.<br />
This project was stimulated and encouraged by the Nonindigenous Species Working<br />
Group <strong>of</strong> RCAC, chaired by Robert Benda (Prince William Sound Community College) and<br />
Gary Sonnevil (USF&WS). We thank Joel Kopp <strong>of</strong> RCAC for his able oversight and friendly<br />
assistance at all stages <strong>of</strong> this project, and Marilyn Leland and Linda Hyce <strong>of</strong> RCAC for their<br />
support. We are grateful to Rex Brown and the managers and staff <strong>of</strong> the Alyeska Pipeline<br />
Service Company and the Valdez Marine Terminal for access and help in sampling. The Masters,<br />
Officers, and crew members <strong>of</strong> the tankers gave willingly <strong>of</strong> their time and help; and we<br />
especially thank those <strong>of</strong> the ships SR Baytown, SR Long Beach, and SR Benicia, ARCO<br />
Independence, and ARCO Spirit for their extensive help on ballast water exchange experiments.<br />
Thanks also to W.P. (Pete) Rupp <strong>of</strong> SeaRiver Maritime and Victor Goldberg <strong>of</strong> ARCO Marine<br />
for their support <strong>of</strong> the exchange experiments. We especially thank the following agents in<br />
Valdez for their logistical assistance, advice and suggestions, and support: Kurt Hallier and<br />
Wayne Brandenburger <strong>of</strong> ARCO Marine; Bill Deppe, Phil Eichenberger, and John Poulos <strong>of</strong><br />
SeaRiver Maritime, and Tom Colby <strong>of</strong> BP. The full cooperation <strong>of</strong> the oil companies operating<br />
these ships is gratefully acknowledged.<br />
Melissa Frey and George Smith <strong>of</strong> SERC provided key technical assistance for sampling<br />
ships, processing samples, conducting experiments, analysis, and many other crucial components<br />
at SERC’s laboratories in Valdez and in Maryland. Exchange experiments were assisted on<br />
board tankers by additional technicians <strong>of</strong> the SERC <strong>Invasions</strong> Biology Program: Linda<br />
McCann, Kim Philips, Safra Altman, Lynn Takata, Cathleen Coss, Kate Murphy, Dani Lipski,<br />
Laura Rodriguez and Brian Steves. Sorting <strong>of</strong> samples was assisted by Melissa Frey, George<br />
Smith, Sara Chaves, Kim Philips, Dani Lipski and Safra Altman. Data management and analysis<br />
were assisted by: Safra Altman, Melissa Frey, Midge Kramer, Kim Philips, George Smith, and<br />
Brian Steves.<br />
For assistance on the NIS cruises in Prince William Sound, special appreciation goes to:<br />
Todd Miller, for his assistance on the trip; Capt. Richard Vodicka who skippered the F/V<br />
Kristina in 1998 and helped in many ways large and small throughout the survey; Milos Falta,<br />
owner <strong>of</strong> the F/V Kristina and skipper during the 1999 surveys; Joel Kopp, RCAC, for loan <strong>of</strong><br />
his zodiac, searching for a charter, and other logistical and support assistance; and the<br />
community <strong>of</strong> Tatitlek for housing the survey team, especially village leader Gary Kompk<strong>of</strong>f and<br />
boat operator/guide Steve Totem<strong>of</strong>f for logistical assistance in the survey <strong>of</strong> Valdez Arm and<br />
Tatitlek areas. Peter Jordan and Bryna Dunaway, high school students in Oregon, assisted in<br />
curating the dried plant collection. Mike Stekoll <strong>of</strong> the University <strong>of</strong> Alaska provided
page xi<br />
microscopes for the study. Lavern Weber, Director <strong>of</strong> the Hatfield Marine Science Center<br />
provided <strong>of</strong>fice and laboratory space for G.I. Hansen. These people all helped make the cruises a<br />
success.<br />
For assistance with the 1998 & 1999 Fouling Plate Surveys, we thank Gary Sonnevil (US<br />
Fish & Wildlife Service), Peter Armato (US Park Service and Alaska SeaLife Center), Chuck<br />
Pratt (Armin F. Koenig Fish Hatchery), Ken Morgan (Valdez Fish Hatchery), Andrea Tesch<br />
(Nuremburg Fish Hatchery), Stan Stephens (Stan Stephens Tours), and the pilots <strong>of</strong> Cordova Air<br />
Service for field assistance in deploying and retrieving plates. We appreciate use <strong>of</strong> facilities and<br />
further assistance from Alyeska Pipeline Service Company, Prince William Sound Science<br />
Center, and Prince William Sound Community College. In addition, we thank Conrad and<br />
Carmen Field for their help at Homer, especially for showing us “an unusual sea star” (Asterias<br />
amurensis).<br />
In addition to participation in the project by taxonomic experts (see cover page), species<br />
identifications were assisted by Klaus Ruetzler for sponges; Frank Ferrari, Olga Kalata and Chad<br />
Walter for copepods; Judy Winston for bryozoans; Lea Ann Henry for hydroids. Charles<br />
Lambert assisted with field collections <strong>of</strong> ascidians during the 1999 field survey.<br />
The cooperative support by all <strong>of</strong> these individuals and organizations made this project<br />
possible.
page xii<br />
Title Page<br />
Executive Summary<br />
Acknowledgements<br />
Table <strong>of</strong> Contents<br />
Table <strong>of</strong> Contents<br />
page<br />
i<br />
iii<br />
ix<br />
xi<br />
1. Introduction 1-1<br />
1A. Project Goals 1-1<br />
1B. Structure & History <strong>of</strong> Project 1-1<br />
1C. Background 1-2<br />
1C1. Invasive Species & Ballast <strong>Water</strong> 1-2<br />
1C2. NIS in High Latitude/<strong>Cold</strong> <strong>Water</strong> <strong>Ecosystems</strong> 1-2<br />
1C3. Risk Factors for Prince William Sound 1-5<br />
2. Ballast <strong>Water</strong> Delivery Patterns 2-1<br />
3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong> in Oil Tankers 3-1<br />
4. Predicting Initial Survival <strong>of</strong> Ballast <strong>Water</strong> Organisms 4-1<br />
5. Ballast <strong>Water</strong> Exchange Experiments 5-1<br />
6. Organisms in Sediments <strong>of</strong> Tanker Ballast Tanks 6-1<br />
7. Fouling Organisms on Hulls and in Sea Chests <strong>of</strong> Oil Tankers 7-1<br />
8. Summary <strong>of</strong> NIS in Prince William Sound and Alaska 8-1<br />
9. NIS Surveys 9A-1<br />
9A. Overview 9A-1<br />
9B. Rapid Community Assessments 9B-1<br />
9C. Focal Taxonomic Collections 9C1-1<br />
9C1. Marine Plants 9C1-1<br />
9C2. Planktonic Medusae, Ctenophores, and Pelagic Molluscs 9C2-1<br />
9C3. Polychaete Worms 9C3-1<br />
9C4. Peracaridan Crustaceans 9C4-1<br />
9C5. Copepod Crustaceans 9C5-1<br />
9C6. Decapod Crustaceans 9C6-1<br />
9C7. Shelled Molluscs 9C7-1<br />
9C8. Opisthobranch Molluscs 9C8-1
page xiii<br />
9C9. Echinoderms 9C9-1<br />
9C10. Ascidians 9C10-1<br />
9D. Fouling Communities 9D-1<br />
9E. Museum, Voucher and Reference Specimens <strong>of</strong> Existing Collection 9E-1<br />
10. Biodiversity <strong>of</strong> Prince William Sound 10-1
Chapt 1. Introduction, page 1- 1<br />
Chapter 1. Introduction<br />
Anson H. Hines, Smithsonian Environmental Research Center<br />
Gregory M. Ruiz, Smithsonian Environmental Research Center<br />
1A. Project Goals<br />
The overall goal <strong>of</strong> this project was to assess the risk <strong>of</strong> biological invasion by<br />
nonindigenous species (NIS) introduced into Port Valdez / Prince William Sound, Alaska.<br />
Currently, ballast water is the major vector for introductions <strong>of</strong> NIS in coastal ecosystems, where<br />
ballast-mediated biological invasions are causing severe ecological and economic impacts. While<br />
the significance <strong>of</strong> ballast-mediated invasions has focused on temperate zone ports, little<br />
consideration has been given to NIS invasions at high latitudes, despite the volume <strong>of</strong> shipping<br />
and critical importance <strong>of</strong> certain cold-water ports to the world economy and especially US<br />
energy interests. Port Valdez is a high latitude-cold water port receiving the third largest annual<br />
volume <strong>of</strong> ballast water in the USA. Moreover, our recent review <strong>of</strong> NIS invasions at high<br />
latitude (Ruiz & Hines 1997) indicates that such cold water ecosystems have been invaded by a<br />
diverse array <strong>of</strong> marine and estuarine species. The specific objectives <strong>of</strong> the project were:<br />
• To analyze the delivery patterns, biological characteristics, and management practices <strong>of</strong><br />
ballast water and other ships arriving to Port Valdez from coast-wise versus foreign voyages.<br />
• To assess viability <strong>of</strong> selected organisms arriving in tanker ballast water to Port Valdez.<br />
• To conduct experiments on the effectiveness <strong>of</strong> ballast water exchange procedures <strong>of</strong> tankers.<br />
• To evaluate organisms occurring in entrained sediments at the bottom <strong>of</strong> ballast tanks <strong>of</strong><br />
crude oil tankers.<br />
• To evaluate fouling organisms on hulls and in sea chests <strong>of</strong> tankers as potential sources <strong>of</strong><br />
NIS.<br />
• To analyze and search for NIS currently invading or already established in coastal waters <strong>of</strong><br />
south central Alaska, using literature searches, an array <strong>of</strong> field sampling methods (field<br />
collections, fouling plates, and plankton sampling), and examination <strong>of</strong> existing preserved<br />
samples from Prince William Sound.<br />
The purpose <strong>of</strong> this report is to summarize the research conducted during 1997-1999 to<br />
assess the risk <strong>of</strong> biological invasions in Prince William Sound, especially with regard to oil<br />
tankers as a vector for transporting NIS into marine ecosystems. Progress during the project was<br />
reported in Ruiz & Hines, 1997 and Hines et al., 1998. Modified elements <strong>of</strong> the earlier reports<br />
are included in the present report, so as to provide a complete overview <strong>of</strong> the project within one<br />
document. Certain limited elements <strong>of</strong> research will be completed during 2000, including further<br />
analysis <strong>of</strong> existing collections at the University <strong>of</strong> Alaska Museum and Institute <strong>of</strong> Marine<br />
Sciences, and work-up <strong>of</strong> sample from ballast water exchange experiments conducted during<br />
summer 1999. These last elements will be reported separately upon completion.<br />
1B. Structure & History <strong>of</strong> Project<br />
This research project has built upon a Pilot Study conducted in 1997 (see Ruiz & Hines,<br />
1997), and includes an expanded scope <strong>of</strong> work conducted during 1998-1999. The multi-faceted<br />
approach to the research required a team <strong>of</strong> diverse CoPrincipal Investigators and subcontracted<br />
taxonomic experts. Drs. Anson Hines and Gregory Ruiz (SERC) have served as over-all project<br />
leaders, providing over-all administrative and scientific oversight for the team. In addition, Drs.<br />
Ruiz and Hines (SERC) had primary responsibilities for: analysis <strong>of</strong> ballast water delivery
Chapt 1. Introduction, page 1- 2<br />
patterns; biological characteristics <strong>of</strong> ballast water; experimental analysis <strong>of</strong> initial survival <strong>of</strong><br />
ballast water organisms; ballast water exchange experiments; analysis <strong>of</strong> ballast tank sediments<br />
and tanker hull fouling; fouling community analysis, and management <strong>of</strong> most <strong>of</strong> the focal<br />
taxonomic studies; as well many aspects <strong>of</strong> field surveys <strong>of</strong> Prince William Sound. To sample<br />
and analyze ballast water <strong>of</strong> tankers arriving to Port Valez, two <strong>Biological</strong> Technicians (Melissa<br />
Frey and George Smith, <strong>of</strong> SERC) alternately rotated at about four month intervals between<br />
SERC’s temporary laboratory established in Valdez, Alaska and the SERC <strong>Biological</strong> <strong>Invasions</strong><br />
Laboratory in Edgewater, Maryland. Ballast water exchange experiments were conducted with<br />
participation by several technicians and students from the SERC <strong>Biological</strong> <strong>Invasions</strong> Laboratory<br />
(see Acknowledgments). Co-PIs Nora Foster (UAF) and Dr. Howard Feder (UAF) had primary<br />
responsibility for analysis <strong>of</strong> existing samples in museum, reference and voucher collections in<br />
the UA Museum and UAF Institute <strong>of</strong> Marine Science. Nora Foster also participated actively in<br />
rapid community assessment surveys <strong>of</strong> Prince William Sound, and focal taxonomic analysis <strong>of</strong><br />
molluscs. Dr. Howard Feder provided oversight to subcontracted focal taxonomic analysis <strong>of</strong><br />
polychaetes. CoPI Dr. John Chapman (OSU) had primary responsibility to conduct rapid<br />
community assessment surveys <strong>of</strong> invertebrates <strong>of</strong> Prince William Sound, with focal taxonomic<br />
analysis <strong>of</strong> pericaridean crustaceans. CoPI Dr. Gayle Hansen (OSU) had primary responsibility<br />
for focal taxonomic field surveys <strong>of</strong> marine plants (especially macro-algae) <strong>of</strong> Prince William<br />
Sound. Dr. James Carlton (Mystic Seaport, Williams College) had primary responsibility for<br />
surveys <strong>of</strong> fouling communities <strong>of</strong> Prince William Sound. In addition, an array <strong>of</strong> systematic<br />
experts was subcontracted to analyze several focal taxonomic groups (see Chapter 9 below).<br />
Authorship <strong>of</strong> the chapters and subsections <strong>of</strong> this report indicate primary responsibilities for<br />
each major element.<br />
Throughout the 1997 Pilot Study and the 1998-1999 expanded phase, the project received<br />
guidance and comment from the Alaska NIS Working Group, which was organized by the<br />
RCAC <strong>of</strong> Prince William Sound and composed <strong>of</strong> academic scientists, resource managers from<br />
state and federal agencies, representatives <strong>of</strong> the oil and shipping industries, and concerned<br />
citizens <strong>of</strong> Alaska (see also Acknowledgments).<br />
Initial funding for the Pilot Study was provided by PWS RCAC, US Fish & Wildlife<br />
Service, and the US Coast Guard (Ruiz & Hines, 1997). Expanded research for the present<br />
project was extended with a proposal submitted in 1997 to the National Sea Grant Program, with<br />
co-funding from RCAC, USF&WS, Alyeska Pipeline Service Company, and in-kind support<br />
from the oil shipping companies (especially ARCO Marine, BP and SeaRiver Maritime). Funds<br />
awarded from the National Sea Grant Program were distributed to the Oregon Sea Grant<br />
Program for John Chapman at Hatfield Marine Science Center and to Alaska Sea Grant Program<br />
for Nora Foster and Howard Feder at University <strong>of</strong> Alaska Fairbanks and UA Museum.<br />
However, funding from Alaska Sea Grant was delayed for one year, so that funding was actually<br />
available beginning in 1999 and will carry through 2000. In 1998 SERC obtained supplemental<br />
funding from the American Petroleum Institute, supported by ARCO Marine and SeaRiver<br />
Maritime, to conduct ballast water exchange experiments on tankers. SERC also received<br />
further funding from USF&WS for these ballast water exchange experiments during 1999<br />
through a proposal submitted to the National Sea Grant Program. The work plan for the ballast<br />
water exchange experiments specifies that sample processing and analysis continue into 2000.<br />
SERC’s technical staff worked in close coordination with the shipping agents, masters,<br />
<strong>of</strong>ficers and crews <strong>of</strong> the oil tankers, and with Alyeska staff <strong>of</strong> the Valdez Marine Terminal. All
Chapt 1. Introduction, page 1- 3<br />
<strong>of</strong> these industry participants provided in-kind contributions and worked actively and<br />
cooperatively to assist project operations to sample ballast water and to conduct ballast water<br />
exchange experiments (see also Acknowledgments).<br />
Significantly more work than originally proposed was accomplished in nearly all<br />
components <strong>of</strong> the project. Moreover, we have been successful in gaining additional resources to<br />
support expanded elements <strong>of</strong> the project, including contributions in kind, and added external<br />
funds for ballast water exchange experiments. Most importantly, a hallmark <strong>of</strong> the project was<br />
the enthusiastic support <strong>of</strong> the project by the full array <strong>of</strong> private citizens, scientific institutions,<br />
governmental agencies, and industry, which served as cooperative partners.<br />
1C. Background<br />
1C1. Invasive Species & Ballast <strong>Water</strong><br />
<strong>Aquatic</strong> nuisance species have invaded many, perhaps most, freshwater and marine ports<br />
around the world. Ballast water from commercial shipping is increasingly recognized as the<br />
most significant vector currently for those invasions occurring (Carlton and Geller, 1993).<br />
Ballast water consists <strong>of</strong> water pumped into dedicated tanks or cargo holds/tanks for trim and<br />
stability during oceanic voyages, especially when the vessel is empty or only partially full <strong>of</strong><br />
cargo. Ballast water is usually taken from coastal water containing a rich diversity <strong>of</strong> planktonic<br />
organisms. Ballast water is <strong>of</strong>ten discharged into a receiving port prior to loading cargo,<br />
inoculating the ecosystem with exotic species. If any <strong>of</strong> the plankton are viable and become<br />
established, these non-indigenous species (NIS) can cause major ecological and economic<br />
disruption in the coastal ecosystem, with numerous examples in San Francisco Bay (Cohen and<br />
Carlton, 1995), the Great Lakes (Mills et al., 1993), Chesapeake Bay (Ruiz et al.,1999), Hawaii<br />
(Coles et al. 1999), and elsewhere (Ruiz et al., 1997). In San Francisco Bay, the rate <strong>of</strong> invasion<br />
has increased to about one new NIS invasion every 16 weeks, probably as a result <strong>of</strong> increased<br />
ballast water discharge (Cohen and Carlton, 1995). Whether the invasion is Eurasian zebra<br />
mussels in the Great Lakes, Asian clams in San Francisco Bay, or North American ctenophores<br />
in the Black Sea, impacts <strong>of</strong> ballast introductions have been devastating and irreversible. Despite<br />
the pr<strong>of</strong>ound impact <strong>of</strong> ballast-mediated invasions, the biological characteristics <strong>of</strong> ballast water<br />
and the factors that regulate invasion success are little studied and poorly understood. In the<br />
USA, biological characteristics <strong>of</strong> ballast water have only been studied in two port systems: Coos<br />
Bay (Carlton and Geller, 1993) and Chesapeake Bay (Smith et al., 1996; 1999; Ruiz et al.,<br />
unpubl. data); and in other countries the biology <strong>of</strong> ballast water has similarly received little<br />
quantitative analysis (Carlton, 1989; however see Williams et al., 1988; Hallegraeff and Bolsch,<br />
1992).<br />
1C2. NIS in High Latitude/<strong>Cold</strong> <strong>Water</strong> <strong>Ecosystems</strong><br />
Although there has been no significant analysis <strong>of</strong> NIS in polar marine ecosystems, there<br />
have been a limited number <strong>of</strong> NIS surveys in high temperate latitudes between 40 o - 60 o and a<br />
study <strong>of</strong> the Baltic Sea, which includes a major bay that extends substantially above 60 o . These<br />
studies include three regions in the northern hemisphere (Baltic Sea, Wadden Sea, and United<br />
Kingdom)(Reise et al., 1999, Lepapakoski 1984) and one region in the southern hemisphere<br />
(Tasmania/New Zealand)(Hayward 1997, R. Thresher, 1999 pers. comm.). Together, these<br />
studies clearly demonstrate that invasions are not limited to lower latitudes. The number <strong>of</strong><br />
known NIS at these locations ranges between 32 and 80 species. For each region, the species<br />
include a broad range <strong>of</strong> taxonomic groups, and some <strong>of</strong> the invasions have generated serious
Chapt 1. Introduction, page 1- 4<br />
concerns about their ecological and economic impacts. As with most invasions, the actual<br />
impacts remain unmeasured (e.g., Ruiz et al., 1999). Nonetheless, based upon reported<br />
abundances and known ecology, species such as the green crab Carcinus maenas (on the North<br />
American east and west coasts, Tasmania), the seastar Asterias amurensis (in Tasmania), and the<br />
laminarian kelp Undaria sp. appear likely to cause significant and irreversible changes.<br />
Furthermore, the cumulative effects <strong>of</strong> the entire NIS assemblage may cause many changes in<br />
ecosystem function that are not easily identified with any single invasion event (Cohen and<br />
Carlton, 1995).<br />
The numbers <strong>of</strong> NIS at high latitudes may be lower than those for temperate regions,<br />
although it is not clear whether low numbers <strong>of</strong> documented NIS reflect lack <strong>of</strong> invasion in high<br />
latitude ecosystems or lack <strong>of</strong> research focused on the invasion biology <strong>of</strong> these areas. At the<br />
outset <strong>of</strong> this study, the number <strong>of</strong> NIS documented in Alaskan waters appeared to be lower than<br />
other high latitude/cold water ecosystems with more extensive analysis, despite the extensive<br />
environmental studies associated with the Exxon Valdez oil spill in Prince William Sound and<br />
other ecological research throughout the region (Ruiz & Hines, 1997).<br />
We advanced two hypotheses why NIS have not been as evident at high latitude as at<br />
mid-latitudes:<br />
(1) NIS are truly rare at high latitude. High latitude communities may be resistant to invasion<br />
(e.g., severe seasonal stress requires specialized evolutionary adaptations not possessed by<br />
non-native species). Transport patterns may not have been conducive to inoculation.<br />
Shipping/ballast water is major source <strong>of</strong> rapidly escalating invasions in temperate latitudes,<br />
but perhaps neither the relatively recent (20 yrs) surge in tanker traffic to Alaska with very<br />
large ballast capacity, nor the current shift in tanker traffic to foreign ports has had time to<br />
produce invasions. Note, however, that NIS invasions mediated by ballast water have been<br />
common over the past 20 years in some cold temperate ports such as San Francisco Bay<br />
(Cohen and Carlton, 1995).<br />
(2) NIS are actually common at high latitude, but have not yet received concentrated study by<br />
experts <strong>of</strong> invasion biology. For example, Carlton (1979) identified some 160 NIS in San<br />
Francisco Bay and Pacific northwest coast, but as <strong>of</strong> 1995 the number <strong>of</strong> NIS documented in<br />
San Francisco Bay was 212 species (Cohen & Carlton, 1995) and is now nearly 250 species<br />
(J.T. Carlton, personal communication). Three years ago, the number <strong>of</strong> NIS in Chesapeake<br />
Bay was considered to be only about 25 species; yet pursuant to our on-going literature<br />
search <strong>of</strong> the historical records, we have documented >140 NIS, and the list is still growing<br />
with continuing research. Despite extensive biological/ecological assessments <strong>of</strong> coastal<br />
ecosystems associated with oil spills in Alaska, NIS probably remain inadequately studied.<br />
The existing and on-going surveys from oil spill work and other studies in Prince William<br />
Sound are probably not adequate because those surveys were designed for purposes other<br />
than detecting introduced species. Also, they mainly focused on rocky shores rather than on<br />
s<strong>of</strong>t-bottom and fouling communities that are most invaded in other regions (e.g., Cohen &<br />
Carlton, 1995). Most introduced species have been discovered by taxonomic experts<br />
systematically examining specimens previously identified by non-specialists or conducting<br />
field surveys <strong>of</strong> their own.
Chapt 1. Introduction, page 1- 5<br />
1C3. Risk Factors for Prince William Sound<br />
Prince William Sound is a relatively pristine, cold water ecosystem at high latitude.<br />
Approximately 20% <strong>of</strong> US domestic oil production is shipped from Port Valdez at the head <strong>of</strong><br />
the Sound (Fig. 1.1). Tankers arriving to Port Valdez come primarily from domestic source ports<br />
<strong>of</strong> the west coast <strong>of</strong> North America and Hawaii; but in the past three years tankers also have been<br />
traveling from foreign ports, especially in eastern Asia and rarely other locations (Fig. 1.2).<br />
Tankers arriving to Prince William Sound discharge two types <strong>of</strong> ballast water: (1) Segregated<br />
ballast water from tanks dedicated solely to ballast water and (2) non-segregated ballast water<br />
from tanks which are used to carry petroleum products. Approximately 20 million metric tons <strong>of</strong><br />
segregated ballast water are discharged annually by tankers into the port and sound, a quantity <strong>of</strong><br />
domestic ballast water that greatly exceeds the volumes <strong>of</strong> foreign ballast water released in other<br />
U.S. West Coast ports and is the third largest volume for all U.S. ports (behind port systems <strong>of</strong><br />
New Orleans and Chesapeake Bay). All non-segregated, oily water (about 50% <strong>of</strong> total)<br />
discharged by tankers in Port Valdez must pass through the Ballast <strong>Water</strong> Treatment Facility<br />
located on shore at the Valdez Marine Terminal. Effects <strong>of</strong> the treatment plant on NIS were<br />
unknown prior to our Pilot Study (Ruiz & Hines, 1997). The Pilot Study showed that nonsegregated<br />
ballast water contained few live planktonic organisms upon leaving tankers and<br />
entering the treatment plant, and that there is little risk <strong>of</strong> NIS in the discharge <strong>of</strong> water from the<br />
treatment plant (Hines et. al., in press). Accordingly, all <strong>of</strong> our analyses in subsequent research<br />
presented in this report focus on segregated ballast water.<br />
Figure 1.1.Map <strong>of</strong> Prince William Sound, Alaska, showing location <strong>of</strong> Valdez Marine Terminal.<br />
Segregated ballast water from tankers is discharged directly into the Sound/Port without<br />
treatment. This volume is many orders <strong>of</strong> magnitude greater than the ballast water released by<br />
other types <strong>of</strong> ships traveling to Prince William Sound. Release <strong>of</strong> ballast water into Prince<br />
William Sound increased markedly with the opening <strong>of</strong> the trans-Alaska Pipeline in 1977.<br />
Tankers have made more than 15,000 voyages through Prince William Sound to Port Valdez<br />
since the startup <strong>of</strong> the terminal in 1977. From 1987-1994, tanker arrivals to Valdez averaged
Chapt 1. Introduction, page 1- 6<br />
799 per year but have declined to less than 600 per year currently. Since 1996, oil shipping<br />
patterns authorized by the US Congress have changed to allow sale <strong>of</strong> crude oil on foreign as<br />
well as domestic markets. Tankers from foreign ports are required to conduct mid-ocean<br />
exchange <strong>of</strong> their coastal ballast water, which is expected to reduce numbers <strong>of</strong> organisms<br />
transported from foreign coastal ecosystems to Alaskan waters. However, the effectiveness <strong>of</strong><br />
mid-ocean exchange is poorly measured. Moreover, the greatest volume <strong>of</strong> ballast water coming<br />
to Alaska derives from domestic ports <strong>of</strong> the west coast <strong>of</strong> North America, which themselves are<br />
highly invaded by NIS. Tankers on these domestic, relatively short coast-wise voyages are not<br />
required to exchange ballast water.<br />
To determine whether the known NIS in Alaska provide an accurate indicator <strong>of</strong> the<br />
probability for biological invasions, we considered 6 factors which contribute elements <strong>of</strong> risk<br />
for invasions <strong>of</strong> Prince William Sound:<br />
1. Huge volume <strong>of</strong> ballast water. The greatest quantities <strong>of</strong> ballast water are transported by<br />
bulk cargo carriers and tankers (Carlton et al., 1995; Smith et al., 1996). Chesapeake Bay<br />
receives 10-fold more ballast water than other ports on the east and west coasts <strong>of</strong> the U.S.<br />
because <strong>of</strong> the high volume <strong>of</strong> bulkers arriving to the ports <strong>of</strong> Baltimore and Norfolk. The<br />
tanker traffic to Port Valdez releases the third largest volume <strong>of</strong> ballast water into a US port.<br />
Other things being equal, larger ballast volumes mean larger inoculations <strong>of</strong> NIS.<br />
2. Short voyage time. Our analysis <strong>of</strong> biological characteristics <strong>of</strong> ballast water arriving to<br />
Chesapeake Bay shows a marked inverse relationship between densities <strong>of</strong> organisms and<br />
length <strong>of</strong> voyage, such that ballast water after voyages <strong>of</strong> 14-24 days had more than 10-fold<br />
fewer organisms than voyages <strong>of</strong> 5-13 days (Smith et al., 1996; 1999). However, these<br />
effects may be confounded by differences in the source <strong>of</strong> ballast, which co-varies with the<br />
length <strong>of</strong> voyage (Smith et al., 1996; 1999). Voyages <strong>of</strong> tankers delivering ballast water to<br />
Prince William Sound average only 3-6 days, quite short compared to most trans-oceanic<br />
voyages that average 12-22 days. Short voyages mean that many larvae and other organisms<br />
are likely to be in good health when they are discharged (see Chapt 2 and 3 below).<br />
3. Pattern <strong>of</strong> repeated delivery from same donor locations. Although Chesapeake Bay receives<br />
about 10-fold more ballast water than does San Francisco Bay, San Francisco Bay appears to<br />
be invaded by many more ballast-mediated NIS. This greater risk could be due to San<br />
Francisco Bay receiving repeated ballast inoculations delivered from relatively fewer ports<br />
than does Chesapeake Bay. Similarly, Prince William Sound could be at increased risk not<br />
only by the large volume <strong>of</strong> ballast water, but also by the repeated inoculation <strong>of</strong> ballast from<br />
a small set <strong>of</strong> west coast ports (Fig. 1.2, Chapt 2 below).<br />
4. The match <strong>of</strong> environmental conditions <strong>of</strong> source and receiving ports. Environmental<br />
conditions in Alaska are <strong>of</strong>ten perceived as being harsh and inhospitable to most potential<br />
invaders from temperate latitudes where moderate conditions prevail. Obviously,<br />
temperature, light and other conditions during winter are indeed more extreme than those in<br />
temperate regions <strong>of</strong> North America and Asia. However, temperature-salinity conditions in<br />
Prince William Sound during spring and summer <strong>of</strong>ten approximate conditions in source<br />
ports <strong>of</strong> northwest North America, especially during productive periods <strong>of</strong> cold water<br />
up-welling. In fact, many <strong>of</strong> the native marine/estuarine species in Alaska have geographic<br />
ranges which extend to British Columbia, Washington, Oregon, and Northern California.<br />
Temperature-salinity conditions in segregated ballast water <strong>of</strong> tankers arriving from several<br />
west coast source ports is shown to be similar to the waters <strong>of</strong> Port Valdez (see Chapt 4<br />
below). In the fjords <strong>of</strong> Prince William Sound, such as Port Valdez, heavy loads <strong>of</strong>
Chapt 1. Introduction, page 1- 7<br />
suspended sediment during summer snow/glacial melt also may be a major stress on marine<br />
organisms in surface waters.<br />
Figure 1.2. Major tanker routes to Port Valdez, Alaska.<br />
5. Lack <strong>of</strong> mid-ocean exchange <strong>of</strong> ballast water delivered to Prince William Sound. Mid-ocean<br />
exchange <strong>of</strong> ballast water reduces concentrations <strong>of</strong> larvae and plankton by 50-90% (Smith et<br />
al., 1996). Exchange presumably limits the risk <strong>of</strong> invasion, as mid-ocean species are<br />
generally thought to be incapable <strong>of</strong> invading near-shore habitats. Delivery <strong>of</strong> ballast from<br />
coastal ports <strong>of</strong> the U.S. West Coast without oceanic exchange before release into Prince<br />
William Sound poses an elevated risk. Further experimental assessment <strong>of</strong> this role <strong>of</strong><br />
mid-ocean exchange in reducing plankton abundance and diversity in ballast water is<br />
presented below in “Ballast <strong>Water</strong> Exchange Experiments”. While ballast water exchange is<br />
required for tankers from foreign ports, the National Invasive Species Act <strong>of</strong> 1996 considers<br />
tankers from U.S. west coast ports to be domestic, coast-wise traffic that does not require<br />
exchange.<br />
6. High frequency <strong>of</strong> known NIS - especially those transported by ballast water - in source<br />
regions <strong>of</strong> ballast coming to Prince William Sound. Some workers consider that there may<br />
be "hotspots" <strong>of</strong> invasion or donation <strong>of</strong> NIS. If such hotspots exist, certainly San Francisco<br />
Bay and other ports <strong>of</strong> the U.S. west coast qualify as having among the highest prevalences<br />
<strong>of</strong> documented ballast-mediated invasions. These, in turn, form the sources donating much<br />
<strong>of</strong> the ballast water delivered to Prince William Sound. The 310+ known NIS <strong>of</strong> the west<br />
coast <strong>of</strong> North America vary considerably in abundance among 6 latitudinally separate<br />
regions (southern California, San Francisco Bay, northern California, Coos Bay Oregon,<br />
northwest region from the Columbia River estuary to British Columbia, and Alaska) (Ruiz &<br />
Hines, 1997). The number <strong>of</strong> known NIS varies from about 80+ species in southern<br />
California to about 40+ species in the northwest region <strong>of</strong> Washington and British Columbia,<br />
with the largest number <strong>of</strong> nearly 250 species occurring in San Francisco Bay. At each<br />
location along the west coast, NIS are common in a diverse array <strong>of</strong> taxonomic groups, with<br />
arthropods, mollusks, and annelids comprising major fractions <strong>of</strong> NIS at most locations. In<br />
several locations (San Francisco Bay, Northern California, Oregon), vascular plants and<br />
chordates also comprise major portions <strong>of</strong> the NIS. Much <strong>of</strong> the variation in number <strong>of</strong> NIS
Chapt 1. Introduction, page 1- 8<br />
probably reflects the level <strong>of</strong> study and state <strong>of</strong> knowledge for each location, especially since<br />
the highest numbers occur at two locations (San Francisco Bay and Coos Bay) where J.T.<br />
Carlton has focused his past research. Many NIS occur in several locations along the west<br />
coast, indicating that invasions by the same species have occurred widely across latitudinally<br />
separate sites. The similarity <strong>of</strong> NIS at less studied sites may be expressed as a percent<br />
overlap with NIS in well-studied San Francisco Bay. The overlap <strong>of</strong> NIS at west coast<br />
source ports with those in San Francisco Bay is high, ranging from about 60-75%. For the<br />
limited sample known for Alaska, the overlap <strong>of</strong> NIS with San Francisco Bay is substantially<br />
lower at about 25%. However, it is not clear whether this lower overlap reflects reduced<br />
compatibility with the Alaskan region or the small sample size, or both.<br />
7. History <strong>of</strong> other vectors for NIS in Prince William Sound. Although ballast water from<br />
tankers is currently a major vector for introductions <strong>of</strong> NIS in Prince William Sound, several<br />
other vectors have been, and continue to be, active in Alaska for long periods <strong>of</strong> time and in<br />
the present. These transport mechanisms include:<br />
• Ballast water from other types <strong>of</strong> ships, especially bulk carriers <strong>of</strong> such products as wood<br />
(logs, wood chips), ore, and coal, which come from foreign or domestic ports to Alaska<br />
in ballast to load. These may provide inoculations in ports nearby Prince William Sound<br />
(e.g., Homer, Seward), that may be spread by coast-wise traffic.<br />
• Fouling <strong>of</strong> ship hulls, which was especially important historically in wooden ships and<br />
before anti-fouling paints. However, fouling continues to be common, and may be<br />
especially important in coast-wise transport to and within Alaskan waters. This vector<br />
may include all types <strong>of</strong> private, recreational and commercial vessels.<br />
• Intentional and accidental release from aquaculture activities. Both species that are<br />
cultured and species that may be coincident with the aquaculture (including disease<br />
organisms) may be released. Oyster culture, salmon culture, and cultured herring roe on<br />
kelp are especially common and potential sources <strong>of</strong> NIS in Prince William Sound.<br />
Mussel culture may be initiated in the area.<br />
• Fishery release has occurred commonly in the past through efforts <strong>of</strong> state and federal<br />
agencies to improve stocks.<br />
• Aquarium and pet trades have resulted in NIS invasions at many places around the world.<br />
• Large-scale changes in current patterns may transport NIS into Alaskan waters. During<br />
1998 the very strong El Niño along the eastern Pacific may have brought warm-water<br />
species much further north than their typical distribution. These shifts may also allow<br />
species transported by human activities to become established.<br />
In many ecosystems, NIS invade over time as a series <strong>of</strong> vectors shift in importance, and this<br />
accumulation <strong>of</strong> NIS can result in major changes in the diversity and function <strong>of</strong> coastal<br />
ecosystems (Cohen & Carlton 1995).<br />
1D. References<br />
Carlton, J.T. 1979. History, biogeography, and ecology <strong>of</strong> the introduced marine and estuarine<br />
invertebrates <strong>of</strong> the Pacific coast <strong>of</strong> North America. Ph.D. Thesis, Univ. Calif., Davis. 904 pp.<br />
Carlton, J.T. 1989. Man's role in changing the face <strong>of</strong> the ocean: biological invasions and the<br />
implications for conservation <strong>of</strong> near-shore environments. Conserv. Biol. 3(3): 265-273.
Chapt 1. Introduction, page 1- 9<br />
Carlton, J.T. and J. B. Geller. 1993. Ecological roulette: the global transport <strong>of</strong> nonindigenous<br />
marine organisms. Science 261: 78-82.<br />
Carlton, J.T., D. Reid and H. van Leeuwen. 1995. The role <strong>of</strong> shipping in the introduction <strong>of</strong><br />
nonindigenous aquatic organisms to the coastal waters <strong>of</strong> the United States (other than the Great<br />
Lakes) and an analysis <strong>of</strong> control options. Report to U.S. Coast Guard, Marine Environment<br />
Protection Division, Washington, DC. 215 pp.<br />
Cohen, A.N. and J.T. Carlton. 1995. Nonindigenous aquatic species in a United States estuary: a<br />
case study <strong>of</strong> the biological invasion <strong>of</strong> San Francisco Bay and delta. Report to U.S. Fish &<br />
Wildlife Service, Washington, DC and National Sea Grant College Program, Connecticut Sea<br />
Grant. 246 pp.<br />
Coles, S. L., R.C. DeFelice, L.G. Eldredge and J.T. Carlton. 1999. Historical and recent<br />
introductions <strong>of</strong> non-indigenous marine species into Pearl Harbor, Oahu, Hawaiian Islands. Mar.<br />
Biol. 135(1): 147-158.<br />
Hallegraeff, G.M. and C.J. Bolch. 1992. Transport <strong>of</strong> diatom and din<strong>of</strong>lagellate resting spores in<br />
ships’ ballast water: implications for plankton biogeography and aquaculture. J. Plankton Res.<br />
14: 1067-1084.<br />
Hayward, B.W. 1997. Introduced marine organisms in New Zealand and their impact in the<br />
Waitemata Harbour, Auckland. Tane 36:197-223.<br />
Hines, A.H., G.M. Ruiz, J. Chapman, J. Carlton and N. Foster. 1998. <strong>Biological</strong> invasions in<br />
cold-water coastal ecosystems: Ballast-mediated introductions in Port Valdez/Prince William<br />
Sound, Alaska. Progress Report to the Regional Citizens' Advisory Council <strong>of</strong> Prince William<br />
Sound.<br />
Leppakoski, E. 1984. Introduced species in the Baltic Sea and its coastal ecosystems. Ophelia,<br />
suppl 3: 123-135.<br />
Mills, E.L., J.H. Leach, J.T. Carlton and C.L. Secor. 1993. Exotic species in the Great Lakes: A<br />
history <strong>of</strong> biotic crises and anthropogenic introductions. J. Gt. Lakes Res. 19(1): 1-54.<br />
Reise, K., S. Gollasch and W.J. Wolff. 1999. Introduced marine species <strong>of</strong> the North Sea coasts.<br />
Helgol. Meeresunters. 52: 219-234.<br />
Ruiz, G.M., J.T. Carlton, A. H. Hines and E.D. Grosholz. 1997(a). Global invasions <strong>of</strong> marine<br />
and estuarine habitats by non-indigenous species: Mechanisms, extent, and consequences. Am.<br />
Zool. 37: 621-632<br />
Ruiz, G.M. and A.H. Hines. 1997. Patterns <strong>of</strong> nonindigenous species transfer and invasion in<br />
Prince William Sound, Alaska: Pilot Study. Report Submitted to the Prince William Sound<br />
Citizens’ Advisory Council. 80pp.
Chapt 1. Introduction, page 1- 10<br />
Ruiz, G.M., P. F<strong>of</strong>on<strong>of</strong>f and A.H. Hines. 1999. Non-indigenous species as stressors in estuarine<br />
and marine communities: Assessing invasion impacts and interactions. Limnol. Oceanogr. 44(3,<br />
part 2): 950-972<br />
Smith, L.D., M.J. Wonham, L.D. MCann, D.M. Reid, G.M. Ruiz and J.T. Carlton. 1996.<br />
<strong>Biological</strong> invasions by nonindigenous species in United States waters: Quantifying the role <strong>of</strong><br />
ballast water and sediments. Parts I and II. Final Report to the U.S. Coast Guard and the U.S.<br />
Department <strong>of</strong> Transportation. 246 pp.<br />
Smith, L.D., M.J. Wonham, L.D. McCann, G.M. Ruiz, A.H. Hines and J.T. Carlton. 1999.<br />
Invasion pressure to a ballast-flooded estuary and an assessment <strong>of</strong> inoculant survival. Biol.<br />
<strong>Invasions</strong> 1: 67-87.<br />
Williams, R.J., F.B. Griffiths, E.J. Van der Wal and J. Kelly. 1988. Cargo vessel ballast water as<br />
a vector for the transport <strong>of</strong> non-indigenous marine species. Est. Coast. Shelf Sci. 26: 409-420.
Chapt 2. Ballast <strong>Water</strong> Delivery Patterns, page 2- 1<br />
Chapter 2. Ballast <strong>Water</strong> Delivery Patterns<br />
Gregory M. Ruiz, Smithsonian Environmental Research Center<br />
Anson H. Hines, Smithsonian Environmental Research Center<br />
George Smith, Smithsonian Environmental Research Center<br />
Melissa A. Frey, Smithsonian Environmental Research Center<br />
2A. Purpose<br />
The primary goal <strong>of</strong> this portion <strong>of</strong> the study was to characterize the traffic patterns and<br />
volumes <strong>of</strong> ballast water discharged into Port Valdez and Prince William Sound (PWS) by oil<br />
tankers. Since ballast water is a major mechanism for the transfer <strong>of</strong> NIS, we wished to describe<br />
the delivery patterns <strong>of</strong> the ballast water to the region by season, source region, voyage duration.<br />
Analysis <strong>of</strong> the biota associated with the tankers’ ballast water is discussed in the next chapter.<br />
2B. Methods<br />
We obtained data on ship arrivals and ballast water histories in two ways. First, we<br />
obtained information about the long-term (10-year) pattern <strong>of</strong> arrivals to Port Valdez from<br />
Alyeska and RCAC. Second, to characterize current patterns, we collected detailed data from<br />
vessels arriving to Port Valdez over the one-year period <strong>of</strong> 1998. Our goal in this latter approach<br />
was to collect comprehensive information on the origin (i.e., last port <strong>of</strong> call), date <strong>of</strong> arrival, and<br />
ballast water histories for as many arriving vessels as possible. Most <strong>of</strong> these data were collected<br />
by SERC staff, during interviews aboard vessels (see below). Additional data were sent to us by<br />
the ships’ personnel and shipping agents.<br />
Beginning December 1997, we implemented a sampling scheme to estimate the amount<br />
<strong>of</strong> segregated ballast water delivered to Prince William Sound and Port Valdez by source port<br />
and season. For tankers arriving to Port Valdez from each <strong>of</strong> the three primary domestic source<br />
port systems (Los Angeles, San Francisco Bay, Puget Sound), we boarded approximately 3<br />
tankers per month (i.e., 10 per quarter x 3 source ports = 30 per quarter). In addition, we<br />
attempted to board most tankers arriving to Port Valdez from foreign ports.<br />
Upon boarding, we conducted an interview <strong>of</strong> the ships’ personnel to collect information<br />
on the quantity, age, source region, and management <strong>of</strong> all ballast water. Following the<br />
interview, we proceeded to sample the segregated ballast water to characterize temperature,<br />
salinity, and resident biota (see Chapter 3).<br />
We excluded non-segregated ballast water from most <strong>of</strong> our current analyses. Although<br />
this can account for roughly 50% <strong>of</strong> the total ballast water aboard tankers arriving to Prince<br />
William Sound (see below), previous analyses indicated that very few viable organisms were<br />
present in this ballast water, which <strong>of</strong>ten includes some residual oil. Furthermore, the<br />
nonsegregated ballast water is pumped to an on-shore treatment facility at the Alyeska terminal.<br />
For review <strong>of</strong> previous results, as well as description <strong>of</strong> the treatment process, see Ruiz and<br />
Hines 1997.<br />
Since vessels with double bottoms are difficult to sample for biota, we focused our sampling<br />
effort primarily on vessels without double bottoms. Thus, to characterize the entire fleet (i.e., all
Chapt 2. Ballast <strong>Water</strong> Delivery Patterns, page 2- 2<br />
arrivals), we obtained additional data on ballast water histories <strong>of</strong> nearly all oil tankers arriving<br />
to Port Valdez in 1998. Ships’ personnel and shipping agents generously provided these data.<br />
2C. Results<br />
2C1. Number and Source <strong>of</strong> Tanker Arrivals to PWS<br />
Over the past decade (1989-98), tanker arrivals to Port Valdez have averaged 713<br />
(se=37.2) ships per year, ranging from 870 to 549 (Fig. 2.1). There has been a noticeable decline<br />
in arrivals since 1991, with each year having fewer arrivals than the previous one.<br />
Figure 2.1. Annual number <strong>of</strong> oil tankers arriving to Port Valdez, 1989-1998. Data as provided by Alyeska.<br />
1000<br />
NUMBER OF ARRIVALS<br />
800<br />
600<br />
400<br />
200<br />
0<br />
89 90 91 92 93 94 95 96 97 98<br />
YEAR<br />
Using 1998 to examine spatial and temporal patterns, tanker arrivals to Port Valdez were<br />
both distributed evenly among seasons and dominated by arrivals from U.S. domestic ports (Fig.<br />
2.2). An average <strong>of</strong> 137.3 (s.e.=2.98) vessels arrived each quarter, and 95.8% (s.e. = 0.82 %) <strong>of</strong><br />
all arrivals came directly from a U.S. port. Of all tanker arrivals, 82.7% came from one <strong>of</strong> three<br />
domestic ports (Fig. 2.3): Puget Sound, Washington (43.0%); San Francisco Bay, California<br />
(28.8.%); and Long Beach, California (10.9%). The residual came from Oregon, Hawaii,<br />
Alaska, or foreign ports. Among arrivals from foreign ports, most (69.6%) came directly from<br />
Korea (Fig. 2.4).<br />
Figure 2.2. Number <strong>of</strong> oil tankers arriving to Port Valdez from foreign and domestic source ports by season<br />
in 1998. Seasons include: Winter (January-March), Spring (April-June), Summer (July-September), and Fall<br />
(October-December). Data based upon boarding interviews and reports from ships’ personnel (see text).<br />
160<br />
SOURCE PORT<br />
Foreign<br />
Domestic<br />
# VESSELS<br />
120<br />
80<br />
40<br />
0<br />
WINTER SPRING SUMMER FALL<br />
SEASON
Chapt 2. Ballast <strong>Water</strong> Delivery Patterns, page 2- 3<br />
Figure 2.3 Number <strong>of</strong> oil tankers arriving to Port Valdez from each domestic source port by season in 1998.<br />
Source ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); Columbia River,<br />
Oregon (OR); Cook Inlet, Alaska (AK); and Barbers Point, Hawaii (HI). Seasons include: Winter (January-March),<br />
Spring (April-June), Summer (July-September), and Fall (October-December). Data based upon boarding<br />
interviews and reports from ships’ personnel (see text).<br />
SOURCE PORT<br />
# VESSELS<br />
80<br />
60<br />
40<br />
20<br />
PS<br />
SF<br />
LB<br />
OR<br />
AK<br />
HI<br />
0<br />
WINTER SPRING SUMMER FALL<br />
SEASON<br />
Figure 2.4. Number <strong>of</strong> oil tankers arriving to Port Valdez from each foreign source port by season in 1998.<br />
Seasons Include: Winter (January-March), Spring (April-June), Summer (July-September), and Fall (October-<br />
December). Data based upon boarding interviews and reports from ships’ personnel (see text).<br />
# VESSELS<br />
6<br />
4<br />
2<br />
SOURCE PORT<br />
Korea<br />
China<br />
Taiwan<br />
Japan<br />
Singapore<br />
0<br />
WINTER SPRING SUMMER FALL<br />
SEASON<br />
2C2. Volume <strong>of</strong> Ballast <strong>Water</strong> delivered to PWS<br />
During 1998, oil tankers arriving to PWS carried an estimated average <strong>of</strong> 65,775m 3<br />
(s.e.=1,252; n=472) <strong>of</strong> total ballast water, the combination <strong>of</strong> segregated and nonsegregated<br />
ballast water. Segregated ballast water comprised an average <strong>of</strong> 54.7% (s.e.=2.1%; n=472) <strong>of</strong><br />
the total among tankers.<br />
Across all vessels, tankers discharged an average <strong>of</strong> 32,715 m 3 (s.e.= 645; n=472) <strong>of</strong><br />
segregated ballast water upon arrival to PWS (Table 2.1). Although there were no seasonal<br />
differences in the average amount <strong>of</strong> ballast water per tanker, there was a significant difference<br />
by source port (Fig. 2.5; 2-way ANOVA, F (3 (seasons), 5 (port source), 517 obs) =3.52, P = 0.004).<br />
Specifically, the mean volume was significantly greater for arrivals from foreign ports compared<br />
to arrivals from all other ports, and the mean volume was significantly lower for tankers from<br />
Hawaii relative to all other sources.
Chapt 2. Ballast <strong>Water</strong> Delivery Patterns, page 2- 4<br />
Table 2.1. Estimated volume <strong>of</strong> ballast water delivered by oil tankers to Port Valdez and PWS in 1998.<br />
Shown by source port and season are: (1) the estimated mean volume <strong>of</strong> ballast water arriving per tanker, including<br />
the standard error and sample size (n=number <strong>of</strong> vessels for which we have volume estimates), (2) the total number<br />
<strong>of</strong> tanker arrivals which is shown as N, (3) the total estimated volume <strong>of</strong> ballast water (calculated as mean volume X<br />
total arrivals). The bottom row (Overall) estimates the total ballast water volume and number <strong>of</strong> arrivals for all<br />
ships combined. Source ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB);<br />
Foreign port with open-ocean exchange (EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI).<br />
Seasons include: Winter (January-March), Spring (April-May), Summer (July-September), and Fall (October-<br />
December). Source <strong>of</strong> data on volumes and arrivals as described in text.<br />
BW vol. (m³)<br />
Port/Source Season Mean(se) n N Total<br />
Winter 28163(1734) 56 64 1,802,432<br />
Spring 31421(2095) 51 60 1,885,260<br />
PS Summer 34448(1901) 53 59 2,032,432<br />
Fall 33894(2199) 46 53 1,796,382<br />
Grand total 206 236 7,516,506<br />
Winter 31841(2798) 31 34 1,082,594<br />
Spring 32809(2984) 40 41 1,345,169<br />
SF Summer 35765(2361) 40 44 1,573,660<br />
Fall 34371(2372) 37 39 1,340,469<br />
Grand total 148 158 5,341,892<br />
Winter 32526(3237) 18 18 585,468<br />
Spring 30399(2902) 12 13 395,187<br />
LB Summer 36045(3633) 14 14 504,630<br />
Fall 28850(1346) 15 15 390,180<br />
Grand total 59 60 1,875,465<br />
Winter 42153(4093) 9 9 379,377<br />
Spring 44294(3775) 6 6 265,764<br />
EX Summer 29856(3689) 5 5 149,280<br />
Fall 47056(5199) 3 3 141,168<br />
Grand total 23 23 935,589<br />
Winter 29182(6929) 4 9 262,638<br />
Spring 28250(2169) 4 4 113,000<br />
OR Summer 32429(1631) 2 3 97,287<br />
Fall 39138(5740) 6 7 273,966<br />
Grand total 16 23 746,891<br />
Winter 27142(2279) 6 7 189,994<br />
Spring 22229(1651) 4 5 111,145<br />
HI Summer 23766(4197) 6 6 142,596<br />
Fall 21729(1544) 4 5 108,370<br />
Grand Total 20 23 552,105<br />
Overall 32,715 472 523 16,968,448<br />
Note: Not included in the table are arrivals from Nikiski, AK (19) which provided no data, and other arrivals (7) for<br />
which port data are unavailable.
Chapt 2. Ballast <strong>Water</strong> Delivery Patterns, page 2- 5<br />
Figure 2.5. Mean volume <strong>of</strong> segregated ballast water per tanker arriving to Port Valdez and PWS by source<br />
port and season, 1998. The mean volumes are estimated for: (A) Each source port across all seasons; (B) Each<br />
season across all source ports. Standard error and sample size is shown above each bar; see Table 2.1 for further<br />
information. Source ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB);<br />
Foreign port with open-ocean exchange (EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI).<br />
Seasons include: Winter (January-March), Spring (April-June), Summer (July-September), and Fall (October-<br />
December). Data based upon boarding interviews and reports from ships’ personnel (see text).<br />
23<br />
40<br />
206<br />
148<br />
59<br />
16<br />
30<br />
20<br />
CUBIC METERS (thousands)<br />
20<br />
10<br />
0<br />
40<br />
30<br />
PS SF LB EX OR HI<br />
PORT<br />
119 113<br />
117<br />
123<br />
20<br />
10<br />
0<br />
WINTER SPRING SUMMER FALL<br />
SEASON<br />
We estimated the total amount <strong>of</strong> segregated ballast water discharged into Prince William<br />
Sound during 1998 was approximately 17,000,000 m 3 (no. arrivals x average ballast water<br />
volume) for source port by season (see Table 2.1). The relative contribution <strong>of</strong> different source<br />
ports to the total varied greatly, reflecting variation in the number <strong>of</strong> arrivals (Fig. 2.6; Table<br />
2.1). As a result, Puget Sound contributed approximately 44% <strong>of</strong> the total, followed by San<br />
Francisco 31% and Long Beach 11%.
Chapt 2. Ballast <strong>Water</strong> Delivery Patterns, page 2- 6<br />
Figure 2.6. Cumulative volume <strong>of</strong> ballast water arriving to Port Valdez and PWS by source port and season,<br />
1998. The cumulative volumes are estimated for season by source port (see Table 2.1 for further detail). Source<br />
ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); Foreign port with openocean<br />
exchange (EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI). Seasons include: Winter<br />
(January-March), Spring (April-June), Summer (July-September), and Fall (October-December). Data based upon<br />
boarding interviews and reports from ships’ personnel (see text).<br />
CUBIC METERS (millions)<br />
2.0<br />
1.5<br />
1.0<br />
0.5<br />
0.0<br />
WINTER SPRING SUMMER FALL<br />
SEASON<br />
SOURCE PORT<br />
PS<br />
SF<br />
LB<br />
EX<br />
OR<br />
HI<br />
2C3. Age and Management <strong>of</strong> Ballast <strong>Water</strong> delivered to PWS<br />
The average age <strong>of</strong> ballast water arriving in tankers varied among source ports, ranging<br />
between 4.8 to 10.2 days (Fig. 2.7). The mean age among all arrivals was 6.6 days (s.e.= 0.2).<br />
For domestic source ports, the age <strong>of</strong> water was correlated with distance from Port Valdez to the<br />
source port, as ballast water came directly from the last port <strong>of</strong> call (the exception was for<br />
experiments conducted at our request, as described in Chapter 4). In contrast, all foreign arrivals<br />
exchanged their ballast water at sea, so the age <strong>of</strong> water was less than the voyage duration. Thus,<br />
for foreign arrivals, the actual source was considered open ocean exchange (EX) instead <strong>of</strong> the<br />
last port <strong>of</strong> call.<br />
Figure 2.7. Mean age (voyage duration) for ballast water arriving to Port Valdez by source port. The mean<br />
age is estimated for each source port based upon all boarding data (December 1997-July 1999). Standard error and<br />
sample size is shown above each bar. Source ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF);<br />
Long Beach, CA (LB); Foreign port with open-ocean exchange (EX); Columbia River, Oregon (OR); and Barbers<br />
Point, Hawaii (HI).<br />
BALLAST WATER AGE (Days)<br />
12<br />
8<br />
4<br />
0<br />
48<br />
50<br />
46<br />
PS SF LB EX OR HI<br />
SOURCE PORT<br />
2D. Discussion<br />
Over the past decade, Prince William Sound and Port Valdez together have received<br />
approximately 23.3 million m 3 (= 713 arrivals/yr x 32,610 m 3 /arrival) <strong>of</strong> segregated ballast water<br />
19<br />
2<br />
4
Chapt 2. Ballast <strong>Water</strong> Delivery Patterns, page 2- 7<br />
each year from oil tankers arriving to the Alyeska terminal. Although most <strong>of</strong> this water is<br />
released in Port Valdez, ships will sometimes begin discharging upon entering Prince William<br />
Sound en route for the Port. Thus, organisms released with this ballast water may experience a<br />
broader range <strong>of</strong> conditions outside <strong>of</strong> Port Valdez than we had originally considered.<br />
The total volume <strong>of</strong> ballast water delivered to Prince William Sound by tankers greatly<br />
exceeds the estimated quantity <strong>of</strong> ballast water arriving to other western U.S. ports (Carlton et al.<br />
1995). However, it is important to recognize that existing estimates for the other ports have<br />
included only the ballast water from foreign sources. In contrast, our estimates for Prince<br />
William Sound included both foreign and domestic sources, but were dominated by the latter.<br />
The amount <strong>of</strong> domestic ballast water released in other U.S. ports is only now being estimated.<br />
Nonetheless, even when we can include data on domestic sources for all ports, it appears likely<br />
that the total volume <strong>of</strong> ballast water released to PWS will still exceed that for the other western<br />
ports, due to the absence <strong>of</strong> extensive domestic tanker and bulker traffic (i.e., those vessels that<br />
discharge the greatest quantities <strong>of</strong> ballast water) at the other ports. Instead, the domestic traffic<br />
for other western U.S. ports is dominated by container ships, which release relatively small<br />
amounts <strong>of</strong> ballast water (Carlton et al. 1995; National Ballast <strong>Water</strong> Information Clearinghouse,<br />
unpubl. data).<br />
The total amount <strong>of</strong> ballast water arriving to Prince William Sound in tankers is also<br />
relatively large on a global scale. Within the U.S., PWS is third only to Chesapeake Bay and<br />
New Orleans in estimated ballast water discharge for 1991 (Carlton et al. 1995, Smith et al.<br />
1999). In a similar estimate <strong>of</strong> ballast water discharged to 46 Australian ports in 1991, only that<br />
for the port <strong>of</strong> Dampier exceeded the volume for PWS (Kerr 1994). As above, estimates for the<br />
both the U.S. and Australian ports were restricted to arrivals from foreign ports. Although these<br />
totals would clearly increase when including domestic arrivals, the overall patterns provide a<br />
useful context, suggesting PWS is on the extreme end <strong>of</strong> the spectrum for amount <strong>of</strong> ballast<br />
water discharge.<br />
It is also important to recognize the present level <strong>of</strong> tanker activity, and the magnitude <strong>of</strong><br />
ballast water delivery, as a recent development in Port Valdez. The terminal began transporting<br />
oil via tankers in 1977. Based upon the arrivals rate and discharge volumes observed in this<br />
decade, we estimate over 700 million m 3 <strong>of</strong> segregated ballast water have been delivered over the<br />
past 3 decades <strong>of</strong> operation. This large cumulative volume underscores the potential importance<br />
<strong>of</strong> ballast water as a vector for the transfer <strong>of</strong> species. Unlike many other commercial ports,<br />
however, the volume <strong>of</strong> ballast water prior to oil exportation was virtually absent, as very few<br />
other vessels currently deliver ballast water to the Port (Ruiz et al., unpubl. data).<br />
Furthermore, the foreign export <strong>of</strong> oil from Port Valdez has only occurred since 1996,<br />
following authorization by U.S. Congress. Prior to this time, all oil export was only to domestic<br />
U.S. ports, which were therefore the source <strong>of</strong> ballast water delivery to PWS and Port Valdez.<br />
Although tankers now export oil to foreign ports from PWS, the delivery <strong>of</strong> ballast water from<br />
foreign traffic remains a small fraction (
Chapt 2. Ballast <strong>Water</strong> Delivery Patterns, page 2- 8<br />
2E. References<br />
Carlton, J.T., D. Reid and H. van Leeuwen. 1995. The role <strong>of</strong> shipping in the introduction <strong>of</strong><br />
nonindigenous aquatic organisms to the coastal waters <strong>of</strong> the United States (other than the Great<br />
Lakes) and an analysis <strong>of</strong> control options. Report to U.S. Coast Guard, Marine Environment<br />
Protection Division, Washington, DC. 215 pp.<br />
Kerr, S. 1994. Ballast water ports and shipping study. Australian Quarantine and Inspection<br />
Service, Report No. 5, Canberra.<br />
Ruiz, G.M. and A.H. Hines. 1997. Patterns <strong>of</strong> nonindigenous species transfer and invasion in<br />
Prince William Sound, Alaska: Pilot Study. Report Submitted to the Prince William Sound<br />
Citizens’ Advisory Council. 80pp.<br />
Smith, L.D., M.J. Wonham, L.D. MCann, D.M. Reid, G.M. Ruiz and J.T. Carlton. 1996.<br />
<strong>Biological</strong> invasions by nonindigenous species in United States waters: Quantifying the role <strong>of</strong><br />
ballast water and sediments. Parts I and II. Final Report to the U.S. Coast Guard and the U.S.<br />
Department <strong>of</strong> Transportation. 246 pp.<br />
Smith, L.D., M.J. Wonham, L.D. McCann, G.M. Ruiz, A.H. Hines and J.T. Carlton. 1999.<br />
Invasion pressure to a ballast-flooded estuary and an assessment <strong>of</strong> inoculant survival. Biol.<br />
<strong>Invasions</strong> 1: 67-87.
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 1<br />
Chapter 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong> in Oil Tankers<br />
Gregory M. Ruiz, Smithsonian Environmental Research Center<br />
Anson H. Hines, Smithsonian Environmental Research Center<br />
George Smith, Smithsonian Environmental Research Center<br />
Melissa A. Frey, Smithsonian Environmental Research Center<br />
3A. Purpose<br />
The overall goal <strong>of</strong> this research component was to characterize the biota associated with<br />
segregated ballast water <strong>of</strong> tankers arriving to Port Valdez. For this analysis, we designed a<br />
sampling program to measure temporal (seasonal, annual) and spatial (source port) variation in<br />
the biota associated with the ballast water.<br />
We focused primarily on the mid-large (>80 micron) zooplankton resident in the water<br />
column <strong>of</strong> ballast tanks, and present the results <strong>of</strong> this analysis here. We have included some<br />
information on the phytoplankton concentrations present in our samples, but the sampling<br />
methods (below) were not designed to characterize these organisms and many <strong>of</strong> the other taxa<br />
(e.g., bacteria, viruses, and other microorganisms) that are small in size. This choice does not<br />
imply that small organisms are not significant from an invasion standpoint, as the potential<br />
effects <strong>of</strong> toxic din<strong>of</strong>lagellate blooms are very evident (see Introduction). Instead, we simply did<br />
not have the resources to include all taxonomic groups in our analyses.<br />
We chose to focus on the mid-large zooplankton for multiple reasons. First, the<br />
taxonomic resolution is relatively good compared to many <strong>of</strong> the other (smaller) organisms.<br />
Second, most known NIS that are established at the source ports <strong>of</strong> oil tankers, such as San<br />
Francisco Bay, occur (for some portion <strong>of</strong> their life history) in this zooplankton community.<br />
Third, we could readily gain access to the plankton community (as opposed to the bottom<br />
sediments). Finally, the sample analysis for larger zooplankton is not as technically difficult or<br />
time consuming as that necessary for the smaller organisms, allowing us to analyze samples from<br />
a large number <strong>of</strong> ships for statistical comparisons.<br />
The analysis <strong>of</strong> zooplankton presented here is one <strong>of</strong> the most comprehensive and<br />
quantitative studies <strong>of</strong> ballast water in the world. Additional data on the biota associated with<br />
ballast water also appear in other chapters. Our analysis <strong>of</strong> ballast water exchange (Chapter 5)<br />
includes survivorship <strong>of</strong> plankton during 8 separate voyages, effects <strong>of</strong> exchange on<br />
zooplankton, and some information on bacteria and ciliate protozoans. Biota associated with the<br />
bottom sediments <strong>of</strong> ballast tanks, as well as the hulls and seachests, are also examined (Chapters<br />
6 and 7, respectively).<br />
3B. Methods<br />
3B1. Source and Number <strong>of</strong> Sampled Ships<br />
For a 13-month period (December 1997 – December 1998, hereafter 1998), we conducted<br />
an intensive sampling program <strong>of</strong> ballast water on tankers that was stratified by source port and<br />
season. Most tankers and ballast water arriving to Port Valdez came from three U.S. domestic
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 2<br />
port systems: Puget Sound, San Francisco Bay, and Long Beach (see Chapter 2). To characterize<br />
biota for these ports by seasons, we sampled a minimum <strong>of</strong> 3 tanker arrivals per month from<br />
each <strong>of</strong> the three domestic source port systems (i.e., 10 per quarter x 3 source ports = 30 per<br />
quarter). Although relatively few (23) tankers arrived from foreign ports in Port Valdez during<br />
this year, we sampled as many as possible (n=19) to compare the biota arriving from foreign<br />
versus domestic sources. We also sampled a limited number <strong>of</strong> arrivals from the other two<br />
domestic ports: Oregon and Hawaii (which comprised < 10% all arrivals; see Chapter 2).<br />
In June <strong>of</strong> three consecutive years (1997, 1998, 1999), we collected samples from<br />
approximately 10 tankers arriving to Port Valdez from the domestic ports <strong>of</strong> Puget Sound, San<br />
Francisco Bay, and Long Beach. Samples from 1997 were collected as part <strong>of</strong> a Pilot Study that<br />
we conducted for RCAC (Ruiz and Hines 1997), and the samples for 1998-1999 were collected<br />
in the present study. Together, these samples were used to characterize annual variation in the<br />
ballast water biota arriving to Alaska.<br />
3B2. Sample Collection and Analysis<br />
We boarded and sampled tankers immediately upon their arrival to Port Valdez. As<br />
described above, we sampled approximately 2-3 vessels per week over the 13-month period.<br />
Although we attempted to collect ballast water from every tanker boarded, vessels with double<br />
bottoms could not be sampled easily without disruption <strong>of</strong> ship operations and modification <strong>of</strong><br />
our standard sampling protocol. In the present analysis, we therefore have included primarily<br />
(but not exclusively) vessels without double bottoms.<br />
We applied our established methods for qualitative and quantitative analysis <strong>of</strong> biota<br />
transported in ballast water, which evolved from methods developed by J.T. Carlton (e.g.,<br />
Carlton and Geller, 1993; LaVoie et al.1999; Smith et al., 1999). Our protocol consisted <strong>of</strong><br />
collecting the following information and samples:<br />
• Ship and ballast management information: Last port <strong>of</strong> call, number <strong>of</strong> tanks by type,<br />
capacity <strong>of</strong> tanks, amount <strong>of</strong> segregated and non-segregated ballast water on board, source(s)<br />
<strong>of</strong> ballast water, age <strong>of</strong> ballast water, date <strong>of</strong> arrival, ballast management practices;<br />
• Physical variables <strong>of</strong> ballast water: <strong>Water</strong> temperature and salinity were measured (surface<br />
and 10m depth) for each tank sampled (as below), collecting ballast water with a Niskin<br />
bottle through the Butterworth hatches; oxygen (O 2 ) concentration was not measured<br />
because previous extensive analysis <strong>of</strong> ballast water tanks in other cargo ships indicated that<br />
O 2 concentrations rarely varied and were not appreciably lower than saturation (Smith et al.,<br />
1996).<br />
• <strong>Biological</strong> samples <strong>of</strong> ballast water: Plankton samples were collected by towing a standard<br />
plankton net (80 micron mesh, 30 cm diameter) vertically through the entire height <strong>of</strong> the<br />
water column in each ballast tank; access to ballast tanks was obtained through the<br />
Butterworth hatches. A single tank was sampled for each ship, when ballast water was<br />
present and accessible, and two plankton tows were collected for each tank; the height <strong>of</strong><br />
each plankton tow was measured to the nearest 10 cm.<br />
• Additional observations and opportunistic samples: Upon initiating sampling <strong>of</strong> ballast<br />
tanks, we routinely examined the surface waters to look for large, mobile biota (e.g., fish)<br />
and organisms attached to the sides <strong>of</strong> tanks; we <strong>of</strong>ten took opportunities to collect any such
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 3<br />
organisms observed, as well as bottom sediments, since these are usually missed in our<br />
plankton tows.<br />
• Physical variables <strong>of</strong> port water: Shipside water temperature and salinity were measured<br />
(surface and 10m depth) usually within an hour <strong>of</strong> sampling ballast water <strong>of</strong> most vessels; the<br />
samples were collected from the berth platform (within 50m <strong>of</strong> the ship), using a Niskin<br />
bottle.<br />
Most plankton samples were returned to the laboratory at Valdez and examined initially<br />
within an hour <strong>of</strong> collection to assess condition <strong>of</strong> organisms present. More specifically, we<br />
examined each plankton sample with our dissecting microscopes (10-40x), to provide a<br />
qualitative assessment <strong>of</strong> plankton viability. Each sample was washed carefully into a finger<br />
bowl for examination, and the presence <strong>of</strong> each morphologically distinct taxonomic group was<br />
noted. For each taxon identified, the percent <strong>of</strong> individuals alive was estimated by evaluating<br />
their morphological integrity, movement, and activity; although status <strong>of</strong> some organisms (e.g.,<br />
diatoms or eggs) was difficult to discern with confidence during a brief screening. After initial<br />
microscopic examination, the plankton samples were preserved in 5% buffered formalin for later<br />
identification and enumeration <strong>of</strong> organisms (as below).<br />
We used two different methods to characterize the plankton samples, as follows:<br />
• Coarse Analysis. All samples were characterized by Coarse Analysis, consisting <strong>of</strong> a direct<br />
count <strong>of</strong> individuals according to general taxonomic groups, usually phyla (e.g., molluscs,<br />
crustaceans, echinoderms). The minimum <strong>of</strong> number <strong>of</strong> distinctly different taxa were also<br />
estimated in Coarse Analysis <strong>of</strong> each sample.<br />
• Fine Analysis. For a subset <strong>of</strong> samples (roughly 1/3 <strong>of</strong> the ships from domestic ports), Fine<br />
Analysis was used to enumerate all morphologically distinct taxa at the lowest taxonomic<br />
level possible. For many groups that included larval invertebrates (e.g., bivalves,<br />
gastropods), identification could not progress beyond gross taxonomic groups; further<br />
identification can only be accomplished with intensive culture <strong>of</strong> larvae to adult stages, upon<br />
which taxonomy is based, or the use <strong>of</strong> molecular probes. For other groups that include<br />
adult stages (e.g., copepods), we sought species-level identifications.<br />
The two methods were selected to provide different types <strong>of</strong> information. The Coarse<br />
Analysis allowed us to test for patterns in the biota across all ships, increasing the statistical<br />
power <strong>of</strong> the analysis. Since many species were not present on each ship, such an approach was<br />
not feasible with finer taxonomic resolution. In contrast, the Fine Analysis allowed us to<br />
quantify the densities <strong>of</strong> particular taxa, usually crustacean groups with adult forms (see results),<br />
and test for the presence <strong>of</strong> nonindigenous species known from the source ports. These data also<br />
allow us to characterize the frequency and density <strong>of</strong> particular nonindigenous species in the<br />
ballast water.<br />
For both analyses, samples were concentrated on an 80 micron sieve and washed into a<br />
finger bowl for identification and enumeration. Each whole sample was examined using a stereo<br />
microscope, and all morphologically distinct taxa were identified to the desired taxonomic level<br />
(as above). For abundant taxa (> 100 individuals/sample), samples were split using a Folsom<br />
plankton splitter to achieve counts between 10-100 individuals per subsample (usually splits <strong>of</strong><br />
1/8 to 1/32). For organisms in split samples, two subsamples were counted.
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 4<br />
Taxonomic identification <strong>of</strong> plankton followed a standard protocol. For those groups <strong>of</strong><br />
organisms that can be identified using the life stages present in ballast water samples (as<br />
discussed above), we made an initial identification based upon our current knowledge and<br />
literature that was immediately available to us at SERC. For many copepods, we were able to<br />
discern genera without much difficulty. Enumeration proceeded based upon the lowest<br />
discernible taxonomic units, and representative specimens were vouchered (in Fine Analysis) for<br />
taxonomic verification and, wherever possible, species-level identification. These voucher<br />
specimens were sent to taxonomists at the Smithsonian Institution’s National Museum <strong>of</strong> Natural<br />
History and elsewhere for verification and identification.<br />
3B3 Data Analysis<br />
Throughout our analyses, we use “ship” as the level <strong>of</strong> replication within a class variable<br />
(e.g., source port, season, year), because multiple samples from the same ship are not<br />
independent <strong>of</strong> each other. Although our replicate plankton samples per ship provide some<br />
important information on variation within ships, these are not statistically independent (since the<br />
ballast water originates from the same source and time) and mainly provide greater confidence in<br />
estimating plankton communities per ship. Thus, we estimated density per ship as the mean <strong>of</strong><br />
replicate tows.<br />
We derive most <strong>of</strong> the results reported from the Coarse Analysis. All enumeration is also<br />
completed for the Fine Analysis, but identification <strong>of</strong> only a portion <strong>of</strong> the voucher specimens<br />
has been finished to date. Although we cannot yet discuss the frequency and density <strong>of</strong><br />
individual taxa (as described above), we confirm the presence <strong>of</strong> numerous nonindigenous<br />
species in the Fine Analysis, and these are reported here. We will include full results <strong>of</strong> the Fine<br />
Analysis, when completed, in a future publication and provide a copy to RCAC.<br />
Virtually all organisms collected in the ballast water samples were alive and appeared to<br />
be in good condition. Indeed, many <strong>of</strong> the organisms collected from these samples performed<br />
well in laboratory culture and experiments, as described in Chapter 4. Thus, we considered all<br />
organisms counted in fixed samples to be alive at the time <strong>of</strong> sampling.<br />
In most <strong>of</strong> the analyses, we have excluded the chain-forming diatoms. Although we<br />
enumerated these organisms to the full extent possible, quantitative counts are particularly<br />
problematic and unreliable, because the chains break apart during sample collection and<br />
processing. We have therefore included information on their prevalence and the counts made but<br />
excluded these data from most estimates <strong>of</strong> organism densities.<br />
Finally, the presentation <strong>of</strong> water temperature and salinity, for both the ballast water <strong>of</strong><br />
oil tankers and Port Valdez, are presented in Chapter 4.<br />
3C Results<br />
Source and Number <strong>of</strong> Sampled Ships<br />
During this study, we sampled the ballast water <strong>of</strong> 169 tankers arriving to Port Valdez<br />
(Table 3.1). Our samples included 8-15 arrivals per quarter for each <strong>of</strong> the three major domestic
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 5<br />
ports (Puget Sound, San Francisco Bay, and Long Beach) and 3-7 arrivals per quarter from<br />
foreign ports (primarily Korea; see Chapter 2).<br />
Table 3.1. Prevalence and densities <strong>of</strong> taxa in the ballast water arriving to Port Valdez for each source port.<br />
Shown for each taxon and source port are the prevalence, density among all ships, and density for ships only when<br />
taxon was present. Standard errors are shown in parentheses with each density measure. Source ports include: Puget<br />
Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); Foreign port with open-ocean exchange<br />
(EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI). The data include all sample dates. Sample<br />
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).<br />
(%) (density/m³)<br />
Phylum Taxa source n Prevalence mean(se) all ships mean(se) when present<br />
DINOFLAGELLATA<br />
PS 48 88 775(223) 902(245)<br />
SF 50 54 66(21) 114(34)<br />
LB 46 65 71(30) 107(45)<br />
EX 19 84 729(658) 886(780)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 100 3.0(1.0) 3.0(1.0)<br />
DIATOMACEA<br />
PROTOZOA<br />
CNIDARIA<br />
CTENOPHORA<br />
PS 48 100 13866(3270) 13866(3270)<br />
SF 50 100 11683(4384) 11683(4384)<br />
LB 46 100 1170(241) 1170(241)<br />
EX 19 100 5346(2184) 5346(2184)<br />
OR 2 100 9409(6120) 9409(6120)<br />
HI 4 100 277(107) 277(107)<br />
PS 48 83 316(122) 361(139)<br />
SF 50 90 5506(3638) 6120(4037)<br />
LB 46 93 210(57) 220(59)<br />
EX 19 74 82(58) 110(78)<br />
OR 2 100 31(13) 31(13)<br />
HI 4 100 4.0(1.3) 4.0(1.3)<br />
PS 48 63 19.5(11.9) 55(32)<br />
SF 50 22 1.8(0.7) 8.3(2.3)<br />
LB 46 87 45(18) 53(21)<br />
EX 19 5 0.3(0.3) 5.7(0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
PS 48 17 0.2(0.1) 1.2(0.3)<br />
SF 50 4 0.01(0.008) 0.5(0.0)<br />
LB 46 37 1.6(0.6) 4.0(1.5)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 6<br />
Table 3.1 continued<br />
Phylum Taxa source n Prevalence (%) mean(se) all ships mean(se) when present<br />
PLATYHELMINTHES<br />
PS 48 27 3.4(1.4) 13(4.3)<br />
SF 50 28 10.5(4.6) 37(14)<br />
LB 46 80 11(2.2) 13(2.4)<br />
EX 19 16 0.1(0.07) 0.9(0.1)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
NEMATODA<br />
ROTIFERA<br />
SIPUNCULA<br />
NEMERTEA<br />
PS 48 6 0.2(0.1) 2.5(0.9)<br />
SF 50 8 0.5(0.3) 6.5(2.4)<br />
LB 46 4 0.1(0.07) 2.5(0.5)<br />
EX 19 5 0.1(0.1) 1.4(0.0)<br />
OR 2 100 4.4(3.0) 4.4(3.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
PS 48 2 0.3(0.3) 12.5(0)<br />
SF 50 2 0.7(0.7) 33(4.5)<br />
LB 46 2 0.6(0.6) 27(0.0)<br />
EX 19 5 0.3(0.3) 5.6(0.0)<br />
OR 2 50 0.2(0.2) 0.4(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
PS 48 0 0(0.0) 0(0.0)<br />
SF 50 4 0.8(0.7) 19.8(11)<br />
LB 46 4 0.7(0.6) 15(4.1)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
PS 48 8 1.0(0.6) 12.4(5.3)<br />
SF 50 8 41(27) 516(275)<br />
LB 46 15 23(12) 150(66)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
ANNELIDA PS 48 90 370(136) 404(148)<br />
SF 50 80 199(55) 251(68)<br />
LB 46 100 33(6.5) 33(6.5)<br />
EX 19 2 2.1(1.3) 6.3(3.6<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 25 0.31(0.31) 1.2(0.0)<br />
MOLLUSCA<br />
Bivalvia PS 48 90 371(100) 396(106)<br />
SF 50 82 319(79) 389(93)<br />
LB 46 98 240(72) 246(74)<br />
EX 19 53 8.0(3.5) 15.2(5.5)<br />
OR 2 50 0.5(0.5) 1.0(0.0)<br />
HI 4 75 1.3(0.7) 2.0(0.5)
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 7<br />
Table 3.1 continued<br />
Phylum Taxa source n Prevalence (%) mean(se) all ships mean(se) when present<br />
Gastropoda PS 48 71 139(39) 191(44)<br />
SF 50 54 34(18) 58(38)<br />
LB 46 91 35(7.6) 37(6.8)<br />
EX 19 37 26.6(17.6) 72(38)<br />
OR 2 50 0.2(0.2) 0.4(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
Other Mollusca PS 48 8 8.0(7.7) 105(83)<br />
SF 50 6 0.6(0.5) 10(7.6)<br />
LB 46 2 0.2(0.2) 9.0(0.0)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
ARTHROPODA/CRUSTACEA<br />
Amphipoda PS 48 29 1.9(1.3) 6.1(3.8)<br />
SF 50 30 0.61(0.16) 1.9(0.3)<br />
LB 46 48 0.7(0.2) 1.4(0.3)<br />
EX 19 19 0.07(0.05) 0.7(0.2)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
Anomura PS 48 29 0.91(0.64) 3.7(2.2)<br />
SF 50 24 0.2(0.08) 0.9(0.3)<br />
LB 46 63 3.8(1.4) 5.9(2.1)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
Brachyura PS 48 44 3.5(1.3) 9.2(2.8)<br />
SF 50 14 0.13(0.07) 1.1(0.5)<br />
LB 46 76 23.6(13.9) 30.2(18.6)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
Caridea PS 48 15 0.2(0.10) 1.3(1.2)<br />
SF 50 2 0.02(0.01) 0.2(0.0)<br />
LB 46 22 1.2(0.60) 3.8(1.5)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 8<br />
Table 3.1 continued<br />
Phylum Taxa source n Prevalence (%) mean(se) all ships mean(se) when present<br />
Cirripedia PS 48 85 832(559) 951(637)<br />
SF 50 66 96(34) 108(47)<br />
LB 46 63 18(5.4) 37(7.2)<br />
EX 19 16 0.8(0.5) 6.9(1.7)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 50 3.1(1.9) 4.8(4.3)<br />
Cladocera PS 48 15 2.7(1.2) 21.7(5.1)<br />
SF 50 16 1.9(0.8) 11.7(3.7)<br />
LB 46 7 0.04(0.02) 0.6(0.1)<br />
EX 19 5 0.3(0.3) 5.7(0.0)<br />
OR 2 100 3.5(1.9) 3.5(1.9)<br />
HI 4 0 0(0.0) 0(0.0)<br />
Copepoda PS 48 100 2395(664) 2395(664)<br />
SF 50 100 9416(2060) 9416(2060)<br />
LB 46 100 5116(685) 5116(685)<br />
EX 19 100 2345(645) 2345(645)<br />
OR 2 100 43(6.0) 43(6.0)<br />
HI 4 100 8.2(4.4) 8.2(4.4)<br />
Cumacea PS 48 10 0.05(0.03) 0.6(0.2)<br />
SF 50 30 0.70(0.2) 2.1(0.5)<br />
LB 46 22 0.13(0.05) 0.6(0.2)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
Decapoda (misc.) PS 48 4 0.05(0.04) 1.3(0.7)<br />
SF 50 8 0.02(0.02) 0.5(0.2)<br />
LB 46 4 0.05(0.02) 2.5(0.0)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
Isopoda PS 48 25 0.70(0.40) 2.1(1.3)<br />
SF 50 18 0.34(0.26) 2.1(1.5)<br />
LB 46 9 0.06(0.03) 0.7(0.1)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
Mysidacea PS 48 2 0.007(0.007) 0.3(0)<br />
SF 50 54 2.48(0.6) 4.6(0.9)<br />
LB 46 78 3.52(0.8) 4.4(0.9)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 25 0.15(0.15) 0.6(0.1)
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 9<br />
Table 3.1 continued<br />
Phylum Taxa source n Prevalence (%) mean(se) all ships mean(se) when present<br />
Ostracoda PS 48 25 0.35(0.16) 1.5(0.6)<br />
SF 50 8 0.40(0.20) 3.9(2.0)<br />
LB 46 17 0.28(0.14) 1.6(0.7)<br />
EX 19 16 0.93(0.54) 5.9(1.4)<br />
OR 2 50 0.6(0.6) 1.2(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
Tanaidacea PS 48 2 0.006(0.006) 0.3(0)<br />
SF 50 0 0(0.0) 0(0.0)<br />
LB 46 2 0.006(0.006) 0.3(0)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
BRYOZOA<br />
PHORONIDA<br />
CHAETOGNATHA<br />
ECHINODERMATA<br />
PS 48 35 16.1(5.4) 40.7(11.5)<br />
SF 50 40 59(33) 147(80)<br />
LB 46 22 1.5(0.5) 6.7(1.7)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
PS 48 2 0.01(0.01) 0.5(0.0)<br />
SF 50 2 0.01(0.01) 0.7(0.0)<br />
LB 46 15 0.5(0.4) 4.2(2.7)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
PS 48 33 0.7(0.4) 2.3(1.1)<br />
SF 50 12 0.5(0.3) 4.1(2.5)<br />
LB 46 72 5.8(2.2) 7.9(3.0)<br />
EX 19 21 0.2(0.1) 1.5(0.3)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
PS 48 33 32(15) 94(43)<br />
SF 50 12 6.3(4.8) 53(37)<br />
LB 46 26 5.8(3.5) 22(12)<br />
EX 19 5 0.3(0.3) 6.0(0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
CHORDATA<br />
Cephalochordata PS 48 0 0(0.0) 0(0.0)<br />
SF 50 0 0(0.0) 0(0.0)<br />
LB 46 9 0.1(0.01) 2.8(2.5)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 10<br />
Table 3.1 continued<br />
Phylum Taxa source n Prevalence (%) mean(se) all ships mean(se) when present<br />
Fish PS 48 10 0.05(0.02) 0.3(0.1<br />
SF 50 6 0.02(0.01) 0.5(0.1<br />
LB 46 7 0.04(0.02) 0.5(0.1)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
Other Chordata PS 48 21 4.9(2.2) 23.8(8.4)<br />
(incl. larvacea) SF 50 8 5.2(3.5) 85(41)<br />
LB 46 63 59.1(30.6) 107(55)<br />
EX 19 5 0.06(0.06) 1.2(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
OTHER<br />
Eggs PS 48 73 87(39) 113(50)<br />
SF 50 66 39(16) 56(22)<br />
LB 46 61 141(41) 232(148)<br />
EX 19 21 6.3(4.1) 15(9.2)<br />
OR 2 100 12(11) 12(11)<br />
HI 4 20 7.3(2.4) 7.3(2.4)<br />
Trochophore<br />
PS 48 33 20(8.2) 65(23)<br />
SF 50 16 16(10) 115(65)<br />
LB 46 30 11(4.6) 36(13)<br />
EX 19 0 0(0.0) 0(0.0)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
Unidentified larvae PS 48 0 0(0.0) 0(0.0)<br />
SF 50 4 95(93) 2376(1388)<br />
LB 46 9 0.02(0.02) 0.4(0)<br />
EX 19 11 27(26) 253(178)<br />
OR 2 0 0(0.0) 0(0.0)<br />
HI 4 0 0(0.0) 0(0.0)<br />
For June <strong>of</strong> 1997-1999, we sampled the ballast water <strong>of</strong> 31 tankers arriving to Port<br />
Valdez from the domestic ports <strong>of</strong> Puget Sound (n=14), San Francisco Bay (n=9), and Long<br />
Beach (n=9). The data for 1998 and 1999 are included in the total 169 vessels sampled during<br />
this study, and the data for 1997 are derived from our previous work (as above).<br />
Figure 3.1. Number <strong>of</strong> oil tankers sampled upon arrival to Port Valdez. Shown are the number <strong>of</strong> ships from<br />
which ballast water samples were collected by source port (i.e., last port <strong>of</strong> call) and season. Source ports include:<br />
Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); Foreign port with open-ocean<br />
exchange (EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI). Seasons include: Winter (January-<br />
March), Spring (April-June), Summer (July-September), and Fall (October-December). Data were collected<br />
primarily from December 1997 – December 1998, and some additional data were collected in May-June 1999.<br />
# VESSELS SAMPLED<br />
15<br />
10<br />
5<br />
0<br />
PORT SOURCE<br />
PS<br />
SF<br />
LB<br />
EX<br />
OR<br />
HI<br />
WINTER SPRING SUMMER FALL<br />
SEASON
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 11<br />
3C2. Abundance <strong>of</strong> Organisms in Ballast <strong>Water</strong><br />
(a) Total Density by Source<br />
We measured an average <strong>of</strong> 12,637 (s.e. = 5,533) total organisms per m 3 in the ballast<br />
water for all 169 vessels from domestic and foreign source ports. This estimate excludes the<br />
chain-forming diatoms, which were detected in the samples from 16 domestic tankers. Although<br />
the chain-forming diatoms are difficult to quantify (as above), we estimated average densities in<br />
excess <strong>of</strong> one million organisms/m 3 , approximately 100 fold higher than the densities <strong>of</strong> solitary<br />
diatoms and all other taxa measured for either domestic or foreign tankers (Fig. 3.2). We have<br />
excluded the chain-forming diatoms from the subsequent analyses and discussion <strong>of</strong> total density<br />
or abundance (i.e., abundance <strong>of</strong> all organisms) below.<br />
Figure 3.2. Densities <strong>of</strong> organisms in ballast water arriving to Port Valdez from foreign and domestic source<br />
ports. The estimated mean densities (#/m³ and standard errors) are shown separately for chain-forming diatoms,<br />
solitary diatoms, and all other taxa in ballast water <strong>of</strong> ships arriving from each domestic ports and foreign ports.<br />
Chain-forming diatoms were only detected on domestic ships (n=16), whereas the other groups were present on all<br />
domestic (n=150) and foreign (n=19) arrivals that were sampled. The data include all sample dates. Since arrivals<br />
from foreign ports all underwent ballast water exchange in open ocean, the source is indicated as exchange.<br />
DENSITY (#m³)<br />
1.E+07 16<br />
1.E+06 10 6<br />
1.E+05<br />
10 4<br />
1.E+04<br />
1.E+03<br />
10 2<br />
1.E+02<br />
150<br />
150<br />
CHAIN DIATOMS<br />
SOLITARY DIATOMS<br />
OTHER TAXA<br />
19<br />
19<br />
1.E+01<br />
0<br />
1.E+00 10<br />
DOMESTIC EXCHANGE<br />
SOURCE<br />
The average total density was significantly greater in ballast water from domestic sources<br />
compared to that for foreign sources (Fig. 3.2; 1-way ANOVA, F (1,168) = 3.63, P = 0.048). The<br />
chain-forming diatoms were only evident in the ballast water <strong>of</strong> domestic arrivals, increasing the<br />
magnitude <strong>of</strong> density differences between foreign and domestic traffic.<br />
The total abundance <strong>of</strong> organisms in ballast water differed among domestic sources. Of<br />
the three major domestic ports, arrivals from Puget Sound and San Francisco Bay had<br />
significantly greater densities than those from Long Beach and foreign sources, whereas the<br />
latter two were not different (Fig. 3.3; ANOVA, F (2,143) = 3.71, P = 0.027). We excluded both<br />
Oregon and Hawaii from this comparison, due to the limited sample size and absence <strong>of</strong> data for<br />
some seasons. Although average density from Oregon arrivals was similar to that for Long<br />
Beach and foreign arrivals, density for Hawaii arrivals was over 10-fold lower than all others.
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 12<br />
Figure 3.3. Densities <strong>of</strong> organisms in ballast water arriving to Port Valdez for each source port. The<br />
estimated mean densities (#/m³ and standard errors) are shown for all organisms by source port. The data include all<br />
sample dates (sample size indicated above bars) but exclude chain-forming diatoms. Source ports include: Puget<br />
Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB); Foreign port with open-ocean exchange<br />
(EX); Columbia River, Oregon (OR); and Barbers Point, Hawaii (HI).<br />
DENSITY (#m³)<br />
1.E+05<br />
1.E+04<br />
1.E+03<br />
1.E+02<br />
1.E+01<br />
48<br />
50<br />
46 19 2<br />
4<br />
1.E+00<br />
PS SF LB EX OR HI<br />
SOURCE<br />
The average total densities among domestic source ports corresponded to voyage<br />
duration, with densities decreasing with voyage duration (see Fig. 2.7 <strong>of</strong> Chapt 2). Arrivals from<br />
Puget Sound and San Francisco Bay had the shortest voyage duration (average <strong>of</strong> 4.3 and 6.0<br />
days, respectively) and highest densities. Tankers arriving from Hawaii had the longest voyage<br />
duration (average <strong>of</strong> 10.2 days) and lowest densities, whereas arrivals from Oregon and Long<br />
Beach were intermediate to the other domestic ports in both respects.<br />
(b) Density by Taxonomic Group and Source<br />
All ballast water samples contained living organisms, but the prevalence and density<br />
varied among taxonomic groups (Table 3.1). Copepods and diatoms were detected in ballast<br />
water from 100% <strong>of</strong> the ships sampled, and protozoans (primarily tintinnids) were found in<br />
nearly all samples. These three groups also exhibited the highest densities, dominating the<br />
plankton community in ballast water (see below).<br />
As with the total density measures, most taxonomic groups also occurred at greater<br />
average densities in ballast water from domestic sources, when pooled, compared to that from<br />
foreign sources (Table 3.1 and Fig 3.4). For most groups, this difference was 10- to 100-fold.<br />
The magnitude <strong>of</strong> density differences between domestic and foreign sources were much less for<br />
crustaceans, primarily due to the presence <strong>of</strong> copepods (see Table 3.1) and solitary diatoms.<br />
Din<strong>of</strong>lagellates were a notable exception to the general pattern, as average density was greatest<br />
in ballast water <strong>of</strong> the foreign arrivals.
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 13<br />
Figure 3.4. Densities <strong>of</strong> major taxonomic groups in ballast water arriving to Port Valdez from foreign and<br />
domestic source ports. The estimated mean densities (#/m³ and standard errors) are shown separately for 10<br />
different major groups <strong>of</strong> organisms in ballast water <strong>of</strong> ships arriving from each domestic ports (n=150) and foreign<br />
ports (n=19). Eight groups are distinct phyla that were most abundant in the ballast water, and two are composed <strong>of</strong><br />
multiple phyla, including eggs and trochophores (which were abundant but could not be classified by phylum) and<br />
all other invertebrates. The data include all sample dates. Since arrivals from foreign ports all underwent ballast<br />
water exchange in open ocean, the source is indicated as exchange.<br />
Echinodermata<br />
Chordata<br />
Other Inverts<br />
SOURCE<br />
Exchange<br />
Domestic<br />
Eggs & Trochophores<br />
TAXA<br />
Annelida<br />
Mollusca<br />
Din<strong>of</strong>lagellata<br />
Protozoa<br />
Crustacea<br />
Diatomacea<br />
0 10 2 10 4<br />
1 10 100 1000 10000 10000<br />
DENSITY (#/m³)<br />
On a finer scale, significant variation existed in the abundance <strong>of</strong> taxonomic groups<br />
among specific source port systems (Fig. 3.5 and Table 3.1). More specifically, differences were<br />
present when comparing mean densities for each taxonomic group among Puget Sound, San<br />
Francisco Bay, Long Beach, and foreign arrivals (ANOVA, see Fig. 3.5 for statistically<br />
significant differences). Among these four ports, the densities <strong>of</strong> most taxa were relatively low<br />
for foreign arrivals, with the exception <strong>of</strong> diatoms and din<strong>of</strong>lagellates. This pattern resulted from<br />
both the prevalence and densities <strong>of</strong> taxa among vessels. For example, the prevalence <strong>of</strong> most<br />
taxa was low in the ballast water <strong>of</strong> foreign arrivals, although the densities may not been<br />
particularly low for the few ships where a taxon was detected (Table 3.1). Across all arrivals,<br />
mean densities <strong>of</strong> the various taxa were generally lowest for both Oregon and Hawaii, although<br />
the limited sample size precludes any formal analysis.<br />
In contrast to total organism density, the greatest average densities for taxonomic groups<br />
did not always correspond to the shortest voyage duration (Fig. 3.5 and Table 3.1). For example,<br />
the highest average densities <strong>of</strong> protozoans, crustaceans, and bryozoans were measured for San<br />
Francisco Bay arrivals, and the brachyuran crabs were most abundant in samples from Long<br />
Beach.<br />
(c) Seasonal Variation in Density<br />
Significant differences among months were present in the total densities <strong>of</strong> organisms<br />
present in domestic ballast water (Fig. 3.6; ANOVA, F (11,168) = 1.98, P = 0.033). This resulted<br />
primarily from a relative increase during the spring and summer months.
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 14<br />
Figure 3.5. Densites <strong>of</strong> major taxonomic groups in ballast water arriving to Port Valdez for each source port.<br />
The estimated mean densities (#/m³ and standard errors) are shown separately for 10 different major groups <strong>of</strong><br />
organisms in ballast water <strong>of</strong> ships by source port. Eight groups are distinct phyla that were most abundant in the<br />
ballast water, and two are composed <strong>of</strong> multiple phyla, including eggs and trochophores (which were abundant but<br />
could not be classified by phylum) and all other invertebrates. Source ports include: Puget Sound, WA (PS); San<br />
Francisco Bay, CA (SF); Long Beach, CA (LB); Foreign port with open-ocean exchange (EX); Columbia River,<br />
Oregon (OR); and Barbers Point, Hawaii (HI). The data include all sample dates. Sample sizes for each source port<br />
as shown in Figure 3.3. Indicated by * are those taxa where mean density among ports, excluding HI and OR, is<br />
significantly different by ANOVA with confidence >95%.<br />
100000<br />
DINOFLAGELLATA*<br />
100000<br />
10000<br />
10000<br />
1000<br />
1000<br />
10100<br />
2<br />
10 100<br />
10<br />
10<br />
0 1<br />
01<br />
PS SF LB EX OR HI<br />
DIATOMACEA*<br />
PS SF LB EX OR HI<br />
100000<br />
10000<br />
1000<br />
10100<br />
2<br />
10<br />
0 1<br />
10 4 0<br />
PROTOZOA*<br />
PS SF LB EX OR HI<br />
100000<br />
10000<br />
4<br />
1000<br />
10100<br />
2<br />
10<br />
01<br />
ANNELIDA*<br />
PS SF LB EX OR HI<br />
DENSITY (#/m³)<br />
100000<br />
MOLLUSCA<br />
10000<br />
1000<br />
10100<br />
2<br />
10<br />
1<br />
PS SF LB EX OR HI<br />
100000<br />
10000<br />
10 4<br />
10 4 0<br />
1000<br />
10 2<br />
100<br />
10<br />
1<br />
CRUSTACEA*<br />
PS SF LB EX OR HI<br />
100000<br />
10000<br />
ECHINODERMATA*<br />
100000<br />
10000<br />
10 4<br />
1000<br />
1000<br />
10100<br />
2<br />
100<br />
10 2<br />
10<br />
10<br />
01<br />
0 1<br />
PS SF LB EX OR HI<br />
OTHER INVERTS*<br />
PS SF LB EX OR HI<br />
100000<br />
100000<br />
CHORDATA*<br />
10000<br />
10 10000<br />
1000<br />
1000<br />
10100<br />
2<br />
10100<br />
2<br />
10<br />
10<br />
0 1<br />
0 1<br />
PS SF LB EX OR HI<br />
SOURCE<br />
EGGS AND TROCHOPHORES<br />
PS SF LB EX OR HI
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 15<br />
Figure 3.6. Monthly densities <strong>of</strong> organisms in ballast water arriving to Port Valdez from domestic source<br />
ports. The estimated mean densities (#/m³ and standard errors) are shown for all organisms, except chain-forming<br />
diatoms, by month. The data for all domestic source ports are included in each month (sample size shown above<br />
bars).<br />
DENSITY (thousands/m³)<br />
80<br />
60<br />
40<br />
20<br />
0<br />
11<br />
10<br />
17 15 11 11<br />
10<br />
10 11 11 13 20<br />
Jan Mar May Jul Sep Nov<br />
SEASON<br />
The seasonal pattern in total density varied among arrivals from the 3 major domestic<br />
ports (Fig. 3.7). Arrivals from San Francisco Bay exhibited a strong spring peak in total<br />
plankton density, whereas the peak appeared later in arrivals from Puget Sound. In contrast, the<br />
density <strong>of</strong> organisms arriving from Long Beach was relatively stable throughout the year.<br />
Figure 3.7. Monthly densities <strong>of</strong> organisms in ballast water arriving to Port Valdez by domestic source ports.<br />
For each <strong>of</strong> the three major domestic source ports, the estimated mean densities (#/m³ and standard errors) are<br />
shown for all organisms, except chain-forming diatoms, by month. Sample size for each source port as follows,<br />
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 –<br />
5,3,3,2,5,6,5,4,4,3,2,6.<br />
DENSITY<br />
(thousands/m³)<br />
60<br />
40<br />
20<br />
0<br />
LONG BEACH<br />
Jan Mar May Jul Sep Nov<br />
DENSITY<br />
(thousands/m³)<br />
60<br />
40<br />
20<br />
0<br />
98.9<br />
SAN FRANCISCO<br />
Jan Mar May Jul Sep Nov<br />
DENSITY<br />
(thousands/m³)<br />
60<br />
40<br />
20<br />
0<br />
PUGET SOUND<br />
Jan Mar May Jul Sep Nov<br />
SEASON
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 16<br />
For the individual taxonomic groups, densities varied both by source port and month (Fig.<br />
3.8). In general, peak densities occurred between spring and summer months for all taxonomic<br />
groups, but the timing <strong>of</strong> these peaks differed among groups. Summed across the three major<br />
domestic ports (Puget Sound, San Francisco Bay, and Long Beach; Fig. 3.8a):<br />
• Din<strong>of</strong>lagellates, echinoderm larvae, chordates, and the combined eggs and trochophores<br />
exhibited peak mean densities in late summer;<br />
• Protozoans and diatoms exhibited a spring to early summer peak in mean density.<br />
• Molluscs and crustaceans for these combined ports were relatively high from spring through<br />
summer, exhibiting a bimodal distribution with spring and later summer peaks;<br />
• Annelids exhibited peaks in density during the summer months <strong>of</strong> June and August;<br />
• All other invertebrates, when combined, had relatively high densities from early spring<br />
through fall compared to the remainder <strong>of</strong> the year.<br />
The relative contributions <strong>of</strong> the three port systems to the overall temporal patterns varied<br />
significantly (Fig 3.8b-d). The general seasonal patterns (i.e., spring-summer peaks) in density<br />
were similar among ports, but clear differences existed in the magnitude and month <strong>of</strong> peak<br />
densities among ports. As noted previously for total density across the entire year (Fig. 3.5), the<br />
magnitude <strong>of</strong> peaks for the taxonomic groups did not correspond consistently to voyage duration.<br />
(d) Annual Variation in Density<br />
There were significant differences among years in the total density <strong>of</strong> plankton arriving<br />
from the three major ports for June 1997-1999 (Fig. 3.9). A 2-way ANOVA revealed differences<br />
among years (F (2,32) = 3.28, P = 0.055) but not ports (P>0.05), and the interaction was not<br />
significant (P>0.05).<br />
The magnitude <strong>of</strong> variation among years was more pronounced for the individual<br />
taxonomic groups in each <strong>of</strong> these ports (Fig. 3.10). Over half <strong>of</strong> the taxonomic groups exhibited<br />
significant differences among years, when analyzed individually for each port source (1-way<br />
ANOVA, see Fig 3.10 for statistically significant differences). As discussed above, some groups<br />
(e.g., echinoderm larvae and chordates) could not be compared statistically, due to low<br />
prevalence among ships.<br />
Interestingly, the changes among years were not consistent among port sources. For<br />
example, din<strong>of</strong>lagellates increased in successive years for Puget Sound and Long Beach arrivals,<br />
but was greatest in 1998 and virtually absent in the other two years for San Francisco arrivals.<br />
While peak years in protozoan and crustacean densities were similar among the port sources, the<br />
peak years for all other taxa were highly divergent among the port sources.
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 17<br />
Figure 3.8. Monthly densities <strong>of</strong> major taxonomic groups in ballast water arriving to Port Valdez from domestic source ports. The estimated monthly<br />
mean densities (#/m³ and standard errors) are shown separately for 10 different major groups <strong>of</strong> organisms in ballast water <strong>of</strong> ships arriving from domestic ports.<br />
Eight groups are distinct phyla that were most abundant in the ballast water, and two are composed <strong>of</strong> multiple phyla, including eggs and trochophores (which<br />
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<br />
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. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 18
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 19
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 20<br />
Figure 3.9. Densities <strong>of</strong> organisms in the ballast water arriving to Port Valdez from domestic ports in June <strong>of</strong><br />
three consecutive years, 1997-1999. The estimated mean densities (#/m³ and standard errors) are shown for all<br />
organisms, excluding chain-forming diatoms, by source port in each year (sample size indicated above bars). Source<br />
ports include: Puget Sound, WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB). Densities are also<br />
shown by year for all three source ports combined (Overall).<br />
3<br />
5<br />
3<br />
10<br />
1.E+04<br />
2<br />
3<br />
4<br />
3<br />
4<br />
6<br />
12<br />
11<br />
1997<br />
1998<br />
1999<br />
Density (#/m³)<br />
1.E+02<br />
1.E+00<br />
LB SF PS Overall<br />
SOURCE<br />
Figure 3.10. Densities <strong>of</strong> major taxonomic groups in the ballast water arriving to Port Valdez from domestic<br />
source ports in June <strong>of</strong> three consecutive years, 1997-1999. The estimated mean densities (#/m³ and standard<br />
errors) are shown for 10 different major groups <strong>of</strong> organisms in ballast water <strong>of</strong> ships arriving by year from three<br />
different source ports. Eight groups are distinct phyla that were most abundant in the ballast water, and two are<br />
composed <strong>of</strong> multiple phyla, including eggs and trochophores (which were abundant but could not be classified by<br />
phylum) and all other invertebrates. Sample size for each source port and year as indicated in Figure 3.9. Indicated<br />
by * are instances where ANOVA showed significant differences in density among years (above bars) and among<br />
ports (below taxa labels) with confidence >95%.<br />
100000<br />
10000<br />
1000<br />
100<br />
LB (JUNE)<br />
1997<br />
1998<br />
*<br />
1999<br />
* * *<br />
DENSITY (#/m³)<br />
10<br />
1<br />
100000<br />
Din<strong>of</strong>lagellata<br />
10000<br />
1000<br />
100<br />
10<br />
Diatomacea<br />
* *<br />
Protozoa<br />
Annelida<br />
SF (JUNE)<br />
Mollusca<br />
Crustacea<br />
Echinodermata<br />
Other Inverts<br />
Chordata<br />
1997<br />
1998<br />
1999<br />
Eggs & Trocophores<br />
*<br />
*<br />
1<br />
100000<br />
10000<br />
1000<br />
100<br />
10<br />
1<br />
*<br />
*<br />
*<br />
PS (JUNE)<br />
1997<br />
1998<br />
1999<br />
Din<strong>of</strong>lagellata<br />
Diatomacea<br />
Protozoa<br />
Annelida<br />
Mollusca<br />
Crustacea<br />
Din<strong>of</strong>lagellata<br />
Echinodermata<br />
*Diatomacea<br />
Other Inverts<br />
*Protozoa<br />
Chordata<br />
Annelida<br />
Eggs & Trocophores<br />
Mollusca<br />
Crustacea<br />
Echinodermata<br />
Other Inverts<br />
Chordata<br />
*Eggs & Trochophores
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 21<br />
3C3. Diversity <strong>of</strong> Organisms in Ballast <strong>Water</strong><br />
In a preliminary analysis <strong>of</strong> our Fine Analysis data (see Methods), there was a significant<br />
difference in the minimum number <strong>of</strong> taxa, and/or species richness, detected among arrivals from<br />
the three main domestic ports (Fig. 3.11). The average species richness was greatest for Long<br />
Beach arrivals and lowest for San Francisco Bay arrivals, and it does not correspond to voyage<br />
duration.<br />
Figure 3.11. Minimum number <strong>of</strong> taxa detected in the ballast water arriving to Port Valdez by domestic<br />
source ports. Shown are the mean number (including standard errors and sample size, above bars) <strong>of</strong> distinctly<br />
different taxa observed in plankton samples <strong>of</strong> ships from each source port. Source ports include: Puget Sound, WA<br />
(PS); San Francisco Bay, CA (SF); Long Beach, CA (LB). All sample dates included.<br />
The season <strong>of</strong> peak species richness differed among port sources (Fig. 3.12). Arrivals<br />
from Puget Sound and San Francisco Bay exhibited peaks in mean species richness in the spring<br />
and summer, whereas those from Long Beach had their highest species richness in the fall and<br />
spring. Subsequent analyses indicated differences among source ports for each season, as well as<br />
differences among seasons for each source port (1-way ANOVA, Fig. 3.12 indicates statistically<br />
significant differences).<br />
To date, we have identified 14 different nonindigenous species arriving to Port Valdez in<br />
the ballast water <strong>of</strong> oil tankers (Table 3.2). One is a fish species and all the other species are<br />
crustaceans (copepods and amphipods, which have successfully invaded the respective source<br />
ports <strong>of</strong> arriving tankers. To date, all <strong>of</strong> these identified NIS have been in ballast water from San<br />
Francisco and Long Beach.<br />
We expect the cumulative list will increase, as final identifications are still underway for<br />
the Fine Analysis data. Upon completion, we will report the frequency and density <strong>of</strong> these NIS<br />
in ballast water arriving from the respective source ports.
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 22<br />
Figure 3.12. Minimum number <strong>of</strong> taxa detected in the ballast water arriving to Port Valdez among domestic<br />
source ports and seasons. Shown are the mean number (including standard error above bars) <strong>of</strong> distinctly different<br />
taxa observed in plankton samples <strong>of</strong> ships from each source port and season. Source ports include: Puget Sound,<br />
WA (PS); San Francisco Bay, CA (SF); Long Beach, CA (LB). Seasons include: Winter, (December – February);<br />
Spring (March – May); Summer (June – August); and Fall (September – November). Indicated by * are significant<br />
differences (ANOVA with confidence >95%) in diversity among seasons within port (see legend) and among ports<br />
within season (see x-axis labels).<br />
Minimum Number <strong>of</strong> Taxa<br />
40 LB<br />
PS<br />
SF*<br />
30<br />
20<br />
10<br />
0<br />
*Fall<br />
*Spring *Summer *Winter<br />
Season<br />
Table 3.2. Nonindigenous species identified in ballast water arriving to Port Valdez. The source <strong>of</strong> ballast<br />
water is indicated in which each species was detected; when two sources are indicated, the species was found in<br />
ballast water from each source port. Source ports are: San Francisco Bay, CA (SF); Long Beach, CA (LB).<br />
Broad Taxa Species Ballast Source<br />
Amphipoda Ampelisca abdita SF and LB<br />
Monocorophium acherusicum<br />
SF<br />
Sinocorophium heteroceratum<br />
LB<br />
Gammarus daiberi<br />
SF and LB<br />
Grandidierella japonica<br />
SF<br />
Copepoda Limnoithona tetraspina SF<br />
Oithona davisae<br />
LB<br />
Acartiella sinensis<br />
SF<br />
Pseudodiaptimus marinus<br />
SF<br />
Pseudodiaptimus forbesi<br />
SF<br />
Sinocalanus doerrii<br />
SF<br />
Tortanus dextrilobatus<br />
SF<br />
Mysidacea Acanthomysis bowmani SF<br />
Chordata Acanthogobius flavimanus SF<br />
3D. Discussion<br />
Our analysis indicates that significantly greater numbers <strong>of</strong> organisms are discharged into<br />
Port Valdez and PWS in oil tankers arriving from domestic ports compared to those from foreign<br />
ports. This results from the number <strong>of</strong> arrivals and the density <strong>of</strong> organisms in their ballast<br />
water, as both are greatest for the domestic arrivals. Accounting for number <strong>of</strong> arrivals and<br />
density (by source port and season), Table 3.3 estimates the total supply <strong>of</strong> plankton that we
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 23<br />
sampled to be roughly 264 billion organisms in 1998. Of this, approximately 3% arrived from<br />
foreign traffic.<br />
The differences observed in total density, as well as taxon-specific density, among arrivals<br />
from different source ports may result from a combination <strong>of</strong> multiple factors, including (a)<br />
differences in initial densities, (b) differences in survivorship, and (c) effects <strong>of</strong> ballast water<br />
exchange (conducted for foreign but not domestic arrivals).<br />
Table 3.3. Estimated number <strong>of</strong> large, planktonic organisms delivered in tankers’ ballast water to PWS and<br />
Port Valdez in 1998. Shown by source port and season are (1) the estimated total ballast water volumes, (2) mean<br />
densities <strong>of</strong> planktonic organisms, including standard errors and sample size, and (3) total number <strong>of</strong> planktonic<br />
organisms arriving in the ballast water <strong>of</strong> oil tankers. Total Volumes are derived from Table 2.1. Mean densities<br />
were estimated from analysis <strong>of</strong> plankton samples, which were collected by 80micron net, and exclude chainforming<br />
diatoms (see text for description). Where no samples were available for a season (e.g., Hawaii and Oregon),<br />
the grand mean across all samples <strong>of</strong> that source port was used. The bottom row (Overall) estimates the total ballast<br />
water volumes and total organisms delivered as a sum. Source ports include: Puget Sound, WA (PS); San Francisco<br />
Bay, CA (SF); Long Beach, CA (LB); Foreign port with open-ocean exchange (EX); Columbia River, Oregon (OR);<br />
and Barbers Point, Hawaii (HI). Seasons include: Winter (January-March), Spring (April-June), Summer (July-<br />
September), and Fall (October-November).<br />
Density organisms (#/m³)<br />
Phyto.+Zoopl. Zoopl. Phyto.+Zoopl. Delivered Zoopl. Delivered<br />
Port/Source Season Total BW (m³) Mean(se) Mean(se) n Billions Billions<br />
Winter 1,802,432 5963(1432) 879(343) 11 10.75 1.58<br />
Spring 1,885,260 17602(9543) 3427(1423) 11 33.18 6.46<br />
PS Summer 2,032,432 24116(8198) 8539(3062) 12 49.01 17.35<br />
Fall 1,796,382 8546(2636) 1269(384) 8 15.35 2.28<br />
Grand total 7,516,506 42 108.30 27.68<br />
Winter 1,082,594 24686(13263) 13435(9497) 8 26.72 14.54<br />
Spring 1,345,169 48504(23134) 29876(14848) 13 65.25 40.19<br />
SF Summer 1,573,660 14572(2914) 10696(1959) 10 22.93 16.83<br />
Fall 1,340,469 8840(1961) 7162(1488) 12 11.85 9.60<br />
Grand total 5,341,892 43 126.75 81.17<br />
Winter 585,468 4298(1094) 3426(954) 11 2.52 2.01<br />
Spring 395,187 4848(667) 3734(490) 10 1.92 1.48<br />
LB Summer 504,630 15951(3274) 15145(3175) 6 8.05 7.64<br />
Fall 390,180 6574(1613) 5508(1639) 12 2.57 2.15<br />
Grand total 1,875,465 39 15.05 13.27<br />
Winter 379,377 8791(3127) 1852(423) 7 3.34 0.70<br />
Spring 265,764 3466(2144) 1634(775) 3 0.92 0.43<br />
EX Summer 149,280 18447(14784) 3393(3141) 5 2.75 0.51<br />
Fall 141,168 3571(1398) 3179(1100) 2 0.50 0.45<br />
Grand total 935,589 17 7.51 2.09<br />
Winter 262,638 - - 0 2.49 0.02<br />
Spring 113,000 - - 0 1.07 0.01<br />
OR Summer 97,287 - - 0 0.92 0.01<br />
Fall 273,966 9474(6138) 65(13) 2 2.60 0.02<br />
Grand total 746,891 7.08 0.05<br />
Winter 189,994 772 17 1 0.15 0.00<br />
Spring 111,145 - - 0 0.05 0.00<br />
HI Summer 142,596 102(72) 14(2) 2 0.01 0.00<br />
Fall 108,370 - - 0 0.05 0.00<br />
Grand Total 552,105 3 0.26 0.01<br />
Overall 16,968,448 145 264.94 124.26
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 24<br />
Among the domestic arrivals, many <strong>of</strong> the observed differences in prevalence and density<br />
probably result from initial differences at the locations where ballast tanks are filled. For<br />
example, this may explain the especially strong differences observed in densities <strong>of</strong> some<br />
organisms, such as din<strong>of</strong>lagellates and protozoans (Fig. 3.8), among source ports. Although we<br />
have very limited data on the initial densities within ballast tanks at the start <strong>of</strong> the tankers’<br />
voyages (see Chapter 5), the published literature indicates that significant variation in the density<br />
and diversity <strong>of</strong> plankton communities among these and other source ports should be expected.<br />
In this context, it is perhaps important to recognize some conspicuous differences that existed<br />
among the domestic source ports. Certainly there are many differences in the habitat<br />
characteristics (e.g., composition, extent, quality, proximity to ports, etc), which may influence<br />
what is initially entrained in the tankers’ ballast tanks. However, there are also two<br />
physical/chemical characteristics that are widely recognized to influence the composition and<br />
dynamics <strong>of</strong> biotic communities: temperature and salinity. Temperature clearly differed among<br />
domestic port systems, increasing from north to south. Among the major domestic ports, salinity<br />
was extremely low for San Francisco Bay compared to Puget Sound and Long Beach in 1998, in<br />
which rainfall was relatively high (due to El Nino Southern Oscillation) and had a<br />
disproportionately large effect on salinity in San Francisco Bay.<br />
Despite any initial differences in plankton communities, it is evident that survivorship<br />
during transit can contribute strongly to the observed differences in biota arriving from various<br />
source ports. A variety <strong>of</strong> studies have now shown a significant decline in the density <strong>of</strong><br />
planktonic organisms in ballast tanks during voyages, and the magnitude <strong>of</strong> decline is timedependent,<br />
increasing significantly with voyage duration (Wonham et al. 1996, LaVoie et al<br />
1999, Smith et al. 1999; however see below for possible exceptions). We have obtained similar<br />
results aboard oil tankers arriving to Port Valdez (Chapter 5). For most taxa, the decline has been<br />
attributed to mortality. However, for a few groups included in our analysis, such as diatoms and<br />
din<strong>of</strong>lagellates, it is possible for the organisms to develop dormant stages that can accumulate on<br />
the bottom <strong>of</strong> ballast tanks.<br />
Ballast water exchange undoubtedly had a significant effect on the plankton community<br />
associated with foreign arrivals, contributing to the major differences in biota between foreign<br />
and domestic arrivals. Exchange can significantly reduce the concentration <strong>of</strong> many organisms<br />
within ballast tanks, and it can also entrain additional organisms from the oceanic site <strong>of</strong><br />
exchange (Ruiz et al. 1997, 1999; see also Chapter 5). In our study, the combination <strong>of</strong> ballast<br />
water exchange and voyage duration (which was relatively high for foreign ports) would both<br />
operate reduce initial densities <strong>of</strong> coastal plankton and contribute to the lower abundance <strong>of</strong><br />
many taxonomic groups in ballast water <strong>of</strong> foreign arrivals compared to that from domestic<br />
arrivals. In contrast, the domestic arrivals did not undergo ballast water exchange and arrived to<br />
Port Valdez with the initial coastal water, following a relatively short voyage.<br />
We hypothesize that the combined effects <strong>of</strong> ballast water exchange and voyage duration,<br />
instead <strong>of</strong> initial densities, were responsible for observed differences in abundance <strong>of</strong> coastal<br />
organisms between domestic and foreign arrivals. More specifically, we suggest that these<br />
forces reduced the densities <strong>of</strong> predominantly coastal organisms such as cnidarians, flatworms,<br />
annelids, molluscs, chordates, echinoderms, bryozoans, barnacles, and many other crustacean<br />
groups (see Chapter 5 for further discussion).
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 25<br />
The effects <strong>of</strong> ballast water exchange for some taxonomic groups, and its contribution to<br />
observed differences in their abundance between foreign and domestic arrivals, is not so well<br />
resolved. Unlike the low abundance <strong>of</strong> coastal organisms, foreign arrivals had relatively high<br />
densities <strong>of</strong> din<strong>of</strong>lagellates, copepods, and solitary diatoms in their ballast water. Most <strong>of</strong> these<br />
organisms were probably oceanic in origin and were entrained during the exchange process.<br />
This is certainly the case for the copepods, for which the species were recognized as oceanic and<br />
the generation time is in excess <strong>of</strong> the voyage duration. However, there is some suggestion that<br />
an increase <strong>of</strong> phytoplankton can result from ballast water exchange, as generation times are<br />
relatively short and the organisms may respond rapidly to changes in water quality following<br />
exchange (Gollasch et al.1998, LaVoie et al. 1999). The extent to which populations <strong>of</strong> these<br />
taxa, either <strong>of</strong> coastal or oceanic origin, may have increased following exchange is uncertain.<br />
The temporal variation observed in plankton densities was largely expected. In general,<br />
the seasonal peaks in density corresponded to seasonal production and density variation<br />
measured for plankton in north temperate estuaries. The magnitude <strong>of</strong> variation observed among<br />
years also is evident in field studies, including especially an El Niño event such as that for 1998.<br />
The heavy rainfall in that year was associated with especially high densities <strong>of</strong> protozoans,<br />
solitary diatoms, and copepods in the arrivals from San Francisco Bay.<br />
Although we have identified 14 nonindigenous species in the ballast water arriving to Port<br />
Valdez, and have provided some comparative data on species richness, these results must be<br />
viewed with caution. Clearly these numbers are minimum estimates. Although both estimates<br />
will increase upon completion <strong>of</strong> the voucher identification (see results), the measures can only<br />
be applied to a subset <strong>of</strong> the taxa and will always represent a minimum value. More specifically,<br />
most <strong>of</strong> the larval invertebrates (e.g., molluscs, barnacles) include many different species, which<br />
cannot be readily distinguished as larvae. All bivalve larvae are therefore treated as one species<br />
in our analysis, masking the diversity that most certainly exists. Thus, this approach is useful<br />
primarily in describing minimum diversity <strong>of</strong> native and exotic species arriving in ballast water,<br />
and does not necessarily reflect actual diversity patterns in space or time.<br />
It is also important to recognize that our conclusions about patterns <strong>of</strong> abundance and<br />
diversity are focused on the large (>80 micron) segment <strong>of</strong> the plankton community within<br />
ballast tanks. We have provided some additional qualitative information about the macr<strong>of</strong>auna<br />
found on the bottom <strong>of</strong> tankers’ ballast tanks (Chapt 6, this report). Thus, our data do not<br />
address density or diversity <strong>of</strong> microorganisms and taxa missed by an 80 micron mesh. The<br />
dynamics <strong>of</strong> these groups are very much in debate, as few good data exist to discern the potential<br />
for population changes (either declines or increases), due especially to mortality, dormancy and<br />
cyst formation, or ballast water exchange.<br />
Beyond the variation in plankton delivery by time and source port, our data underscore<br />
that both the concentration and cumulative amount <strong>of</strong> plankton arriving in tankers’ ballast water<br />
to Port Valdez and PWS is relatively high compared to that estimated for other ports (e.g.,<br />
Carlton and Geller 1993, Smith et al. 1999). This results from both the volume <strong>of</strong> water<br />
delivered (Chapter 2) and the concentration <strong>of</strong> plankton, as both values are relatively high. We<br />
hypothesize that the abundant plankton results from the short voyage duration for domestic<br />
traffic, accounting for approximately 97% <strong>of</strong> the total tanker arrivals at present. In contrast,
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 26<br />
although Chesapeake Bay receives more total ballast water per year than PWS, most <strong>of</strong> the<br />
ballast water comes from Europe, arriving in 10-14 days with an average concentration <strong>of</strong> 200<br />
organisms / m 3 (Smith et. al. 1999; Ruiz et al., unpubl. data). Furthermore, it appears that<br />
ballast water arriving to the Chesapeake from domestic ports also has a lower density than that<br />
arriving to PWS from domestic ports.<br />
Finally, from an invasion perspective, there are three unusual features <strong>of</strong> our analysis that<br />
deserve explicit mention:<br />
• This is the most comprehensive analysis <strong>of</strong> domestic, coastwise ballast transfer. Most<br />
organisms that arrive in ballast water to PWS come from domestic source ports, which are<br />
themselves highly invaded. Thus, our study examines the opportunity for sequential<br />
invasions, which can “leapfrog” up the coast following initial colonization <strong>of</strong> North America.<br />
• The delivery <strong>of</strong> ballast water by tankers to Port Valdez is a relatively recent development,<br />
beginning in 1977. Although many features may influence the risk <strong>of</strong> invasion, it is <strong>of</strong>ten<br />
considered to increase with the frequency, density, and duration <strong>of</strong> inoculation. Our results<br />
indicate that the risk associated with the first two <strong>of</strong> these is relatively high. However, the<br />
operation <strong>of</strong> this transfer mechanism has only existed for three decades. Even at the current<br />
rates <strong>of</strong> organism delivery (see above), invasion success may be influenced strongly by<br />
duration. In contrast, many other ports have been receiving ballast water and ballast<br />
materials for a century or more.<br />
• Delivery <strong>of</strong> ballast water from foreign sources by tankers is even a more recent development,<br />
beginning in 1996. Although this accounted for only 3% <strong>of</strong> the total volume <strong>of</strong> ballast water<br />
delivered in 1998, all <strong>of</strong> the ballast water delivered by foreign arrivals had undergone<br />
exchange. Some coastal organisms remained in the exchanged tanks (see Chapter 5);<br />
however, the total supply <strong>of</strong> organisms from foreign ports is both relatively small and<br />
extremely recent compared to other port systems.<br />
3E. References<br />
Carlton, J.T. and J. B. Geller. 1993. Ecological roulette: the global transport <strong>of</strong> nonindigenous<br />
marine organisms. Science 261: 78-82.<br />
Gollasch, S., M. Dammer, J. Lenz and H.G. Andres. 1998. Non-indigenous organisms introduced<br />
via ships into German waters. Pp. 50-64 in Carlton, J.T. (ed.), Ballast <strong>Water</strong>: Ecological and<br />
Fisheries Implications. International Council for the Exploration <strong>of</strong> the Sea (ICES), Denmark.<br />
Lavoie, D. M., L.D. Smith and G.M. Ruiz. 1999. The potential for intracoastal transfer <strong>of</strong> nonindigenous<br />
species in the ballast water <strong>of</strong> ships. Est. Coast. Shelf Sci. 48: 551-564.<br />
Ruiz, G.M. and A.H. Hines. 1997. Patterns <strong>of</strong> nonindigenous species transfer and invasion in<br />
Prince William Sound, Alaska: Pilot Study. Report, Prince William Sound Regional Citizens’<br />
Advisory Council. 80pp.<br />
Ruiz, G.M., J.T. Carlton, A. H. Hines and E.D. Grosholz. 1997. Global invasions <strong>of</strong> marine and<br />
estuarine habitats by non-indigenous species: Mechanisms, extent, and consequences. Am. Zool.<br />
37: 621-632.
Chapt 3. <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>, page 3- 27<br />
Ruiz, G.M., P. F<strong>of</strong>on<strong>of</strong>f and A.H. Hines. 1999. Non-indigenous species as stressors in estuarine<br />
and marine communities: Assessing invasion impacts and interactions. Limnol. Oceanogr. 44:<br />
950-972.<br />
Smith, L.D., M.J. Wonham, L.D. MCann, D.M. Reid, G.M. Ruiz and J.T. Carlton. 1996.<br />
<strong>Biological</strong> invasions by nonindigenous species in United States waters: Quantifying the role <strong>of</strong><br />
ballast water and sediments. Parts I and II. Final Report to the U.S. Coast Guard and the U.S.<br />
Department <strong>of</strong> Transportation. 246 pp.<br />
Smith, L.D., M.J. Wonham, L.D. McCann, G.M. Ruiz, A.H. Hines and J.T. Carlton. 1999.<br />
Invasion pressure to a ballast-flooded estuary and an assessment <strong>of</strong> inoculant survival. Biol.<br />
<strong>Invasions</strong> 1: 67-87.<br />
Wonham. M.J., W.C. Walton, A.M. Frese and G.M. Ruiz. 1996. Transoceanic transport <strong>of</strong><br />
ballast water: <strong>Biological</strong> and physical dynamics <strong>of</strong> ballasted communities and the effectiveness<br />
<strong>of</strong> mid-ocean exchange. Final Report to the U.S. Fish and Wildlife Service and the Compton<br />
Foundation.
Chapt 4. Predicting Initial Survival <strong>of</strong> Ballast <strong>Water</strong> Organisms, page 4- 1<br />
Chapter 4. Predicting Initial Survival <strong>of</strong> Ballast <strong>Water</strong> Organisms<br />
Anson H. Hines, Smithsonian Environmental Research Center<br />
Gregory M. Ruiz, Smithsonian Environmental Research Center<br />
George Smith, Smithsonian Environmental Research Center<br />
Melissa Frey, Smithsonian Environmental Research Center<br />
4A. Purpose<br />
When ballast water is discharged into the receiving waters, the associated plankton<br />
encounters new conditions without time to acclimate. Survival may depend on short-term<br />
tolerances to accute variation in salinity-temperature combinations. If temperature-salinity<br />
conditions <strong>of</strong> ballast water closely match those <strong>of</strong> the receiving waters, then initial survival is<br />
predicted to be higher than when the conditions do not match closely. To determine if NIS<br />
arriving in ballast water can survive the initial exposure to temperature-salinity conditions in<br />
Prince William Sound, we tested the match <strong>of</strong> conditions between ballast water and ship-side<br />
water, and the short-term survival <strong>of</strong> ballast organisms in representative combinations.<br />
4B. Temperature & Salinity: Match <strong>of</strong> Source and Receiving Ports<br />
4B1. Methods<br />
Samples <strong>of</strong> ballast water were collected from segregated ballast tanks and from ambient<br />
waters adjacent to each ship sampled for ballast water plankton. A small niskin bottle lowered<br />
through hatch covers into the ballast tanks and lowered <strong>of</strong>f the end <strong>of</strong> the ship’s berth to collect<br />
samples from the water surface and 10 m depth, which was determined to be below a potential<br />
thermocline or pycnocline. Salinity was determined to the nearest ppt with a refractometer and<br />
temperature to the nearest 0.5 o C with a hand-held thermometer.<br />
4B2. Results<br />
Temperature and salinity <strong>of</strong> the receiving waters <strong>of</strong> Port Valdez exhibit a distinct<br />
seasonal pattern (Fig. 4.1a, b). <strong>Water</strong> temperatures <strong>of</strong> Port Valdez at 10 m depth cycle<br />
seasonally from a low <strong>of</strong> 4 o C in February to a high <strong>of</strong> 13 o C in July. Surface water temperatures<br />
are more variable and 1-5 o C warmer than deep water in the spring. Salinity during December to<br />
April was about 31 ppt and the water column was well mixed. <strong>Water</strong> in Port Valdez was sharply<br />
stratified by depth as snow melted from late April to September, with salinities <strong>of</strong> surface waters<br />
dropping to 4-15 ppt while salinities at 10m depth declined only to about 25 ppt.<br />
<strong>Water</strong> in the segregated ballast tanks rarely exhibited much depth stratification.<br />
Temperatures <strong>of</strong> segregated ballast water varied seasonally with a winter mean low <strong>of</strong> about 7.5<br />
o C (+-3 o C) and a summer high <strong>of</strong> about 16 o C (+-3 o C) (Fig. 4.1c). Salinities <strong>of</strong> ballast water did<br />
not exhibit a seasonal pattern, but salinities fell into two distinct ranges, depending on the source<br />
port <strong>of</strong> the tanker (Fig. 4.1d). Most tankers delivered high salinity ballast water (ca. 30 ppt,<br />
range 20-36 ppt). In contrast, about 20% <strong>of</strong> the tankers throughout the year released ballast water<br />
<strong>of</strong> low salinity (ca. 4 ppt, range 0-14 ppt, mainly from Benicia in San Francisco Bay, especially<br />
during the heavy El Niño rains in 1998).
Chapt 4. Predicting Initial Survival <strong>of</strong> Ballast <strong>Water</strong> Organisms, page 4- 2<br />
Figure 4.1 Seasonal Cycles <strong>of</strong> Temperatures and Salinities <strong>of</strong> Receiving and Ballast <strong>Water</strong>. Shown above are<br />
temperatures (A) and salinities <strong>of</strong> Port Valdez, Alaska at 0m and 10m depth. Shown below are average<br />
temperatures (C) and salinities (D) <strong>of</strong> ballast water discharged into Port Valdez by tankers. Averages are for two<br />
tank depths (0m and 10m depth) combined.<br />
A. Port Valdez<br />
C. Ballast <strong>Water</strong><br />
Temperature (ºC)<br />
25<br />
20<br />
15<br />
10<br />
5<br />
0<br />
Dec Feb Apr Jun Aug Oct Dec<br />
0 m<br />
10 m<br />
Temperature (ºC)<br />
25<br />
20<br />
15<br />
10<br />
5<br />
0<br />
Dec Feb Apr Jun Aug Oct Dec<br />
B. Port Valdez<br />
40<br />
D. Ballast <strong>Water</strong><br />
40<br />
Salinity (‰)<br />
30<br />
20<br />
10<br />
0 m<br />
10 m<br />
Salinity (‰)<br />
0<br />
Dec Feb Apr Jun Aug Oct<br />
SEASON<br />
Dec<br />
30<br />
20<br />
10<br />
0<br />
Dec Feb Apr Jun Aug Oct Dec<br />
SEASON<br />
Annual variation in temperature and salinity among source ports and receiving waters <strong>of</strong><br />
Port Valdez was compared in June <strong>of</strong> 1997, 1998, and 1999 (Fig. 4.2). Temperature <strong>of</strong> ballast<br />
water from Long Beach (13-14 o C) and San Francisco(13-14 o C) was about a degree warmer than<br />
from Puget Sound (11-12 o C), but temperature <strong>of</strong> ballast water from Puget Sound was similar to<br />
the surface water <strong>of</strong> Port Valdez (11-12 o C), which was 1-3 o C warmer than water at 10 m depth<br />
(7-9 o C). However, there were no significant differences in temperatures among years. Salinity <strong>of</strong><br />
ballast water from Long Beach (33ppt) was highest <strong>of</strong> the source ports, while that from San<br />
Francisco (10-14 ppt) was the lowest, and Puget Sound was intermediate (29-30 ppt). Surface<br />
salinity at the surface <strong>of</strong> Port Valdez (10-21ppt) was similar to ballast water from San Francisco<br />
Bay, while salinity at 10 m depth in Port Valdez (29-30 ppt) was similar to ballast water from<br />
Puget Sound. Salinity <strong>of</strong> ballast water from Long Beach and Puget Sound, as well as deep water<br />
at Port Valdez, did not differ among years. However, salinity <strong>of</strong> ballast water from San<br />
Francisco Bay and at the surface in Port Valdez was lowest in 1998, highest in 1997 and<br />
intermediate in 1999.<br />
Thus, there was <strong>of</strong>ten a good correspondence <strong>of</strong> physical characteristics between ballast<br />
water and receiving water, depending on the source port, time <strong>of</strong> year and water depth in Port<br />
Valdez. Temperatures <strong>of</strong> ballast water were a bit higher than <strong>of</strong> receiving waters, but the<br />
differences were not great, and there was considerable overlap between ballast and receiving<br />
water throughout the year. Higher salinities <strong>of</strong> ballast water from most source ports were similar<br />
to deeper water <strong>of</strong> Port Valdez throughout the year and similar to surface water during winter<br />
and early spring. During summer the vertical stratification in Port Valdez resulted in ballast<br />
water from both high and low salinities having good correspondence in major areas <strong>of</strong> the<br />
receiving waters. Based on these physical characteristics <strong>of</strong> ballast and receiving water,<br />
temperatures <strong>of</strong> Port Valdez would not appear to prevent survival <strong>of</strong> organisms from most source<br />
ports. Nonindigenous species from nearly fresh water, estuarine and full-strength sea water may<br />
also find corresponding salinities in Port Valdez.
Chapt 4. Predicting Initial Survival <strong>of</strong> Ballast <strong>Water</strong> Organisms, page 4- 3<br />
Figure 4.2 Annual Variation in Temperature and Salinity <strong>of</strong> Ballast and Receiving <strong>Water</strong>. Shown are<br />
temperatures (top) and salinities (bottom) <strong>of</strong> ballast water arriving to Port Valdez in tankers from west-coast source<br />
ports (PS = Puget Sound, WA; SF = San Francisco, CA; LB = Long Beach, CA), and <strong>of</strong> recieving waters <strong>of</strong> Port<br />
Valdez at surface (V-0) and 10m depth (V-10). Bars indicate means and S.E. for June 1997, 1998, 1999.<br />
15.0<br />
June’97<br />
June’98<br />
June’99<br />
DEGREES (ºC)<br />
10.0<br />
5.0<br />
0.0<br />
PS SF LB V-0 V-10<br />
30.0<br />
SALINITY (‰)<br />
20.0<br />
10.0<br />
0.0<br />
PS SF LB V-0 V-10<br />
SOURCE<br />
4C. Temperature-Salinity Tolerance Experiments <strong>of</strong> Ballast <strong>Water</strong> Plankton<br />
4C1. Methods<br />
We conducted experiments at the SERC laboratory in Valdez to test for temperature x<br />
salinity tolerance <strong>of</strong> selected planktonic organisms arriving in segregated ballast water. Based on<br />
the two salinity categories <strong>of</strong> segregated ballast water released into Port Valdez (see above, Fig.<br />
4.1a, b), we grouped the experimental organisms into "freshwater taxa" and "seawater taxa" (Fig.<br />
4.3).<br />
Figure 4.3. Survivorship <strong>of</strong> Ballast <strong>Water</strong> Organisms in Salinity x Temperture Experiments. Survivorship <strong>of</strong><br />
ballast water organisms at 96 hour exposure to 9 combinations <strong>of</strong> salinity and temperature in laboratory experiments.<br />
Three salinities (10, 20, 30 ppt) and three temperatures (3, 9, 15ºC) were tested to represent the range <strong>of</strong> seasonal<br />
variation in Port Valdez. (A) Trials with organisms from ballast water with fresh water sources (n = 9). (B) Trials<br />
with organisms from ballast water with seawater sources (n = 15).<br />
A<br />
B<br />
Mean Survivorship<br />
1.0<br />
0.8<br />
0.6<br />
0.4<br />
0.2<br />
0.0<br />
1.0<br />
0.8<br />
"Freshwater" Taxa<br />
3 C<br />
9 C<br />
15 C<br />
10 20 30<br />
"Seawater" Taxa<br />
0.6<br />
0.4<br />
0.2<br />
0.0<br />
10 20 30<br />
Salinity (ppt)
Chapt 4. Predicting Initial Survival <strong>of</strong> Ballast <strong>Water</strong> Organisms, page 4- 4<br />
Nine combinations <strong>of</strong> three temperatures (3, 9, and 15 o C) and three salinities (10, 20, and<br />
30 ppt) were selected to represent the seasonal range <strong>of</strong> conditions for Port Valdez. Organisms<br />
used in the experiments are collected from the common species <strong>of</strong> plankton arriving in tanker<br />
ballast water. For each experiment, 10 individuals were placed in each <strong>of</strong> 3 replicate culture<br />
dishes at each <strong>of</strong> the 9 treatment combinations. Thus, there were 27 trials in each experiment (9<br />
treatments x 3 replicates). The test organisms were sorted in the lab and transferred directly to<br />
culture dishes maintained in incubators for 96 hrs, simulating release <strong>of</strong> ballast water into<br />
conditions <strong>of</strong> the Sound. Phytoplankton or brine shrimp nauplii were supplied as food to the<br />
cultures during the test period.<br />
Experiments (n = 24) were completed with organisms from the following taxonomic<br />
categories:<br />
Tintinnid protistan<br />
Nemertean worm larvae<br />
Spionid polychaete worm larvae<br />
Gastropod veliger larvae<br />
Copepods<br />
calanoid<br />
harpacticoid<br />
cyclopoid (Oithona spp.)<br />
Barnacle nauplii<br />
Crab zoea<br />
Mysid shrimp<br />
1 experiment<br />
2 experiments<br />
3 experiments<br />
1 experiment<br />
4 experiments<br />
1 experiment<br />
6 experiments<br />
3 experiments<br />
1 experiment<br />
2 experiments<br />
Nine <strong>of</strong> these were "freshwater taxa" and 15 were "seawater taxa". We intentionally selected<br />
copepods especially Oithona spp., for many <strong>of</strong> our experiments, because we recognized these as<br />
NIS arriving in apparently good condition from San Francisco Bay.<br />
4C2. Results<br />
Short-term survivorship <strong>of</strong> these ballast water organisms was high (>50%) for fresh water<br />
taxa at 10 ppt, and for seawater taxa at 20-30 ppt. These short-term experiments also showed that<br />
the ballast organisms had distinct, but quite broad tolerances that clearly overlap conditions <strong>of</strong><br />
temperature and salinities in Port Valdez. For example, although there was considerable<br />
variation in survivorship among individual experiments, mean survivorship <strong>of</strong> calanoid copepods<br />
varied from about 20-80% for each test salinity, but survivorship <strong>of</strong> calanoids in most<br />
experiments increased with salinity (Fig. 4.3). Oithona spp. (which include known NIS<br />
copepods) were able to tolerate salinity-temperature conditions they would encounter in the<br />
receiving waters <strong>of</strong> Port Valdez.<br />
The freshwater and seawater taxa differed substantially in their patterns<br />
<strong>of</strong> temperature x salinity tolerance (Fig. 4.3). The survivorship <strong>of</strong> seawater taxa at any <strong>of</strong> the 3<br />
test temperatures generally increased with increasing salinity, and there were not great<br />
differences in survivorship among temperatures. However, survivorship <strong>of</strong> freshwater taxa at all<br />
temperatures generally declined sharply with increasing salinity, and survivorship at 3 o C was<br />
markedly lower than at 9 or 15 o C.
Chapt 4. Predicting Initial Survival <strong>of</strong> Ballast <strong>Water</strong> Organisms, page 4- 5<br />
4D. Conclusions<br />
Planktonic species arriving to Port Valdez in ballast water have high potential <strong>of</strong><br />
surviving the salinity-temperature conditions that they encounter during initial discharge from<br />
the ship. Although some taxa will not tolerate some salinity layers in the seasonally stratified<br />
conditions in the Port, the overlap <strong>of</strong> ballast water with Port conditions at some strata is high.<br />
Plankton in the ballast water, including known NIS such as Oithona spp., should be able to<br />
tolerate these conditions. Conditions other than initial salinity-temperature combinations<br />
probably determine whether or not these organisms survive to become established within Prince<br />
William Sound.
Chapt 5 Ballast <strong>Water</strong> Exchange Experiments, page 5- 1<br />
Chapter 5. Ballast <strong>Water</strong> Exchange Experiments on Tankers<br />
Gregory M. Ruiz, Smithsonian Environmental Research Center<br />
Anson H. Hines, Smithsonian Environmental Research Center<br />
George Smith, Smithsonian Environmental Research Center<br />
Melissa Frey, Smithsonian Environmental Research Center<br />
Safra Altman, Smithsonian Environmental Research Center<br />
Kimberly Philips, Smithsonian Environmental Research Center<br />
Tami Huber, Smithsonian Environmental Research Center<br />
Sara Chaves, Smithsonian Environmental Research Center<br />
5A. Purpose<br />
The primary objective <strong>of</strong> this research component is to measure the efficacy <strong>of</strong> ballast<br />
water exchange in removing various types <strong>of</strong> taxa from ballast tanks <strong>of</strong> oil tankers. There are very<br />
few quantitative studies that have measured the effects <strong>of</strong> exchange, and these are restricted to<br />
just a few vessel types and measure the effect on a small subset <strong>of</strong> entrained taxa. It is likely,<br />
however, that the efficacy <strong>of</strong> exchange varies by vessel type, tank design, and organism type. To<br />
date, there have been no measures <strong>of</strong> ballast water exchange for oil tankers.<br />
Ballast water exchange is the most widely used national and international management<br />
strategy to limit new invasions associated with ships’ ballast water (Hallegraeff 1998, Zhang &<br />
Dickman 1999, Dickman and Zhang 1999,. Moreover, exchange is currently the only treatment<br />
method available for commercial ships to reduce the quantities <strong>of</strong> non-indigenous coastal plankton<br />
in ballast water (National Research Council 1996). This practice is recommended by the<br />
International Maritime Organization (IMO) to reduce the risk <strong>of</strong> invasion by shipping.<br />
Furthermore, the U.S. Congress passed the National Invasive Species Act <strong>of</strong> 1996 (NISA) to<br />
encourage ballast water exchange. Specifically, NISA requests that vessels arriving from outside<br />
<strong>of</strong> the Exclusive Economic Zone (EEZ) voluntarily conduct open-ocean exchange <strong>of</strong> ballast tanks<br />
to be discharged in U.S. ports<br />
Commercial ships practice two basic types <strong>of</strong> ballast water exchange to replace coastal with<br />
oceanic water. Flow-Through (FT) Exchange is conducted by pumping oceanic sea water<br />
continuously through a ballast tank to flush out the ballast water originating from a coastal source<br />
port. Empty-Refill (ER) Exchange is performed by emptying a ballast tank <strong>of</strong> its coastal water and<br />
refilling it with oceanic water.<br />
Each exchange method may vary in efficacy due to the amount and circulation <strong>of</strong> water<br />
being removed, independent <strong>of</strong> any tank- or vessel-specific effects on efficacy. For example, FT<br />
Exchange initially has the effect <strong>of</strong> dilution but not complete replacement <strong>of</strong> ballast water.<br />
Alternatively, organisms may differ in their distribution or response to water turbulence. Some<br />
taxa may swim against currents or always reside near the bottom <strong>of</strong> tanks, which could greatly<br />
influence the effect <strong>of</strong> ballast water exchange on removal.<br />
To maximize the degree <strong>of</strong> exchange, multiple exchanges are <strong>of</strong>ten recommended. The<br />
current IMO standard recommendation is 300% exchange for Flow-Through, while 100-200%
Chapt 5 Ballast <strong>Water</strong> Exchange Experiments, page 5- 2<br />
exchange is common for Empty-Refill. These recommended standards provide a theoretical level<br />
<strong>of</strong> at least 90% replacement <strong>of</strong> coastal water by oceanic water, but this is largely untested among<br />
the broad range <strong>of</strong> vessel types and tank configurations. Specifically, there are almost no<br />
experimental analyses which quantify the efficacy <strong>of</strong> alternative exchange methods and multiple<br />
tank exchanges, even though (a) this is the present national and international management strategy<br />
being implemented and (b) the cost <strong>of</strong> such exchanges is substantial in ship fuel and operations<br />
time.<br />
We initiated a rigorous quantitative comparison <strong>of</strong> ballast water exchange methods on oil<br />
tankers arriving to PWS. Across two years, we conducted replicated exchange experiments,<br />
allowing us to measure the effects <strong>of</strong> both exchange methods on reduction <strong>of</strong> entrained organisms<br />
and standard physical and biological tracers.<br />
We hypothesize that (1) Empty-Refill exchange will have the highest efficiency, (2)<br />
relatively little reduction in density occurs after the first exchange event, and (3) a significant<br />
difference exists among taxa on the effect <strong>of</strong> ballast water exchange. Our experiments were<br />
designed to directly test these hypotheses and provide needed quantitative data on this management<br />
practice.<br />
This work was initiated in two phases, extending the duration <strong>of</strong> the analyses and<br />
allowing us to increase the replication (and therefore strengthen the statistical power and value <strong>of</strong><br />
this analysis). The first phase was initiated in summer <strong>of</strong> 1998. Through the cooperation and<br />
financial support from the American Petroleum Institute, SeaRiver and ARCO, we conducted the<br />
ballast water exchange experiments on four separate voyages <strong>of</strong> tankers to Alaska in June/July <strong>of</strong><br />
that year. The second phase was initiated in spring <strong>of</strong> 1999, when we received additional<br />
funding from U.S. Fish and Wildlife Service to conduct similar experiments (with increased<br />
measurements) on another four voyages in summer. In addition to analysis <strong>of</strong> these experiments,<br />
we agreed (with additional funding from phase two) to provide a review <strong>of</strong> existing data on the<br />
efficacy <strong>of</strong> ballast water exchange, allowing us to examine our results from oil tankers to those<br />
reported for other vessel types.<br />
All <strong>of</strong> the experiments have been completed, but we have not yet completed the full<br />
analysis <strong>of</strong> all samples. We report here on the experiments conducted, including the status <strong>of</strong><br />
sample analysis and initial results. We will provide a comprehensive report <strong>of</strong> the results across<br />
both phases upon completion <strong>of</strong> our analyses. We anticipate that these results will be available<br />
by June 2000.<br />
5B. Methods<br />
Although the overall goal <strong>of</strong> this research was to measure the efficacy <strong>of</strong> ER and FT<br />
methods <strong>of</strong> ballast water exchange in removing coastal plankton from ballast water, our<br />
experimental design allowed us to address three specific objectives:<br />
• Compare the efficacy <strong>of</strong> ER and FT exchange methods in removing a range <strong>of</strong> different<br />
materials (biotic and abiotic);<br />
• Measure the effect <strong>of</strong> repeated exchanges: comparing 100, 200, and 300% exchange <strong>of</strong> the<br />
tanks.
Chapt 5 Ballast <strong>Water</strong> Exchange Experiments, page 5- 3<br />
• Measure the survivorship <strong>of</strong> organisms in ballast water over the course <strong>of</strong> routine voyages.<br />
Since the density <strong>of</strong> organisms can change during a voyage (see Chapter 3 for discussion), it<br />
was important to control for such changes in experimental tanks that were independent <strong>of</strong> the<br />
exchange treatment. For this purpose, we included identical measures for an unexchanged<br />
control tank on every vessel. Although key to the experimental analysis, this also provided<br />
an opportunity to address this third objective.<br />
All experiments were conducted aboard oil tankers during regular operations, en route to<br />
Port Valdez. Each tanker served as an experimental platform. Each ship was boarded by a pair <strong>of</strong><br />
SERC staff at a domestic source port (San Francisco, Puget Sound, or Long Beach), where ballast<br />
water used to fill the segregated ballast tanks just in advance <strong>of</strong> departure for Port Valdez. The<br />
tanks underwent various treatments and were sampled repeatedly during the voyage (below).<br />
Experimental Design<br />
The experiment consisted <strong>of</strong> a replicated, factorial, and paired design. On each ship, we<br />
sought to use 3 different ballast tanks that were each subjected to a different treatment: No<br />
exchange (=Control), ER Exchange, and FT Exchange. Each Treatment tank was sampled as<br />
many as 5 time points, coinciding with: initial ballast loading, 100% exchange, 200% exchange,<br />
300% exchange, and final at arrival to PWS.<br />
Ballast water was loaded in accordance with standard operating procedures at dockside. All<br />
exchanges occurred in open, oceanic conditions well outside the influence <strong>of</strong> coastal waters (>75<br />
miles <strong>of</strong>fshore). Exchanges <strong>of</strong> tanks were managed by ships’ crews in coordination with the desired<br />
sampling schedule. For FT Exchange, sea water was pumped into the ballast tanks, causing ballast<br />
water to overflow through the top <strong>of</strong> the tanks and onto the deck. After a volume <strong>of</strong> water equal to<br />
the volume <strong>of</strong> the tank was pumped, the exchange was interrupted and samples were collected. For<br />
ER Exchange, ballast tanks were drained initially by gravity and then by pumping before refilling<br />
with sea water. The process required approximately 12 hours for each multiple <strong>of</strong> exchange.<br />
The specific details <strong>of</strong> implementation and sampling are described below in various<br />
sections.<br />
Biotic and Abiotic Tracers.<br />
In addition to quantifying the effect <strong>of</strong> ballast water exchange on entrained plankton<br />
communities in the ballast tanks, we used four different types <strong>of</strong> tracers for parallel measures <strong>of</strong><br />
efficacy. One <strong>of</strong> these (salinity <strong>of</strong> the resident water) simply involved collecting and measuring<br />
attributes <strong>of</strong> the resident water. The other three involved materials that we added directly to the<br />
ballast tanks, including: Rhodamine dye to trace the fate <strong>of</strong> the initial water; 1 um Fluorescent<br />
Microspheres that simulate passive particles such as cysts; and newly hatched Artemia (brine<br />
shrimp) nauplii, native to San Francisco Bay, as a living particle.<br />
Each tracer can provide information about different components <strong>of</strong> the ballast tank<br />
environment during exchange (as indicated above). Moreover, we were interested in developing<br />
some standardized measures for comparisons across ships. Since the resident community within<br />
each ship’s ballast tanks may differ considerably (see Chapter 3), the tracers could provide a
Chapt 5 Ballast <strong>Water</strong> Exchange Experiments, page 5- 4<br />
common currency for comparing exchange performance among many ships in a way that is simply<br />
not possible for the entrained plankton communities.<br />
Tracers were added to ballast tanks in a standardized way, following approval for use in<br />
these experiments by the U.S. Environmental Protection Agency. The quantity <strong>of</strong> tracer was<br />
chosen to produce desired concentrations for the specific volumes <strong>of</strong> each tank, such that<br />
anticipated dilution during exchanges would allow us to detect at least a 100-fold reduction in<br />
measurable concentrations <strong>of</strong> each tracer.<br />
Tracers were added to at least two locations in each tank during early stages <strong>of</strong> ballast tank<br />
filling (i.e., before the tank was 25-50% full), so as to increase the opportunity for mixing<br />
throughout each tank. Rhodamie and microspheres were added directly to all tanks. In contrast,<br />
Artemia were initially cultured (i.e., the cysts were added to salt water and hatched in buckets in<br />
advance <strong>of</strong> boarding the ship), and the resulting organisms were used to inoculate ballast tanks.<br />
Sample Collection.<br />
Replicate samples were collected at 2 –3 different locations (i.e., tank access points) from<br />
each tank, for up to 5 different sampling periods (as above). Sampling procedures for the plankton<br />
community followed our established protocol for characterization <strong>of</strong> ballast water (see Chapter 3<br />
for description); Artemia abundance was measured as a component <strong>of</strong> the plankton (see below).<br />
Replicate whole water samples were collected from two depths (0m and 10m), using a Niskin<br />
bottle. Whole water samples were used to measure salinity, temperature, and concentrations <strong>of</strong> dye<br />
and microspheres.<br />
For each sampling location and period, we collected 2 replicate samples for all measures.<br />
Thus, for each tank and sampling period, we obtained at least: 4 plankton samples (2 locations x 2<br />
samples); 8 rhodamine samples (2 locations x 2 depths x 2 samples); 8 mircosphere samples (2<br />
locations x 2 depths x 2 samples). Temperature and salinity measures were made immediately<br />
upon all replication Niskin samples (at least 8 per tank and sampling period, as above).<br />
Although we followed the same general sampling protocol for all voyages (in both years),<br />
we collected additional whole water samples during the 1999 experiments to measure changes in<br />
the abundance <strong>of</strong> total bacteria and ciliate protozoans. For both measures, we collected at least 8<br />
samples per tank and sampling period (2 locations x 2 depths x 2 samples). Samples were<br />
collected from all experiments in 1999 to measure total bacteria. However, we only included<br />
samples from one vessel for the protozoans, due to the time-intensive nature <strong>of</strong> analysis for this<br />
group.<br />
Sample processing.<br />
<strong>Water</strong> temperature and salinity were measured immediately, using a hand-held<br />
thermometer and refractometer, respectively. Plankton/Artemia samples were examined aboard<br />
ship initially to assess general condition (live/dead, active, lethargic) soon after collection, using<br />
dissecting microscopes; these samples were then preserved in 5% buffered formalin. The preserved<br />
plankton samples were sorted and enumerated in the laboratory as described in Chapter 3 for Fine
Chapt 5 Ballast <strong>Water</strong> Exchange Experiments, page 5- 5<br />
Analysis. Thus, densities were estimated for each taxon, and voucher samples were sent to experts<br />
to verify the taxonomic identity.<br />
The tracers in whole water samples are being quantified in the laboratory at SERC (dye<br />
concentration with a fluorometer, micro-spheres with direct counts under a fluorescent compound<br />
microscope). Total bacteria are also being estimated by direct count with a compound microscope,<br />
using standard techniques. The protozoan have been sent to a colleague (Dr. Richard Pierce,<br />
expert in ciliate protozoa) for direct counts by taxon.<br />
5C. Results<br />
The experiments were conducted on 8 different voyages, which were divided evenly<br />
between the two years (Table 5.1). All experiments were conducted from June to mid July, to<br />
control for seasonal variation and to occur during a period <strong>of</strong> high plankton abundance (see<br />
Chapter 3). Six <strong>of</strong> the 8 ships included all three treatments: ER exchange, FT exchange, and<br />
control. These were SeaRiver ships, departing from the ports <strong>of</strong> San Francisco Bay and Puget<br />
Sound. However, the large ARCO tankers were not able to perform ER exchange; experiments<br />
aboard these remaining 2 ships included only FT exchange and control tanks.<br />
Table 5.1. Overview and status <strong>of</strong> ballast water exchange experiments conducted aboard oil tankers arriving to<br />
Port Valdez, 1998-1999. For each <strong>of</strong> 8 replicate experiments: (A) The upper table indicates the vessel, start date,<br />
source port, exchange methods, and number collected samples (physical/chemical and biological); (B) The lower table<br />
indicates the status <strong>of</strong> the respective samples. Physical/chemical tracers include salinity, rhodamine, and flourescent<br />
microspheres (shown in lower table). <strong>Biological</strong> tracers include resident zooplankton and brine shrimp (Artemia) in<br />
both years, as well as total bacteria in1999 only. . Source ports: Puget Sound, WA (PS); San Francisco Bay, CA (SF);<br />
Long Beach, CA (LB). Exchange types: Empty-Refill (ER) and Flow-Through (FT). See text for experimental<br />
design.<br />
A.<br />
# <strong>of</strong> samples # <strong>of</strong> samples<br />
Ship Date Port Source Exchange type(s) Physical tracers Biol. Tracers<br />
S/R Baytown 27-Jun-98 SF ER+FT 120 60<br />
S/R Benicia 01-Jul-98 SF ER+FT 120 60<br />
S/R Long Beach 08-Jul-98 SF ER+FT 120 60<br />
ARCO Independence 18-Jul-98 LB FT 80 40<br />
S/R Baytown 11-Jun-99 PS ER+FT 60 144<br />
ARCO Spirit 12-Jun-99 LB FT 64 164<br />
S/R Baytown 08-Jul-99 PS ER+FT 60 144<br />
S/R Long Beach 19-Jul-99 SF ER+FT 120 252<br />
B.<br />
status <strong>of</strong> proccessing (ip=in progress)<br />
Ship Date Salinity Rhodamine Microspheres Zooplankton/Artemia Bacteria<br />
S/R Baytown 27-Jun-98 done done ip done -<br />
S/R Benicia 01-Jul-98 done done done done -<br />
S/R Long Beach 08-Jul-98 done done done done -<br />
ARCO Independence 18-Jul-98 done done done done -<br />
S/R Baytown 11-Jun-99 ip done done done ip<br />
ARCO Spirit 12-Jun-99 ip done done done ip<br />
S/R Baytown 08-Jul-99 ip done done ip ip<br />
S/R Long Beach 19-Jul-99 ip done done ip ip
Chapt 5 Ballast <strong>Water</strong> Exchange Experiments, page 5- 6<br />
We distributed our experiments among the three source ports to maximize the range <strong>of</strong><br />
conditions (e.g., taxonomic groups, voyage duration, vessel types, and salinity), allowing us to test<br />
for general patterns among oil tankers. For example, diversity (and salinity) was generally greatest<br />
for ships from Long Beach and Puget Sound, and voyage duration differed among source ports<br />
(see Chapters 2 and 3). However, the traffic from each source port presented some unique<br />
constraints to the overall design:<br />
(1) Ships from Long Beach could not perform ER Exchange;<br />
(2) Ships from San Francisco contained low salinity waters, especially during the 1998 El Nino<br />
year, creating a possible physiological stress for organisms when exposed to exchange (not<br />
present at the other ports);<br />
(3) Ships from Puget Sound were not able to complete as many exchanges during the short voyage<br />
duration, limiting the total exchange volume to 100% or 200% (instead <strong>of</strong> the 300% possible<br />
for the longer voyages from other source ports).<br />
For all experiments on all 8 vessels, we have measured the effect <strong>of</strong> at least one full (100%)<br />
exchange for the exchange methods and tracers indicated in Table 5.1. In addition, for the majority<br />
<strong>of</strong> vessels we have also measured the effect <strong>of</strong> multiple exchange events.<br />
Table 5.1 also indicates the number <strong>of</strong> samples taken for each voyage and the status <strong>of</strong><br />
these samples. The analysis for physical tracers is actually twice the number shown, as the same<br />
sample is used for analysis <strong>of</strong> rhodamine and microspheres.<br />
Although the samples are now at various stages <strong>of</strong> analysis (Table 5.1), our initial<br />
analyses suggest a significant difference between ER and FT exchange in the reduction <strong>of</strong><br />
rhodamine dye. For example, Figure 5.1 shows the average change in concentration <strong>of</strong><br />
rhodamine for the respective treatments across multiple exchange events in 1998. The<br />
concentration <strong>of</strong> dye was reduced by 80 and 99% (for FT and ER exchange, respectively)<br />
compared to the initial concentrations. Interestingly, the concentration <strong>of</strong> dye in the control tank<br />
increased between the first and second measures, and this change is attributed to inadequate<br />
mixing at time T o for some ships (as evidenced by vertical stratification that was present in our<br />
raw data). Similar patterns exist for the rhodamine data collected in 1999.<br />
Despite the rhodamine results, demonstrating relatively high levels <strong>of</strong> exchange, it is<br />
premature to draw conclusions about the efficacy <strong>of</strong> exchange to remove organisms. We are<br />
now analyzing the samples to measure removal rates for both biological and physical tracers<br />
(i.e., microspheres), and we expect to complete these analyses by June 2000. At present, it is<br />
evident that some taxa declined in abundance (following exchange) to the same extent as<br />
rhodamine dye. Figure 5.2 shows a decline in the abundance <strong>of</strong> Limnoithona sp. for both ER and<br />
FT exchange on one vessel. The density actually increased in the control tank, and this was due<br />
most likely to growth <strong>of</strong> copepodite stages instead <strong>of</strong> mixing. Changes in the abundance <strong>of</strong> other<br />
taxa appear to be much less striking, although analysis <strong>of</strong> the overall pattern (including variation<br />
among taxa) must await completion <strong>of</strong> all quantitative counts and taxonomic verification.
Chapt 5 Ballast <strong>Water</strong> Exchange Experiments, page 5- 7<br />
Figure 5.1. Effect <strong>of</strong> ballast water exchange on rhodamine dye concentrations. Data are from exchange<br />
experiments conducted in the ballast tanks <strong>of</strong> oil tankers arriving to Port Valdez. Shown for 1998 voyages (n=4<br />
vessels) is the mean percent change in rhodamine dye concentration (compared to the initial time measure) at each<br />
<strong>of</strong> four successive time points for 3 different treatments: Control – ballast tanks that did not undergo exchange; ER<br />
Exchange – ballast tanks that underwent Empty-Refill Exchange; FT Exchange – ballast tanks that underwent Flow-<br />
Through Exchange. See text for experimental design.<br />
200%<br />
CONTROL<br />
ER EXCHANGE<br />
FT EXCHANGE<br />
DYE REMAINING<br />
150%<br />
100%<br />
50%<br />
0%<br />
T0 T1 T2 T3 Tf<br />
TIME POINT<br />
Figure 5.2. Effect <strong>of</strong> ballast water exchange on Limnoithona sp.density. Data are from exchange experiments<br />
conducted in the ballast tanks <strong>of</strong> an oil tanker arriving to Port Valdez. Shown for one voyage is the mean density <strong>of</strong><br />
the copepod SPECIES at each <strong>of</strong> five successive time points for 3 different treatments: Control – ballast tanks that<br />
did not undergo exchange; ER Exchange – ballast tanks that underwent Empty-Refill Exchange; FT Exchange –<br />
ballast tanks that underwent Flow-Through Exchange. See text for experimental design.<br />
DENSITY (#m³)<br />
4000<br />
3000<br />
2000<br />
1000<br />
TREATMENT<br />
CONTROL<br />
ER EXCHANGE<br />
FT EXCHANGE<br />
0<br />
T0 T1 T2 T3 Tf<br />
TIME POINT<br />
Changes in the density <strong>of</strong> entrained biota within the control tanks <strong>of</strong> each vessel will<br />
measure survivorship over time (i.e., during transit). This will allow us to test our hypothesis<br />
(above) about the effect <strong>of</strong> voyage duration on survivorship, and whether differences among port<br />
sources are due to such time-dependent survivorship.<br />
5D. Discussion<br />
This is the first study to compare the relative efficiency <strong>of</strong> exchange methods (ER and FT<br />
exchange) for any vessel type or taxon.
Chapt 5 Ballast <strong>Water</strong> Exchange Experiments, page 5- 8<br />
Quantitative and experimental analyses <strong>of</strong> ballast water exchange have been very limited to<br />
date, and these can be classified into 3 general types:<br />
1. Comparison <strong>of</strong> ballast water in ships that have or have not exchanged ballast water.<br />
These data indicate that, compared to ships that have not conducted mid-ocean ballast water<br />
exchange, ships with exchanged ballast water have reduced abundance <strong>of</strong> plankton. However,<br />
with this approach, it is not possible to (a) compare directly methods <strong>of</strong> exchange (FT vs. ER) ,<br />
(b) control for initial plankton densities or the percentage <strong>of</strong> water exchanged (as below). Thus,<br />
the data are highly variable and interpretation is limited (e.g., Smith et al. 1996).<br />
2. Comparison <strong>of</strong> ballast water in tanks <strong>of</strong> the same ship that have not exchanged ballast<br />
water, with measurements made only after exchange is complete and upon arrival to port.<br />
These data suggest a reduction <strong>of</strong> roughly 90% occurred, but interpretation is also limited with<br />
this design (Ruiz and Hines 1997; see below). Initial variation among tanks can be<br />
considerable, depending upon the timing (e.g., day vs. night) and sequence <strong>of</strong> ballasting, which<br />
creates potentially large differences among tanks independent <strong>of</strong> exchange treatment.<br />
Furthermore, it is not possible to compare efficiency between methods <strong>of</strong> exchange, or for<br />
multiples <strong>of</strong> exchange, because ships usually only perform one method and volume <strong>of</strong><br />
exchange.<br />
3. Comparison <strong>of</strong> ballast water in tanks <strong>of</strong> the same ship before and after exchange <strong>of</strong> ballast<br />
water, with measurements made on board ship at various stages <strong>of</strong> the exchange process.<br />
These data provide a clear measure <strong>of</strong> efficiency within a single tank, and we have conducted<br />
this analysis on approximately 5 military vessels and 1 commercial bulk carrier (Ruiz et al.<br />
1999, Wohnam et al. 1996). However, the sample size is small (and taxa included in ships to<br />
date are limited), and comparison between exchange methods or multiples <strong>of</strong> exchange (on the<br />
same ship) has not been included or possible to date.<br />
The 1997 Pilot Study provided initial data comparing the end result <strong>of</strong> FT Exchange (300%<br />
and 100%) on plankton abundance. These data suggested that approximately 70- 90% <strong>of</strong> coastal<br />
plankton was removed by FT exchange, compared to control tanks from the same source.<br />
Interestingly, it was not clear that an increased level <strong>of</strong> exchange (100 vs. 300 %) produced a<br />
parallel reduction in key taxonomic groups.<br />
In both the Pilot Study and the current study (Chapter 3), it was evident that abundance <strong>of</strong><br />
coastal organisms was 10-100 fold lower in tankers from foreign ports (that underwent ballast<br />
water exchange) compared to domestic arrivals (that do not undergo exchange). Although this<br />
difference may result from the exchange, it is confounded by differences in the initial<br />
concentrations (i.e., source ports) and voyage duration that can also have a strong influence.<br />
The results <strong>of</strong> this study – the most comprehensive and rigorous to date - will<br />
significantly advance our understanding <strong>of</strong> the strengths and limitations <strong>of</strong> ballast water<br />
exchange, providing multiple quantitative measures for the two exchange methods, both for oil<br />
tankers specifically but for commercial ships more generally.
Chapt 5 Ballast <strong>Water</strong> Exchange Experiments, page 5- 9<br />
Importantly, when completed, our study will also provide a set <strong>of</strong> standards for<br />
evaluating ballast water management in two ways. First, we have developed and tested a<br />
standard set <strong>of</strong> assays to measure exchange efficiency across vessel types, vessel tanks, and<br />
under various conditions. This will be useful in comparing efficiency among studies. Second,<br />
the results obtained by this and future studies will provide a benchmark against which to assess<br />
the efficacy <strong>of</strong> emerging technologies.<br />
5E. References<br />
Carlton, J.T. and J.B. Geller. 1993. Ecological roulette: The global transport <strong>of</strong> nonindigenous<br />
marine organisms. Science 261: 78-82.<br />
Dickman, M. and F. Zhang. 1999. Mid-ocean exchange <strong>of</strong> container vessel ballast water. 2:<br />
Effects <strong>of</strong> vessel type in the transport <strong>of</strong> diatoms and din<strong>of</strong>lagellates from Manzanillo, Mexico,<br />
to Hong Kong, China. Mar. Ecol. Prog. Ser. 176: 253-262.<br />
Hallegraeff, G.M. 1998. Transport <strong>of</strong> toxic din<strong>of</strong>lagellates via ships’ ballast water: bioeconomic<br />
risk assessment and efficacy <strong>of</strong> possible ballast water management strategies. Mar. Ecol. Prog.<br />
Ser. 168:297-309.<br />
National Research Council. 1996. Stemming the Tide: Controlling Introductions <strong>of</strong><br />
Nonindigenous Species by Ships’ Ballast <strong>Water</strong>. National Academy Press, Washington, D.C.<br />
Ruiz, G.M. and A.H. Hines. 1997. Patterns <strong>of</strong> nonindigenous species transfer and invasion in<br />
Prince William Sound, Alaska: Pilot Study. Report Submitted to the Prince William Sound<br />
Citizens’ Advisory Council. 80pp.<br />
Ruiz, G.M., L. S. Godwin, J. T<strong>of</strong>t, L.D. Smith, A.H. Hines and J.T. Carlton. 1999. Ballast water<br />
transfer and management by U.S. Navy vessels. Final Report to the U.S. Dept. <strong>of</strong> Defense. 24pp.<br />
Smith, L.D., M.J. Wonham, L.D. McCann, D.M. Reid, G.M. Ruiz and J.T. Carlton. 1996.<br />
<strong>Biological</strong> invasions by nonindigenous species in United States waters: Quantifying the role <strong>of</strong><br />
ballast water and sediments. Parts I and II. Final report to the U.S. Coast Guard and the U.S.<br />
Department <strong>of</strong> Transportation.<br />
Wonham, M.J., W.C. Walton, A.M. Frese and G.M. Ruiz. 1996. Transoceanic transport <strong>of</strong><br />
ballast water: <strong>Biological</strong> and physical dynamics <strong>of</strong> ballasted communities and the effectiveness<br />
<strong>of</strong> mid-ocean exchange. Final Report to the U.S. Fish & Wildlife Service and the Compton<br />
Foundation.<br />
Zhang, F. and M. Dickman. 1999. Mid-ocean exchange <strong>of</strong> container vessel ballast water. 1:<br />
Seasonal factors affecting the transport <strong>of</strong> harmful diatoms and din<strong>of</strong>lagellates. Mar. Ecol. Prog.<br />
Ser. 176: 243-251.
Chapt 6. Organisms in Sediments <strong>of</strong> Tanker Ballast Tanks, page 6- 1<br />
Chapter 6. Organisms in Sediments <strong>of</strong> Tanker Ballast Tanks<br />
Anson H. Hines, Smithsonian Environmental Research Center<br />
Gregory M. Ruiz, Smithsonian Environmental Research Center<br />
6A. Purpose<br />
At certain times and source ports, appreciable quantities <strong>of</strong> bottom sediment are taken up<br />
by tankers during ballasting. The entrained sediment potentially includes bottom dwelling<br />
organisms, which may be discharged and introduced into a receiving port (Smith et al. 1996).<br />
Few studies <strong>of</strong> such entrained sediment exist, but our samples <strong>of</strong> bulk carriers in Chesapeake Bay<br />
revealed that the bottoms <strong>of</strong> ballast tanks <strong>of</strong>ten hold a wide variety <strong>of</strong> large crabs, fish, shrimp, as<br />
well as many small organisms. To determine whether tankers arriving to Prince William Sound<br />
transported organisms associated with sediment in the bottoms <strong>of</strong> segregated ballast water tanks,<br />
we sampled a subset <strong>of</strong> ships traveling between Port Valdez and west coast ports, and between<br />
west coast ports and Asian ports.<br />
6B. Methods<br />
During 1998-99 we supplied 13 ships with “sediment sampling kits”, which we<br />
developed in cooperation with the shipping agents. The ships’ mates collected core samples and<br />
evident organisms in the sediment during routine cleaning operations, which usually occurred on<br />
voyages from Valdez to west coast ports, when ballast tanks were empty and open for<br />
maintenance, and in Asian ports, when ships were in dry-dock. The samples were preserved in<br />
10% formaldehyde sea water, labeled and returned to Valdez. Samples were then sent to the<br />
SERC lab in Maryland for processing and identification. Subsamples <strong>of</strong> whole sediment were<br />
sent to Mary McGann in USGS, Menlo Park for identification <strong>of</strong> foramenifera. Remaining<br />
sediment was washed through a 0.5 mm mesh sieve and identified under a dissecting<br />
microscope.<br />
6C. Results<br />
Sediment samples <strong>of</strong> the 13 tankers contained a diverse array <strong>of</strong> taxa, including fish,<br />
polychaete worms, mollusks, adult crabs and other crustaceans, cnidarians, and other<br />
invertebrates (Fig. 6.1,Tables 6.1, 6.2). The ships averaged 2.8 taxa per ship, ranging from 0-6<br />
taxa, with annelid worms occurring in about 90% <strong>of</strong> the ships. The number <strong>of</strong> individuals per<br />
sample varied widely from 1-147 individuals, with a mean <strong>of</strong> 47 individuals. Small crustaceans<br />
(particularly cumaceans) were the most abundant taxa, however polychaete worms were the most<br />
prevalent. The sediments also contained several species <strong>of</strong> Foraminfera, including Trochammina<br />
hadai, an NIS that has invaded many west coast ports and is very common in San Francisco Bay<br />
(McGann, pers. comm.), and which is reported from Prince William Sound in samples collected<br />
from deep sediments following the ExxonValdez oil spill. Organisms were abundant in<br />
sediments taken up in both San Francisco Bay (Benicia) and in Long Beach, where the ship<br />
intakes are near the port bottom. Ships sampled in dry dock in Asia tended to have few<br />
organisms, perhaps as a result <strong>of</strong> longer voyage time across the Pacific. However, the diversity<br />
<strong>of</strong> higher taxonomic groups present in sediments <strong>of</strong> ballast tanks did not show any obvious<br />
pattern by source port.
Chapt 6. Organisms in Sediments <strong>of</strong> Tanker Ballast Tanks, page 6- 2<br />
Table 6.1. Taxa Recovered From Ballast Tank Sediments <strong>of</strong> 12 Tankers.<br />
Foraminifera Annelida Mollusca Crustacea Chordata Other inverts.<br />
Ammonia hadai Capitellidae Mytilidae Alpheidae Engraulidae Bryozoa<br />
Bulimina sp. Nereis sp. Nudibranchia Amphipoda Sciaenidae Sipuncula<br />
Elphidium sp. Oligochaeta Balanus balanoides Turbellaria<br />
Globigerina sp. Spionidae Calanus<br />
Haglophragmoides sp. Syllidae Canuellidae<br />
Jadammina macrescens Caridea<br />
Lagena sp. Cirripedia<br />
Rosalina globularis Crangonidae<br />
Trochammina hadai Cumacea<br />
Trochammina inflata Grapsidae<br />
Trochammina pacifica Harpacticoida<br />
Hyperiidae<br />
Majidae<br />
Ostracoda<br />
Tanaidacea<br />
Table 6.2. Presence/Absence <strong>of</strong> Taxa in Ballast Tank Sediment Samples Presented by Source Region(s).<br />
Source(s) Diatomacea Foraminifera Annelida Mollusca Crustacea Chordata Other Inverts n<br />
Korea P P P A A A A 1<br />
LB & Korea P P P P P P P 2<br />
PS A A A P P A P 1<br />
PS & SF P P P P P P P 1<br />
SF A P P P P P P 6<br />
SF&China A A P A P A A 1
Chapt 6. Organisms in Sediments <strong>of</strong> Tanker Ballast Tanks, page 6- 3<br />
Figure 6.1. Prevalence (A) and numbers (B) <strong>of</strong> organisms recovered from sediments <strong>of</strong> tanker ballast tanks.<br />
Bars indicate means for 12 tankers.<br />
A<br />
Cnidaria<br />
Chordata<br />
Diatomacea<br />
Other Inverts<br />
Foraminifera<br />
Mollusca<br />
Crustacea<br />
Annelida<br />
0 20 40 60 80<br />
PERCENT SHIPS<br />
n=12<br />
B<br />
Cnidaria<br />
Chordata<br />
Diatomacea<br />
Other Inverts<br />
Foraminifera<br />
Mollusca<br />
Annelida<br />
Crustacea<br />
NA<br />
NA<br />
0 20 40 60 80<br />
# OF ORGANISMS RECOVERED/SHIP<br />
6D. Conclusions<br />
Sediment that accumulated in the bottom <strong>of</strong> ballast tanks <strong>of</strong>ten contained organisms from<br />
a diverse array <strong>of</strong> taxa. Many <strong>of</strong> these were adults in full reproductive condition. At least one<br />
NIS (the foraminiferan Trochammina hadai) found in these samples appears to be established in<br />
Prince William Sound, although the current status <strong>of</strong> this invasion is not known (McGann, pers.<br />
comm. 1999).<br />
In future work, it would be valuable to sample sediment in ballast tanks that have<br />
undergone mid-ocean exchange. It is not clear if bottom-dwelling organisms will be affected by<br />
exchange in the same ways as planktonic organisms in the water column.<br />
6E. References<br />
McGann, M. 1999. Personal communication.<br />
Smith, L.D., M.J. Wonham, L.D. McCann, D.M. Reid, G.M. Ruiz and J.T. Carlton. 1996.<br />
<strong>Biological</strong> invasions by nonindigenous species in United States waters: Quantifying the role <strong>of</strong><br />
ballast water and sediments. Parts I and II. Final report to the U.S. Coast Guard and the U.S.<br />
Department <strong>of</strong> Transportation.
Chapt 7. Organisms Fouling Hulls and Sea Chests, page 7- 1<br />
Chapter 7. Organisms Fouling Hulls and Sea Chests <strong>of</strong> Tankers<br />
Anson H. Hines, Smithsonian Environmental Research Center<br />
Gregory M. Ruiz, Smithsonian Environmental Research Center<br />
7A. Purpose<br />
Historically, fouling organisms on ships have been a major source <strong>of</strong> introduced species<br />
(Carlton 1979a, 1979b, 1987, 1989). Modern anti-fouling paints and high ship speeds greatly<br />
reduce the amount <strong>of</strong> fouling today. However, fouling is <strong>of</strong>ten common in sea chests and at<br />
certain points on the bottom. We sampled tankers during routine maintenance in dry dock,<br />
selected to estimate the potential range <strong>of</strong> fouling and diversity <strong>of</strong> fouling organisms.<br />
7B. Methods<br />
We sampled the fouling communities <strong>of</strong> two ships in dry dock: the S/R Baytown (in San<br />
Francisco Bay), which had not been cleaned in dry dock for approximately 2 years; and the S/R<br />
Benicia (in Portland), which had been cleaned in dry dock within about 6 months. The S/R<br />
Baytown had remained within San Francisco Bay for several months without making an ocean<br />
voyage prior to haul out, providing time for further accumulation <strong>of</strong> fouling organisms.<br />
Representative patches <strong>of</strong> fouling communities were scraped from the bottoms <strong>of</strong> the ships<br />
within 6 hours <strong>of</strong> haul out and before any cleaning had commenced. The sea chests and strainers<br />
<strong>of</strong> the ambient water intakes were also sampled. All samples were preserved in 10%<br />
formaldehyde and returned to the laboratory for sorting and identification using a dissecting<br />
microscope.<br />
7C. Results<br />
The two ships exhibited divergent extremes in the quantity and diversity <strong>of</strong> organisms<br />
(Table 7.1). The ship that had not been in dry dock for approximately 2 years exhibited<br />
extensive fouling communities, with abundant mussels and associated worms, crustaceans, and<br />
sediments. At least one NIS for the west coast (the mussel Musculista senhousia) was identified<br />
specifically on this ship. In contrast, the ship that had been hauled recently had a relatively<br />
sparse number <strong>of</strong> organisms, with most <strong>of</strong> the hull completely clean <strong>of</strong> fouling communities, and<br />
only organisms present in the sea chest. However, even this ship had organisms that are NIS for<br />
the west coast (e.g., the striped bass Morone saxatilis) in its water intake strainers.<br />
7D. Conclusions<br />
We hypothesize that these two vessels represent the extremes in fouling communities,<br />
corresponding to the length <strong>of</strong> time since the last entry into dry dock for bottom cleaning.<br />
However, there are two other features that may contribute to these overall patterns. First, the S/R<br />
Baytown had been resident in San Francisco Bay for over 6 months, and may have developed an<br />
unusually rich fouling community. Second, the other vessel entered relatively fresh water <strong>of</strong> the<br />
Columbia River that may have had an adverse effect on the resident community <strong>of</strong> fouling<br />
organisms. To distinguish the effect <strong>of</strong> dry dock schedule on fouling community structure (from<br />
these other confounding variables), it would be valuable to sample more ships which differ in<br />
time since last haul out, but preferably sampled at the same dry dock to control for potential<br />
effects <strong>of</strong> different salinity. Nevertheless, both ships carried NIS, which indicates that this is an<br />
active mechanism <strong>of</strong> transport and introduction.
Chapt 7. Organisms Fouling Hulls and Sea Chests, page 7- 2<br />
Table 7.1. Organisms from hulls and sea chests <strong>of</strong> two oil tankers in dry dock (*=NIS).<br />
S/R Baytown<br />
S/R Benicia<br />
Algae<br />
Cnidaria<br />
Ulva sp . Garveia franciscana *<br />
Diatomacea<br />
Mollusca, Bivalvia<br />
Protozoa Mytlilus sp.<br />
Folliculina sp.<br />
Crustacea, Cirripedia<br />
Cnidaria Balanus sp.<br />
Cordylophora caspia *<br />
Crustacea, Amphipoda<br />
Garveia franciscana * Corophium sp.<br />
Nematoda<br />
Pisces<br />
Unidentified sp. Morone saxatlis *<br />
Nemertea Sardinopsis sagax<br />
Unidentified sp.<br />
Polychaeta<br />
Neries sp .<br />
Ophelidae, unidentified sp.<br />
Polydora sp.<br />
Mollusca, Bivalvia<br />
Musculista senhousia *<br />
Mytilus sp .<br />
Crustacea/Copepoda<br />
Cyclopoida, unidentified sp.<br />
Harpacticoida, unidentified sp.<br />
Crustacea/Amphipoda<br />
Corophium sp.<br />
Gammaridae, unidentified sp.<br />
Crustacea/Isopoda<br />
Unidentified sp.<br />
Crustacea/Brachyura<br />
Unidentified sp.<br />
Bryozoa<br />
Bowerbankia<br />
Membrenipora<br />
Victorella sp.<br />
7E. References<br />
Carlton, J.T. 1979a. History, biogeography, and ecology <strong>of</strong> the introduced marine and estuarine<br />
invertebrates <strong>of</strong> the Pacific coast <strong>of</strong> North America. Ph.D. Thesis, Univ. Calif., Davis. 904 pp.<br />
---. 1979b. Introduced invertebrates <strong>of</strong> San Francisco Bay. Pp. 427-444 in Conomos, T. J. (ed.),<br />
San Francisco Bay: The Urbanized Estuary. California Academy <strong>of</strong> Sciences, San Francisco.<br />
---. 1987. Patterns <strong>of</strong> transoceanic marine biological invasions in the Pacific Ocean. Bull. Mar.<br />
Sci. 41(2): 452-465.<br />
---. 1989. Man’s role in changing the face <strong>of</strong> the ocean: biological invasions and the implications<br />
for conservation <strong>of</strong> near-shore environments. Conserv. Biol. 3(3): 265-273.
Chapt 7. Organisms Fouling Hulls and Sea Chests, page 7- 3<br />
Table 7.1. Organisms from hulls and sea chests <strong>of</strong> two oil tankers in dry dock (* = NIS).<br />
S/R Baytown<br />
S/R Benicia<br />
Diatomacea<br />
Cnidaria<br />
Protozoa Garveia franciscana *<br />
Folliculina sp.<br />
Crustacea, Cirripedia<br />
Cnidaria<br />
Balanus sp.<br />
Garveia franciscana *<br />
Crustacea, Amphipoda<br />
Cordylophora caspia *<br />
Corophium sp.<br />
Hydroid – unident sp.<br />
Mollusca, Bivalvia<br />
Bryozoa<br />
Mytilus sp.<br />
Bowerbankia sp.<br />
Pisces<br />
Canopeum sp. Morone saxatilis *<br />
Victorella sp.<br />
Sardinopsis sagax<br />
Nemertea<br />
Unidentified sp.<br />
Nematoda<br />
Unidentified spp.<br />
Polychaeta<br />
Ophellidae, unidentified sp.<br />
Polydora sp.<br />
Nereis sp<br />
Crustacea/Copepoda<br />
Harpacticoida, unidentified sp.<br />
Cyclopoida, unidentified sp.<br />
Crustacea/Amphipoda<br />
Gammaridae, unidentified sp.<br />
Corophium sp.<br />
Crustacea/Isopoda<br />
Unidentified sp.<br />
Crustacea/Brachyura<br />
Unidentified sp.<br />
Mollusca, Bivalvia<br />
Musculista senhousia *<br />
Mytilus sp.<br />
Algae<br />
Ulva sp.
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 1<br />
Chapter 8. Summary <strong>of</strong> NIS in Prince William Sound and Alaska<br />
Anson H. Hines, Smithsonian Environmental Research Center<br />
Gregory M. Ruiz, Smithsonian Environmental Research Center<br />
Paul W. F<strong>of</strong>on<strong>of</strong>f, Smithsonian Environmental Research Center<br />
8A. Purpose<br />
To summarize our knowledge <strong>of</strong> marine NIS in Prince William Sound specifically and<br />
Alaska generally, we extracted information obtained from the literature, our field surveys, focal<br />
taxonomic research by systematists, and analysis <strong>of</strong> existing specimens in museum and reference<br />
collections (see detailed reports in Chapt 9). Because prior ecological and systematic work in<br />
Alaska has not focused on NIS, we also wish to establish a baseline for the status <strong>of</strong> NIS in<br />
Alaskan waters, against which future introductions may be measured. We partitioned the species<br />
records into 5 categories:<br />
(1) Definite & probable NIS, along with particularly suspicious cryptogenic species;<br />
(2) Cryptogenic species;<br />
(3) New, undescribed species discovered by this study.<br />
(4) Species with range extensions into south central Alaska discovered by this study; and<br />
(5) Species that were reported/suspected as NIS, but which we dismissed upon further analysis.<br />
The sudden appearance <strong>of</strong> apparently new or undescribed species in an ecosystem is<br />
<strong>of</strong>ten a good indicator <strong>of</strong> a biological invasion. Similarly, analysis <strong>of</strong> species’ range extensions<br />
is an important tool in detecting NIS, which may be introduced from distant biogeographic<br />
provinces or from adjoining provinces. However, where the native biota is as poorly studied as<br />
in Alaska, it may be difficult to distinguish NIS from native species that are new to science, or<br />
from new records <strong>of</strong> species within their normal range. Discovery <strong>of</strong> undescribed species and<br />
range extensions needs to be evaluated in the context <strong>of</strong> other indicators <strong>of</strong> biological invasions,<br />
such as association with sites <strong>of</strong> human activities and particular transport mechanisms. Thus,<br />
designating species as native or NIS requires a series <strong>of</strong> graded criteria (see Methods below), but<br />
the origin <strong>of</strong> many species may remain unknown, i.e., “cryptogenic”. These cryptogenic species<br />
may be further categorized into species that have particular, suspicious attributes in some<br />
criteria, or species that have not received adequate research to evaluate their origin. In other<br />
cases, species initially may be designated or suspected as NIS, but further consideration by<br />
experts may refute the initial concern.<br />
8B. Methods<br />
The graded criteria (derived from J.T. Carlton, e.g., Carlton 1979a, Chapman & Carlton<br />
1991) used to determine whether each species in our database is introduced, native, or<br />
cryptogenic are described below. "Cryptogenic species" cannot be identified clearly as native or<br />
introduced, and thus have unknown origin (Carlton 1996). In Alaska, the marine biota in many<br />
groups have received little systematic and biogeographic analysis, and a large portion <strong>of</strong> species<br />
in these groups may be cryptogenic in origin due to lack <strong>of</strong> study without particular suspicions <strong>of</strong><br />
invasive characteristics. Further discussion <strong>of</strong> criteria for identifying species as introductions are<br />
given in Chapman (1988), Chapman and Carlton (1991) and Eno (1996). Often a single criterion<br />
is not sufficient to designate a species as being introduced, but combinations <strong>of</strong> several factors
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 2<br />
increase the probability <strong>of</strong> an accurate reconstruction <strong>of</strong> introductions and invasions. In several<br />
cases, we have indicated cryptogenic species that have some suspicious characteristics <strong>of</strong> NIS.<br />
• Paleontological - NIS are absent from fossil record even though they are present in other<br />
locations; native species are found locally as recent fossils; cryptogentic species are not in the<br />
local fossil record, but they are not reliably fossilized generally.<br />
• Archeological - NIS are absent from shell middens and other archeological deposits; native<br />
species are in local deposits; cryptogenic species would not be expected to be found in<br />
archeological deposits.<br />
• Historical - NIS are not recorded by direct observation at early periods, especially by trained<br />
naturalists, but suddenly appear where trained observers did not find them previously; native<br />
species are recorded in the earliest observations <strong>of</strong> trained observers; cryptogenic species are<br />
species that were not studied by early trained observers.<br />
• Biogeographic - NIS exhibit grossly disjunct patterns <strong>of</strong> distribution (we took care to<br />
evaluate artifacts <strong>of</strong> the distribution <strong>of</strong> biologists/taxonomists); native species have<br />
continuous geographic ranges which include Alaska/Prince William Sound or other high<br />
latitudes; cryptogenic species have poorly known distributions or "cosmopolitan"<br />
distributions.<br />
• Ecological - NIS have habitats in close association with other NIS (co-evolved species;<br />
specialized predator-prey, commensal or host-parasite relations); native species are closely<br />
associated with other native species; cryptogenic species are more generalized, lacking close,<br />
specialized association with other species.<br />
• Dispersal Mechanisms - NIS presence cannot be plausibly explained by natural dispersal<br />
mechanisms and have documented human-mediated mechanisms which could effect their<br />
distributions; native species have natural dispersal mechanisms and lack known<br />
human-mediated mechanisms <strong>of</strong> introduction; cryptogenic species have both natural and<br />
human-mediated mechanisms <strong>of</strong> dispersal that could account for their distribution.<br />
• Evolutionary/Genetic - NIS have isozyme or DNA frequencies which match distant proposed<br />
source populations and are significantly different from adjacent natural populations; native<br />
species have population genetics which blend with adjacent natural populations; cryptogenic<br />
species have not been studied with molecular techniques.<br />
We also researched all published and anecdotal reports <strong>of</strong> NIS or range extensions <strong>of</strong> species that<br />
we were able to find in the scientific and informed popular literature for the region. We use<br />
these reports interactively with our field and museum work, both to direct our field surveys and<br />
re-examination <strong>of</strong> existing collections, and to determine the history <strong>of</strong> suspicious species that we<br />
collected in the field.<br />
8C. Results<br />
A diverse array <strong>of</strong> 24 species <strong>of</strong> plants and animals has been introduced into Alaskan<br />
waters, with 15 <strong>of</strong> these species being recorded in Prince William Sound (Table 8.1; see also<br />
Species Notes below). Of these definite/probableNIS, we collected 12 species in our Focal<br />
Taxonomic Analyses (Chapt 9), including 5 species <strong>of</strong> algae (Ceramium sinicola, Croodactylon<br />
ramosum, Fucus cottoni, Macrocystis integrifolia, Codium fragile tomentosoides), 1 species <strong>of</strong><br />
sponge (Cliona thosina), 1 hydroid at Homer (Garveia franciscana),1 polychaete worm<br />
(Heteromastus filiformis), 2 molluscs (Mya arenaria, Crassostrea gigas), 1 bryozoan<br />
(Schizoporella unicornis), and 1 tunicate (Botrylloides violaceus). Our findings include 7 “first
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 3<br />
TABLE 8.1 Definite/Probable NIS for Alaska<br />
* Found by this project<br />
Species Common Name Probable AK Regions Date 1st Invasion Population Ecological References<br />
Region <strong>of</strong> Origin Record Status Status Impacts<br />
Rhodophyta<br />
* Ceramium sinicola a red alga NE Pacific (CA) Prince William Sound 1998 Probable Established Fouling Hansen 1998<br />
* Chroodactylon ramosum a red alga NW Pacific Prince William Sound 1998 Probable Established Fouling Hansen 1998<br />
Phaeophyta<br />
* Fucus cottoni (=muscoides)<br />
Probable Established Unknown Hansen 1998; South and<br />
a rockweed NE Atlantic Prince William Sound;<br />
Kenai Peninsula<br />
Tittley 1986<br />
* Macrocystis integrifolia a kelp NE Pacific (SE AK) Prince William Sound 1979 Definite<br />
Not<br />
reproducing NIS vector Hansen 1998<br />
* Microspongium globosum a brown alga NW Pacific Prince William Sound 1998 Probable Established Fouling Hansen 1998<br />
Sargassum muticum<br />
Chlorophyta<br />
Japanese brown<br />
alga<br />
NW Pacific SE Alaska
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 4<br />
TABLE 8.1 continued<br />
Species Common Name Probable AK Regions Date 1st Invasion Population Ecological References<br />
Region <strong>of</strong> Origin Record Status Status Impacts<br />
Chordata- Ascidiacea<br />
* Botrylloides violaceus (=Botrylus<br />
aurantius )<br />
a tunicate NW Pacific Prince William Sound 1999 Definite Established Fouling G.Lambert 1999pers.comm.<br />
Chordata-Osteichthyes<br />
Alosa sapidissima American Shad NW Atlantic N to Cook Inlet; Kodiak I. 1896 Definite Migrant Predator on Chapman 1942; McPhail<br />
salmonid<br />
fry<br />
and Lindsey 1986; USGS<br />
1999<br />
Dallia pectoralis (FW) Alaska Blackfish Arctic Slope (FW) Anchorage area (FW) 1950s Definite Established Predator on Morrow 1980; USGS 1999<br />
salmonid<br />
fry<br />
Esox lucius (FW) Northern Pike Northern N.<br />
America (FW)<br />
Salmo salar Atlantic Salmon N Atlantic<br />
(Anadromous)<br />
Salvelinus fontinalis (FW) Brook Trout Eastern N. America<br />
(FW)<br />
Anchorage area (FW) 1970s Definite Established Predator on<br />
salmonid<br />
fry<br />
SE Alaska-Prince William<br />
Sound<br />
1990 Definite Unknown Predator/<br />
competitor<br />
<strong>of</strong> salmonids<br />
SE Alaska 1920 Definite Established Predator on<br />
salmonid<br />
fry<br />
Morrow 1980; USGS 1999<br />
Wing et al. 1992; Freeman<br />
1998 pers. comm.; USGS<br />
1999<br />
Morrow 1980; Alaska<br />
Department <strong>of</strong> Fish and<br />
Game 1994; USGS 1999
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 5<br />
records” for NIS in the region. Our Rapid Community Assessment (Chapt 9) found 1 NIS<br />
species (the s<strong>of</strong>t-shelled clam Mya arenaria ) to be widely distributed in intertidal sediments<br />
throughout Prince William Sound and the Kenai Peninsula. Two species (the oyster Crassostrea<br />
gigas, and the kelp Macrocystis integrifolia) are not established as self-sustaining, reproducing<br />
populations within the Sound; but these aquaculture introductions are being sustained by ongoing<br />
inputs that serve as a potentially important mechanism <strong>of</strong> transport for many other<br />
associated species. Further notes on each <strong>of</strong> these NIS are provided below.<br />
The literature reports 11 other NIS species, including 1 algal species (Sargassum<br />
muticum), 1 marsh plant (Cotula coronopifolia), 1 foraminiferan (Trochammina hadai), an<br />
amphipod crustacean (Jassa marmorata), 1 bryozoan (Cryptosula pallasiana), 1 brittle star<br />
(Ophiothrix koreana), and 5 species <strong>of</strong> fish (Alosa sapidissima, Dallia pectoralis, Esox lucius,<br />
Salmo salar, Salvelinus fontinalis). Several <strong>of</strong> these fish species were intentionally introduced in<br />
fresh water to augment fisheries, and we have included them here because they potentially have<br />
important impacts on native salmonid species in the region. Further notes on each <strong>of</strong> these NIS<br />
are provided below.<br />
In addition, we consider two cryptogenic species to be particularly suspicious as NIS,<br />
because <strong>of</strong> their new appearance at harbor areas (i.e., Homer, Cordova)(Table 8.2). These<br />
species include a sea star (Asterias amurensis) that is native to Alaska in the Bering Sea, but<br />
which has a history <strong>of</strong> invading other regions (probably via ballast water transport), and which<br />
appears to have suddenly extended its range to Homer in south central Alaska. Despite surveys<br />
<strong>of</strong> the area by good naturalists, this large animal has not been recorded at Homer/Katchemak Bay<br />
until now. We also discovered a new, undescribed species <strong>of</strong> ascidian (Dispalia sp. nov.) in the<br />
fouling communities <strong>of</strong> Homer and Cordova, but it was not present at other locations with rich<br />
fouling communities but lacking intense boat/ship traffic. Further notes on each <strong>of</strong> these<br />
suspicious species are provided below.<br />
A large portion <strong>of</strong> Alaskan marine species is cryptogenic in origin due to inadequate<br />
biogeographic and taxonomic study. However, many cryptogenic species also either exhibit wide<br />
distributions that may reflect global spread by early shipping traffic (Carlton 1996) or have other<br />
suspcicious traits <strong>of</strong> NIS (Table 8.3). During this project we collected at least 24 such species in<br />
Prince William Sound, and identified at least 5 others found elsewhere in Alaska.<br />
During our study we discovered several apparently new/undescribed species (Table 8.4)<br />
and documented range extensions for many other species. (Table 8.5), which also highlights the<br />
need for more analysis <strong>of</strong> Alaskan marine biodiversity. We found specimens <strong>of</strong> 10 apparently<br />
new/undescribed species in Prince William Sound, including 1 brown alga, 6 polychaete worms,<br />
2 molluscs, and 1 tunicate (Table 8.4). Formal species description <strong>of</strong> the tunicate species<br />
(Distaplia sp. nov.) is proceeding. We documented range extensions or first records for Prince<br />
William Sound or Cook Inlet (although some are known in the Bering Sea and further north) for<br />
74 species (4 algae, 11 hydrozoan cnidarians, 2 ctenophores, 24 polychaete worms, 20 molluscs,<br />
7 crustaceans, 2 bryozoans, 1 echinoderm, 2 tunicates, and 1 fish) (Table 8.5).
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 6<br />
TABLE 8.2 Highly Suspicious Cryptogenic Species<br />
* Found by this project<br />
Species Common Name Probable AK Regions Date 1st Invasion Population Ecological References<br />
Region <strong>of</strong> Origin Record Status Status Impacts<br />
Echinodermata - Asteroidea<br />
* Asterias amurensis<br />
Asian Sea Star<br />
NW Pacific; Bering<br />
Sea<br />
Homer Spit 1999 Suspicious<br />
Range<br />
extension<br />
Established<br />
Predator on Baranova 1976; Ward and<br />
molluscs & Andrew 1995; Foster et al.<br />
other inverts 1999 (Chapt 9, this report)<br />
Chordata - Ascidiacea<br />
* Distaplia sp. nov.<br />
a tunicate unknown Homer, Prince William<br />
Sound (Cordova)<br />
1998 Suspicious<br />
New<br />
Established Fouling G.Lambert 1999 pers. comm.
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 7<br />
TABLE 8.3 Examples <strong>of</strong> Cryptogenic Species<br />
Species Common Name Probable AK Regions Date 1st Invasion Population Ecological References<br />
Region <strong>of</strong> Origin Record Status Status Impacts<br />
Rhodophyta<br />
Porphyra miniata a red alga Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998<br />
Phaeophyta<br />
Demaraeaea attenuata a brown alga NW Pacific Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998<br />
Punctaria latifolia NE Pacific Prince William Sound 1998 Range<br />
Hansen 1998<br />
extension<br />
Punctaria plantaginea a brown alga Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998<br />
Heterokontophyta-Xanthophycaeae<br />
Vaucheria longicaulis a golden-brown alga NE Pacific Prince William Sound 1998 Range<br />
Hansen 1998<br />
extension,<br />
overlooked<br />
Chlorophyta<br />
Blidingia marginata a green alga NE Pacific Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998<br />
Caposiphon fulvescens a green alga NE Pacific Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998<br />
Halochlorococcum moorei a green alga NE Pacific Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998<br />
Kornmannia leptoderma non zostericola a green alga NE Pacific Prince William Sound 1998 Cryptogenic Established Unknown Hansen 1998<br />
Angiospermophyta<br />
Atriplex patula (=A. p. var. littoralis) Orach; Spearscale Eurasia SE Alaska 1883 Cryptogenic Established Unknown Meehan 1884; Hulten 1968<br />
Atriplex prostrata (=A. patula var. Halberd-Leaved Orach Eurasia SE Alaska Cryptogenic Established Unknown Hulten 1968<br />
hastata)<br />
Cnidaria<br />
Protohydra sp. a worm-like hydroid Cosmopolitan<br />
(CA-BC )<br />
Prince William Sound Cryptogenic Established Unknown Kozl<strong>of</strong>f 1987; Foster 1999 pers.<br />
comm.; UAF collections<br />
Annelida- Polychaeta<br />
Barantolla (americana species complex) a capitellid polychaete Circumboreal Prince William Sound 1988 Cryptogenic Established Unknown Kozl<strong>of</strong>f 1987; Kudenov 1998,<br />
pers. comm.<br />
Amphitrite (cirrata species complex) a terebellid polychaete Circumboreal Prince William Sound 1998 Cryptogenic Established Unknown Kozl<strong>of</strong>f 1987; Kudenov 1998,<br />
pers. comm.<br />
Capitella (capitata species complex) a capitellid polychaete Cosmopolitan Prince William Sound 1980 Cryptogenic Established Unknown Jewett 1998; Cohen and Carlton<br />
1995; Kudenov 1998, pers.<br />
comm.<br />
Decamastus sp. a capitellid polychaete Cosmopolitan<br />
(WA-BC)<br />
Prince William Sound 1998 Cryptogenic Established Unknown Jewett 1998; Kozl<strong>of</strong>f 1987; Cohen<br />
and Carlton 1995<br />
Eteone (longa species complex) a phyllodocid polychaete Circumboreal Prince William Sound 1980 Cryptogenic Established Unknown Pettibone 1963; Kozl<strong>of</strong>f 1987;<br />
Kudenov 1998, pers. comm.<br />
Eumida (sanguinea species complex) a phyllodocid polychaete Circumboreal Prince William Sound 1998 Cryptogenic Established Unknown Kozl<strong>of</strong>f 1987; Kudenov 1998,<br />
pers. comm.<br />
Harmathoe (imbricata species complex) a polynoid polychaete Circumboreal Prince William Sound 1980 Cryptogenic Established Unknown Kozl<strong>of</strong>f 1987; Cohen and Carlton<br />
1995; Kudenov 1998, pers.<br />
comm.<br />
Mediomastus sp. a capitellid polychaete Cosmopolitan Prince William Sound 1988 Cryptogenic Established Unknown [Jewett 1998]; Cohen and Carlton<br />
1995<br />
Pholoe (minuta species complex) a sigalionid polychaete Circumboreal Prince William Sound 1979 Cryptogenic Established Unknown Jewett 1998; Cohen and Carlton<br />
1995; Kudenov 1998, pers.<br />
comm.<br />
Polydora quadrilobata a spionid polychaete NE Pacific<br />
(British<br />
Columbia)<br />
Prince William Sound Cryptogenic Established Unknown Kozl<strong>of</strong>f 1987; Foster 1999 pers.<br />
comm.; UAF collections
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 8<br />
TABLE 8.3 continued<br />
Species Common Name Probable AK Regions Date 1st Invasion Population Ecological References<br />
Region <strong>of</strong> Origin Record Status Status Impacts<br />
Crustacea- Copepoda<br />
Leimia vaga a harpactacoid copepod NW Atlantic Prince William Sound 1999 Cryptogenic Established Unknown Cordell 1999 pers. comm.<br />
Mollusca- Bivalvia<br />
Macoma balthica Baltic Clam Northern<br />
oceans, NW<br />
Atlantic cryptic<br />
Bryozoa<br />
Alcyonidium "polynoum" or "mytili" a bryozoan Unknown<br />
(Pacific, NW<br />
Atlantic)<br />
Alaska Pacific coast<br />
before<br />
1924<br />
Cryptogenic Established Likely Carlton 1979; Meehan et al. 1989;<br />
Cohen & Carlton 1995<br />
Kachemak Bay Cryptogenic Established Unknown Carlton 1979; Cohen & Carlton<br />
1995; Winston 1999 pers. comm.<br />
Callopora lineata a bryozoan Unknown Prince William Sound Cryptogenic Established Unknown Foster 1999 pers. comm.;<br />
Winston 1999 pers. comm.<br />
Celleporella hyalina a bryozoan Unknown Resurrection Bay Cryptogenic Established Unknown Foster 1999 pers. comm.;<br />
Winston 1999 pers. comm.<br />
Cellepora craticula a bryozoan Unknown Prince William Sound Cryptogenic Established Unknown Foster 1999 pers. comm.;<br />
Winston 1999 pers. comm.<br />
Cribilina corbicula a bryozoan Unknown Prince William Sound Cryptogenic Established Unknown Foster 1999 pers. comm.;<br />
Winston 1999 pers. comm.<br />
Parasmittina trispinosa a bryozoan Unknown Prince William Sound Cryptogenic Established Unknown Foster 1999 pers. comm.;<br />
Winston 1999 pers. comm.<br />
TABLE 8.4 New or Undescribed Species<br />
Species Common Name Probable AK Regions Date 1st Invasion References<br />
Region <strong>of</strong> Origin Record Status<br />
Phaeophyta<br />
Coilodesme n. sp. a brown alga NE Pacific Prince William Sound 1998 New species Hansen 1998<br />
Annelida- Polychaeta<br />
Eumida sp. a phyllodocid polychaete Unknown Prince William Sound 1998 undescribed species Kuden<strong>of</strong>f 1998 pers. comm.<br />
Exogone sp. a syllid polychaete Unknown Prince William Sound 1998 undescribed species Kuden<strong>of</strong>f 1998 pers. comm.<br />
Glycera sp. a glycerid polychaete Unknown Prince William Sound 1998 undescribed species Kuden<strong>of</strong>f 1998 pers. comm.<br />
Nephtys sp. a nephtyid polychaete Unknown Prince William Sound 1998 undescribed species Kuden<strong>of</strong>f 1998 pers. comm.<br />
Polygordius sp. an archiannelid polychaete Unknown Prince William Sound 1998 undescribed species Foster 1999 pers. comm.;<br />
UAF collections<br />
Scolopos sp. an orbiniid polychaete Unknown Prince William Sound 1998 undescribed species Kuden<strong>of</strong>f 1998 pers. comm.<br />
Mollusca -<br />
Gastropoda<br />
*Adalaria sp 1. a nudibranch, Adalaria sp. 1 Unknown Prince William Sound 1999 undescribed species Goddard 1999 pers. comm.<br />
<strong>of</strong> Behrens (1991)<br />
Adalaria sp. 2 a nudibranch Unknown Prince William Sound 1999 Unidentified species Goddard 1999 pers. comm.<br />
Chordata - Asciiacea<br />
*Diastaplia n. sp. a tunicate Unknown Homer, Prince William<br />
Sound (Cordova)<br />
1998 New species G. Lambert 1999 pers. comm.<br />
* This species has been known from West coast <strong>of</strong> US for several years, but is not yet described (Goddard, 1999 pers. comm.)
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 9<br />
TABLE 8.5 Species Range Extensions or First Records for Cook Inlet / Prince Wiliam Sound (sc AK)<br />
Species Common Name Probable AK Regions Date 1st Invasion References<br />
Region <strong>of</strong> Origin Record Status<br />
Rhodophyta<br />
Polysiphonia senticulosa a red alga NE Pacific Prince William Sound 1998 Range extension Hansen 1998<br />
Phaeophyta<br />
Ectocarpus acutus a brown alga NE Pacific Prince William Sound 1998 Range extension Hansen 1998<br />
Ectocarpus dimorphus a brown alga NE Pacific Prince William Sound 1998 Range extension Hansen 1998<br />
Chlorophyta<br />
Codium fragile spp. fragile a green alga NE Pacific Prince William Sound 1998 Range extension Hansen 1998<br />
Cnidaria-Hydrozoa<br />
Aequorea aequorea a hydromedusa NE Pacific (SE AK) Prince William Sound,<br />
Bering Sea north<br />
1999 First Record sc AK Mills, Chapt 9C2, this report<br />
Aequorea victoria a hydromedusa NE Pacific (SE AK) Prince William Sound 1999 First Record sc AK Mills, Chapt 9C2, this report<br />
Clytia gregaria (=Phialidium a hydromedusa NE Pacific (SE AK) Prince William Sound, 1999 First Record sc AK Mills, Chapt 9C2, this report<br />
gregarium)<br />
Bering Sea north<br />
Eperetmus typus a hydromedusa NE Pacific (SE AK) Prince William Sound,<br />
Bering Sea north<br />
1999 First Record sc AK Mills, Chapt 9C2, this report<br />
Euphysa sp. a hydromedusa NE Pacific (SE AK) Prince William Sound,<br />
Bering Sea north<br />
Gonionemus vertens a hydromedusa NE Pacific (SE AK) Prince William Sound,<br />
Bering Sea north<br />
Halitholus sp. a hydromedusa NE Pacific (SE AK) Prince William Sound,<br />
Bering Sea north<br />
Melicertum octocostatum a hydromedusa NE Pacific (SE AK) Prince William Sound,<br />
Bering Sea north<br />
Proboscidactyla flavicirrata a hydromedusa NE Pacific (SE AK) Prince William Sound,<br />
Bering Sea north<br />
Sarsia spp. a hydromedusa NE Pacific (SE AK) Prince William Sound,<br />
Bering Sea north<br />
Tiaropsis multicirrata a hydromedusa NE Pacific (SE AK) Prince William Sound,<br />
Bering Sea north<br />
Ctenophora:<br />
Bolinopsis infundibulum a ctenophore NE Pacific (SE AK) Prince William Sound,<br />
Bering Sea north<br />
1999 First Record sc AK Mills, Chapt 9C2, this report<br />
1999 First Record sc AK Mills, Chapt 9C2, this report<br />
1999 First Record sc AK Mills, Chapt 9C2, this report<br />
1999 First Record sc AK Mills, Chapt 9C2, this report<br />
1999 First Record sc AK Mills, Chapt 9C2, this report<br />
1999 First Record sc AK Mills, Chapt 9C2, this report<br />
1998 First Record sc AK Mills, Chapt 9C2, this report<br />
1999 First Record sc AK Mills, Chapt 9C2, this report<br />
Pleurobrachia bachei () a ctenophore NE Pacific (SE AK) Prince William Sound, 1999 Range extension N Mills, Chapt 9C2, this report<br />
Dutch Harbor<br />
Annelida- Polychaeta<br />
Chaetozone senticosa a cirratulid polychaete NE Pacifc Prince William Sound 1980 Range extension N Kudenov 1998, pers. comm.<br />
Cirratulus cirratulus a cirratulid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Dodecaria sp. a spionid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Drilonereis falcata minor a lumbrinereid polychaete NE Pacific (BC Canada) Prince William Sound 1980 Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Drilonereis minor () a lumbrinereid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Flabelligera mastigophora a lumbrinereid polychaete NW Pacific (Chukchi Sea) Prince William Sound 1980 Range extension S Foster 1999 pers. comm.; UAF<br />
Hesperonoe complanata a polynoid polychaete NE Pacific Prince William Sound 1980 Range extension S Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Lumbrineris limicola a lumbrinereid polychaete NE Pacific Prince William Sound Range extension Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Lumbrineris luti a lumbrinereid polychaete NE Pacific (BC Canada) Prince William Sound 1988 Range extension N Kozl<strong>of</strong>f 1987; Kudenov 1998 pers. comm.<br />
Magelona berkleyi a magelonid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Magelona hobsoni a magelonid polychaete NE Pacific (BC Canada) Prince William Sound 1988 Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Magelona sacculata a magelonid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Mesochaetopterus taylori a chaetopterid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Microphthalmus sczelkowi a hesionid polychaete NE Pacific (CA) Prince William Sound 1980 Range extension N Foster 1999 pers. comm.; UAF<br />
Mysta barbata a hesionid polychaete NW Pacific (Chukchi Sea) Prince William Sound Range extension S Foster 1999 pers. comm.; UAF<br />
Onuphis (=Nothria, an onuphid polychaete NE Pacific Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Oriopsis sp. a sabellid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 10<br />
TABLE 8.5 continued<br />
Species Common Name Probable AK Regions Date 1st Invasion References<br />
Region <strong>of</strong> Origin Record Status<br />
Nemidia sp. a polynoid polychaete Bering Sea Prince William Sound Range extension S Foster 1999 pers. comm.; UAF<br />
Nemidia tamarae a polynoid polychaete Bering Sea Prince William Sound Range extension S Foster 1999 pers. comm.; UAF<br />
Phyllodoce medipalpa a phyllodocid polychaete NE Pacific Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Kuden<strong>of</strong>f 1998 pers. comm.<br />
Rhynchospio gluteae a spionid polychaete Unknown Prince William Sound Range extension Kudenov 1998 pers. comm.<br />
Syllis (Typosyllis) harti a syllid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Foster 1999 pers. comm.; UAF<br />
Syllis (Typosyllis) harti a syllid polychaete NE Pacific (BC Canada) Prince William Sound Range extension N Foster 1999 pers. comm.; UAF<br />
Tharyx secundus a cirratulid polychaete NE Pacific (BC Canada) Prince William Sound 1980 Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
UAF collections<br />
Mollusca- Gastropoda- Prosobranchia<br />
Barleeia acuta Acute Barleysnail NE Pacific (BC Canada) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Turgeon et al. 1988; Foster<br />
1999 pers. comm.; UAF collections<br />
Mollusca- Gastropoda- Opisthobranchia<br />
Acanthodoris nanaimoensis Wine-Plumed Spiny Doris NE Pacific (BC Canada) Prince William Sound 1999 Range extension N Kozl<strong>of</strong>f 1987; Turgeon et al. 1988; Foster<br />
1999 pers. comm.; UAF collections;<br />
Goddard 1999 pers. comm.<br />
Adalaria jannae Janna’s Adalaria NE Pacific (BC Canada) Prince William Sound 1999 Range extension N Kozl<strong>of</strong>f 1987; Goddard 1999 pers. comm<br />
Adalaria sp.1 <strong>of</strong> Behrens (1991) Armed Adalaria NE Pacific (SE AK) Prince William Sound 1999 Range extension N Goddard 1999 pers. comm<br />
Alderia modesta Modest Alderia NE Pacific (BC Canada) Prince William Sound 1999 Range extension N Kozl<strong>of</strong>f 1987; Turgeon et al. 1988;<br />
Goddard 1999 pers. comm<br />
Ancula pacifica Pacific Ancula NE Pacific (SE AK) Prince William Sound 1999 Range extension N Kozl<strong>of</strong>f 1987; Turgeon et al. 1988;<br />
Goddard 1999 pers. comm<br />
Cuthona albocrusta White-Crust Cuthona NE Pacific (BC Canada) Prince William Sound 1999 Range extension N Kozl<strong>of</strong>f 1987; Turgeon et al. 1988;<br />
Goddard 1999 pers. comm<br />
Cuthona pustulata NE Pacific (BC Canada) Homer 1999 Range extension N Kozl<strong>of</strong>f 1987; Turgeon et al. 1988;<br />
Goddard 1999 pers. comm<br />
Eubranchus olivaceus Green Balloon Aeolis NE Pacific (BC Canada) Prince William Sound<br />
and Cook Inlet<br />
Range extension N<br />
Kozl<strong>of</strong>f 1987; Turgeon et al. 1988; Foster<br />
1999 pers. comm.; UAF collections;<br />
Goddard 1999 pers. comm<br />
Geitodoris heathi Heath’s Dorid NE Pacific (SE AK) Prince William Sound 1999 Range extension N Goddard 1999 pers. comm<br />
Janolus fuscus NE Pacific (SE AK) Cook Inlet 1999 Range extension N Foster 1999 pers. comm.; Goddard 1999<br />
pers. comm<br />
Albatross Aglaja NE Pacific (SE AK) Prince William Sound 1980 Range extension N Kozl<strong>of</strong>f 1987; Turgeon et al. 1988; Foster<br />
1999 pers. comm.; UAF collections<br />
Melanochlamys diomedeum NE Pacific (SE AK) Prince William Sound 1998 Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
Melanochlamys ocelliger Arctic Odostome Bering Sea Prince William<br />
Sound,Shumagin Is.,<br />
Range extension S Berh 1894; Lee & Foster 1985; Foster<br />
1999 pers. comm.; UAF collections<br />
Kodiak Is.<br />
Odostomia arctica Hansine Seaslug NE Pacific (BC Canada) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Turgeon et al. 1988;<br />
Goddard 1999 pers. comm<br />
Olea hansineensis Banded Polycera NE Pacific (N to Hawkins<br />
Island, Prince William<br />
Sound)<br />
Prince William Sound<br />
Range extension W Foster 1999 pers. comm.; Goddard, pers.<br />
comm., 1999<br />
Palio zosterae Arctic Barrel-Bubble Bering Sea Prince William Sound Range extension S Turgeon et al. 1988; Foster 1999 pers.<br />
comm.; UAF collections<br />
Retusa obtusa<br />
Mollusca- Bivalvia Glacial Mussel Bering Sea Prince William Sound Range extension S Foster 1999 pers. comm.; UAF<br />
collections<br />
Musculus glacialis<br />
Crustacea- Copepoda Unidentified copepod NE Pacific (subtropical) Prince William Sound Range extension S Ted Cooney 1998 pers. comm.<br />
Unidentified Calanoid<br />
Crustacea- Leptostraca<br />
Nebalia sp. a nebaliacean NE Pacific (BC Canada) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
UAF collections<br />
Crustacea- Isopoda<br />
Gnathia tridens an isopod NE Pacific (CA) Prince William Sound Range extension N Foster 1999 pers. comm.; UAF<br />
collections
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 11<br />
TABLE 8.5 continued<br />
Species Common Name Probable AK Regions Date 1st Invasion References<br />
Region <strong>of</strong> Origin Record Status<br />
Munna chromocephala an isopod NE Pacific (WA) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
UAF collections<br />
Munna ubiquita an isopod NE Pacific (WA) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
UAF collections<br />
Pleurogonium sp. an isopod NE Pacific (WA) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
UAF collections<br />
Synodidotea ritteri an isopod NE Pacific (CA) Prince William Sound Range extension N Smith and Carlton 1975; Foster 1999<br />
pers. comm.; UAF collections<br />
Brachiopoda<br />
Terebratalia crossei a brachiopod NW, NE Pacific Prince William Sound Range extension N, Foster 1999 pers. comm.; UAF<br />
NE<br />
collections<br />
Bryozoa<br />
Cribilina annulata a bryozoan Bering Sea Prince William Sound Range extension S Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
UAF collections<br />
Filicrisia smithi a bryozoan Bering Sea Prince William Sound Range extension S Foster 1999 pers. comm.; UAF<br />
collections<br />
Echinodermata-Asteroida<br />
Asterias amurensis Asian Sea Star NW Pacific; Bering Sea Homer Spit 1999 Range extension REFS<br />
Chordata- Ascidiacea<br />
Chelysoma columbianum a tunicate NE Pacific (BC Canada) Prince William Sound Range extension N Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
UAF collections<br />
Halocynthia hilgendorfi igaboja a tunicate NE Pacific (BC Canada) Prince William Sound Range extension Kozl<strong>of</strong>f 1987; Foster 1999 pers. comm.;<br />
UAF collections<br />
Chordata- Osteichthyes<br />
Sphyraena argentea Pacific Barracuda NE Pacific (BC Canada) Prince William Sound 1998 Range extension Valdez Vanguard newspaper, 1998
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 12<br />
We also considered several reports and specimens that were initially considered as<br />
possible NIS, but which we reject primarily as misidentifications <strong>of</strong> similar native species (Table<br />
8.6).<br />
TABLE 8.6 Species/Specimens Misidentified as NIS<br />
Putative Species Common Name Probable AK Regions Date 1st Invasion References<br />
Region <strong>of</strong> Origin<br />
Record Status<br />
Rhodophyta<br />
Porphyra redidiva a red alga NE Pacific Prince William Sound 1998 Misidentified earlier Hansen 1998<br />
Chlorophyta<br />
Monostroma fractum a green alga NE Pacific Prince William Sound 1998 Misidentification, overlooked Hansen 1998<br />
Angiospermophyta<br />
Myriophyllum spicatumEurasian <strong>Water</strong>milfoil Eurasia SE Alaska (FW) Probable misidentification USGS 1998<br />
Cnidaria-Hydrozoa<br />
Halitholus sp. a hydromedusa NE Pacific Prince William Sound 1998 Not identifiable to species Mills, Chap9C2,this rept<br />
Leuckartiara sp. a hydromedusa NE Pacific Prince William Sound 1998 Not identifiable to species Mills, Chap9C2,this rept<br />
Annelida- Polychaeta<br />
Anaspio boreas a spionid polychaete Gulf <strong>of</strong> Alaska Prince William Sound Uncertain identification Foster 1999 pers.<br />
comm.; UAF<br />
collections<br />
Polydora cf. P.<br />
brachycephalata<br />
a spionid polychaete<br />
NE Pacific<br />
(Oregon)<br />
Prince William Sound Misidentification Kozl<strong>of</strong>f 1987; Foster<br />
1999 pers. comm.;<br />
UAF collections<br />
There are few over-arching ecological traits that characterize marine NIS in Alaska. NIS<br />
were variable in their local distributions in the region, with distributions <strong>of</strong> most species<br />
apparently limited to particular sites, but with many sites having some NIS. Although NIS were<br />
frequently associated with harbor areas and aquaculture sites, some species (e.g., Mya arenaria)<br />
occurred widely wherever the appropriate habitat was present. NIS occurred in a wide range <strong>of</strong><br />
habitats from coastal marshes (Cotula coronopifolia) and the high intertidal zone (Fucus<br />
cottonii) to deep subtidal waters (Trochammina hadai), and from variable and low salinity areas<br />
(Mya arenaria, Heteromastus filiformis) to stenohaline high salinities (Botryloides violaceus).<br />
NIS included species inhabiting hard and s<strong>of</strong>t substrates. NIS also include species from a wide<br />
range <strong>of</strong> motility, from migratory fish to sessile plants and invertebrates; and they included a full<br />
range <strong>of</strong> trophic modes from autotrophs (algae) to suspension feeders to predators. While many<br />
<strong>of</strong> the NIS have life cycles with a dispersal stage (especially echinoderms and bivalves with<br />
long-lived planktonic larvae), others had little motility (e.g., sessile tunicates with short-lived<br />
planktonic larval stages). Thus, although NIS were most common in habitats most impacted by<br />
human activities, there were few sites or habitats within the region and few ecological niches that<br />
were immune from invasion.<br />
Although oil tankers transport great quantities <strong>of</strong> abundant and diverse plankton<br />
(including known NIS) into Prince William Sound, we have not identified any established NIS<br />
that is clearly attributable to introduction via tanker ballast water. However, analysis <strong>of</strong> probable<br />
transport mechanisms for species introductions is difficult in most regions where multiple<br />
transfer agents have been active. In south central Alaska, NIS were commonly found at harbor<br />
areas (e.g., Homer), where ballast water and hull fouling associated with cargo ships and the full<br />
range <strong>of</strong> fishing and other vessels are potential vectors. Bulk carriers like wood chip and log<br />
ships arriving in ballast to Homer, as well as tankers arriving to Port Valdez, are sources <strong>of</strong> the<br />
largest volumes <strong>of</strong> ballast water (Smith et al., 1999). Fishing and recreational vessels <strong>of</strong>ten have<br />
extensive fouling communities which may be transported coastwise within the region. NIS were<br />
also found at sites <strong>of</strong> associated with aquaculture (e.g., Tatitlek) and with fishery introductions
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 13<br />
(e.g., Atlantic salmon). Oyster (Crassostrea gigas) culture imports spat from Washington and<br />
Oregon hatcheries as “clean” seed for grow-up in the field. However, associated parasitic,<br />
commensal and fouling organisms frequently could be transported unintentionally with the spat<br />
(Carlton, 1992). Similarly, transfer <strong>of</strong> kelp (Macrocystis integrifolia), however “clean” in<br />
appearance, from Oregon and Washington (in the past) and southeast Alaska (in the present)<br />
could also serve as a vector for many fouling species, epiphytes, or organisms hiding in<br />
holdfasts. Several species <strong>of</strong> fish have been introduced intentionally into Alaskan freshwaters,<br />
where they may impact salmonids at key stages <strong>of</strong> their migratory life cycle. Also, escapes <strong>of</strong><br />
Atlantic salmon from pen culture have resulted in established populations <strong>of</strong> Salmo salar in<br />
British Columbia, as well as in increasingly frequent instances <strong>of</strong> this NIS fish being caught in<br />
Prince William Sound and throughout south central to southeast Alaska.<br />
The number <strong>of</strong> marine NIS in Alaska appears to be significantly lower than other marine<br />
ecosystems along the west coast <strong>of</strong> North America, where numbers <strong>of</strong> NIS range from about 50<br />
species in Puget Sound (Cohen et al., 1998) to 250 species in San Francisco Bay (Cohen &<br />
Carlton, 1995; Carlton, pers. comm.).<br />
Species Notes:<br />
RHODOPHYTA<br />
Ceramium sinicola- This red alga was found as an epiphyte <strong>of</strong> Codium fragile (tomentosoides)<br />
near Green Island. It has not been found previously north <strong>of</strong> southern California, and is strongly<br />
suspected <strong>of</strong> being an introduction (Hansen 1998).<br />
Chroodactylon ramosum- This microscopic, primitive, red alga was found growing on oyster<br />
floats at Tatilek. This species is previously known from Japan, Australia, and southern California<br />
(and the Great Lakes, where it was introduced, Mills et al. 1993) but has not been found in the<br />
well-studied waters <strong>of</strong> British Columbia and Washington. It may have been introduced with<br />
oysters (Hansen 1998).<br />
PHAEOPHYTA<br />
Fucus cottoni (=muscoides)- This brown seaweed is known from European coasts from northern<br />
Spain to Scandinavia (South and Tittley 1986). In the northeast Pacific, it was first found by G.<br />
I. Hansen on Vancouver Island in 1981, and subsequently found to be abundant in high marsh<br />
and mudflat areas along Prince William Sound (Hansen, 1998; Chapt 9 Hansen). Its status as a<br />
separate species has been questioned by Fletcher (1987), who considers this species to be an<br />
ecotype <strong>of</strong> F. vesiculosus adapted to marsh and mudflat habitats. Specimens from Prince<br />
William Sound have been sent to Esther Serrao, Portugal, who is studying the phylogenetic<br />
relationships <strong>of</strong> Fucus using molecular techniques. The widespread distribution <strong>of</strong> this plant in<br />
British Columbia and Alaska, suggests that it is not a recent introduction (Hansen 1998; Chapt. 9<br />
Hansen).<br />
Macrocystis integrifolia- This giant kelp is found from California to southeast Alaska. Since<br />
1979, this kelp had been transported by plane from southeast Alaska to Prince William Sound to<br />
be used as substrate for the Herring-Roe-on-Kelp fishery. Blades <strong>of</strong> kelp are placed in<br />
impoundment nets with gravid herring, which deposit their eggs on the kelp. The egg-laden
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 14<br />
blades are then harvested and shipped to Japan, as a delicacy. Blades and holdfasts <strong>of</strong> kelp are<br />
commonly found in Prince William Sound, but attached plants have not been found, indicating<br />
that this kelp has not become established. While “clean” kelp blades are selected for the fishery,<br />
the practice represents a potential vector for transport <strong>of</strong> microscopic developing stages <strong>of</strong> algae<br />
and invertebrates into Prince William Sound (Jay Johnson, Alaska Fish and Game, pers. comm.<br />
to G. I. Hansen ; Hansen 1998). Our examination <strong>of</strong> several large plants including blades, stipes<br />
and holdfasts at Knight Island in the Sound during June 1998 showed that a variety <strong>of</strong><br />
gastropods, ophiuroids, amphipods and bryozoans were present. It was not clear whether these<br />
associated organisms colonized the plants or were present at the time <strong>of</strong> release into the Sound.<br />
Microspongium globosum- This tiny brown alga is known previously from the North Atlantic<br />
and Japan, but it has not been found in the waters <strong>of</strong> British Columbia and Washington. It grows<br />
epiphytically on the cryptogenic brown alga Demaraeaea attenuata, attached to oyster floats at<br />
Tatilek (Hansen 1998).<br />
Sargassum muticum- This Japanese seaweed was first observed on the U.S. west coast in 1947,<br />
in Coos Bay, Oregon. By 1986, it was well established from southern California to southeast<br />
Alaska. It was probably transported across the Pacific on the shells <strong>of</strong> Pacific Oysters from<br />
Japan, and then transported along the coast by currents, shipping, and oyster transplants (Scagel<br />
1956; Scagel et al. 1986; Cohen and Carlton 1995; US Fish & Wildlife Service, Nonindigenous<br />
<strong>Aquatic</strong> Species Database 1999).<br />
CHLOROPHYTA<br />
Codium fragile (ssp. tomentosoides), Dead Man’s Fingers- The green algal species Codium<br />
fragile occurs on the West Coast as a species complex consisting <strong>of</strong> several unnamed subspecies,<br />
presumably native (Cynthia Trowbridge, pers. comm. to G. I. Hansen), as well as the introduced<br />
C. f. tomentosoides. The latter is native to the Northwest Pacific, and now widely introduced in<br />
temperate waters (Farnham 1980; Carlton and Scanlon 1985; Trowbridge 1995). On the West<br />
Coast, this seaweed has previously been known only from San Francisco Bay, where it was first<br />
collected in 1977, and probably was introduced on ship fouling (Cohen and Carlton 1995). A<br />
form <strong>of</strong> Codium nearly identical to C. f. tomentosoides was found in 1998, at Green Island, in<br />
Prince William Sound, together with a more typically native Codium. According to experts on<br />
the genus consulted by Hansen, both forms lie within the morphological range <strong>of</strong> the native<br />
populations, but molecular studies will be needed to determine their identity and relationships.<br />
In any event, the occurrence <strong>of</strong> Codium in Prince William Sound represents a range extension<br />
from southeastern Alaskan waters, and a possible introduction.<br />
ANGIOSPERMOPHYTA<br />
Cotula coronopifolia, Brass Buttons- This attractive flowering plant <strong>of</strong> the aster family is native<br />
to South Africa. It was first reported on the Pacific Coast in 1878, along San Francisco Bay and<br />
now occurs in coastal marshes from southern California to southeast Alaska (Hultén 1968;<br />
Cohen and Carlton 1995). Brass Buttons was probably transported in the dry ballast <strong>of</strong> ships to<br />
San Francisco Bay and other Pacific ports, as well as to scattered sites on the Atlantic coast <strong>of</strong><br />
North America and Europe (Hultén 1968). Seeds <strong>of</strong> this plant (Cohen and Carlton 1995) are a<br />
favorite food <strong>of</strong> waterfowl, which may be how this species reached Alaska.
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 15<br />
Additional flowering plants, identified by Hultén (1968) as “introduced weeds” <strong>of</strong> “waste<br />
places” and roadsides, probably occur at the edges <strong>of</strong> seashores and salt-to-fresh tidal marshes on<br />
the Pacific coast <strong>of</strong> Alaska, based on their habits and distribution elsewhere in North America.<br />
The following species are likely to occur in tidal marsh and shore habitats, especially disturbed<br />
ones: Agrostis gigantea (Redtop); Polypogon monspeliensis (Beard Grass); Puccinellia distans<br />
(Alkali Grass); Rumex crispus (Curly Dock); Rumex obtusifolius (Round-Leaved Dock)); Rumex<br />
maritimus (Golden Dock); Polygonum prolificum (Prolific Knotweed); Spergularia rubra (Sand<br />
Spurrey); Plantago major (English Plantain) (e.g. Fernald 1950; Gleason and Cronquist 1991;<br />
Cohen and Carlton 1995). Polygonum prolificum is native to eastern North America; the other<br />
species are <strong>of</strong> Eurasian origin (Hultén 1968). Many <strong>of</strong> these species were present on the coast <strong>of</strong><br />
southeast Alaska by 1883 (Meehan 1884), and may have been introduced in ship’s ballast.<br />
PROTOZOA- FORAMINIFERA<br />
Trochammina hadai- This foraminiferan is native to Japan, and was first found in North America<br />
in San Francisco Bay in 1990-1993 (Cohen and Carlton 1995; McGann and Sloan 1996). It was<br />
subsequently found in many Pacific Coast estuaries, from San Diego Bay to Puget Sound (Cohen<br />
et al. 1998; McGann 1998 pers. comm.). In San Francisco Bay it forms very dense populations<br />
and it processes large amounts <strong>of</strong> carbon in the benthic communities throughout the estuary. T.<br />
hadai was also found in EVOS samples taken from deep (300 ft) water <strong>of</strong> Prince William Sound<br />
(McGann 1998 pers. comm.). This benthic protozoan inhabits the sediments (preferably muddy)<br />
<strong>of</strong> brackish-marine estuaries (Matsushita and Kitazato 1990, Kitazato and Matsuchita 1996). It<br />
probably has been introduced in ballast water, and was common in sediments in the ballast tanks<br />
<strong>of</strong> oil tankers travelling between west coast ports and Port Valdez (McGann and Sloan 1996;<br />
McGann 1998 pers. comm.). However, sediment samples collected from low intertidal to<br />
shallow subtidal zones throughout Prince William Sound during 1998-1999 did not contain T.<br />
hadai, so the extent <strong>of</strong> this population in the Sound remains unclear (Hines & McGann, pers.<br />
comm.).<br />
PORIFERA<br />
Cliona thosina- This boring sponge was originally described in 1888 using specimens on oyster<br />
shells from an unknown locality (possibly France or Mexico). C thosina was found boring in<br />
field cultured oysters (Crassostrea gigas) in Prince William Sound in 1998 (Hines, 1998;<br />
Ruetzler pers. comm. 1998). Its boring activities weaken oyster shells and can cause shell<br />
deformation, breakage and increase vulnerability to predators (such as crabs). The larval stages<br />
<strong>of</strong> C. thosina are short lived (1-2 days), limiting its ability to be transported in ballast water.<br />
Oysters cultured in the Sound arrive as “clean” spat derived from laboratory cultures in Oregon<br />
and Washington. However, oyster spat is not always as “clean” as the suppliers claim, and many<br />
associated species may be found in these types <strong>of</strong> aquaculture sources (Carlton, 1992). Cliona is<br />
common in oysters <strong>of</strong> the lower west coast, so it is possibly derived from these populations.<br />
CNIDARIA- HYDROZOA<br />
Garveia franciscana (Rope Grass Hydroid)- This hydroid has been found in many estuaries<br />
around the world, but its origin is uncertain. The Indo-Pacific and the Black--Caspian Sea basin<br />
have been suggested as possible native regions (Cohen and Carlton 1995; Calder 1997 pers.<br />
comm.) It was first described from San Francisco Bay in 1902, which was its only known<br />
location on the west coast <strong>of</strong> North America (Cohen and Carlton 1995), until we found it near
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 16<br />
Homer in 1999 (Lee-Anne Henry pers. comm. 1999; Chapt 9 Fouling Communities). In other<br />
regions <strong>of</strong> the world, this hydroid has been an economically important fouling organism,<br />
adversely affecting ships, power plants and fishing gear (Simkina 1963; Andrews 1973; McLean<br />
1972).<br />
ANNELIDA- POLYCHAETA<br />
Heteromastus (filiformis)- This sediment-dwelling, free-burrowing polychaete, <strong>of</strong> the family<br />
Capitellidae, was first described from Europe, but it is now widely distributed in coastal waters<br />
around the world. On the west coast <strong>of</strong> North America, H. filiformis was first reported in 1936,<br />
from San Francisco Bay, and subsequently has been found north to British Columbia and Prince<br />
William Sound. Its introduction to the Pacific Coast could have occurred with Atlantic or Pacific<br />
oysters, or in the ballast water <strong>of</strong> ships (Carlton 1979; Cohen and Carlton 1995). Heteromastus<br />
“filiformis”, as with some other capitellid species, may constitute a complex <strong>of</strong> several<br />
morphologically similar species (Cohen and Carlton 1995). H. filiformis was collected<br />
commonly in Port Valdez in 1971-1972, 7 years after the 1964 earthquake that disrupted the<br />
benthic system, indicating that it was established well before initiation <strong>of</strong> tanker traffic to the<br />
Port (Feder et al. 1973).<br />
Lumbrineris heteropoda- This infaunal and polychaete is known from the Sahkalin and Japan,<br />
and from two Alaskan specimens, one from Resurrection Bay, and another from Glacier Bay.<br />
The wide gap between the known range and the Alaska records is suggestive <strong>of</strong> an introduction<br />
(Nora Foster, 1999 pers. comm.).<br />
MOLLUSCA- BIVALVIA<br />
Crassostrea gigas (Pacific Oyster; Japanese Oyster)- The Pacific Oyster was first planted in<br />
North American waters in 1902, in Puget Sound. By 1939, it was cultivated in Ketchikan,<br />
Alaska, and it is now reared in Prince William Sound, Katchemak Bay and other locations.<br />
Alaskan waters are too cold for natural reproduction <strong>of</strong> C. gigas, so spat must be transferred from<br />
southern waters (Quayle 1969; Carlton 1979;R. Piorkowski 1999 pers. comm.).<br />
Mya arenaria (S<strong>of</strong>tshell Clam)- The S<strong>of</strong>tshell Clam has a complex biogeographical history. This<br />
species evolved in the Pacific, in the Miocene Period, and subsequently invaded the Atlantic, but<br />
became extinct in the Eastern Pacific (Strasser 1999). Living populations <strong>of</strong> Mya arenaria<br />
remain in the Bering Sea, but on the Eastern Pacific Coast, shells <strong>of</strong> s<strong>of</strong>tshell clams are absent<br />
from subfossil deposits and shell middens, including those recently examined for M. arenaria<br />
(Foster, Chapt 9C7, this report). Mya arenaria was re-introduced to the Pacific Coast in San<br />
Francisco Bay in 1874, probably with plantings <strong>of</strong> Eastern Oysters (Crassostrea virginica). It<br />
was soon widely transplanted along the coast, reaching Alaska by the 1960s-1970s (Carlton<br />
1979). The clam has been widely established for decades in Prince William Sound and Port<br />
Valdez (Feder et al. 1973, Feder and Paul 1973), and was heavily impacted by benthos upheaval<br />
in the 1964 earthquake (Baxter 1971).<br />
CRUSTACEA-AMPHIPODA<br />
Jassa sp.; Jassa marmorata- A tube-dwelling amphipod <strong>of</strong> the genus Jassa from Prince William<br />
Sound was found in University <strong>of</strong> Alaska collections (Nora Foster pers. comm.). Specimens are<br />
being examined by John Chapman, but are not yet identified to species. Jassa marmorata, native
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 17<br />
to the northwest Atlantic, has been widely introduced in the world’s oceans, and has been<br />
collected from Alaska waters (Point Slocum, Conlan 1989; Cohen and Carlton 1995; Chapman<br />
1998 pers. comm.). Amphipods <strong>of</strong> this genus build tubes on hard surfaces, including ship hulls,<br />
but also have been collected from ballast water (Cohen and Carlton 1995).<br />
BRYOZOA<br />
Cryptosula pallasiana- This bryozoan is apparently native to the Atlantic Ocean, but is now<br />
widely distributed in the Pacific. An early (1925) record <strong>of</strong> C. pallasiana from Homer, Alaska<br />
was a misidentified specimen <strong>of</strong> C. okadai, but in 1944-46 it was found in Sitka (U. S. Navy<br />
1951; Carlton, pers. comm.), as well as San Francisco Bay, and Newport Harbor, California. It<br />
was probably transported in ship fouling (Cohen and Carlton 1995).<br />
Schizoporella (unicornis)- This northwest Pacific bryozoan was first collected in the Eastern<br />
Pacific in 1927, in Puget Sound (Carlton 1979; Cohen and Carlton 1995). Its first Alaska<br />
collection was made between 1944 and 1949, in Kodiak (U. S. Navy 1951; Powell 1970; Dick<br />
and Ross 1988; Carlton, pers. comm.). Schizoporella unicornis may have been introduced in ship<br />
fouling or with plantings <strong>of</strong> Pacific Oysters (Cohen and Carlton 1995). In 1999, it was found in<br />
Tatitlek. (This form, while definitely introduced to the Pacific coast, may actually be a complex<br />
<strong>of</strong> several species (Winston 1999 pers. comm.).<br />
ECHINODERMATA- OPHIUROIDEA<br />
Ophiothrix koreana- A single brittlestar from Southeast Alaska (Juneau) has been tentatively<br />
identified as O. koreana (Kyte 1998, pers. comm.). If this identification is correct, this collection<br />
would be the first record <strong>of</strong> this northwest Pacific ophiuroid from the eastern Pacific. Since only<br />
a single specimen has been collected, the existence <strong>of</strong> established populations is unknown. Most<br />
brittlestars have long-lived planktonic larvae, so ballast-water transport is likeliest, but transport<br />
with oysters or ship fouling can not be ruled out.<br />
CHORDATA- ASCIDIACEA<br />
Botrylloides violaceus (=Botryllus aurantius)- This colonial tunicate is native to the northwest<br />
Pacific (Japan), and may have been first found on the West Coast in 1973, in San Francisco Bay<br />
(Cohen and Carlton 1995). It is now widespread, from southern California to British Columbia<br />
(Cohen et al. 1998; Lambert and Lambert 1998). Botrylloides violaceus was abundant on fouling<br />
plates in Prince William Sound in 1999 (G. Lambert 1999 pers. comm.).<br />
CHORDATA-OSTEICHTHYES<br />
Alosa sapidissima (American Shad)- This anadromous fish, native to the Atlantic coast <strong>of</strong> North<br />
America, was introduced in 1871, to the Sacramento River. It rapidly spread along the Pacific<br />
coast, and was first collected in Alaska in the Stikine River in 1896. Shad spawn in freshwater<br />
rivers from San Francisco Bay, north to the Columbia River, but feeding adult and juvenile fish<br />
wander as far north as Cook Inlet and the Kamchatka Peninusla (Chapman 1942; Cohen and<br />
Carlton 1995). A specimen <strong>of</strong> this species from Port Moller, Alaska Peninsula resides in the<br />
University <strong>of</strong> Alaska Museum collections (Foster 2000, pers. comm.). American Shad have been<br />
captured by seines and gill nets in Southeast Alaska during strong El Niño years, e.g., 1969 and<br />
1983 (J. Karinen, 2000 pers. comm.)
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 18<br />
Dallia pectoralis (Alaska Blackfish)- This small freshwater fish is native to the North slope and<br />
Yukon-Kuskakwim Delta <strong>of</strong> Alaska and eastern Siberia, but it was introduced in 1950 to Hood<br />
and Spenard Lakes in Anchorage, in the Susitna River drainage, and has spread to other lakes in<br />
the vicinity (Morrow 1980). We are unaware <strong>of</strong> records <strong>of</strong> this fish in brackish or tidal waters,<br />
but we are including it here because <strong>of</strong> concerns <strong>of</strong> adverse impacts on Rainbow Trout<br />
(Oncorhynchus mykiss) and other salmonid populations in the Anchorage area (Morrow 1980).<br />
Esox lucius (Northern Pike)- This large predatory, freshwater gamefish is native to most <strong>of</strong> the<br />
glaciated regions <strong>of</strong> North America and Eurasia (Scott and Crossman 1973). In Alaska, the<br />
native range includes the Bering Sea drainage, and North Slope, but not Pacific watersheds.<br />
Northern Pike were illegally introduced to the Susitna River valley in the 1970s (Morrow 1980).<br />
This species is known to enter brackish waters (Scott and Crossman 1973), though we are<br />
unaware <strong>of</strong> estuarine occurrences in Alaska. Pike have a reputation as predators <strong>of</strong> salmonids, so<br />
their introduction has long been discouraged on the West Coast (Lampman 1946; Dill and<br />
Cordone 1997).<br />
Salmo salar (Atlantic Salmon)- Atlantic Salmon are native to both sides <strong>of</strong> the North Atlantic,<br />
and spawn in rivers <strong>of</strong> Europe and eastern North America. Many unsuccessful attempts were<br />
made to stock this species on the West Coast, beginning in the Sacramento River in 1874 (Dill<br />
and Cordone 1997), but the extensive use <strong>of</strong> S. salar in net-pen aquaculture has again raised the<br />
possibility <strong>of</strong> its establishment in Pacific waters. Rearing <strong>of</strong> Atlantic Salmon is illegal in<br />
Alaskan waters, but occurs in British Columbia and Washington (USGS, Nonindigenous <strong>Aquatic</strong><br />
Species Database 1999). In Alaska, the Atlantic Salmon was first caught <strong>of</strong>f Cape Cross, in<br />
southeastern waters, in 1990 (Wing et al. 1992). Since then, many S. salar have been caught in<br />
the state’s marine waters, and in 1998, the first one was caught in Alaska freshwater (Freeman<br />
1998 pers. comm.; USGS, Nonindigenous <strong>Aquatic</strong> Species Database 1999), and some have been<br />
caught in Prince William Sound with landings reported at Port Valdez and Cordova (Benda 1997<br />
pers. comm., Freeman 1998 pers. comm.). Many escaped cultured fishes are in poor condition<br />
(USGS, Nonindigenous <strong>Aquatic</strong> Species Database 1999), but successful reproduction has been<br />
documented on Vancouver Island (Volpe 1999), and could well occur in Alaska waters.<br />
Salvelinus fontinalis (Brook Trout)- This eastern North American fish was introduced into<br />
southeast Alaska in the 1920s, and continued to be stocked into the 1950s. In its native range,<br />
the Brook Trout has anadromous populations in coastal regions, from Massachusetts to Labrador<br />
(Morrow 1980). We have not found documentation <strong>of</strong> sea-running fish in Alaskan waters, but<br />
estuarine occurrences <strong>of</strong> this trout are possible. However, few populations occur in coastal lakes,<br />
and none are known from streams or rivers (Alaska Department <strong>of</strong> Fish and Game 1994) The<br />
Brook Trout may hybridize with the native Dolly Varden, but the impact <strong>of</strong> this crossing on<br />
native populations is unknown (Morrow 1980).<br />
Suspicious Species<br />
ECHINODERMATA- ASTEROIDEA<br />
Asterias amurensis (Common Asian Sea Star)- This sea star is native to the Northwest Pacifc,<br />
including the coasts <strong>of</strong> Japan and Russia north to the Tatarskii Inlet and the southern Kuril<br />
Islands, and to the Bering Sea Coasts <strong>of</strong> Russia and Alaska (Baranova 1976; Ward and Andrew
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 19<br />
1995, Jewett and Feder 1981). Its recent appearance at Homer in Cook Inlet could represent a<br />
natural range extension. However, this species has long-lived planktonic larvae and could also<br />
be carried in ship ballast water. Asterias amurensis has successfully invaded the coast <strong>of</strong><br />
Tasmania, where it poses a threat to shellfisheries (Ward and Andrew, 1995). Shipping traffic<br />
into Homer by bulk carriers <strong>of</strong> logs and wood chips increased markedly recently as spruce trees<br />
killed by a beetle outbreak have been forested. These ships probably bring large quantities <strong>of</strong><br />
ballast water to Homer from Asian ports within the established <strong>of</strong> range A. amurensis. The<br />
recent appearance <strong>of</strong> A. amurensis in the low intertidal zone at the tip <strong>of</strong> Homer Spit was noted<br />
as a sudden arrival by experienced naturalists (C. & C. Field, 1999 pers. comm.), who have been<br />
studying the area for several years. This large conspicuous sea star would not be overlooked<br />
easily and was not recorded previously in the many benthic trawl surveys <strong>of</strong> the Gulf Alaska (N.<br />
Foster, UA Museum 1999 pers. comm.). Specimens <strong>of</strong> A. amurenis are not represented in the<br />
University <strong>of</strong> Alaska Museum’s collections for trawl surveys in either the Gulf <strong>of</strong> Alaska (Foster,<br />
1999 pers. comm.) or Cook Inlet (Feder and Paul 1981, Feder et al. 1981). The survey <strong>of</strong> Cook<br />
Inlet also included scuba surveys. It would have been very unusual for such collections to miss<br />
a large conspicuous sea star (30 cm from ray tip to ray tip). We consider this species to be<br />
cryptogenic in Cook Inlet, with characteristics that are very suspicious <strong>of</strong> an NIS. Because it is a<br />
voracious predator, it could have a major impact on benthic communities.<br />
ASCIDIACEA<br />
Distaplia n. sp.- This tunicate is a new, undescribed species (G. Lambert, 2000, Chapt 9C10, this<br />
report), which is very abundant in fouling communities on floats and man-made substrates in<br />
marinas at Homer and Cordova. It was first collected in 1998 in Homer and was found in both<br />
Homer and Cordova in 1999. It was not found at other sites within Prince William Sound where<br />
other, native species <strong>of</strong> tunicates were common in fouling communities but lack similar<br />
shipping/boating traffic (e.g., Tatitlek, Chenega, Port Chalmers). These sites were sampled by<br />
the same expert taxonomists and systematists who found the species at Homer and Cordova.<br />
Also, sites with and without the new species were sampled at the same times with an equivalent<br />
sampling effort (fouling plates, Chapt 9D, this report). Its appearance is also suspicious, because<br />
it was not identified in 1901 when tunicates were collected in the region at nearby sites (Ritter<br />
1903), but this ascidian taxonomist tended to “lump” species <strong>of</strong> Distaplia (G. Lambert, 1999<br />
pers. comm.). This tunicate could be a formerly rare native species that has taken advantage <strong>of</strong><br />
the newly created marina habitat (G. Lambert 1999 pers. comm.), or a recent introduction.<br />
CRYPTOGENIC SPECIES<br />
PHAEOPHYTA<br />
Demaraeaea attenuata- This is a possible introduction from the Northwest Pacific, but G.<br />
Hansen treated this alga as cryptogenic, although she considered its epiphyte Ceramium sinicola<br />
to be a more likely introduction (Chapt 9C1, this report).<br />
ANGIOSPERMOPHYTA<br />
Atriplex patula & A. prostrata- These flowering plants commonly occur on marshes and<br />
beaches, but Hultén (1968) refers to them (lumped as A. patula) as an “introduced weed”.<br />
Botanists are divided on their status in North America, and on the East Coast they can be traced<br />
back to the early 1700’s. Here, we designate them “cryptogenic”.
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 20<br />
ANNELIDA-POLYCHAETA<br />
Capitella capitata and other polychaete species complexes- Many cosmopolitan polychaete<br />
“species” are believed to represent groups <strong>of</strong> sibling species, with little morphological<br />
differentiation, but possibly with differing life histories and environmental adaptations. This has<br />
been shown for the pollution-tolerant worm Captitella capitata (Grassle and Grassle 1976), and<br />
is suspected for many others. Cryptic invasions by foreign sibling species could be common for<br />
species with planktonic larvae, especially in newly polluted harbors, where less-tolerant natives<br />
could be replaced by better adapted invaders. Such invasions could only be detected by genetic<br />
methods, or by very exacting morphological studies.<br />
Polydora quadrilobata- This spionid polychaete has a wide distribution, including both sides <strong>of</strong><br />
the North Atlantic, the Northwest Pacific, and the Northeast Pacific coast from California to<br />
Puget Sound (Blake 1971; Kozl<strong>of</strong>f 1987). At least 7 species <strong>of</strong> spionids have been introduced to<br />
the Northeast Pacific (Cohen and Carlton 1995; Cohen et al. 1998). Foster (N. Foster, UA<br />
Museum, 1999 pers. comm.) considers P. quadrilobata’s wide distribution to be suspicious, in<br />
view <strong>of</strong> the numerous introductions <strong>of</strong> this group: “The suspicious designation results from my<br />
perception that Spionidae do seem to make up a large proportion <strong>of</strong> the NIS listed by the Puget<br />
Sound expedition.”<br />
BRYOZOA<br />
Alcyonidium (polyoum) Native and introduced cryptic species are presumed to exist on the<br />
Pacific Coast. However, Alaska animals may be more likely to represent native forms, while<br />
San Francisco Bay bryozoans are more likely to be introduced (Cohen and Carlton 1995).<br />
MOLLUSCA- BIVALVIA<br />
Macoma balthica- Native and introduced cryptic species are presumed to exist on the Pacific<br />
Coast. However, Alaska animals may be more likely to represent the native forms (Meehan et<br />
al. 1989; Cohen and Carlton 1995).<br />
CRUSTACEA-COPEPODA<br />
Leimia vaga- This benthic harpacticoid copepod was first described from Nova Scotia, but has a<br />
limited distribution in the North Atlantic and is found in several Oregon and Washington<br />
estuaries (Chapt 9C5; Jeff Cordell 1999 pers. comm.). In 1999, our surveys found it in Prince<br />
William Sound (Chapt 9C5, this report). Its disjunct distribution is suggestive <strong>of</strong> transport<br />
between coasts, but the origin <strong>of</strong> this species is unknown.<br />
8D. Conclusions<br />
We have identified 24 species <strong>of</strong> plants and animals comprising the NIS <strong>of</strong> marine related<br />
ecosystems in Alaska, including 15 species recorded from Prince William Sound. In addition, 2<br />
cryptogenic species have highly suspicious characteristics <strong>of</strong> NIS. These species represent a<br />
diverse array <strong>of</strong> taxa that occupy a wide range <strong>of</strong> ecological niches and habitats, although there<br />
appear to be more NIS associated with boat harbors and with aquaculture activities. Several <strong>of</strong><br />
these NIS are first records for Alaska. We also recorded several new, previously undescribed<br />
species as well as numerous range extensions for species, which probably reflect the poor level<br />
<strong>of</strong> study and understanding <strong>of</strong> taxonomy and biogeography in Alaskan marine ecosystems. Many<br />
<strong>of</strong> the Alaskan NIS have larval stages which could be transported in ballast water; however,
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 21<br />
other vectors (including intentional and incidental release for fisheries and aquaculture) are<br />
obvious possibilities. None are clearly associated with ballast water <strong>of</strong> oil tankers as a primary<br />
mechanism, even though many NIS are frequently found in ballast water arriving to Port Valdez<br />
(see Chapt 3, this report). The number <strong>of</strong> marine NIS in Alaska appears to be significantly lower<br />
than other marine ecosystems along the west coast <strong>of</strong> North America, where numbers <strong>of</strong> known<br />
NIS range from about 50 to 250 species. The complexities and uncertainties <strong>of</strong> the native biota<br />
and the history <strong>of</strong> vectors in the south-central Alaskan region will inevitably result in an evolving<br />
analysis, typically revealing previously hidden importance and impacts <strong>of</strong> NIS.<br />
8E. References<br />
Alaska Department <strong>of</strong> Fish and Game. 1994. Wildlife Notebook Series. Web Address:<br />
http://www.state.ak.us/local/akpages/FISH.GAME/notebook/fish/b^trout.htm<br />
Andrews, J. D. 1973. Effect <strong>of</strong> tropical storm Agnes on epifaunal invertebrates in Virginia<br />
estuaries. Chesapeake Sci. 14(4): 223-234.<br />
Baranova, Z. I. 1976. Phylum Echinodermata. Pp. 114-120 in Golikov, A. N., A.V. Zhimunski,<br />
E.V. Krasnol, O.G. Kusakin, A.F. Makienko, E.V. Markivskaya, O.A. Skarlato and A.A. Strekov<br />
(ed.), Animals and Plants <strong>of</strong> Peter the Great Bay. Nauka, Leningrad. (In Russian).<br />
Baxter, R. 1971. Earthquake effects on clams <strong>of</strong> Prince William Sound. In The Great Alaska<br />
Earth Quake <strong>of</strong> 1964. Biology. National Academy <strong>of</strong> Sciences, Washington, DC. 287 p.<br />
Benda, R. 1997. Biology Dept, Prince William Sound Community College, Valdez, AK.<br />
Personal communication.<br />
Blake, J. A. 1971. Revision <strong>of</strong> the genus Polydora from the East Coast <strong>of</strong> North America<br />
Polychaeta: Spionidae). Smithson. Contrib. Zool. 75: 1-32.<br />
Calder, D. 1997. Personal communication.<br />
Carlton, J.T. 1979. History, biogeography, and ecology <strong>of</strong> the introduced marine and estuarine<br />
invertebrates <strong>of</strong> the Pacific Coast <strong>of</strong> North America. PhD. Thesis, Univ. Calif., Davis. 904 pp.<br />
Carlton, J.T.. 1992. The dispersal <strong>of</strong> living organisms into aquatic ecosystems as mediated by<br />
aquaculture and fisheries activities. Pp. 13-45 in Rosenfield, A. and R. Mann (ed.), Dispersal <strong>of</strong><br />
Living Organisms into <strong>Aquatic</strong> <strong>Ecosystems</strong>. Maryland Sea Grant Publications, College Park,<br />
Maryland.<br />
Carlton, J.T.. 1996. <strong>Biological</strong> invasions and cryptogenic species. Ecology 77(6): 1653-1655.<br />
Carlton, J. T. and J.A. Scanlon. 1985. Progression and dispersal <strong>of</strong> an introduced alga: Codium<br />
fragile ssp. tomentosoides (Chlorophyta) on the Atlantic Coast <strong>of</strong> North America. Bot. Mar. 28:<br />
155-165.
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 22<br />
Chapman J.W. 1988. <strong>Invasions</strong> <strong>of</strong> the northeast Pacific by Asian and Atlantic Gammaridean<br />
amphipod crustaceans, including a new species <strong>of</strong> Corophium. J. Crustac. Biol. 8(3): 362-382.<br />
Chapman, J.W. and J.T. Carlton. 1991. A test <strong>of</strong> criteria for introduced species: the global<br />
invasion by the isopod Synidotea laevidorsalis (Miers, 1881). J. Crustac. Biol. 11(3): 386-499.<br />
Chapman, W.M. 1942. Alien fishes in the waters <strong>of</strong> the Pacific Northwest. Calif. Fish Game 28:<br />
9-15.<br />
Cohen, A.N. and J.T. Carlton. 1995. Nonindigenous aquatic species in a United States estuary: a<br />
case study <strong>of</strong> the biological invasion <strong>of</strong> San Francisco Bay and delta. Report to U.S. Fish &<br />
Wildlife Service, Washington, DC and National Sea Grant College Program, Connecticut Sea<br />
Grant. 246 pp.<br />
Cohen, A., C. Mills, H. Berry, M. Wonham, B. Bingham, B. Bookheim, J. Carlton, J. Chapman,<br />
J. Cordell, L. Harris, T. Klinger, A. Kohn, C. Lambert, G. Lambert, K. Li, D. Secord and J. T<strong>of</strong>t.<br />
1998. Puget Sound expedition: A rapid assessment survey <strong>of</strong> non-indigenous species in the<br />
shallow waters <strong>of</strong> Puget Sound. Washington State Department <strong>of</strong> Natural Resources. Olympia,<br />
Washington.<br />
Conlan, K. E. 1989. Revision <strong>of</strong> the crustacean amphipod genus Jassa (Corophioidea:<br />
Ischyroceridae). Can. J. Zool. 68: 2031-2075.<br />
Cooney, T. 1998. Personal communication.<br />
Dick, M.H. and J.R.P. Ross. 1988. Intertidal Bryozoa (Cheilostomata) <strong>of</strong> the Kodiak vicinity,<br />
Alaska. Center for Pacific Northwest Studies, Western Washington University, Occasional<br />
Paper No. 23.<br />
Dill, W.A. and A.J. Cordone. 1997. History and status <strong>of</strong> introduced fishes in California, 1871-<br />
1996. Calif. Dep. Fish Game Fish Bull. 178: 1-414.<br />
Eno, N.C. 1996. Non-native marine species in British waters: effects and controls. Aquat.<br />
Conserv. Mar. Freshw. Ecosyst. 6: 215-228.<br />
Farnham, W. F. 1980. Studies on aliens in the marine flora <strong>of</strong> southern England. Pp. 875-914 in<br />
Price, J. H., D.E.G. Irvine and W.F. Farnham (ed.), The Shore Environment. Academic Press,<br />
London.<br />
Feder, H.M., G.J. Mueller, H.H. Dick and D. B. Hawkins. 1973. Preliminary benthos survey. Pp.<br />
305-391 in Hood, D.W., E. Shiels and E.J. Kelley (ed.), Environmental Studies <strong>of</strong> Port Valdez.<br />
Inst. Mar. Science Occas. Publ. No. 3. University <strong>of</strong> Alaska, Fairbanks.<br />
Feder, H.M. and A.J. Paul. 1973. Age, growth and size-weight relationships <strong>of</strong> the s<strong>of</strong>t-shell<br />
clam, Mya arenaria, in Prince William Sound, Alaska. Proc. National Shellfish. Assoc. 64:45-<br />
52.
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Feder, H. M. and A.J. Paul. 1980. Seasonal trends in mei<strong>of</strong>aunal abundance on two beaches in<br />
Port Valdez, Alaska. Syesis 13: 27-36.<br />
Feder. H.M. and A.J. Paul. 1981. Distribution and abundance <strong>of</strong> some epibenthic invertebrates<br />
<strong>of</strong> Cook Inlet, Alaska. Univ. Alaska Fairbanks, Institute <strong>of</strong> Marine Science Tech. Rept. R80-3,<br />
154 p.<br />
Feder, H.M., A.J. Paul, M. Hoberg, and S. Jewett. 1981. Distribution, abundance, community<br />
structure and trophic relationships <strong>of</strong> the nearshore benthos <strong>of</strong> Cook Inlet. In: Environmental<br />
Assessment <strong>of</strong> the Alaskan Continental Shelf. Final Reports, <strong>Biological</strong> Studies 14:45-676.<br />
Fernald, M. L. 1950. Gray’s Manual <strong>of</strong> Botany. Van Nostrand, New York.<br />
Fields, C. and C. Fields. 1999. Personal communication.<br />
Fletcher, R.L. 1987. Seaweeds <strong>of</strong> the British Isles. Vol. 3. Fucophyceae (Phaeophyceae). Part 1.<br />
British Museum (Natural History), London. 359 pp.<br />
Freeman, G. 1998. Alaska. Personal Communication.<br />
Gleason, H. A. and A. Cronquist. 1991. A Manual <strong>of</strong> Vascular Plants <strong>of</strong> Northeastern United<br />
States and Adjacent Canada, Second Edition. New York Botanical Garden, Bronx, New York.<br />
Grassle, J.P. and J.F. Grassle. 1976. Sibling species in the marine pollution indicator Capitella<br />
(Polychaeta). Science 192: 567-569.<br />
Hansen, G.I. 1998. Field survey <strong>of</strong> marine plants in Prince William Sound, Alaska. Pp. 21-31 in<br />
Hines, A.H., G.M. Ruiz, J. Chapman, J. Carlton and N. Foster. <strong>Biological</strong> invasions in coldwater<br />
coastal ecosystems: Ballast-mediated introductions in Port Valdez/Prince William Sound,<br />
Alaska. Progress Report to the Regional Citizens’ Advisory Council <strong>of</strong> Prince William Sound.<br />
Henry, L.-A. 1999. Personal communication.<br />
Hines, A.H., G.M. Ruiz, J. Chapman, J. Carlton and N. Foster. 1998. <strong>Biological</strong> invasions in<br />
cold-water coastal ecosystems: Ballast-mediated introductions in Port Valdez/Prince William<br />
Sound, Alaska. Progress Report to the Regional Citizens’ Advisory Council <strong>of</strong> Prince William<br />
Sound.<br />
Hultén, E. 1968. Flora <strong>of</strong> Alaska and Neighboring Territories: A Manual <strong>of</strong> the Vascular Plants.<br />
Stanford University Press. Stanford, California.<br />
Jewett, S.C. 1998. Personal communication.<br />
Jewett, S.C. and H. M Feder. 1981. Epifaunal invertebrates <strong>of</strong> the continental shelf <strong>of</strong> the Bering<br />
and Chukchi Seas. Pp. 1131-1153 in Hood, D.W. and J. Clader (ed.), The Eastern Bering Sea
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 24<br />
Shelf: Oceanography and Resources. Vol. II, U.S. Dept <strong>of</strong> Commerce. Distributed by the<br />
University <strong>of</strong> Washington Press, Seattle. 1339 pp.<br />
Johnson, J., Alaska Department <strong>of</strong> Fish and Game. 1998. Personal communication to G. I.<br />
Hansen.<br />
Karinen, J., NOAA. 2000. Personal communication.<br />
Kitazato, H. and S. Matsushita. 1996. Laboratory observations <strong>of</strong> sexual and asexual<br />
reproduction <strong>of</strong> Trochammina hadai Uchio. Trans. Proc. Paleontol. Soc. Japan. N.S. No. 182:<br />
454-466.<br />
Kozl<strong>of</strong>f, E.N. 1987. Marine invertebrates <strong>of</strong> the Pacific Northwest. University <strong>of</strong> Washington<br />
Press, Seattle.<br />
Kyte. 1998. Personal communication.<br />
Lambert, C. C. and G. Lambert. 1998. Non-indigenous ascidians in southern California harbors<br />
and marinas. Mar. Biol. 130: 675-688.<br />
Lampman, B. H. 1946. The Coming <strong>of</strong> the Pond Fishes. Binfords & Mort, Portland, Oregon.<br />
Lambert, G. 1999. Personal communication.<br />
Matsushita, S. and H. Kitazato. 1990. Seasonality in the benthic foraminiferal community and<br />
the life history <strong>of</strong> Trochammina hadai Uchio in Hamana Lake, Japan. Pp. 695-715 in Hemleben,<br />
C. et. al. (ed.), Paleoecology, biostratigraphy, paleoceanography and taxonomy <strong>of</strong> agglutinated<br />
foraminifera. Kluwer Academic Publishers, Netherlands.<br />
McGann, M. 1998. Personal communication.<br />
McGann, M. and D. Sloan. 1996. Recent introduction <strong>of</strong> the foraminifer Trochammina hadai<br />
Uchio into San Francisco Bay, California, USA. Mar. Micropaleontol. 28: 1-3.<br />
McLean, R.I. 1972. Chlorine tolerance <strong>of</strong> the colonial hydroid Bimeria franciscana. Chesapeake<br />
Sci. 13: 229-230.<br />
McPhail, J. D. and C.C. Lindsey. 1986. Zoogeography <strong>of</strong> the freshwater fishes <strong>of</strong> Cascadia (the<br />
Columbia system and rivers north to the Stikine). Pp. 615-637 in Hocutt, C.H. and E.O. Wiley<br />
(ed.), Introduction to the Zoogeography <strong>of</strong> North American Fishes. John Wiley & Sons, New<br />
York.<br />
Meehan, B.W., J.T. Carlton and R. Wenne. 1989. Genetic affinities <strong>of</strong> the bivalve Macoma<br />
balthica from the Pacific coast <strong>of</strong> North America: evidence for recent introduction and historical<br />
distribution. Mar. Biol. 102(2): 235-241.
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 25<br />
Meehan, T. 1884. Catalogue <strong>of</strong> plants collected in July, 1883, during an excursion along the<br />
Pacific Coast in southeastern Alaska. Proc. Acad. Nat. Sci. Phila. 36: 76-96.<br />
Mills, E.L., J. H. Leach, J.T. Carlton and C.L. Secor. 1993. Exotic species in the Great Lakes: a<br />
history <strong>of</strong> biotic crises and anthropogenic introductions. J. Gt. Lakes Res. 19 (1): 1-54.<br />
Morrow, J.E. 1980. The Freshwater Fishes <strong>of</strong> Alaska. Alaska Northwest Publishing Company.<br />
Anchorage.<br />
Pettibone , M.H. 1963. Marine polychaete worms <strong>of</strong> the New England region. 1. Aphroditidae<br />
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Piorkowski, R., Alaska Department <strong>of</strong> Fish and Game. 1999. Personal communication.<br />
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Quayle, D. B. 1969. Pacific oyster culture in British Columbia. Can. Fish. Res. Board Bull. 169:<br />
1-192<br />
Ruetzler, K. 1998. Personal communication.<br />
Scagel, R.F. 1956. Introduction <strong>of</strong> a Japanese alga, Sargassum muticum, into the Northeast<br />
Pacific. Wash. Dep. Fish., Fish. Res. Pap. 1(4): 49-58.<br />
Scagel, R.F., D.J. Garbary, L. Golden and M.W. Hawkes. 1986. A synopsis <strong>of</strong> the benthic<br />
marine algae <strong>of</strong> British Columbia, northern Washington and southeast Alaska. Phycological<br />
Contribution No. 1. Department <strong>of</strong> Botany, The University <strong>of</strong> British Columbia, Vancouver.<br />
Scott, W. B. and E.J. Crossman. 1973. Freshwater Fishes <strong>of</strong> Canada. Fisheries Research Board <strong>of</strong><br />
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Simkina, R. G. 1963. On the ecology <strong>of</strong> the hydroid polyp Perigonimus megas Kinne- a new<br />
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California Coast, Third Edition. University <strong>of</strong> California Press, Berkeley. 716 pp.<br />
Smith, L.D., M.J. Wonham, L.D. McCann, G.M. Ruiz, A.H. Hines and J.T. Carlton. 1999.<br />
Invasion pressure to a ballast-flooded estuary and an assessment <strong>of</strong> inoculant survival. Biol.<br />
<strong>Invasions</strong> 1: 67-87.
Chapt. 8. Summary <strong>of</strong> NIS, page 8- 26<br />
South, G. R., and I. Tittley. 1986. A checklist and distributional index <strong>of</strong> the benthic marine<br />
algae <strong>of</strong> the North Atlantic Ocean. Huntsman Marine Laboratory and British Museum (Natural<br />
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Strasser, M. 1999. Mya arenaria- an ancient invader <strong>of</strong> the North Sea Coast. Helgol.<br />
Meeresunters. 52: 309-324.<br />
Trowbridge, C. 1998. Personal communication to G. I. Hansen.<br />
Trowbridge, C. D. 1995. Establishment <strong>of</strong> the green alga Codium fragile ssp. tomentosoides on<br />
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Turgeon, D.D., A.E. Bogan, E.V. Coan, W.K. Emerson, W.G. Lyons, W.L. Pratt, E.F.E. Roper,<br />
A. Scheltema, F.G. Thompson and J.D. Williams. 1988. Common and Scientific Names <strong>of</strong><br />
<strong>Aquatic</strong> Invertebrates from the United States and Canada: Mollusks. American Fisheries Society,<br />
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USGS, Nonindigenous <strong>Aquatic</strong> Species Database 1998. Sargassum muticum. Florida Caribbean<br />
Science Center, U.S. Geological Survey, <strong>Biological</strong> Resources Division. Gainesville FL.<br />
Web Address: http://nas.er.usgs.gov/nas/algae/sa_mutic.html<br />
U. S. Navy 1951. Report on marine borers and fouling organisms in 56 important harbors and<br />
tabular summaries <strong>of</strong> marine borer data from 160 widespread locations. U. S. Bureau <strong>of</strong> Yards<br />
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Volpe, J. P. 1999. Atlantic salmon (Salmo salar) in British Columbia and the biology <strong>of</strong><br />
invasion. 1 st National Conference on Marine Bioinvasions (January 24-27), Abstracts.<br />
Massachusetts Institute <strong>of</strong> Technology, Cambridge Massachusetts. Unpaged.<br />
Ward, R. D. and Andrew, J. 1995. Population genetics <strong>of</strong> the northern Pacific seastar Asterias<br />
amurensis (Echinodermata: Asteriidae): allozyme differentiation among Japanese, Russian, and<br />
recently introduced Tasmanian populations. Mar. Biol. 124(1): 99-109.<br />
Wing, B. L., C.M. Guthrie and A. Gharrett. 1992. Atlantic salmon in marine waters <strong>of</strong><br />
southeastern Alaska. Trans. Am. Fish. Soc. 121: 814-818.<br />
Winston, J. 1999. Personal communication.
Chapt 9A. Overview <strong>of</strong> NIS Surveys, page 9A- 1<br />
Chapter 9A. Overview <strong>of</strong> NIS Surveys<br />
Anson H. Hines, Smithsonian Environmental Research Center<br />
Gregory M. Ruiz, Smithsonian Environmental Research Center<br />
9A1. Purpose<br />
A central goal <strong>of</strong> this project was to determine whether NIS have been, or are becoming,<br />
established within Prince William Sound. Because <strong>of</strong> the diversity <strong>of</strong> potential vectors and<br />
changing patterns <strong>of</strong> transport mechanisms, our purpose was to provide as broad and<br />
comprehensive a search for NIS as possible with our resources. Since previous ecological and<br />
systematic work in Alaska has not focused on NIS, we also wished to provide a baseline for the<br />
status <strong>of</strong> NIS in Alaskan waters, against which future introductions could be measured. Recent<br />
and on-going work on NIS along the temperate west coast <strong>of</strong> North America indicates that many<br />
invasive species appear to be spreading northward from a peak <strong>of</strong> NIS diversity in San Francisco<br />
Bay, as well as other highly invaded source ports for ballast water arriving to Prince William<br />
Sound. Some <strong>of</strong> these NIS (e.g., Carcinus maenas) have moved rapidly from central California<br />
to Washington and British Columbia, and may be expected to reach Alaskan waters in coming<br />
years. We focused on Prince William Sound for our field surveys and analysis <strong>of</strong> existing<br />
samples; but because NIS <strong>of</strong>ten spread coast-wise, we also sampled ports on the adjacent Kenai<br />
Peninsula, and we considered scientific reports broadly from Alaskan waters.<br />
9A2. Approach<br />
To detect recent or well-established NIS in Port Valdez / Prince William Sound and<br />
adjoining areas <strong>of</strong> risk for invasion, we used several methods, including:<br />
• Rapid assessment field surveys <strong>of</strong> estuarine and marine invertebrates and plants for Port<br />
Valdez, Prince William Sound, Seward and Homer. The objective was for experienced<br />
general ecologists to survey major habitats and communities, especially for large NIS plants<br />
and animals detectable in the field by experienced naturalists.<br />
• Focal taxonomic field collections in Prince William Sound, Seward and Homer. The<br />
objective was for taxonomic experts to sample and analyze key taxonomic groups that have<br />
known NIS but which are difficult for generalists to identify, providing definitive<br />
identification and careful, authoritative assessments <strong>of</strong> the native, invasive and cryptogenic<br />
status. Whenever possible, we also wished the taxonomic experts to have the opportunity to<br />
sample the sites using their specialized methods and knowledge for collecting the focal<br />
taxon.<br />
• Fouling plate surveys in Prince William Sound, Seward and Homer. The objective was to<br />
provide a replicated standard sampling method <strong>of</strong> assay for NIS in a community that is prone<br />
to invasions, but which has received little prior ecological analysis in Alaska.<br />
• Re-examination <strong>of</strong> museum and reference collections for Prince William Sound. The<br />
objective was to re-examine extensive collections already available in the University <strong>of</strong><br />
Alaska Museum and vouchers samples from Exxon Valdez Oil Spill (EVOS) and other<br />
ecological studies, developing a screening method <strong>of</strong> screening for potential NIS.<br />
Our approach <strong>of</strong> utilizing this array <strong>of</strong> methods served to maximize spatial, temporal, taxonomic,<br />
and habitat coverage, while still focusing our limited resources upon elements known to be <strong>of</strong><br />
highest risk <strong>of</strong> invasion. The field surveys provided broad coverage <strong>of</strong> Prince William Sound, as
Chapt 9A. Overview <strong>of</strong> NIS Surveys, page 9A- 2<br />
will as Anchorage, Homer and Seward as important ports in neighboring Cook Inlet and the<br />
Kenai Peninsula (Fig. 9.A.1).
Chapt 9A. Overview <strong>of</strong> NIS Surveys, page 9A- 3<br />
In any ecosystem, the ability to detect established NIS varies in space and time, because<br />
most aspects <strong>of</strong> species distribution and abundance are probabilistic. Like any population, NIS<br />
populations are typically patchily distributed across the full range <strong>of</strong> habitats, and may undergo<br />
marked seasonal and annual fluctuations. In high latitude ecosystems like Prince William<br />
Sound, this variation is extremely pronounced, due to rapid changes in photoperiod and<br />
fluctuations in surface salinities resulting from warm season run<strong>of</strong>f. By using an array <strong>of</strong><br />
sampling at several times throughout the growing season, we increased the probability <strong>of</strong><br />
detecting NIS.<br />
In addition to the stochastic aspect <strong>of</strong> detecting NIS, detection <strong>of</strong> NIS in Prince William<br />
Sound is difficult because the biota is not well described by taxonomists and biogeographers.<br />
Despite the extensive sampling in Prince William Sound for EVOS and other ecological<br />
programs, there are no comprehensive keys or field guides to the biota <strong>of</strong> the region (a notable<br />
exception is the guide to Alaskan molluscs; Foster 1991). In our Pilot Study (Ruiz & Hines<br />
1997) <strong>of</strong> Port Valdez alone, we found a surprisingly high percentage <strong>of</strong> new species records, and<br />
our literature analysis showed that 20-50% <strong>of</strong> the species are cryptogenic in origin.<br />
It was necessary to sample several habitats <strong>of</strong> Prince William Sound that have received<br />
little scientific study. For example, there are almost no publications on the fouling community <strong>of</strong><br />
floats and pilings, yet NIS are very common in this habitat in west coast source ports, where<br />
some <strong>of</strong> these NIS have been very destructive. S<strong>of</strong>t-bottom habitats <strong>of</strong> Prince William Sound<br />
have received less study than rocky substrates.<br />
Also, in much <strong>of</strong> the previous work associated with the Exxon Valdez Oil Spill,<br />
taxonomic identifications were not carried out to species but were reported at relatively high<br />
taxonomic levels (e.g., Family, Order). Without careful identification to species by expert<br />
taxonomists, NIS are <strong>of</strong>ten confused with similar native species. Therefore, we brought<br />
systematic experts for several focal taxonomic groups to Alaska for field collections or<br />
contracted them to identify selected subsets <strong>of</strong> samples.<br />
9A3. Site selection and sampling design<br />
In addition to a survey <strong>of</strong> Port Valdez and Sawmill Bay during our Pilot Study in<br />
June 1997, we conducted two broad expeditions to survey Prince William Sound for NIS during<br />
low tide series <strong>of</strong> 20-28 June 1998 and 8-16 August 1999. Fouling Plate Surveys were<br />
conducted during 7-17 September 1998 and in 8-16 August 1999.<br />
The surveys in September 1998 and August 1999 included sampling stations at Homer<br />
and Cordova on the Kenai Peninsula adjacent to Prince William Sound. The survey sampling<br />
design focused on invertebrates and plants in a variety <strong>of</strong> habitats <strong>of</strong> shallow water, the intertidal<br />
zone, and accessible man-made surfaces (e.g., floats, pilings, buoys). We selected sampling sites<br />
that were judged to be most susceptible to invasion by NIS:<br />
• areas most likely to be in the path <strong>of</strong> ballast water discharged from tankers (as estimated by<br />
circulation models and the path <strong>of</strong> the Exxon Valdez Oil Spill);<br />
• ports and sites <strong>of</strong> sustained disturbance by human activities (especially Port Valdez, Cordova,<br />
and Whittier, as well as Homer and Seward);<br />
• habitats associated with previously reported NIS and cryptogenic species;
Chapt 9A. Overview <strong>of</strong> NIS Surveys, page 9A- 4<br />
• warmer water areas;<br />
• marinas, floats, buoys, and pilings with accessible fouling communities; and<br />
• sites <strong>of</strong> active aquaculture for the Japanese Oyster Crassostrea gigas.<br />
Together, these habitats comprise a broad area <strong>of</strong> the shallow, nearshore margins and islands <strong>of</strong><br />
Prince William Sound. The survey attempted to gain broad coverage <strong>of</strong> these habitats, including<br />
46 sites in June 1998, 9 sites for fouling plates in September 1998, and 33 sites including a<br />
subset <strong>of</strong> 7sites for fouling plates in August 1999, which were spread throughout the major<br />
regions <strong>of</strong> the Sound and the port sites <strong>of</strong> the Kenai Peninsula (see map Fig. 9A.1, Table 9A.1).<br />
The survey sampled three major habitats: intertidal and shallow rocky substrates; intertidal and<br />
shallow s<strong>of</strong>t sediments; and fouling communities on floats, buoys, pilings and oyster culture<br />
structures. The salinity <strong>of</strong> the array <strong>of</strong> sites ranged from fresh to fully marine areas <strong>of</strong> the Sound.<br />
The rapid assessment survey methods were similar to those employed in NIS surveys <strong>of</strong><br />
San Francisco Bay (Cohen & Carlton 1995) and Puget Sound (Cohen et al. 1998). The approach<br />
utilizes a team <strong>of</strong> experienced naturalists and general ecologists to sample as great a diversity <strong>of</strong><br />
organisms at as broad an array <strong>of</strong> sites as possible within the region <strong>of</strong> concern. The team for the<br />
surveys consisted <strong>of</strong>:<br />
• John Chapman (OSU), general NIS <strong>of</strong> northeast Pacific and peracaridan crustacea;<br />
• Nora Foster (UAF), marine invertebrates <strong>of</strong> Alaska, especially mollusca;<br />
• Anson Hines (SERC), barnacles and decapod crustaceans; and<br />
• Todd Miller (Hatfield Marine Science Center), technical assistance and peracaridan<br />
crustaceans.<br />
The survey teams utilized a variety <strong>of</strong> transportation modes to travel among sites<br />
throughout Prince William Sound, including vans, ferries, float planes, small boats, vessels <strong>of</strong><br />
Stan Stephens Tours, and the Fishing Vessel Kristina. Following collecting, samples were<br />
processed in temporary laboratories provided by USF&WS Refuge in Homer, the Seward<br />
Marine Science Center, University <strong>of</strong> Alaska in Seward, Prince William Science Center in<br />
Cordova, the SERC <strong>Invasions</strong> Biology Laboratory in Valdez, the Prince William Sound<br />
Community College in Valdez, and several hotels. The F/V Kristina also served as laboratory<br />
platform for processing <strong>of</strong> samples while in transit among some <strong>of</strong> the sites. For each sampling<br />
site, the diversity and relative abundance <strong>of</strong> species were recorded. Field notes included GPS<br />
readings, sketches <strong>of</strong> the sites, salinity and temperature readings, and notes on common or<br />
abundant species identified in the field. Samples were collected by hand, by scraper to remove<br />
fouling organisms, and by trowel to collect s<strong>of</strong>t sediment. Samples <strong>of</strong> sediment, algalinvertebrate<br />
turf, and scrapings <strong>of</strong> fouling communities were washed and sieved on 5mm, 1mm<br />
and 0.5 mm mesh. Each sample was “rough sorted” immediately after collection to aid in<br />
identification <strong>of</strong> large or delicate specimens and to preserve voucher specimens for subsequent<br />
work-up in the laboratory. Voucher samples were preserved in either 70% EtOH or 10%<br />
formalin (as appropriate to the type <strong>of</strong> organism). Voucher samples from the survey have been<br />
distributed to appropriate taxonomic experts for definitive identification in the laboratory using<br />
microscopes.
Chapt 9A. Overview <strong>of</strong> NIS Surveys, page 9A- 5<br />
Map<br />
No.<br />
TABLE 9A.1. Collecting Sites for NIS Surveys<br />
(1997,1998,1999).<br />
Site Station 98 Cruise<br />
Station<br />
No.<br />
Latitude Longitude 1997<br />
Pilot<br />
Study<br />
Rapid Community<br />
Assessments<br />
1998<br />
Surveys<br />
1999<br />
Survey<br />
Fouling Plate<br />
Surveys<br />
1998 1999<br />
1 Anchorage Port Anchorage 61º14'N 149º45'W X<br />
Westchester Lagoon 61º13'N 149º50'W X<br />
2 Potter Potter flats 61º05'N 149º40'W X<br />
3 Katchemak Bay Homer small boat harbor X X X<br />
Homer spit X X X<br />
Homer spit mudflat X X X<br />
4 Sadie Cove X<br />
5 Seward Seward small boat harbor X X X<br />
Lowell Point<br />
X<br />
6 Whittier Whittier small boat harbor 60º46'37"N 148º41'24"W X X X<br />
Whittier ferry dock 60º46'25"N 148º40'55"W X X<br />
7 Shotgun Cove Shotgun Cove 60º47'26"N 148º32'30"W X<br />
8 Esther Island Lake Bay buoy 28 60º47'37"N 148º05'01"W X<br />
Lake Bay oysters 30 60º48'00"N 148º05'24"W X<br />
9 Squaw Bay Squaw Bay oysters 36 60º50'00"N 147º49'20"W X<br />
10 Eaglek Bay Eaglek Bay oysters 37 60º51'00"N 147º45'36"W X<br />
11 Fairmont Bay Fairmont Bay oysters X X<br />
12 Growler Island Growler mudflat 38 60º54'15"N 147º07'48"W X X<br />
Growler dock 39 60º54'13"N 147º07'48"W X X<br />
13 Valdez Arm, Sawmill Bay Sawmill Bay shore 9 61º03'15"N 146º47'24"W X X<br />
Sawmill Bay mudflat 10 61º03'23"N 146º47'24"W X X<br />
Navigation buoy 40 61º03'16"N 146º41'39"W X<br />
14 Valdez Arm Potato Point X<br />
15 Port Valdez Anderson Bay X<br />
16 Valdez Duck Flats low intertidal 44 61º07'28"N 146º18'00"W X X<br />
Duck Flats high intertidal 45 61º08''24"N 146º19'30"W X X<br />
Floating cargo dock 43 61º07'25"N 14618'36 "W X X<br />
SERVS dock 61º07'25"N 146º21'15"W X X X<br />
Small boat harbor 46 61º07'25"N 146º21'15"W X X X X X<br />
USCG dock " " X X<br />
Ferry dock " " X<br />
17 Port Valdez, Dayville flats Dayville flats 4 61º04'54"N 146º19'00"W X X<br />
18 Valdez Marine Terminal Alyeska small boat ramp 1 61º05'12N 146º23'30"W X X<br />
Alyeska small boat harbor 2 61º05'10"N 146º22'28"W X X<br />
Alyeska entrance 3 61º05'10"N 146º21'55"W<br />
Terminal floats 42 61º05'20"N 146º24'09"W X X
Chapt 9A. Overview <strong>of</strong> NIS Surveys, page 9A- 6<br />
Map<br />
No.<br />
TABLE 9A.1. (continued) Collecting Sites for NIS<br />
Surveys (1997,1998,1999).<br />
Site Station 98 Cruise<br />
Station<br />
No.<br />
Latitude Longitude 1997<br />
Pilot<br />
Study<br />
Rapid Community<br />
Assessments<br />
1998<br />
Surveys<br />
1999<br />
Survey<br />
Fouling Plate<br />
Surveys<br />
1998 1999<br />
19 Valdez Arm, Rockey Point Rockey Point 11 60º57'36"N 146º45'36"W X<br />
20 Busby Island South reef 5 60º52'55"N 146º46'29"W X<br />
Busby Island 6 60º52'54"N 146º46'24"W X<br />
21 Tatitlek Tatitlek Narrows oysters 8,12 60º52'06"N 146º43'30"W X X X<br />
Village dock 60º52'06"N 146º43'30"W X X X<br />
Ferry dock 60º52'06"N 146º43'28"W X X<br />
22 Bligh Island Cloudman Bay 7 60º50'11"N 146º43'15"W X<br />
23 Sheep Bay Upper Sheep Bay 13 60º52'12"N 146º43'48"W X<br />
Middle Sheep Bay 14 60º40'21"N 145º57'06"W X<br />
24 Cordova Small boat harbor 15,19 60º32'30"N 145º46'28"W X X X<br />
Mudflat S <strong>of</strong> Small Boat Harbor 18 60º32'28"N 145º46'28"W X X X<br />
Marine Science Center 17 60º32'48"N 145º46'27"W X X<br />
Fish Dock & Flats 16 60º32'27"N 145º46'26"W X<br />
Ferry dock 60º32'27"N 145º46'24"W X<br />
25 Hawkins Island Windy Bay 20 60º33'54"N 145º58'38"W X<br />
26 Orca Bay Channel Buoy 21 60º32'22"N 146º55'55"W X<br />
27 Hinchinbrook Island Constantine Harbor X<br />
28 Montague Island Port Chalmers X X<br />
29 Green Island Green Island 22 60º18'19"N 145º58'38"W X<br />
30 Evans Island Sawmill Bay 23 60º03'31"N 147º59'47"W X<br />
Port San Juan 24 60º04'2"N 148º03'36"W X<br />
31 Chenega Island Chenega dock X X<br />
32 Eleanor Island Northwest Bay middle arm 25 60º32'57"N 147º34'48"W X<br />
33 Knight Island Passage Knight Island Passage buoy 26 60º33'52"N 147º49'11W X<br />
34 Main Bay Main Bay 27 60º31'58"N 148º04'41"W X<br />
Main Bay fish hatchery 28 60º31'16"N 148º05'35"W X
Chapt 9A. Overview <strong>of</strong> NIS Surveys, page 9A- 7<br />
The experts for the Focal Taxonomic Collections included:<br />
• Gayle Hansen (Hatfield Marine Science Center, Oregon State University), phycologist with<br />
special expertise in Alaskan macroalgae;<br />
• John Chapman (Hatfield Marine Science Center, Oregon State University), peracarid<br />
crustacea;<br />
• Jeff Cordell (School <strong>of</strong> Fisheries, University <strong>of</strong> Washington), copepod crustaceans;<br />
• Nora Foster (University <strong>of</strong> Alaska Museum), molluscs and other marine invertebrates <strong>of</strong><br />
Alaska;<br />
• Jeffery Goddard (University <strong>of</strong> California, Santa Barbara), opisthobranch molluscs<br />
• Jerry Kudenov (University <strong>of</strong> Alaska, Anchorage), polychaete worms;<br />
• Gretchen Lambert (Friday Harbor Laboratories, University <strong>of</strong> Washington), ascidians;<br />
• Charles Lambert (Friday Harbor Laboratories, University <strong>of</strong> Washington), ascidians;<br />
• Claudia Mills (Friday Harbor Laboratories, University <strong>of</strong> Washington), cnidarian medusae<br />
and ctenophora;<br />
• Lise Schickel (University <strong>of</strong> California, Santa Barbara), decapod crustaceans and parasitic<br />
crustaceans;<br />
• Anson Hines (SERC), barnacles and decapod crustaceans;<br />
• Judith Winston (Virginia Natural History Musem), bryozoans; and<br />
• Lea Ann Henry (University <strong>of</strong> Toronto), hydrozoan cnidarians.<br />
At numerous locations in Port Valdez / Prince William Sound, sediment samples were collected<br />
for identifications <strong>of</strong> foraminiferans, particularly the Asian NIS Trochammina hadai, which is<br />
extensively introduced in San Francisco Bay and other west coast source ports. This NIS was<br />
reported from deep-water samples taken for the Exxon Valdez Oil Spill study. Samples were<br />
processed and sent to Mary McGann <strong>of</strong> US Geological Survey, Menlo Park, CA, for<br />
identification.<br />
Other details <strong>of</strong> the methods are provided below within the individual chapters for each<br />
Focal Taxonomic Collection, the Fouling Community Analysis and the Re-examination <strong>of</strong> the<br />
Museum, Reference and Voucher Collections.<br />
9A4. References<br />
Cohen, A.N. and J.T. Carlton. 1995. Nonindigenous aquatic species in a United States estuary: a<br />
case study <strong>of</strong> the biological invasion <strong>of</strong> San Francisco Bay and delta. Report to U.S. Fish &<br />
Wildlife Service, Washington, DC and National Sea Grant College Program, Connecticut Sea<br />
Grant. 246 pp.<br />
Cohen, A., C. Mills, H. Berry, M. Wonham, B. Bingham, B. Bookheim, J. Carlton, J. Chapman,<br />
J. Cordell, L. Harris, T. Klinger, A. Kohn, C. Lambert, G. Lambert, K. Li, D. Secord, and J. T<strong>of</strong>t.<br />
1998. Puget Sound expedition: A rapid assessment survey <strong>of</strong> non-indigenous species in the<br />
shallow waters <strong>of</strong> Puget Sound. Washington State Department <strong>of</strong> Natural Resources. Olympia,<br />
Washington.<br />
Foster, N.R. 1991. Intertidal Bivalves: A Guide to the Common Marine Bivalves <strong>of</strong> Alaska.<br />
University <strong>of</strong> Alaska Press, Fairbanks. 152 pp.
Chapt 9A. Overview <strong>of</strong> NIS Surveys, page 9A- 8<br />
Ruiz, G.M. and A.H. Hines. 1997. Patterns <strong>of</strong> nonindigenous species transfer and invasion in<br />
Prince William Sound, Alaska: Pilot Study. Report Submitted to the Prince William Sound<br />
Citizens’ Advisory Council. 80pp
Chapt 9B. NIS Surveys: Rapid Community Assessments, page 9B- 1<br />
Chapter 9B. Rapid Community Assessment<br />
John Chapman, Hatfield Marine Science Center, Oregon State University<br />
Collections <strong>of</strong> some taxonomic groups from the 1998 survey were transferred to<br />
systematic experts for focal taxonomic analysis provided in other sections <strong>of</strong> this report- see<br />
sections below for Focal Taxonomic Collections.<br />
Other data and analysis to be provided by John Chapman have not been forthcoming.
Chapt 9C1. Marine Plants, page 9C1- 1<br />
Chapter 9C1. Focal Taxonomic Collections: Marine Plants in Prince William Sound,<br />
Alaska<br />
Gayle I. Hansen, Hatfield Marine Science Center, Oregon State University<br />
Background<br />
Several NIS marine plants with potential for invasion <strong>of</strong> Alaskan waters have been<br />
reported on the west coast <strong>of</strong> North America. For example, the pervasive algae Sargassum<br />
muticum, Lomentaria hakodatensis, and the Japanese eelgrass Zostera japonica are thought to<br />
have been introduced with the aquaculture <strong>of</strong> oysters by the importation <strong>of</strong> spat from Japan. At<br />
least 5 oyster farms occur in Prince William Sound, and all have imported spat. For the herringroe-on-kelp<br />
(HROK) pound fishery, the giant kelp Macrocystis integrifolia is transported to<br />
Prince William Sound via plane from southeast Alaska (the northern limit <strong>of</strong> this species) to be<br />
used as a substrate for herring roe. Although the giant kelp cannot recruit in Prince William<br />
Sound, it seems likely that other species, accidentally co-transported with Macrocystis, could<br />
become established. Our Pilot Study (Ruiz and Hines 1997) also considered several NIS algal<br />
species reported from Alaskan waters, including a report <strong>of</strong> a cosmopolitan species Codium<br />
fragile tomentasoides from Green Island.<br />
Methods<br />
Sample Period. Marine benthic algae, seagrasses, and intertidal lichens were sampled as a part<br />
<strong>of</strong> the cruise aboard the F/V Kristina during 20-28 June 1998, described above for invertebrates.<br />
Site Information. A subset <strong>of</strong> 19 <strong>of</strong> the 46 sites selected for invertebrate sampling were chosen<br />
for the plant study, including 13 intertidal sites (4 within Port Valdez and 9 in Prince William<br />
Sound) and 6 <strong>of</strong>f-shore float sites. Site abbreviations (for tables and figures to follow),<br />
coordinates, temperature, and salinity are given in Table 9C1.1. Please note that the site numbers<br />
in Table 9C1.1 for plants do not correspond to the site numbers on the map (Fig. 9A2) or Table<br />
9A1 for invertebrates. The substratum types, listed for each site, are only those sampled for<br />
algae and seagrasses. For analysis and discussion, the 19 plant sites have been grouped into 5<br />
basic habitat types: harbors, mud bays, rocky headlands and reefs, rocky bays, and floats. These<br />
will be discussed in greater detail in the Results section below.<br />
Surveying Techniques. At each site, intertidal areas accessible by foot within the time period<br />
provided were sampled. Since introduced species could potentially occur in any <strong>of</strong> the marine<br />
plant taxonomic groups, it was important to sample the entire range <strong>of</strong> species present from as<br />
broad an area as possible. Marine algal populations are well-known for being extremely patchy<br />
in distribution, caused primarily by narrow species requirements (and tolerances) for substratum,<br />
tidal height (exposure), salinity, nutrients, and sunlight. Since the species were patchily<br />
distributed, they were encountered and collected sporadically, not uniformly over time. For this<br />
study, abundance was noted only when unusually large patches <strong>of</strong> a particular species were<br />
encountered; it was not documented uniformly for entire sites.<br />
Time Allotment. As shown in Table 9C1.s, sampling times at major sites varied from 10<br />
minutes to 2 hours. At low diversity sites, such as Cloudman Bay, the time provided was<br />
sufficient for complete algal collection; at other sites, such as Green Island, the time was <strong>of</strong>ten
Chapt 9C1. Marine Plants, page 9C1- 2<br />
TABLE 9C1.1. General Site Information, 1998 Collections*<br />
ABBR. DATE LOCATION LAT LON SUBSTR. T SAL<br />
(ºC) (0/00)<br />
Port Valdez (Val)<br />
al-sbr Jun 20 Alyeska, small boat ramp, 61º 05' 12''N 146º 23' 30"W br, co 9 0<br />
Port Valdez<br />
al-pil Jun 20 Alyeska, small boat harbor, 61º 05' 10"N 146º 22' 28"W pi 9 0<br />
Port Valdez<br />
sough Jun 20 Slough, near Alyeska gate, 61º 04' 54"N 146º 19' 00"W mu 11 0<br />
Port Valdez<br />
duckflat Jun 28 Duckflat, Port Valdez 61º 07' 28''N 146º 18' 00''W mf 17-22 0-4<br />
Other Harbors<br />
Cor Jun 23 Cordova, Orca Inlet 60º 32' 28''N 145º 46' 28''W dm, mf 10-16 5-28<br />
Whit Jun 26 Whittier, Passage Canal 60º 46' 25''N 148º 40' 55''W dm, bo, gr, mu 4-10 8-14<br />
Mud/Cobble Bays<br />
CB Jun 21 Cloudman Bay, Bligh I. 60º 50' 11''N 146º 43' 15''W mu, co 10 3<br />
SMB Jun 22 Sawmill Bay, Valdez Arm 61º 03' 15''N 146º 47' 24''W mu, co 8 5<br />
Gro Jun 27 Growler I. 60º 54' 15''N 147º 07' 48''W mu, co 22 11<br />
Rk Headlands and Reefs<br />
RP Jun 22 Rocky Point, Valdez Arm 60º 57' 36''N 146º 45' 36''W br, co 11 15<br />
Bus Jun 21 Busby I., south reef 60º 52' 55''N 146º 46' 29''W br, co 11 23<br />
Green Jun 24 Green I., northwest reef 60º 18' 19''N 147º 23' 47''W bo, br 11 30<br />
Rk Bays<br />
NW Jun 25 Northwest Bay, middle arm, 60º 32' 57"N 147º 34' 48''W gr, co 12 10-27<br />
Eleanor Island<br />
Floats and Buoys<br />
TAT Jun 22 Tatitlek Narrows, Bligh I. 60º 52' 12''N 146º 43' 48''W oy 12 26<br />
WBF Jun 23 Windy Bay, Hawkins I. 60º 33' 54''N 145º 58' 38''W oy 14 28<br />
MBF Jun 25 Main Bay 60º 31' 58''N 148º 04' 41''W bb 14 19-20<br />
EIF Jun 25 Lake Bay, Esther I. 60º 48' 00''N 148º 05' 24''W bb 12 16<br />
SBF Jun 26 Squaw Bay 60º 50' 00''N 147º 49' 20''W oy 14 24<br />
EBF Jun 26 Eaglek Bay 60º 51' 00''N 147º 45' 36''W oy 14 24<br />
Abbreviations:<br />
*= coordinates, temperature, and salinity provided by T. Miller lon=longitude<br />
abbr.=abbreviations<br />
mf=mudflat<br />
bb=barrier buoy<br />
mu=mud<br />
bo=boulders<br />
oy=oysterfloats<br />
br=bedrock<br />
pi=wood pilings<br />
co=cobble<br />
rk=rocky<br />
dm=docks/marina<br />
sal=salinity<br />
gr=gravel<br />
subst=substratum<br />
lat=latitude<br />
t=temperature<br />
seriously inadequate to sample fully the algal diversity present. Differences among sites in<br />
amount <strong>of</strong> time for collecting were not factored out or corrected after sampling was completed;<br />
however, the sites which were judged to be undercollected are designated with an “*” in Table<br />
9C1.2.<br />
TABLE 9C1.2. Collection Efficiency Records<br />
Data Type<br />
Major Collection Sites (without <strong>of</strong>f-shore floats)<br />
Val al-sbr al-pil slough duckflat Cor Whit CB SMB Gro RP Bus Green NW<br />
New Records 5 1 1 1 4 5 2 2 5 4 2 2 6 1<br />
Total Species 47 35 7 7 24 41 56 10 45 63 61 59 71 69<br />
Collection Time 165 40* 10 15 100 120 120 30 45* 105 70* 75* 65* 136<br />
Correlation R R 2 * = undercollected sites<br />
Total Species:Time** 0.896 43% ** = both without Val (Total Valdez) included<br />
New Records:Time** 0.498 7%
Chapt 9C1. Marine Plants, page 9C1- 3<br />
Field Sampling & Processing. All algal sampling was done by hand or with a chisel. Collected<br />
specimens were then placed in plastic bags for transport back to the boat for processing. On<br />
board, the samples were sorted to species and then either pressed in a plant press or preserved in<br />
5% formalin/seawater. Site notes and preliminary species lists were made in the field, and some<br />
final identifications were done on board. However, for most species final determinations were<br />
not made until returning to the Hatfield Marine Science Center in Newport, Oregon, where<br />
compound microscopy was available. The smaller marine algae that require microscopic<br />
examination while still alive for identification were necessarily excluded from this study. After<br />
identification, both liquid and dried specimens were curated, labeled, and deposited in the<br />
herbarium at OSU/HMSC for reference in future Alaskan marine algal studies.<br />
Identifications. Since no marine flora (identification guide) <strong>of</strong> Prince William Sound exists, the<br />
algae collected during this study were identified using a wide variety <strong>of</strong> literature. For common<br />
species, the most important references utilized were Abbott and Hollenberg (1976), Gabrielson et<br />
al. (1993), Perestenko (1994), and Sears (1998). For more obscure species, much <strong>of</strong> the world<br />
taxonomic literature on temperate/arctic marine algae was employed. To confirm the<br />
identification <strong>of</strong> particularly difficult or important species, some specimens were sent out to<br />
colleagues for identification using molecular techniques. These taxa are designated with a "#" in<br />
the species charts. Due to the costs <strong>of</strong> these tests, these results will not be presented here, but<br />
instead will be presented at a later date as part <strong>of</strong> the papers prepared by these experts.<br />
Distributions, Residency Status, and New Records. In determining if species were introduced,<br />
the local and global distributions had to be determined from the literature. Some <strong>of</strong> the<br />
references used for this process were: Scagel et al. (1993), Sears et al.(1998), Selivanova and<br />
Zhigadlova (1997), Lee (1980), Guiry (1998), Rueness (1977), Phillips and Menez (1988),<br />
Yoshida et al. (1995), Adams (1983, 1994), Womersley (1984, 1987, 1994, 1996), Lindstrom<br />
(1977), Hansen et al. (1981), and Hansen (1997). These distributions, summarized in the<br />
abbreviated form explained below, are shown under range (Ra) in the first column <strong>of</strong> the species<br />
site lists (Tables 9C1.3 – 9C1.5). The ranges provided the basis for determining the Residency<br />
Status (St) <strong>of</strong> the species. Residency status rankings include the following 5 categories:<br />
• E (Endemic) = species known only from Alaska<br />
• N (Native) = species native to the North Pacific, including species with ranges limited to the<br />
northeast Pacific (nep) and those that occur in all other areas around the northern Pacific rim<br />
(np).<br />
• C (Cryptogenic) = species with extremely broad distributions that occur circumboreally (cb)<br />
and/or extend to the southern hemisphere (ws).<br />
• I (Introduced) = species that appear to have been introduced to the area.<br />
• F (Failed Introduction) = deliberately introduced species that have failed to colonize the<br />
area
Chapt 9C1. Marine Plants, page 9C1- 4<br />
TABLE 9C1.3. Marine and Estuarine Plants Collected in Port Valdez, Alaska<br />
NIS ANALYSIS<br />
PORT VALDEZ<br />
TAXA<br />
…………..June 1998 Collections……..…. Total<br />
Ra St NR So Val al-sbr al-pil slough duckflat Checklist<br />
RHODOPHYTA, Rhodophyceae<br />
ws C Ahnfeltia fastigiata O<br />
nep N Ahnfeltiopsis gigartinoides O<br />
nep N Antithamnionella pacifica O<br />
ws C Audouinella purpurea O<br />
ws C Bangia atropurpurea X X X<br />
nep N# Ceramium gardneri O<br />
np N Constantinea subulifera O<br />
np N Corallina frondescens O<br />
np N Corallina vancouveriensis O<br />
np N Cryptonemia borealis O<br />
np N Cryptonemia obovata O<br />
nep N Cryptosiphonia woodii X* X* O<br />
cb C Devaleraea ramentacea X X O<br />
cb C Dumontia contorta O<br />
np N Dumontia simplex O<br />
nep N Endocladia muricata O<br />
ws C Erythrotrichia carnea O<br />
np N Glioipeltis furcata X* X* O<br />
np,ar N Halosaccion firmum X X O<br />
np N Halosaccion glandiforme X* X* O<br />
ws C Hildenbrandia rubra O<br />
np N Leachiella pacifica O<br />
np N Lithophyllum dispar O<br />
nep N Mastocarpus papillatus complex X* X* O<br />
np N Mastocarpus cf. pacificus X* X* X<br />
nep N Mazzaella heterocarpa O<br />
np N Mazzaella phyllocarpa X X X<br />
nep N Mazzaella splendens O<br />
nep N Microcladia borealis O<br />
ws C Nemalion helminthoides O<br />
np N Neorhodomela aculeata X X O<br />
np N Neorhodomela larix X X O<br />
np N Neorhodomela oregona X* X* O<br />
nep N Odonthalia floccosa O<br />
np N Odonthalia kamtschatica O<br />
np N Odonthalia setacea (drift) O<br />
nep N Palmaria hecatensis X* X* O<br />
cb C Palmaria mollis/palmata O<br />
np N Phycodrys riggii O<br />
ws C Polysiphonia brodiaei O<br />
nep N Polysiphonia hendryi v. deliquescens X X O<br />
nep N Polysiphonia hendryi v. hendryi O<br />
nep N Polysiphonia hendryi v. luxurians O<br />
nep N Polysiphonia pacifica v. pacifica O<br />
nep N Porphyra cuneiformis O<br />
nep N Porphyra mumfordii O<br />
np N Porphyra perforata O<br />
cb C# NR NAT Porphyra purpureo-violacea O
Chapt 9C1. Marine Plants, page 9C1- 5<br />
Table 9C1.3. continued<br />
nep N NR Wa Porphyra rediviva X* X* X<br />
np N Pterosiphonia bipinnata X* X* O<br />
np N Ptilota filicina O<br />
cb C Ptilota serrata (incl. pectinata ) O<br />
cb C Rhodomela lycopodioides O<br />
cb C Scagelia americana O<br />
np N Tokidadendron kurilensis X X O<br />
nep N Weeksia coccinea X* X* X<br />
HETEROKONTOPHYTA, Phaeophyceae<br />
Val al-sbr al-pil slough duckflat Checklist<br />
cb C Agarum clathratum (cribrosum) O<br />
cb C Chordaria flagelliformis Xun X un O<br />
np N Chordaria gracilis O<br />
cb C Coilodesme bulligera O<br />
np N Costaria costata O<br />
cb C Desmarestia aculeata O<br />
cb C Desmarestia viridis O<br />
cb C Dictyosiphon foeniculaceus X* X* X un O<br />
nep N Ectocarpus parvus O<br />
ws C Ectocarpus siliculosus O<br />
nep N Elachista lubrica X* X* O<br />
cb I NR NAT Fucus cottonii X* X* X<br />
cb C Fucus gardneri/distichus/evanescens X* X* X* un X* O<br />
cb C Fucus spiralis X X X X<br />
np, a N Laminaria "groenlandica"/bongardiana X* X* O<br />
cb C Laminaria saccharina X* X* X O<br />
np N Laminaria yezoensis X* X* O<br />
ws C Leathesia difformis O<br />
cb C Melanosiphon intestinalis X X O<br />
ws C Petalonia fascia O<br />
ws C Pilayella littoralis/washingtonensis X* X* X X* O<br />
ws C Scytosiphon simplicissimus X* X* X un O<br />
np N Soranthera ulvoidea X* X* O<br />
ws C Sphacelaria rigidula O<br />
cb C Spongonema tomentosum O<br />
HETEROKONTOPHYTA, Xanthophyceae<br />
Val al-sbr al-pil slough duckflat Checklist<br />
ws C NR BC Vaucheria longicaulis () mats X* X* X<br />
CHLOROPHYTA, Chlorophyceae<br />
Val al-sbr al-pil slough duckflat Checklist<br />
cb C Acrosiphonia arcta X* X* X* O<br />
nep N Acrosiphonia coalita O<br />
np N Acrosiphonia saxatilis O<br />
cb C Blidingia chadefaudii O<br />
ws C NR BC Blidingia marginata X* X* X* X<br />
ws C Blidingia minima X* X* X* O<br />
cb C Blidingia subsalsa X* X* X* X* O<br />
cb C Chaetomorpha capillaris/cannabina O<br />
nep N NR Wa Chaetomorpha recurva O<br />
ws C Cladophora albida O<br />
ws C Cladophora sericea O<br />
ws C Enteromorpha clathrata O<br />
ws C Enteromorpha in compressa O<br />
ws C Enteromorpha intestinalis X* X* X* O<br />
ws C Enteromorpha linza O<br />
ws C Enteromorpha prolifera/torta X* X* X* X* X
Chapt 9C1. Marine Plants, page 9C1- 6<br />
Table 9C1.3. continued<br />
cb C Gayralia oxyspermum X* X* X<br />
cb C NR BC Halochlorococcum moorei X* X* X* X<br />
cb C Kornmannia zostericola (epiphytic) O<br />
cb C Monostroma grevillei/arcticum O<br />
ws C Rhizoclonium implexum X* X* X* X* O<br />
ws C Rhizoclonium riparium X* X* X* X* O<br />
ws C Rhizoclonium tortuosum O<br />
ws C Ulothrix implexa (non flacca ) X* X* X* X X* O<br />
np C# Ulva fenestrata /expansa/lactuca X* X* X* O<br />
cb C Ulvaria obscura X* X* X* O<br />
ws C Urospora penicilliformis X* X* X<br />
SEAGRASSES<br />
Val al-sbr al-pil slough duckflat Checklist<br />
cb C Zostera marina X* X* O<br />
LICHENS<br />
Val al-sbr al-pil slough duckflat Checklist<br />
cb C Verrucaria maura O<br />
cb C Verrucaria mucosa O<br />
Ra St NR So Val al-sbr al-pil slough duckflat Checklist<br />
TOTALS:<br />
Species 47 35 7 7 24 112<br />
New Records 5 1 1 1 4 7<br />
Abbreviations:<br />
! = abundant or common<br />
# = currently being examined with molecular techniques<br />
= uncertainty <strong>of</strong> identification<br />
al-sbr=Alyeska boat ramp and vicinity<br />
al-pil=Alyeska small boat harbor pilings<br />
BC=British Columbia<br />
C=cryptogenic<br />
cb=circumboreal<br />
Checklist=total records for Port Valdez including literature and the present study<br />
duckflat=Mudflat east <strong>of</strong> the town <strong>of</strong> Valdez<br />
N=native to North Pacific<br />
NAT=North Atlantic<br />
nep= northeast Pacific<br />
np= North Pacific<br />
NR = new record to Alaska<br />
nr = northward range extension within Alaska<br />
O= records from the literature and pilot study<br />
slough=Slough about 1 mile from Alyeska gate<br />
So=Closest source to PWS<br />
Stat=NIS Status (native, cryptogenic, introduced, etc)<br />
un= living unattached<br />
Val=Total records for Port Valdez for the present study<br />
Wa=Washington<br />
ws= widespread, occurring in North Pacific, North Atlantic, and Australia or New Zealand<br />
X*= current record noted with specimen<br />
X= current record noted in the field
Chapt 9C1. Marine Plants, page 9C1- 7<br />
TABLE 9C1.4. Marine and Estuarine Plants Collected at Shore***<br />
Sites in Prince William Sound<br />
NIS ANALYSIS<br />
HARBORS MUD BAYS HEADLANDS RK<br />
TAXA<br />
AND REEFS BAYS<br />
Ra St NR So Val Cor Whit CB SMB Gro RP Bus Green NW<br />
RHODOPHYTA, Rhodophyceae<br />
ws C Ahnfeltia fastigiata X* X*!<br />
nep N Antithamnionella pacifica X* X* X*<br />
ws C Antithamnionella spirographidis X X<br />
ws C Audouinella purpurea X* X*<br />
ws C Bangia atropurpurea X X* X* X*<br />
np N Bossiella cretacea X* X*<br />
nep N Bossiella plumosa X*<br />
nep N Callithamnion acutum X*<br />
nep N Callithamnion pikeanum v. laxum X*<br />
nep N Callithamnion pikeanum v. pikeanum X*<br />
cb C# Ceramium cimbricum X* X*<br />
nep I# NR Cal Ceramium sinicola (on Codium ) X*<br />
nep N# Ceramium gardneri X*<br />
nep N# Ceramium pacificum/washingtonensis X* X* X* X* X* X*!<br />
ws C# Ceramium rubrum/kondoi X* X*<br />
np N Constantinea subulifera X* X X* X*! X*!<br />
np N Corallina frondescens X* X* X* X<br />
np,ch N Corallina <strong>of</strong>ficinalis v. chilensis X* X* X*!<br />
nep N Cryptosiphonia woodii X* X* X* X* X* X* X*! X*<br />
nep N Delesseria decipiens X<br />
cb C Devaleraea ramentacea X X* X*<br />
cb C Dumontia contorta X* X* X* X* X X*<br />
np N Dumontia simplex X<br />
nep N Endocladia muricata X* X*<br />
ws C Erythrotrichia carnea X* X* X* X*<br />
np N Glioipeltis furcata X* X X* X* X* X* X*! X*!<br />
np,ar N Halosaccion firmum X X* X X X<br />
np N Halosaccion glandiforme X* X X* X X* X X*<br />
np N Leachiella pacifica X* X* X* X*<br />
nep N Mastocarpus papillatus complex X* X*<br />
np N Mastocarpus cf. pacificus X* X X* X* X X*<br />
np N Mazzaella phyllocarpa X X* X* X*! X* X*! X!<br />
nep N Mazzaella splendens X<br />
nep N Microcladia borealis X<br />
np N Neorhodomela aculeata X X X* X* X* X*! X*! X!<br />
np N Neorhodomela larix X X* X* X*<br />
np N Neorhodomela oregona X* X*! X X* X* X X*! X*! X*!<br />
np N Neoptilota asplenioides X* X X* X! X<br />
nep N Odonthalia floccosa X! X X*! X* X*! X*!<br />
np N Odonthalia setacea (drift) X*<br />
nep N Opuntiella californica X*<br />
np N Palmaria calophylloides/stenogona X X X X** X*!<br />
nep N Palmaria hecatensis X* X*! X* X* X<br />
cb C Palmaria mollis/palmata X* X! X* X X* X* X!<br />
np N Phycodrys riggii X X* X* X*
Chapt 9C1. Marine Plants, page 9C1- 8<br />
Table 9C1.4. continued<br />
nep N Platythamnion pectinatum X X X*<br />
np N Pleonosporium cf vancouverianum X*<br />
nep N Polysiphonia eastwoodae X*<br />
ws C Polysiphonia brodiaei X<br />
nep N Polysiphonia hendryi v. deliquescens X X<br />
nep N Polysiphonia hendryi v. hendryi X* X*<br />
nep N Polysiphonia pacifica v. determinata X*<br />
nep N Polysiphonia pacifica v. pacifica X* X* X* X*<br />
np,nz N nr SeA Polysiphonia senticulosa X* X* X* X* X* X*<br />
cb C Polysiphonia stricta (urceolata) X* X* X*<br />
nep N Porphyra cuneiformis X* X* X<br />
cb C NR Com Porphyra miniata X*<br />
nep N Porphyra nereocystis X*<br />
nep N NR Wa Porphyra rediviva X*<br />
np N Porphyra torta/abbottae X*<br />
np N Pterosiphonia bipinnata X* X* X* X* X* X* X* X*!<br />
cb C Ptilota serrata (incl. pectinata ) X* X* X* X*<br />
np,ar N Rhodymenia pertusa X X* X X* X!<br />
cb C Scagelia americana X* X* X* X* X* X*<br />
nep N Smithora naiadum X*<br />
np N Tokidadendron kurilensis X X* X*! X*!<br />
nep N Weeksia coccinea X* X X*<br />
HETEROKONTOPHYTA, Phaeophyceae<br />
Val Cor Whit CB SMB Gro RP Bus Green NW<br />
np N# Alaria taeniata/angusta/crispa X* X X<br />
np N# Alaria tenuifolia/pylaii//membranacea X*<br />
np N# Alaria praelonga/marginata X X* X* X*!<br />
cb C Agarum clathratum (cribrosum) X X* X*<br />
np N Analipus japonicus X X* X* X* X* X<br />
cb C Chorda filum X* X* X* X<br />
cb C Chordaria flagelliformis Xun X*! X* X* X* X* X* X*<br />
cb C Coilodesme bulligera X*<br />
ak E NR Coilodesme n. sp. X*<br />
np,nz N Colpomenia bullosa X*<br />
ws C Colpomenia peregrina X* X* X* X*<br />
np N Costaria costata X* X*<br />
np N Cymanthera triplicata X*<br />
np N Cystoseira geminata X*<br />
cb C NR Com Delamarea attenuata X* X*<br />
cb C Desmarestia aculeata X X X X X* X<br />
cb C Desmarestia viridis X X X*<br />
cb C Dictyosiphon foeniculaceus X* X*! X*! X* X* X* X* X* X* X<br />
ws C Ectocarpus siliculosus X<br />
cb C Elachista fucicola X* X X X*<br />
nep N Elachista lubrica X* X* X*<br />
cb C Eudesme virescens X* X* X* X*<br />
cb I NR NAT Fucus cottonii X* X* X* X*<br />
cb C Fucus gardneri/distichus/evanescens X* X*! X*! X X X* un X* X X*! X*<br />
cb C Fucus spiralis X X* X X X X! X!<br />
np,ar N Laminaria "groenlandica"/bongardiana X* X* X* X* X* X* X* X*!<br />
cb C Laminaria saccharina X* X X*! X* X!<br />
np N Laminaria yezoensis X* X*! X*
Chapt 9C1. Marine Plants, page 9C1- 9<br />
Table 9C1.4. continued<br />
ws C Leathesia difformis X X X* X! X<br />
nep N Leathesia nana X* X* X* X*<br />
nep F Macrocystis integrifolia X* drift<br />
cb C Melanosiphon intestinalis X X*! X* X* X* X* X* X*<br />
ws C Pilayella littoralis/washingtonensis X* X* X* X* X* X* X X X*<br />
ws C nr SeA Punctaria latifolia X* X*<br />
ak E Punctaria lobata X*<br />
cb C NR Jap Punctaria plantaginea* X X* X X X*<br />
cb C Punctaria tenuissima X*<br />
cb C Ralfsia fungiformis X* X*<br />
np N Saundersella simplex X* X*<br />
ws C Scytosiphon simplicissimus X* X* X* X* X X* X* X*<br />
np N Soranthera ulvoidea X* X* X X* X X* X* X*<br />
np N Soranthera ulvoidea f. difformis X* X*<br />
cb C Sphacelaria racemosa X* X*<br />
ws C Sphacelaria rigidula X X* X* X*<br />
HETEROKONTOPHYTA, Xanthophyceae<br />
Val Cor Whit CB SMB Gro RP Bus Green NW<br />
ws C NR BC Vaucheria longicaulis () X* X*!<br />
CHLOROPHYTA, Chlorophyceae<br />
Val Cor Whit CB SMB Gro RP Bus Green NW<br />
cb C Acrosiphonia arcta X* X* X*! X X* X* X* X X*<br />
np N Acrosiphonia saxatilis X* X* X*<br />
ws C NR BC Blidingia marginata X*<br />
ws C Blidingia minima X* X* X* X* X X* X* X*<br />
cb C Blidingia subsalsa X* X* X*<br />
cb C NR BC Capsosiphon fulvescens X*<br />
ws C Cladophora albida X* X* X* X* X X* X*<br />
cb C Cladophora hutchinsiae X*<br />
ws C Cladophora sericea X X* X X* X* X* X* X* X*!<br />
np N Cladophora stimpsonii X* X*<br />
nep N nr SeA Codium fragile subsp. fragile X*<br />
ws I# NR Wa Codium fragile subsp. tomentosoides X*<br />
ws C Enteromorpha intestinalis X* X* X X X X X<br />
ws C Enteromorpha linza X*! X X* X* X<br />
ws C Enteromorpha prolifera/torta X* X X* X* X* X X*<br />
cb C Gayralia oxyspermum X* X X X*<br />
cb C NR BC Halochlorococcum moorei X* X*<br />
cb C Kornmannia zostericola (epiphytic) X*<br />
cb C NR NAT Kornmannia leptoderma (epilithic) X*<br />
nep N NR Wa Monostroma fractum X* X*<br />
cb C Monostroma grevillei/arcticum X X X* X*<br />
ws C Percursaria percursa X* X*<br />
ws C Rhizoclonium implexum X*<br />
ws C Rhizoclonium riparium X*<br />
ws C Rhizoclonium tortuosum X* X* X* X* X<br />
ws C Ulothrix implexa (non flacca ) X* X* X<br />
ws C# Ulva fenestrata /expansa/lactuca X* X* X X* X* X* X* X*<br />
cb C Ulvaria obscura X* X X* X X*<br />
np N Ulvella setchellii X*<br />
ws C Urospora penicilliformis X* X
Chapt 9C1. Marine Plants, page 9C1- 10<br />
Table 9C1.4. Continued<br />
SPERMATOPHYTA, Seagrasses<br />
Val Cor Whit CB SMB Gro RP Bus Green NW<br />
cb C Zostera marina X* X* X* X X* X X*<br />
nep N Phyllospadix scouleri X X*<br />
nep N Phyllospadix serrulatus X<br />
LICHENS<br />
Val Cor Whit CB SMB Gro RP Bus Green NW<br />
cb C Verrucaria maura X X X<br />
47 41 56<br />
TOTALS: Species (total =146)<br />
47 41 56 10 45 63 61 59 71 69<br />
New Records (total =17)<br />
5 5 2 2 5 4 2 2 6 1<br />
GROUP TOTALS (with overlap excluded):<br />
Harbors Mud Bays Headlands Rk Bays<br />
Species in Merged Groups 87<br />
78<br />
96<br />
69<br />
New Records in Merged Groups 8<br />
7<br />
8<br />
1<br />
Abbreviations:<br />
Site Abbreviations and Dates:<br />
! = abundant or common Bus=Busby Island south reef, 21 June<br />
# = currently being examined with molecular techniques CB=Cloudman Bay, East Bleigh I, 21 June<br />
***=shore sites include both shore and marina sites<br />
Cor=Cordova, 23 June<br />
= uncertainty <strong>of</strong> identification Green=Green Island, northwest point, 24 June<br />
ak=Alaska<br />
Gro=Growler Island, near resort, 27 June<br />
ar=arctic<br />
NW=Northwest Bay, Knight Island, 25 June<br />
BC=British Columbia<br />
RP=Rocky Point headland, 22 June<br />
C=cryptogenic<br />
SMB=Saw Mill Bay, 22 June<br />
Cal=California<br />
V-Ck=Port Valdez checklist (all records known)<br />
cb=circumboreal Val=all Port Valdez collections, June 1998<br />
Ch=Chile<br />
Whit=Whittier, 26 June<br />
Com=Commander Islands, Russia<br />
Cor=Cordova<br />
drift= dying unattached<br />
E=endemic to Alaska<br />
F=failed introduction<br />
I=possible introduction<br />
Jap=Japan<br />
N=native to North Pacific<br />
NAT=North Atlantic<br />
nep= northeast Pacific<br />
np=North Pacific<br />
NR = new record to Alaska<br />
nr = northward range extension within Alaska<br />
nz=New Zealand<br />
O=Presence known from the Pilot Study and literature<br />
Ra=Distribution Range<br />
RK=rocky<br />
SEA=Southeast Alaska<br />
So=Closest source to PWS<br />
St=range status (see N, C, E, and I)<br />
un= living unattached<br />
Wa=Washington<br />
ws= widespread, occurring in North Pacific, North Atlantic, and Australia or New Zealand<br />
X*=Presence known from the current study; specimens available<br />
X=Presence known from the current study; no specimen taken
Chapt 9C1. Marine Plants, page 9C1- 11<br />
TABLE 9C1.5. Marine Algae Collected from Off-Shore*** Floats<br />
in Prince William Sound, June 1998<br />
NIS ANALYSIS<br />
TAXA Float FLOATS<br />
Only<br />
Range Stat NR So Cklist TAT WBF MBF EIF SBF EBF on floats<br />
CYANOPHYTA, Cyanophyceae<br />
ws C Calothrix crustacea X X* **<br />
ws C Rivularia atra X X* **<br />
RHODOPHYTA, Rhodophyceae<br />
nep N Antithamnionella pacifica X X*<br />
cb I NR SD Chroodactylon ramosum X X* **<br />
nep N nr SeAk Polysiphonia senticulosa X X*<br />
ws C Polysiphonia urceolata X X* **<br />
cb C Scagelia americana X X*<br />
HETEROKONTOPHYTA, Phaeophyceae<br />
nep N Coilodesme californica X X* **<br />
ak E NR Coilodesme n. sp. X X*<br />
np N Cystoseira geminata X X*<br />
cb C NR Com Delamarea attenuata X X*<br />
nep N nr BC Ectocarpus acutus X X* **<br />
nep N nr BC Ectocarpus dimorpha X X* **<br />
nep N Ectocarpus parvus X X* **, V-CK<br />
Ectocarpus sp. (Acinetospora ) X*<br />
Giffordia sp. X*<br />
np N Laminaria groenlandica X X*<br />
ws C Laminaria saccharina X X* X*<br />
np N Laminaria yezoensis X X*<br />
cb C Melanosiphon intestinalis X X*<br />
cb I NR Jap Microspongium globosum X X* **<br />
ws C Pilayella littoralis X X*<br />
Pilayella sp. (elongate X*<br />
intercalary structures)<br />
cb C NR Jap Punctaria plantaginea X X*<br />
cb C nr SeAk Punctaria latifolia (Desmotrichum ) X X*<br />
ws C Scytosiphon simplicissima X X*<br />
CHLOROPHYTA, Chlorophyceae<br />
ws C Cladophora albida X X* X*<br />
ws C Cladophora sericea X X* X* X*<br />
ws C Enteromorpha prolifera/torta X X*<br />
ws C Percursaria percursa X X*<br />
TOTALS: 27 17 1 4 1 4 7 9<br />
New Records 9 4 1 1 0 0 3 4<br />
Abbreviations:<br />
*=Specimen available LJ=LaJolla, California Float Sites and Dates (Coord. with JC)<br />
**=Only on floats in this study N=native EBF=Eaglek Bay floats, 26 June<br />
***=Sites accessed by boat nep=northeast Pacific EIF=Ester Island float, 25 June<br />
BC=British Columbia np=North Pacific MBF=Main Bay barrier buoy, 25 June<br />
C=cryptogenic nr=new record from neighboring area SBF=Squaw Bay float, 26 June<br />
cb=circumboreal NR=new record from remote area TAT=Oyster floats near Tatilek, 21 & 22 June<br />
Com=Commander Islands, SeAk=Southeast Alaska WBF=Windy Bay floats, 23 June<br />
Jap=Japan V-Ck=also known from the Valdez Checklist Float Cklist=species used in this study<br />
ws=widespread
Chapt 9C1. Marine Plants, page 9C1- 12<br />
Species were then categorized as to whether they were new distribution records (NR) to the area.<br />
These included species that had never before been reported from Prince William Sound (nr) or<br />
Alaska (NR). In each <strong>of</strong> these cases, the closest known records to the area were given as the<br />
source (So). Since the new records seemed to be the most likely category in which to find<br />
recognizable NIS, they are highlighted in gray throughout the charts and tables.<br />
This preliminary quantification <strong>of</strong> marine plant species and NIS in Prince William Sound<br />
required a number <strong>of</strong> lengthy, detailed steps. After gathering, identifying, and curating all <strong>of</strong> the<br />
species, site lists had to be prepared and both local and global biogeographic information<br />
compiled. Then, with this information in hand, the residency status and new distribution records<br />
<strong>of</strong> each species were determined. Only after all <strong>of</strong> this was completed could NIS begin to be<br />
recognized. Since many <strong>of</strong> the steps in this process revealed important data that characterized not<br />
only NIS but the marine flora in general, this report presents the site lists in their entirety and<br />
then summarizes the results for comparative purposes in tables and graphs. Since the results are<br />
lengthy, they have been organized into the following 7 major parts that are presented below:<br />
• The Species Lists by Site, including Port Valdez, Shore, and Floats.<br />
• The Total Species Numbers and Composition <strong>of</strong> the Individual Sites.<br />
• Total Species Numbers and Composition in each Habitat Type.<br />
• Native, Cryptogenic, and Introduced Species and their Taxonomic Composition.<br />
• Native, Cryptogenic, and Introduced Species in the Habitat Types.<br />
• New Species Records and Probable Introductions.<br />
• Comments on the Five Probable Introductions and One Important Failed Introduction.<br />
Results<br />
During our 9-day search for NIS in Port Valdez and Prince William Sound, 489 plant<br />
samples were processed (Table 9C1.6). These samples contained 155 different species<br />
dominated by the red (Rhodophyceae), brown (Phaeophyceae), and green (Chlorophyceae)<br />
algae, in that order. Among these species, 21 were found to be new records to the area, and, <strong>of</strong><br />
these, at least 5 appear to be introduced. In addition, 70 species were found to be cryptogenic,<br />
some <strong>of</strong> which have suspicious characteristics <strong>of</strong> NIS.<br />
TABLE 9C1.6. Collection Data*<br />
TAXONOMIC SAMPLES<br />
TOTAL NEW<br />
GROUP Total Herb. Form. SPECIES RECORDS<br />
Rhodophyceae 199 135 64 69 5<br />
Phaeophyceae 162 120 42 49 8<br />
Chlorophyceae 117 99 18 30 7<br />
Xanthophyceae 2 2 0 1 1<br />
Seagrasses 7 4 3 3 0<br />
Lichens 0 0 0 1 0<br />
Cyanophyceae 2 1 1 2 0<br />
Total, June 1998 489 361 128 155 21<br />
Abbreviations:<br />
* =Samples and species counts are only for the June 1998 trip.<br />
Herb.=Pressed herbarium sheets<br />
Form.=Bottles <strong>of</strong> preserved specimens
Chapt 9C1. Marine Plants, page 9C1- 13<br />
The total species and new species records/collecting site were correlated with collection<br />
time (Fig. 9C1.1). Longer collecting periods yielded more species at an R 2 value <strong>of</strong> 43%. New<br />
records, on the other hand, appear to be almost unaffected by collection time, showing an R 2 <strong>of</strong><br />
only 7%.<br />
FIG. 9C1.1. Collection Efficiency<br />
80<br />
180<br />
Total Species<br />
70<br />
60<br />
50<br />
40<br />
30<br />
20<br />
Val<br />
al-sbr<br />
al-pil<br />
slough<br />
duckflat<br />
Cor<br />
Whit<br />
10<br />
0<br />
Major Collection Sites<br />
CB<br />
SMB<br />
Gro<br />
RP<br />
Bus<br />
Green<br />
NW<br />
New Records<br />
Total Species<br />
Collection Time<br />
160<br />
140<br />
120<br />
100<br />
80<br />
60<br />
40<br />
20<br />
0<br />
Collection Time (minutes)<br />
The Site Species Lists. Species <strong>of</strong> marine and estuarine plants identified at all sites sampled<br />
during our June 1998 survey are listed in Tables 9C1.3, 9C1.4, and 9C1.5. The plants sampled<br />
were predominantly macrobenthic marine algae <strong>of</strong> the Rhodophyceae, Phaeophyceae,<br />
Chlorophyceae, Xanthophyceae, and Cyanophyceae along with several species <strong>of</strong> seagrasses and<br />
marine lichens. Species occurrence at the various sites is designated with an X in the lists. If<br />
samples were taken and curated for identification purposes, the species are listed with an X*,<br />
indicating that vouchers are available in the OSU/HMSC herbarium for study. In addition, each<br />
species is categorized for several biogeographic features that are necessary for the NIS Analysis,<br />
explained in the Methods section above.<br />
• Species <strong>of</strong> Port Valdez. Samples from 4 sampling sites in Port Valdez included 47 algal<br />
species and 5 new records (Table 9C1.3). The sites covering the largest areas (the Alyeska<br />
small boat ramp and the duckflat) contained the majority <strong>of</strong> the species. The highest species<br />
count occurred at the Alyeska boat ramp where the greatest amount <strong>of</strong> hard substratum was<br />
available for algal settlement. The highest number <strong>of</strong> new records occurred in the duckflat.<br />
A few <strong>of</strong> these species are good candidates for NIS status. However, their lack <strong>of</strong> earlier<br />
discovery may have an obvious explanation. Mudflats, like the duckflat, are not only<br />
notoriously poor habitats for most marine algae, but they can be dangerous in Alaska.<br />
Therefore, earlier phycologists avoided many <strong>of</strong> these areas. Knowing this to be true, it was<br />
possible to predict the occurrence <strong>of</strong> some new records (e.g., Fucus cottonii and Vaucheria<br />
longicaulis) in the mudflats and sloughs. Also shown in Table 9C1.3 is a Checklist <strong>of</strong> Algal<br />
Species for Port Valdez, which includes the species collected from the 1998 sampling, as
Chapt 9C1. Marine Plants, page 9C1- 14<br />
well as those found previously during the 1997 Pilot Study (Ruiz & Hines, 1997) and the<br />
literature (Calvin and Lindstrom, 1980, and Weigers et al., 1997). In addition to our summer<br />
sampling, this list includes year-round collections taken by the earlier investigators. The<br />
total count for the entire Port Valdez area, including these earlier records, amounts to 112<br />
species.<br />
• Species <strong>of</strong> Shore Sites. Shore sites include all intertidal areas and marinas sampled during<br />
the June 1998 cruise, along with the combined records for Port Valdez (Table 9C1.4). These<br />
10 sites covered a wide range <strong>of</strong> habitats, which were grouped into 4 major habitat types:<br />
Harbors, Mud Bays, Headlands and Reefs, and Rocky Bay (presented in more detail below).<br />
The overall species count for all <strong>of</strong> the shore areas was 146 species with 17 new records.<br />
The highest species diversity occurred at Green Island, Northwest Bay, and Growler Island,<br />
all <strong>of</strong> different habitat types; while the highest number <strong>of</strong> new records occurred at Green<br />
Island, Saw Mill Bay, Cordova, and Port Valdez, also a mixture <strong>of</strong> habitat types. Only two<br />
species (Dictyosiphon foeniculaceus and Fucus gardneri ) were found at all shore sites<br />
sampled. Four others (Cladophora sericea, Acrosiphonia arcta, Pilayella littoralis, and<br />
Neorhodomela oregona) were found at all but one site (and were possibly overlooked there).<br />
Numerous species (31) were common to all <strong>of</strong> the habitat types, but there were also an<br />
extraordinary number <strong>of</strong> species that appeared to be limited to only 1 habitat type (17 were<br />
found only in harbors, 9 only in mud bays, 19 only on headlands and reefs, and 5 only in<br />
rocky bays). Two species restricted to harbors are new records to the area: Porphyra<br />
rediviva, a newly discovered free-floating marsh plant that could be easily transported by<br />
ships, and Vaucheria longicaulis , a species unique to high mudflats that is common to many<br />
southern west coast harbors. Although not a new record, another interesting harbor species is<br />
Antithamnionella spirographidis. This species is reported to be common to harbors in British<br />
Columbia and is thought to be introduced to that area (Lindstrom in DeWreede 1996).<br />
However, its circumboreal and Australian existence leads me to categorize it as cryptogenic<br />
in this paper.<br />
• Species from Floats. Marine plants were sampled from five oyster floats and one barrier buoy<br />
(MBF) (Table 9C1.5). A total <strong>of</strong> 27 different algal species were identified from the floats.<br />
Of these, 9 were not collected at any <strong>of</strong> the other sites during our trip. Most <strong>of</strong> these unique<br />
species are small and could have been overlooked in other areas, but several are species that<br />
probably could only find suitable habitat on the floats. Over half the 27 species collected are<br />
well-known fouling organisms (e.g., Cladophora sericea, Pilayella littoralis, and<br />
Polysiphonia urceolata). Nine new species records, the highest habitat number in our<br />
survey, were also found on the floats. This may be related to the fact that most <strong>of</strong> the floats<br />
sampled are used in aquaculture, which could be a source <strong>of</strong> introductions. The highest<br />
counts for both species and new records occurred on the floats at Tatitlek Narrows (TAT)<br />
and Eaglek Bay (EBF) used in active oyster culture. Two <strong>of</strong> the new records found at these<br />
sites (Chroodactylon ramosum and Microspongium globosum) and possibly more are thought<br />
to be introduced. One species (Polysiphonia senticulosa) is considered to be a range<br />
extension from southeast Alaska, but it is already widespread in Prince William Sound. This<br />
species is presumably native in Washington to Southeast Alaska, and was recently reported<br />
to be introduced and pervasive in New Zealand (Nelson and Maggs, 1996).
Chapt 9C1. Marine Plants, page 9C1- 15<br />
The Total Number <strong>of</strong> Species and Species Composition <strong>of</strong> the Individual Sites. The overall<br />
number <strong>of</strong> species was 155 for all sites, but the numbers <strong>of</strong> species per site ranged from only 10<br />
to 71 species, indicating that there is considerable variation in species composition among<br />
habitats (Table 9C1.7, Fig. 9C1.1, 9C1.2). The 21 new records across all areas ranged from 1 to<br />
9 at the individual sites and was highest on floats. The highest species count (71 species)<br />
occurred at Green Island, the most exposed and highly saline site. At this site, the proportion <strong>of</strong><br />
red algal species was nearly 2 times that <strong>of</strong> the brown algae and 4 times that <strong>of</strong> the greens. The<br />
lowest species count (10 species) occurred at Cloudman Bay, a sheltered, estuarine mud bay. At<br />
this site there were almost no red algae, and the brown algae were more abundant than the<br />
greens.<br />
Table 9C1.7. Total Species Numbers and Composition at Each Site, June 1998<br />
Taxonomic Group Val Cor Whit CB SMB Gro RP Bus Green NW Floats Total* V-Total**<br />
Rhodophyceae 18 12 23 1 15 26 27 27 40 37 5 69 84<br />
Phaeophyceae 13 11 14 5 20 24 23 21 21 22 16 49 52<br />
Chlorophyceae 14 16 17 3 9 12 9 8 10 9 4 30 36<br />
Xanthophyceae 1 1 0 0 0 0 0 0 0 0 0 1 1<br />
Seagrasses 1 1 1 1 1 1 1 3 0 0 0 3 3<br />
Lichens 0 0 1 0 0 0 1 0 0 1 0 1 2<br />
Cyanophyceae 0 0 0 0 0 0 0 0 0 0 2 2 2<br />
TOTALS 47 41 56 10 45 63 61 59 71 69 27 155 180<br />
NEW RECORDS 5 5 2 2 5 4 2 2 6 1 9 21 23<br />
Abbreviations: Sites as in Table 1; Val=the Port Valdez collections combined; Floats=the float collections combined;<br />
Total*=with overlap and earlier collections excluded; V-Total**=Total* with the Port Valdez Checklist species included.<br />
Figure 9C1.2. Total Species Numbers and<br />
Composition at each Site, June 1998<br />
80<br />
70<br />
60<br />
Total Species<br />
50<br />
40<br />
30<br />
20<br />
10<br />
0<br />
Val Cor Whit CB SMB Gro RP Bus Green NW Floats<br />
Collecting Sites<br />
Rhodophyceae Phaeophyceae Chlorophyceae<br />
Xanthophyceae Seagrasses Lichens<br />
Cyanophyceae<br />
Total Species Numbers and Composition in each Habitat Type. Since several sampling sites<br />
had mixed habitats, categorizing the sites into distinct habitat types had some weaknesses.<br />
However, it increased the number <strong>of</strong> species sampled for each category <strong>of</strong> habitat, providing<br />
more power to the data analysis (Table 9C1.8, Fig. 9C1.3a, b).
Chapt 9C1. Marine Plants, page 9C1- 16<br />
TABLE 9C1.8. Habitat Type and Species Composition<br />
Taxonomic Group Valdez Harbors Mud Bays Rk Bays Headlands Floats Total<br />
Rhodophyceae 18 37 31 37 48 5 69<br />
Phaeophyceae 13 22 31 22 30 16 49<br />
Chlorophyceae 14 25 15 9 14 4 30<br />
Xanthophyceae 1 1 0 0 0 0 1<br />
Seagrasses 1 1 1 0 3 0 3<br />
Lichens 0 1 0 1 1 0 1<br />
Cyanophyceae 0 0 0 0 0 2 2<br />
Total 47 87 78 69 96 27 155<br />
Fig 3a. Habitat Type and<br />
Species Composition<br />
120<br />
100<br />
Total Species<br />
80<br />
60<br />
40<br />
20<br />
0<br />
Valdez<br />
Harbors<br />
Mud Bays<br />
Rk Bays<br />
Headlands<br />
Floats<br />
Habitat Type<br />
Rhodophyceae Phaeophyceae Chlorophyceae<br />
Xanthophyceae Seagrasses Lichens<br />
Cyanophyceae<br />
Fig. 3b. Habitat Type and<br />
Percentage Composition<br />
100%<br />
80%<br />
Percentage Composition<br />
60%<br />
40%<br />
20%<br />
0%<br />
Valdez Harbors Mud Bays Rk Bays Headlands Floats<br />
Habitat Type<br />
Rhodophyceae Phaeophyceae Chlorophyceae<br />
Xanthophyceae Seagrasses Lichens<br />
Cyanophyceae
Chapt 9C1. Marine Plants, page 9C1- 17<br />
The 5 habitat types with their features and included sites are:<br />
• Harbors* (Val, Cor, and Whit). Sheltered areas with variable salinities (0-28 ppt) and<br />
variable substrates including mud, cobble, and wood (the pilings). Heavily influenced by<br />
boat traffic and other human activities. [* Note that Port Valdez (Val), also included in the<br />
Harbor group, is included separately in several <strong>of</strong> the tables to show the comparable diversity<br />
<strong>of</strong> this targeted site.]<br />
• Mud Bays (CB, SMB, Gro). Sheltered bays with salinity ranges from 3-11 ppt with a<br />
substratum <strong>of</strong> primarily mud, although cobble and bedrock was <strong>of</strong>ten available.<br />
• Rocky Bays (RB). One semi-sheltered bay with a salinity ranging from 10-27 ppt and a<br />
substratum varying from gravel to cobble to bedrock.<br />
• Headlands and Reefs (RP, Bus, Green). Very exposed habitats with salinity ranges from 15-<br />
30 ppt and a substratum consisting almost totally <strong>of</strong> bedrock and cobble.<br />
• Floats (TAT, WBF, MBF, EIF, SBF, EBF). Exposed to semi-sheltered <strong>of</strong>f-shore habitats.<br />
Salinities ranged from 16-28 ppt and substrata included 5 plastic oyster floats and line and 1<br />
cement buoy (MBF).<br />
Headlands and reefs with high exposure and high salinity had the greatest species<br />
diversity. As would be expected for temperate zones, they are dominated in descending order by<br />
red, brown, and green algae. Surprisingly, the next largest diversity <strong>of</strong> species occurred in the<br />
harbors. Although the red algae also predominated there, harbors had a large number <strong>of</strong> green<br />
algae. In the mud bays and on <strong>of</strong>f-shore floats, there was a tendency for increase in the<br />
percentage <strong>of</strong> brown algae. However, since total species number varies among habitat types,<br />
composition <strong>of</strong> algal groups may be partly an artifact <strong>of</strong> small number <strong>of</strong> species at the low<br />
diversity sites.<br />
The numbers <strong>of</strong> new records among the various habitat types were fairly uniform except<br />
in rocky bays where the sample size (1 bay) was small (Table 9C1.9). The slight increase in<br />
numbers on floats may be significant, but overall, the data indicate that habitat type has little to<br />
do with the discovery <strong>of</strong> new species records.<br />
TABLE 9C1.9. Habitat Type and New Species Records<br />
Valdez Harbors Mud Bays Headlands Rk Bays Floats (pvc/ Totals*<br />
TAXONOMIC (mixed) (mixed) (mud/cob) and Reefs (gravel/cob) concrete)<br />
GROUP SP NR SP NR SP NR SP NR SP NR SP NR SP NR %<br />
Rhodophyceae 18 1 37 2 31 1 48 2 37 1 5 2 69 5 7<br />
Phaeophyceae 13 1 22 2 31 4 30 4 22 0 16 7 49 8 16<br />
Chlorophyceae 14 2 25 3 15 2 14 2 9 0 4 0 30 7 23<br />
Xanthophyceae 1 1 1 1 0 0 0 0 0 0 0 0 1 1<br />
Seagrasses 1 0 1 0 1 0 3 0 0 0 0 0 3 0<br />
Lichens 0 0 1 0 0 0 1 0 1 0 0 0 1 0<br />
Cyanophyceae 0 0 0 0 0 0 0 0 0 0 2 0 2 0<br />
Total 47 5 87 8 78 7 96 8 69 1 27 9 155 21<br />
% Habitat Total 11 9 9 8 1 33 14<br />
% NR Total (21) 24 38 33 38 5 43 100<br />
* = excludes overlapping records<br />
cob= cobble
Chapt 9C1. Marine Plants, page 9C1- 18<br />
Native, Cryptogenic, and Introduced Species and their Taxonomic Composition. Of the 155<br />
total algal species found, 52% are native, 45% are cryptogenic, and 3% (5 species) appear to be<br />
introduced (Table 9C1.10, Fig. 9C1.4). The taxonomic composition <strong>of</strong> these groups parallels the<br />
findings in the Pilot Study survey for Port Valdez. The native species contain a very large<br />
percentage <strong>of</strong> red algae, about 64% <strong>of</strong> the total. The brown algae make up 27% <strong>of</strong> the natives,<br />
and the greens only 6%. The composition <strong>of</strong> the cryptogenic forms is almost the reverse. The<br />
red algae are only about 23% <strong>of</strong> the total count, while the browns and the greens both average<br />
about 35%.<br />
Table 9C1.10. Native, Cryptogenic, and Introduced<br />
Species and their Taxonomic Composition<br />
Taxonomic<br />
Group Total N<br />
Status**<br />
C I NR<br />
Rhodophyceae 69 51 16 2 5<br />
Phaeophyceae 49 22 25 2 8<br />
Chlorophyceae 30 5 24 1 7<br />
Xanthophyceae 1 0 1 0 1<br />
Seagrasses 3 2 1 0 0<br />
Lichens 1 0 1 0 0<br />
Cyanophyceae 2 0 2 0 0<br />
Totals 155 80 70 5 21<br />
% <strong>of</strong> Total 100 52 45 3 14<br />
NR 21 7 9 5<br />
** = For simplification, 2 endemics and 1 failed introduction are included with the natives.<br />
N=native, C=cryptogenic, I=potentially introduced, NR=new records<br />
Fig. 9C1.4. Residency Status and<br />
Taxonomic Composition<br />
180<br />
160<br />
Total Species<br />
140<br />
120<br />
100<br />
80<br />
60<br />
40<br />
Cyanophyceae<br />
Lichens<br />
Seagrasses<br />
Xanthophyceae<br />
Chlorophyceae<br />
Phaeophyceae<br />
Rhodophyceae<br />
20<br />
0<br />
Total N C I NR<br />
Status<br />
Of the 21 new species records across all habitats, 5 were red algae, 8 brown, 7 green, and<br />
1 was a Xanthophyte (Table 9C1.9), reflecting a fairly uniform distribution <strong>of</strong> new records across<br />
at least the 3 major taxonomic groups. However, the percentage <strong>of</strong> new species records by group<br />
increased dramatically from red to brown to green algae. It is possible that this increase relates,<br />
in part, to our overall level <strong>of</strong> taxonomic understanding in each <strong>of</strong> these 3 major classes. Since<br />
stable morphological features usable in taxonomy decrease as one moves from the red to the<br />
brown to the green algae, ease <strong>of</strong> accurate identification likewise decreases. Hence, it is likely
Chapt 9C1. Marine Plants, page 9C1- 19<br />
that our knowledge <strong>of</strong> the Alaskan flora is correspondingly most complete for the reds, then the<br />
browns, and lastly the greens.<br />
Native, Cryptogenic and Introduced Species by Habitat Types. In all habitat types except the<br />
floats, the native and cryptogenic species were fairly evenly distributed and ranged from 44 to<br />
55% <strong>of</strong> the species (Table 9C1.11, Fig. 9C1.5a, b). However, there were some predictable<br />
reversals <strong>of</strong> dominance. In the harbors and mud bays and on the floats, the cryptogenic species<br />
were the most abundant, while on the headlands and reefs and in the rocky bays, the native<br />
species predominated. This reversal reflects the confounding effect <strong>of</strong> variation in groups among<br />
habitats (Figs. 9C1.3a, b). Since red algae predominated on the reefs and rocky bays (and green<br />
algae are relatively low in numbers), native species, consisting mostly <strong>of</strong> red algae, were also<br />
predominant there. On the other hand, in harbors and mud bays red algae were not as common<br />
(and green algae are more abundant); hence the numbers <strong>of</strong> native species were lower in those<br />
habitats. The reefs and rocky bays were under-collected in most cases, weakening the<br />
conclusions about algal species in these areas. On floats the cryptogenic forms consisted <strong>of</strong><br />
55% <strong>of</strong> the species while the native species consisted <strong>of</strong> only 37%. Since most <strong>of</strong> the floats<br />
sampled were from oyster farms, it is likely that they are periodically cleaned. Each cleaning <strong>of</strong><br />
the floats would provide cleared primary substrata for ephemeral (opportunistic) species that are<br />
quick to colonize and reproduce. Since ephemeral species are most <strong>of</strong>ten cryptogenic, their<br />
higher percentage may be understandable.<br />
Table 9C1.11. The Native, Cryptogenic, and Introduced<br />
Species Occurring in Each Major Habitat<br />
Residency<br />
Status Harbors<br />
Habitat Types<br />
Mud Bays Headlands Rk Bays Floats Totals<br />
Native* 38 35 52 36 10 80<br />
Cryptogenic 48 42 42 33 15 70<br />
Introduced 1 1 2 0 2 5<br />
Totals 87 78 96 69 27 155<br />
New Records 8 7 8 1 9 21<br />
*2 endemics and 1 failed introduction (under rk bay browns) are included in natives.<br />
Total Species<br />
Fig 9C1.5a. Species Residency Types<br />
Occurring in Each Major Habitat<br />
120<br />
100<br />
80<br />
60<br />
40<br />
20<br />
0<br />
% <strong>of</strong> Habitat<br />
Fig 9C1.5b. Major Habitats and Residency<br />
Type Composition<br />
100%<br />
80%<br />
60%<br />
40%<br />
20%<br />
Harbors<br />
Mud Bays<br />
Habitat Type<br />
Headlands<br />
Rk Bays<br />
Floats<br />
Introduced<br />
Cryptogenic<br />
Native*<br />
0%<br />
Harbors<br />
Mud Bays<br />
Habitat Types<br />
Headlands<br />
Rk Bays<br />
Floats<br />
Introduced<br />
Cryptogenic<br />
Native*
Chapt 9C1. Marine Plants, page 9C1- 20<br />
New Species Records and Probable Introductions. The 21 new species records for the June<br />
1998 trip are the most likely candidates to be NIS; however, several factors should be considered<br />
further before the status <strong>of</strong> the species can be determined definitively (Table 9C1.12). Since all<br />
<strong>of</strong> the new records appeared to have been overlooked for at least some period <strong>of</strong> time, the<br />
obvious questions related to why this oversight occurred are:<br />
TABLE 9C1.12. New Records <strong>of</strong> Benthic Marine Algae to Prince William Sound<br />
(species overlooked, misidentifications, range extensions, and possible introductions)<br />
Justification Stat Taxon Location Type Ra So C Comments<br />
Rhodophyta, Rhodophyceae<br />
gi I Ceramium sinicola Green Exp nep Cal 3 epiphyte, MB in progress<br />
gi, sol I Chroodactylon ramosum TAT F cb S. Cal. 1 microscopic<br />
rex N Polysiphonia senticulosa RP, Cor, Whit, Bus, All np,nz SeAk 1 easy to recognize<br />
Green, NW, TAT ex M invasive<br />
mid C Porphyra miniata Gro M cb Com. 2 MB in progress<br />
mid N Porphyra redidiva Val (duckflat) M nep Wa 1 recently described<br />
Heterokontophyta, Phaeophyceae<br />
mid E Coilodesme n. sp. Green, TAT Exp, F ak 1 epiphyte, morph. needed<br />
mid C Delamaraea attenuata SMB, Bus, EBF Exp, F cb Com 1 recently illustrated<br />
rex N Ectocarpus acutus MBF, EBF F nep BC 1<br />
rex N Ectocarpus dimorphus EBF F nep BC 1<br />
gi I Fucus cottonii Val (slough), SMB, M cb N. Atl 1 common in marshes<br />
CB, Gro<br />
MB in progress<br />
gi, sol I Microspongium globosum EBF F cb Jap, N. Atl 1 microscopic<br />
rex C Punctaria latifolia TAT, SMB, RP Exp, F cb SeAk 1<br />
mid C Punctaria plantaginea SMB, Cor, Whit, All cb N. Atl. 1 some think cold water<br />
Green, Gro ex F form <strong>of</strong> latifolia<br />
Heterokontophyta, Xanthophyceae<br />
reproductive material<br />
sol, rex C Vaucheria longicaulis Val, Cor M, H ws BC 3 needed to confirm sp.<br />
Chlorophyta, Chlorophyceae<br />
rex C Blidingia marginata Val M ws BC 1<br />
sol, rex C Capsosiphon fulvescens CB M cb BC 1 microscopic<br />
rex N Codium fragile* (NE Pacific form) Green Exp nep SeAk 1 epi=C. sinicola <br />
gi I Codium fragile<br />
subsp. tomentosoides Green Exp ws Wa. 2 MB needed for subsp.<br />
sol, rex C Halochlorococcum moorei Val, Cor H cb BC 1 microscopic, endophytic<br />
mid, sol C Kornmannia leptoderma non<br />
zostericola (epilithic) Cor H cb N. Atl. 2 Culture work needed<br />
mid, sol N Monostroma fractum Gro M nep Wa. 2 Culture work needed<br />
Abbreviations: (see earlier charts)<br />
* = Recently also reported in O’Clair et al. , 1996, morph.=morphological studies<br />
from my earlier EVOS collections<br />
rex=range extension<br />
Category=preliminary decisions based on<br />
sol=species overlooked<br />
morphological and distributional features<br />
Stat=residency status<br />
and the literature available<br />
Type=habitat type<br />
ex=except<br />
Exp=exposed cobble<br />
C=Certainty <strong>of</strong> Idenfication<br />
F=on floats<br />
1=absolute certainty<br />
H=harbor, on cobble<br />
2=Morphological identity but additional<br />
gi=geographic isolation study (eg, MB or cultures) needed<br />
I=likely introduction<br />
3=Vegetative morphology similar, but reproductive<br />
ID=identification or MB data needed for positive identification<br />
M=mud<br />
MB=molecular biological study<br />
mid=earlier misidentification
Chapt 9C1. Marine Plants, page 9C1- 21<br />
• Are any <strong>of</strong> the species taxonomically problematic Such problematic species <strong>of</strong>ten end up in<br />
new records lists; and, indeed, several <strong>of</strong> the new records are problematic species that require<br />
further study for positive identification. Investigations are currently in progress for 5 <strong>of</strong> these<br />
species.<br />
• Could the species have been mistaken for other similar species in the past Species that<br />
resemble one another can be confused easily. Often these mistakes are not revealed until a<br />
species is newly illustrated or described. In these cases, misidentifications and distributions<br />
could easily be corrected with herbarium searches. In the list, at least 6 species fall into this<br />
category, including at least 1 undescribed species.<br />
• Has small size or habitat restriction influenced the species discovery Microscopic species<br />
are frequently overlooked as are species from unusual habitats. On the list, 4 species are<br />
microscopic, and 1 occurs in the unlikely habitat <strong>of</strong> a high marsh.<br />
• Is the species new to the area through range extension or through an actual introduction For<br />
marine plants, historical (baseline and fossil) information, geographic isolation, and<br />
molecular data are appropriate for proving the latter.<br />
The final justification for categorizing a species <strong>of</strong> marine plant as introduced (Table 9C1.12)<br />
was based on many <strong>of</strong> these factors, but remains tentative. All 5 species listed as introduced are<br />
geographically isolated. Nine other species are northward range extensions from southeast<br />
Alaska or British Columbia, and these species are tentatively identified as native. However,<br />
these range extensions could be caused by either natural dispersal, possibly caused by El Niño<br />
events <strong>of</strong> the past few years, or they could be introduced with aquaculture transports.<br />
Of the 21 new records, at least 5 are at this time very strongly supported for NIS status<br />
based on their geographic isolation, and this is a very conservative estimate. To further confirm<br />
the status <strong>of</strong> these, molecular biological pro<strong>of</strong> <strong>of</strong> identifications are currently in progress.<br />
Comments on the 5 Probable Introductions and on 1 Important Failed Introduction.<br />
Additional description <strong>of</strong> each <strong>of</strong> the most probable introduced species and their habitats and<br />
distribution are provided below:<br />
Chroodactylon ramosum. Chroodactylon is a microscopic primitive red alga that is typically<br />
bright bluegreen in color. Its uniseriate, dichotomously branched filaments are unmistakable<br />
under the microscope. Although common to the North Atlantic in both Europe and North<br />
America, in the Pacific it is only known from Japan, southern Australia, and southern California.<br />
Because this alga generally occurs in estuarine or freshwater habitats (Vis and Sheath 1993), its<br />
occurrence in the turf algae <strong>of</strong> the oyster floats at Tatitlek was a surprise, except that it could<br />
have been brought in with oysters. The lack <strong>of</strong> records for this species in the well-worked<br />
marine and estuarine environments <strong>of</strong> British Columbia and Washington indicates that it is truly<br />
an isolated population and in all likelihood introduced.<br />
Codium fragile subsp. tomentosoides and the northeast Pacific complex*. The normal range <strong>of</strong><br />
the native species complex <strong>of</strong> Codium fragile in the northeast Pacific is from Baja California to<br />
southeast Alaska. This complex appears to consist <strong>of</strong> several unnamed subspecies (C.<br />
Trowbridge, pers. comm.). Separate from this is an alien subspecies called tomentosoides that<br />
has been reported to occur in San Francisco Bay. This alien subspecies is differentiated from the
Chapt 9C1. Marine Plants, page 9C1- 22<br />
native complex by having a different branching frequency and more rounded and mucronate<br />
utricle tips. At Green Island, two different subspecies <strong>of</strong> Codium appear to occur. The low<br />
intertidal form appears to be identical to the native complex. Its utricle tips are sharply pointed<br />
and it is fairly tightly branched. The second form occurs in the mid to upper intertidal and is<br />
more loosely branched with very short mucronate tips, nearly identical to subsp. tomentosoides.<br />
However, experts in the field (Silva and Max) have told me after considerable hesitancy, that<br />
neither are truly subsp. tomentosoides, and that both fall clearly within the native complex. This<br />
indicates that the Green Island Codium is probably a range extension or an introduction from<br />
southeast Alaska or from Washington, Oregon, or California. Perhaps studies on its epiphyte<br />
(discussed below) will enable us to detect its true source. (* Recently O’Clair et al., 1996, also<br />
noted that Codium occurs on Green Island. This record appears to be based on G.I. Hansen’s<br />
earlier EVOS project collections, now located in Juneau.)<br />
Ceramium sinicola. This Ceramium species was an epiphyte <strong>of</strong> Codium fragile at Green Island.<br />
The species, unlike Ceramium codicola, does not have bulbous rhizoids. It is completely<br />
corticated except for some slightly broken cortication near the tips like in the southern California<br />
species Ceramium sinicola. The morphology <strong>of</strong> the plants most closely matches the descriptions<br />
<strong>of</strong> Dawson (1950) and Setchell and Gardner (1924) for C. sinicola, and male, female, and<br />
tetrasporic specimens have all been observed. Its occurrence in Alaska is extremely unlikely<br />
unless it is a recent introduction. This past year I have been working with Mr. Tae Oh Cho, who<br />
is monographing the world species <strong>of</strong> Ceramium with both morphological and molecular<br />
techniques. He has agreed to look at this material for me and will also look at my Alaskan<br />
material <strong>of</strong> Ceramium rubrum which he feels may actually be C. kondoi, a Japanese/Korean<br />
species, and possibly another introduction to Alaska.<br />
Fucus cottonii. This species is unrecorded for the North Pacific, and yet it occurred in nearly all<br />
<strong>of</strong> the high mudflat/marsh areas visited during the June 1998 cruise <strong>of</strong> Prince William Sound.<br />
The plant was first observed in unpublished notes by G.I. Hansen on Vancouver Island in 1981<br />
and then at several Prince William Sound and Kenai sites during the EVOS studies. In some<br />
areas it dominated the supralittoral zone extending even into the terrestrial. At Cloudman Bay it<br />
occurred 100 meters away from the bay on stream banks intermixed with mosses and vascular<br />
plants. The plants in Alaska are mat-forming and either loose-lying on mud, entangled with<br />
other algae (such as Fucus gardneri), or intermixed with terrestrial plants. They range from 1-5<br />
cm in height. The blades are dichotomously branched and <strong>of</strong>ten terete and only 1-3 mm in<br />
diameter. In some habitats, they become flattened, still without a visible midrib, and up to 5 mm<br />
in diameter. No receptacles were found during the June trip, but during the EVOS studies G.I.<br />
Hansen found a number <strong>of</strong> plants with relatively small (up to 2 cm long), elongate, somewhat<br />
pointed receptacles with conceptacles and oogonia bearing 8 eggs. In some areas, the extent <strong>of</strong><br />
the mats <strong>of</strong> this small fucoid makes me question its form <strong>of</strong> reproduction. Since receptacles are<br />
so uncommon, propagation <strong>of</strong> the mats must be by fragmentation and vegetative growth, an<br />
advantageous feature for dispersal.<br />
There is some question as to the use <strong>of</strong> Fucus cottonii (= F. muscoides) as a valid species.<br />
Fletcher (1987) considers the species as a high marsh ecad <strong>of</strong> Fucus vesiculosis, an Atlantic and<br />
Arctic species, but others have accepted F. cottonii as a distinct species (Guiry, 1998). To<br />
confirm the validity <strong>of</strong> the species and my designation <strong>of</strong> the Prince William Sound material,
Chapt 9C1. Marine Plants, page 9C1- 23<br />
samples are being sent to Esther Serrao in Portugal, who is studying the phylogenetic<br />
relationships <strong>of</strong> the genus with molecular techniques.<br />
Microspongium globosum. This tiny brown alga was found growing epiphytically on<br />
Delamaraea attenuata on the floats at Tatitlek. The thalli were abundant and bore plurilocular<br />
sporangia that clearly match the diagrams for this species in Fletcher (1987). Known only from<br />
the North Atlantic and Japan, the species makes a surprising appearance in Alaska. It also has<br />
not been reported from the well investigated areas to the south. Its occurrence on the oyster<br />
floats at Tatitlek as an epiphyte on another new record to Alaska indicates to me that this species,<br />
possibly along with its host, is another new introduction to the area. However, its vector could<br />
also have been oysters from Japan.<br />
Macrocystis integrifolia. Since 1979 (Jay Johnson, Alaska Fish and Game, pers com.)<br />
Macrocystis has been imported (by plane) from southeast Alaska to Prince William Sound to be<br />
used as substrate for herring eggs in the lucrative Herring-Roe-On-Kelp (HROK) fishery.<br />
Normally only blades and fronds <strong>of</strong> the giant kelp are transported northward for the fishery.<br />
These are then placed in impoundment nets which house both the kelp and the fish. The eggladen<br />
blades are then harvested and sold primarily to Japan as a gourmet food item.<br />
Theoretically, the blades that are brought up to the Sound are clean (the most desirable for<br />
HROK) and are all harvested for later sale. However, during our June trip and during many <strong>of</strong><br />
Hansen’s earlier trips to the area, blades and holdfasts <strong>of</strong> the kelp that had escaped were found<br />
adrift in Prince William Sound. Perhaps due to the climate, none <strong>of</strong> these plants appear to have<br />
propagated in the area since none have ever been found attached anywhere north <strong>of</strong> southeast<br />
Alaska. Hence, in the site list the species is listed as a failed introduction. However, even with<br />
the transport <strong>of</strong> “eye-clean” blades, it is likely that numerous small algal and animal species are<br />
co-transported accidentally from southeast Alaska to Prince William Sound every year with this<br />
kelp. This may account, as much as the current El Niño, for many <strong>of</strong> our new range extensions.<br />
Discussion<br />
During the search for marine plant NIS in Prince William Sound, it was important to<br />
characterize the flora at each <strong>of</strong> the sites so that the probable introduced species and their impacts<br />
on the community could be recognized. In addition, information on the taxonomic and residency<br />
status composition <strong>of</strong> these communities was absolutely essential to be able to determine<br />
vulnerable sites for future invasions. Although 155 species <strong>of</strong> plants collected during the 1998<br />
cruise is probably only about half that <strong>of</strong> the actual flora <strong>of</strong> Prince William Sound, the data<br />
compiled reveal important trends in community composition. In addition, the final lists <strong>of</strong> new<br />
records and probable introductions give valuable insight into the difficulties <strong>of</strong> recognizing NIS.<br />
Limitations <strong>of</strong> the data. Although 19 sites were visited during the June 1998 survey, the<br />
time allowed for sampling was inadequate at many <strong>of</strong> the beaches. This had a substantial impact<br />
on the overall data. Moreover, the lack <strong>of</strong> year-round collections for the area limits the results in<br />
ways that cannot even be predicted. In terms <strong>of</strong> numbers, this can be shown clearly by<br />
comparing the total count <strong>of</strong> 112 species shown for Port Valdez in the Checklist (which includes<br />
seasonal collections) with the count <strong>of</strong> 47 for the area obtained during this short trip.
Chapt 9C1. Marine Plants, page 9C1- 24<br />
Information derived from additional collections and herbarium specimens from our sites would<br />
help to overcome these difficulties and to improve the resolution <strong>of</strong> the data.<br />
Although temperature and salinity influenced the total species counts for marine algal<br />
species, this could not be demonstrated clearly with the data on hand. Prince William Sound has<br />
regions heavily effected by rain, snow and ice melt, and marked changes in temperature and<br />
salinity occur throughout the year. During summers, salinity is the lowest as run<strong>of</strong>f produces a<br />
freshwater lens on the surface <strong>of</strong> many <strong>of</strong> the bays, including Port Valdez and Whittier. In these<br />
areas intertidal species are subjected to wide salinity fluctuations with the tidal cycle. Since these<br />
physical factors were measured only during the limited sampling periods, they do not reflect the<br />
range <strong>of</strong> conditions encountered over time by the intertidal species sampled.<br />
Limited historical knowledge <strong>of</strong> the flora and new records. Only two floristic papers<br />
on the marine algae (and plants) <strong>of</strong> Prince William sound have ever been published (Calvin and<br />
Lindstrom, 1980; Wiegers et al., 1997). In addition, an overall identification guide to the marine<br />
algae does not exist for Prince William Sound or even for Alaska, and we are left with using an<br />
assortment <strong>of</strong> references from neighboring areas to identify species. This lack <strong>of</strong> both taxonomic<br />
information and baseline data for Prince William Sound is clearly evident in the discovery <strong>of</strong> 21<br />
new records to the area amounting to 13.5% <strong>of</strong> the species collected during our short 9 day<br />
cruise. During our earlier Pilot Study, an additional 3 new records were found in Port Valdez<br />
alone. Though fairly evenly distributed among the 3 major taxonomic groups, there were a few<br />
more new records among brown and green algae than among the red, and the majority <strong>of</strong> the<br />
species appeared to be cryptogenic. In addition, new record species were slightly more abundant<br />
at certain sites. Green Island, the most diverse site in the study, bore 6 new records, while the<br />
float sites combined bore 9. Both <strong>of</strong> these areas (probably along with many others in Prince<br />
William Sound) appear to have been understudied in the past. Since, for this study, our probable<br />
NIS were derived from the new records, it is understandable that each <strong>of</strong> these 2 sites (or site<br />
types) also bore 2 <strong>of</strong> the 5 probable NIS designated in the study.<br />
Taxonomic composition. In nearly all temperate outer-coastal habitats, the red algal<br />
species are the highest in numbers followed by the brown and then the green algae forming a<br />
R>B>G hierarchical pattern <strong>of</strong> dominance. Proceeding from open coasts into protected bays and<br />
estuaries, the ratio changes to reflect a reduction in the number <strong>of</strong> red algae. For instance, <strong>of</strong>f the<br />
coast <strong>of</strong> Oregon, the R:B:G ratio is 61:22:17. In Prince William Sound, the overall ratio <strong>of</strong><br />
R:B:G in the species surveyed was 47:33:20, a ratio probably indicating the influence <strong>of</strong><br />
sheltered and less saline water. However, the overall composition pattern was still R>B>G<br />
(Table 9C1.13). The R>B>G dominance pattern in Prince William Sound occurred only at<br />
Rocky Headlands and Reefs and in Rocky Bays, all areas <strong>of</strong> moderate to high water movement<br />
(exposure) and relatively uniform salinity and temperature supporting established communities<br />
with numerous annuals and perennials. In Harbors, the proportion <strong>of</strong> green algae increased and<br />
the pattern became R>G>B, reflecting the tolerance <strong>of</strong> green algae for lower salinities found in<br />
this habitat. In addition, since many <strong>of</strong> the green and brown algae are ephemeral (opportunistic),<br />
they can survive the wide fluctuations in temperature and salinity. Moreover, since ephemeral<br />
forms are <strong>of</strong>ten fouling organisms, many are repeatedly brought in to seed these areas by boat<br />
traffic. In the mud bays and on the floats, the proportion <strong>of</strong> brown algae increased. In mud bays,<br />
the frequent shifts in mud level smothers many <strong>of</strong> the species, providing niches primarily for
Chapt 9C1. Marine Plants, page 9C1- 25<br />
ephemerals and unattached forms. On the oyster floats, the early successional ephemeral forms<br />
are also encouraged due to the periodic cleaning <strong>of</strong> the habitat. The higher and generally more<br />
uniform salinity <strong>of</strong> both <strong>of</strong> these habitats appears to enable the ephemeral browns to outcompete<br />
the ephemeral greens.<br />
Table 9C1.13. Summary <strong>of</strong> the Hierarchical Composition<br />
and Physical Features <strong>of</strong> each Habitat Type<br />
Observed during the June 1998 Survey<br />
Habitat Types<br />
Harbors Mud Bays Rk Headlands Rk Bays Floats<br />
Composition<br />
and Reefs<br />
Taxonomic* R>G>B B=R>G R>B>G R>B>G B>R>G<br />
variable variable variable<br />
Residency Status C>N C>N N>C N>C C>N<br />
Exposure low low high low med-high<br />
Salinity variable low-med high variable high<br />
Temperature variable variable uniform uniform uniform<br />
Substratum variable s<strong>of</strong>t hard hard hard<br />
* = includes only the 3 major taxonomic groups sampled.<br />
Resident type composition. Cryptogenic species predominated in the more disturbed<br />
and variable habitats <strong>of</strong> the Harbors, Mud Bays, and Floats, while the native species<br />
predominated in the less disturbed and more uniform habitats provided by Rocky Headlands and<br />
Reefs, and Rocky Bays. Cryptogenic algal species appear to contain a high percentage <strong>of</strong><br />
ephemeral forms. Hence, their ability to survive in fluctuating environments and perhaps in<br />
ballast water and on ship bottoms is high.<br />
Introduced species and their impact. The 5 probable plant NIS discovered during our<br />
survey are all isolated (and probably young) populations. Although four <strong>of</strong> these species do not<br />
appear to have wide distribution in Prince William Sound, Fucus cottonii does appear to have an<br />
expanding range. It was found at 4 <strong>of</strong> our sites and appears to be prevalent in the supra-littoral <strong>of</strong><br />
all <strong>of</strong> these areas. Unique to sloughs and the marsh area <strong>of</strong> mudflats, this species does not seem<br />
to be replacing any <strong>of</strong> the known marine or estuarine species. However, in Cloudman Bay, it<br />
may actually be out-competing some terrestrial plants. Fortunately, none <strong>of</strong> the probable NIS<br />
plants found in Prince William Sound appear to be hazardous to the environment. None are as<br />
toxic or as invasive as the Mediterranean introduction Caulerpa taxifolia (Lemee et al., 1993;<br />
Verlaque and Fritayre, 1994).<br />
The transport mechanisms <strong>of</strong> these introductions is only partially clear. The two species<br />
(Chroodactylon ramosum, Microspongium globosum ) found on oyster floats could have been<br />
brought into the area with the transplantation <strong>of</strong> oysters for aquaculture purposes. The vector for<br />
Codium and its epiphyte Ceramium is more debatable. The subspecies Codium fragile fragile<br />
was potenitally transported up from southeast Alaska with Macrocystis for the HROK industry.<br />
But the only method <strong>of</strong> transport for the subspecies Codium fragile tomentasoides would have to<br />
be either ballast water or as fouling on the hulls <strong>of</strong> ships. The importation mechanism <strong>of</strong> Fucus<br />
cottonii is even less clear. Its relatively widespread occurrence in Prince William Sound (and in<br />
patchy spots along the west coast) indicates that it is probably not a recent introduction.<br />
However, since it is a predominantly unattached species, it is also an excellent candidate for<br />
transport by ballast water.
Chapt 9C1. Marine Plants, page 9C1- 26<br />
Other potential introductions and their significance. What <strong>of</strong> the other 70 cryptogenic<br />
species which are possibly introductions, but which have less obvious characteristics <strong>of</strong><br />
invasion These species are, by definition, wide-ranging and many are abundant, <strong>of</strong>ten heavily<br />
impacting the communities in which they occur. Pro<strong>of</strong> <strong>of</strong> the NIS status <strong>of</strong> these prominent<br />
species is possible, but it will require detailed comparative morphological study and world-wide<br />
molecular biological tracking <strong>of</strong> their distributions. Furthermore, knowledge <strong>of</strong> the impacts <strong>of</strong><br />
these species on community structure will demand complex physiological and ecological studies<br />
<strong>of</strong> the species in both their introduced and native habitats. These studies are important projects<br />
for future investigators who are concerned about the conservation <strong>of</strong> our native biodiversity.<br />
References<br />
Abbott, I. A., and G. J. Hollenberg. 1976. Marine Algae <strong>of</strong> California. Stanford University<br />
Press, Stanford. xii+827 pp.<br />
Adams, N. M. 1983. Checklist <strong>of</strong> marine algae possibly naturalized in New Zealand. N. . J.<br />
Bot. 21: 1-2.<br />
Adams, N. M. 1994. Seaweeds <strong>of</strong> New Zealand, an illustrated guide. Canterbury University<br />
Press Publ., New Zealand. 360 pp.<br />
Calvin, N. I., and S. C. Lindstrom. 1980. Intertidal algae <strong>of</strong> Port Valdez, Alaska: species and<br />
distribution with annotations. Bot. Mar. 23: 791-797.<br />
Carlton, J. T. 1996. <strong>Biological</strong> invasions and cryptogenic species. Ecology 77 (6): 1653-1655.<br />
Chapman, J., and G. Hansen. 1997. Surveys <strong>of</strong> nonindigenous aquatic species for Port Valdez,<br />
Alaska. In: Ruiz, G.M. and A.H. Hines. 1997. Patterns <strong>of</strong> nonindigenous species transfer and<br />
invasion in Prince William Sound, Alaska: Pilot Study. Report, Prince William Sound Regional<br />
Citizens’ Advisory Council. 80pp.<br />
Dawson. E. Y. 1950. A review <strong>of</strong> Ceramium along the Pacific coast <strong>of</strong> North America with<br />
special reference to its Mexican representatives. Farlowia 4: 113-138.<br />
DeWreede, R. E. 1996. The impact <strong>of</strong> seaweed introductions on biodiversity. Global<br />
Biodiversity 6: 2-9.<br />
Fletcher, R. L. 1987. Seaweeds <strong>of</strong> the British Isles. Vol. 3. Fucophyceae (Phaeophyceae). Part<br />
1. British Museum (Natural History), London. 359 pp.<br />
Gabrielson, P. W., R. F. Scagel, and T. B. Widdowson. 1989. Keys to the benthic marine algae<br />
and seagrasses <strong>of</strong> British Columbia, southeast Alaska, Washington and Oregon. Phycological<br />
Contribution Number 4. Dept. <strong>of</strong> Botany, University <strong>of</strong> British Columbia, Vancouver. vi+187<br />
pp.<br />
Guiry, R. 1998. An internet accessible “taxonomic database” on “seaweeds” (primarily <strong>of</strong><br />
Europe). http://seaweed.ucg.ie.
Chapt 9C1. Marine Plants, page 9C1- 27<br />
Hansen, G. 1997. A revised checklist and preliminary assessment <strong>of</strong> the macrobenthic marine<br />
algae and seagrasses <strong>of</strong> Oregon. Pp. 175-200 in Kaye, T., A. Liston, R. Love, D. Luoma, R.<br />
Meinke, and M. Wilson (ed.). Conservation and Management <strong>of</strong> Native Flora and Fungi. Native<br />
Plant Society <strong>of</strong> Oregon, Corvallis.<br />
Hansen, G. I., D. J. Garbary, J. C. Oliveira, and R. F. Scagel. 1981. New records and range<br />
extensions <strong>of</strong> marine algae from Alaska. Syesis 14: 115-123.<br />
Lee, R. K. S. 1980. At catalogue <strong>of</strong> the marine algae <strong>of</strong> the Canadian Arctic. Publications in<br />
Botany, No. 9. National Museums <strong>of</strong> Canada, National Museum <strong>of</strong> Natural Sciences, Ottawa,<br />
ON. 82 pp.<br />
Lemee, R., D. Pesando, M. Durand-Clement, A. Dubreuil, A. Meinesz, A. Guerriero, and F.<br />
Pietra. 1993. Preliminary survey <strong>of</strong> toxicity <strong>of</strong> the green alga Caulerpa taxifolia introduced into<br />
the Mediterranean. J. Appl. Phycol. 5:485-493.<br />
Lindstrom, S. C. 1977. An annotated bibliography <strong>of</strong> the benthic marine algae <strong>of</strong> Alaska.<br />
ADF&G Technical Data Report No. 31. Juneau. 172 pp.<br />
Nelson, W. A., and C. A. Maggs. 1996. Records <strong>of</strong> adventive marine algae in New Zealand:<br />
Antithamnionella ternifolia, Polysiphonia senticulosa (Ceramiales, Rhodophyta), and Striaria<br />
attenuata (Dictyosiphonales, Phaeophyta). N. Z. J. Mar. Freshwat. Res. 30: 449-453.<br />
O’Clair, R. M., S. C. Lindstrom, I. R. Brodo. 1996 [1997]. Southeast Alaska’s Rocky Shores:<br />
Seaweeds and Lichens. Plant Press, Auke Bay.<br />
Perestenko, L. P. 1994. Red Algae <strong>of</strong> the Far-Eastern Seas <strong>of</strong> Russia. Komarov Botanical<br />
Institute, Russian Academy <strong>of</strong> Sciences, St. Petersburg. 331 pp. [In Russian].<br />
Phillips, R. C., and E. G. Menez. 1988. Seagrasses. Smithsonian Contributions to the Marine<br />
Sciences 34. v+104 pp.<br />
Rueness, J. 1977. Norsk Algeflora. Universitetsforlaget, Oslo. 266 pp. [In Norwegian].<br />
Ruiz, G. M., and A. H. Hines. 1997. The risk <strong>of</strong> nonindigenous species invasion in Prince<br />
William Sound associated with tanker traffic and ballast <strong>Water</strong> Management: Pilot Study.<br />
Regional Citizens’ Advisory Council <strong>of</strong> Prince William Sound RFP Number 632.97.1. 47 pp +<br />
52 pp tables and graphs.<br />
Scagel, R. F., P. W. Gabrielson, D. J. Garbary, L. Golden, M. W. Hawkes, S. C. Lindstrom, J. C.<br />
Oliveira, and T. B. Widdowson. 1989 [1993]. A Synopsis <strong>of</strong> the Benthic Marine Algae <strong>of</strong><br />
British Columbia, southeast Alaska, Washington and Oregon. Phycological Contribution<br />
Number 3. Dept <strong>of</strong> Botany, University <strong>of</strong> British Columbia, Vancouver, BC. 535 pp.
Chapt 9C1. Marine Plants, page 9C1- 28<br />
Sears, J. R. 1998. NEAS Keys to the Benthic Marine Algae <strong>of</strong> the Northeastern Coast <strong>of</strong> North<br />
America from Long Island sound to the Strait <strong>of</strong> Belle Isle. NEAS Contribution Number 1,<br />
Dartmouth, MA. xi+161pp.<br />
Selivanova, O. N., and G. G. Zhigadlova. 1997. Marine algae <strong>of</strong> the Commander Islands:<br />
preliminary remarks on the revision <strong>of</strong> the flora. Bot. Mar. 40: 1-24.<br />
Setchell, W. A., and N. L. Gardner. 1924. Expedition <strong>of</strong> the California Academy <strong>of</strong> Sciences to<br />
the Gulf <strong>of</strong> California in 1921. Proc. <strong>of</strong> the Cal. Acad. Sci., 4 th Series, 12: 12-88.<br />
Verlaque, M., and P. Fritayre. 1994. Mediterranean algal communities are changing in face <strong>of</strong><br />
the invasive alga Caulerpa taxifolia (Vahl) C. Agardh. Oceanol. Acta 17: 659-672.<br />
Vis, M. L., and R. G. Sheath. 1993. Distribution and systematics <strong>of</strong> Chroodactylon and<br />
Kyliniella (Porphyridiales, Rhodophyta) from North American streams. Jap. J. <strong>of</strong> Phycology 41:<br />
237-241.<br />
Wiegers, J. K., H. M. Feder, W. G. Landis, L. S. Nortensen, D. G. Shaw, V. J. Wilson. 1997. A<br />
regional multiple-stressor ecological risk assessment for Port Valdez, Alaska. IETC No. 9701<br />
and RCAC 1033.102. Inst. <strong>of</strong> Environmental Toxicology and Chemistry, Western Washington<br />
Univ., Bellingham, WA.<br />
Womersley, H. B. S. 1984. The Marine benthic flora <strong>of</strong> southern Australia, Part 1. Woolman,<br />
Government Printer, South Australia, 329 pp.<br />
Womersley, H. B. S. 1987. The marine benthic flora <strong>of</strong> southern Australia, Part 2. Australian<br />
Government Printing Division, Adelaide, 484 pp.<br />
Womersley, H. B. S. 1994. The marine benthic flora <strong>of</strong> southern Australia. Part 3A. Australian<br />
<strong>Biological</strong> Resources Study, Canberra, 508 pp.<br />
Womersley, H. B. S. The marine benthic flora <strong>of</strong> southern Australia. Part 3B. Australian<br />
<strong>Biological</strong> Resources Study, Canberra. 392 pp.<br />
Yoshida, T., K. Yoshinaga, and Y. Nakajima. 1995. Checklist <strong>of</strong> marine algae <strong>of</strong> Japan. Jap. J.<br />
Phycol. 43: 115-171. [In Japanese].
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 1<br />
Chapter 9C2. Focal Taxonomic Collections: Planktonic Cnidaria, Ctenophora, and<br />
Pelagic Mollusca<br />
Claudia Mills, Friday Harbor Laboratories, University <strong>of</strong> Washington<br />
Methods<br />
Medusae, ctenophores, and pelagic molluscs were collected at sites in both Prince William<br />
Sound and Cook Inlet from August 8–14, 1999 using small plankton nets and a water scoop attached<br />
to a long handle. Specimens were examined in the field, relaxed, and fixed for transport to the<br />
laboratory. Specimens were reexamined microscopically at the Friday Harbor Laboratories in<br />
September 1999, in order to verify or assign species names.<br />
Results<br />
All pelagic Hydrozoa, Scyphozoa, Ctenophora and Mollusca are identified by site in Table<br />
9C2.1. Separate species lists for these groups follow for Prince William Sound (Table 9C2.2) and<br />
Cook Inlet (Table 9C2.3). An annotated species list follows (Table 9C2.4), including all species on<br />
both lists. In the time allotted to this project, I do not feel that I completed a comprehensive search<br />
<strong>of</strong> the literature for species previously collected in Prince William Sound and Cook Inlet, but no<br />
other papers came to mind. I also did not search for unpublished data at the University <strong>of</strong> Alaska.<br />
A few coelenterates were conspicuously missing from the region. We saw no<br />
stauromedusae, no Epiactis anemones on eelgrass, no Chrysaora or Phacellophora<br />
scyphomedusae, and no Anthopleura elegantissima or A. xanthogrammica.<br />
No known nonindigenous species <strong>of</strong> planktonic Cnidaria or Ctenophora were collected in<br />
Prince William Sound or Cook Inlet by our scientific teams in either 1998 or 1999. In the 1999<br />
expedition, 15 species <strong>of</strong> Hydrozoa were collected (including 3 hydroids [see section 9D. Fouling<br />
Communities) for more thorough hydroid work-up] and 14 species <strong>of</strong> hydromedusae), two<br />
scyphomedusae and unidentified scyphozoan polyps (scyphistomae), and two species <strong>of</strong><br />
ctenophores. Two molluscan species were also taken in the water column.<br />
The following species appear to be new records in the Prince William Sound region:<br />
Hydromedusae<br />
*Aequorea aequorea<br />
*Aequorea victoria<br />
*Clytia gregaria (=Phialidium gregarium)<br />
Eperetmus typus<br />
Euphysa sp.<br />
Gonionemus vertens<br />
Halitholus sp.<br />
*Melicertum octocostatum<br />
*Proboscidactyla flavicirrata<br />
Sarsia spp.<br />
Tiaropsis multicirrata
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 2<br />
Ctenophora:<br />
*Bolinopsis infundibulum<br />
*Pleurobrachia bachei<br />
New NAME for common Scyphomedusa<br />
Aurelia labiata<br />
* indicates common species whose presence in PWS may be known, but I have not seen reports<br />
in print. Dr. Jennifer Purcell (Horn Point Laboratory, University <strong>of</strong> Maryland) is working with<br />
some <strong>of</strong> these, but her results are unpublished as yet.<br />
Following the annotated species list is a list and discussion <strong>of</strong> nonindigenous cnidarian<br />
species already present in some west coast estuaries that might be positioned to ultimately invade<br />
locations in Alaska. This list is accompanied by an Appendix (following the report) titled<br />
“Commentary on species <strong>of</strong> Hydrozoa, Scyphozoa and Anthozoa (Cnidaria) sometimes listed as<br />
non-indigenous in Puget Sound”, reprinted from Cohen et al. (1998). References are given at the end<br />
<strong>of</strong> the main report as well as the Appendix.<br />
References<br />
Arai, M. N. and A. Brinckmann-Voss, 1980. Hydromedusae <strong>of</strong> British Columbia and Puget Sound.<br />
Can. Bull. Fish Aquat. Sci., 204: 192 pp.<br />
Bigelow, H. B. 1912. The ctenophores. Bull. Mus. Comp. Zool., 56: 369-404, 2 pls.<br />
Bigelow, H. B. 1913. Medusae and siphonophorae collected by the U. S. Fisheries steamer<br />
“Albatross” in the Northwestern Pacific, 1906. Proc. U. S. Nat. Museum, 44: 1-119, 6 pls.<br />
Bigelow, H. B. 1920. Medusae and Ctenophora. Rep. Canadian Arctic Exp. 1913-18, Southern Party<br />
1913-16, Volume VIII: Mollusks, Echinoderms. Coelenterates, etc., Part H: 3H-22H, 2 pls.<br />
Calder, D. R. 1988. Shallow-water hydroids <strong>of</strong> Bermuda: the Athecatae. Royal Ontario Museum Life<br />
Sciences Contributions, 148: 1-107.<br />
Cohen, A., C. Mills, H. Berry, M. Wonham, B. Bingham, B. Bookheim, J. Carlton, J. Chapman, J.<br />
Cordell, L. Harris, T. Klinger, A. Kohn, C. Lambert, G. Lambert, K. Li, D. Secord, and J. T<strong>of</strong>t,<br />
November 1998. Report <strong>of</strong> the Puget Sound Expedition, September 8–16, 1998: a Rapid<br />
Assessment survey <strong>of</strong> non-indigenous species in the shallow waters <strong>of</strong> Puget Sound. Washington<br />
State Department <strong>of</strong> Natural Resources, Olympia, Washington, 37 pages.<br />
Greenberg, N., R. L. Garthwaite and D. C. Potts, 1996. Allozyme and morphological evidence for<br />
a newly introduced species <strong>of</strong> Aurelia in San Francisco Bay. Marine Biology, 125: 401-410.<br />
Harbo, R. M. 1999. Whelks to Whales: <strong>Coastal</strong> Marine Life <strong>of</strong> the Pacific Northwest. Harbour<br />
Publishing, Madeira Park, B.C., Canada.
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 3<br />
Kramp, P. L. 1961. Synopsis <strong>of</strong> the medusae <strong>of</strong> the world. J. Mar. Biol. Assoc. U.K., 40: 1–469.<br />
Mills, C. E. 1981. Seasonal occurrence <strong>of</strong> planktonic medusae and ctenophores in the San Juan<br />
Archipelago (NE Pacific). Wasmann J. Biol., 39: 6-29.<br />
Mills, C. E. 1998 to present. Web Site: http://faculty.washington.edu/cemills/<br />
Mills, C. E. and F. Sommer, 1995. Invertebrate introductions in marine habitats: two species<br />
<strong>of</strong> hydromedusae (Cnidaria) native to the Black Sea, Maeotias inexspectata and Blackfordia<br />
virginica, invade San Francisco Bay. Marine Biology, 122: 279-288.<br />
Murbach, L. and C. Shearer. 1903. On medusae from the coast <strong>of</strong> British Columbia and<br />
Alaska. Proc. Zool. London 2:164-192, pls. 17-22.<br />
Purcell, J. E. 1998 Project report 98163S - Jellyfish as competitors and predators <strong>of</strong> fishes.<br />
On the web at http://www.uaa.alaska.edu/enri/apex/98163S.html.<br />
Ricketts, E. F. and J. Calvin, 1939. Between Pacific Tides. Stanford University Press, Stanford.<br />
Wrobel, D. and C. Mills, 1998. Pacific Coast Pelagic Invertebrates: a Guide to the Common<br />
Gelatinous Animals. Sea Challengers and the Monterey Bay Aquarium, Monterey, California.
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 4<br />
Table 9C2.1. Hydrozoa, Scyphozoa, Ctenophora and pelagic Mollusca by site, 1999.<br />
Collections and identifications by Claudia E. Mills.<br />
1999 PWS EXPEDITION<br />
C.E. MILLS REPORT<br />
Homer<br />
Sadie<br />
Cove<br />
Seward<br />
Marina<br />
Lowell<br />
Pt<br />
Seward<br />
south <strong>of</strong><br />
Esther<br />
Is.<br />
Whittier<br />
Fairmoun<br />
t Bay<br />
HYDROZOA<br />
X<br />
Aequorea aequorea v, albida X X X X X X<br />
Aglantha digitale<br />
X<br />
Bougainvillia superciliaris<br />
x<br />
Clytia gregaria<br />
X X X X X<br />
(=Phialidium gregarium)<br />
Eperetmus typus X X<br />
Euphysa sp.<br />
X<br />
Eutonina indicans<br />
X<br />
Gonionemus vertens<br />
X<br />
Halitholus sp.<br />
X<br />
Leuckartiara sp.<br />
X<br />
Melicertum octocostatum X X X<br />
Mitrocoma cellularia<br />
X<br />
Obelia hydroids X X X<br />
Proboscidactyla flavicirrata X +<br />
polyps<br />
X X X<br />
Sarsia/Coryne sp. hydroids X<br />
Sarsia spp. medusae X X X X<br />
SCYPHOZOA<br />
Aurelia labiata<br />
X<br />
Cyanea capillata X X X X X X X<br />
unidentified scyphistomae<br />
(probably Aurelia sp.)<br />
X<br />
X<br />
Valdez<br />
Mar<br />
ina<br />
Tatitlek<br />
Bus<br />
-by<br />
Is.<br />
Cord<br />
ova<br />
CTENOPHORA<br />
Bolinopsis infundibulum<br />
Pleurobrachia bachei<br />
X<br />
X<br />
MOLLUSCA<br />
Melibe leonina X X X<br />
Clione limacina X
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 5<br />
Table 9C2..2. Prince William Sound species list. See Table 1 for specific locations.<br />
Collections and identifications by Claudia E. Mills, unless otherwise noted.<br />
HYDROZOA<br />
REFERENCE<br />
Aequorea aequorea v. albida PWS 99; Purcell, 1998<br />
Aequorea victoria/ A. aequorea v. aequorea Purcell, personal communication 1999<br />
Catablema multicirrata Bigelow, 1913<br />
Clytia gregaria PWS 99<br />
(=Phialidium gregarium)<br />
Eperetmus typus PWS 98, PWS 99<br />
Euphysa sp. PWS 99<br />
Gonionemus vertens PWS 99<br />
Halitholus sp. PWS 99<br />
Melicertum octocostatum PWS 99<br />
Obelia longissima PWS 98<br />
Obelia spp. hydroids PWS 99<br />
Proboscidactyla flavicirrata PWS 99<br />
Sarsia spp. medusae PWS 99<br />
Staurophora mertensii Bigelow, 1913<br />
Tiaropsis multicirrata PWS 98<br />
SCYPHOZOA<br />
Aurelia"aurita" (Mills quotes) Purcell, 1998<br />
Aurelia labiata PWS 99<br />
Cyanea capillata PWS 99; Purcell, 1998<br />
Unidentified scyphistomae PWS 99<br />
(probably Aurelia sp.)<br />
CTENOPHORA<br />
Bolinopsis infundibulum PWS 99<br />
Pleurobrachia bachei PWS 99; Purcell, 1998<br />
MOLLUSCA<br />
Melibe leonina PWS 99<br />
* PWS 98 refers to specimens collected by Ruiz et al., June 1998 in Cook Inlet.<br />
PWS 99 refers to specimens collected by Greg Ruiz et al., August 1999 in Cook Inlet.
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 6<br />
Table 9C2.3. Cook Inlet species list.<br />
Collections and identifications by Claudia E. Mills, unless otherwise noted.<br />
HYDROZOA<br />
*REFERENCE AND LOCATION<br />
Aequorea aequorea v. albida PWS 99 - Homer Marina<br />
Aglantha digitale<br />
PWS 99 - Sadie Cove, Katchemak Bay<br />
Bougainvillia superciliaris PWS 99 - Sadie Cove, Katchemak Bay<br />
Clytia gregaria<br />
PWS 99 - Sadie Cove, Katchemak Bay<br />
(=Phialidium gregarium)<br />
Eperetmus typus<br />
PWS 99 - Sadie Cove, Katchemak Bay<br />
Eutonina indicans<br />
PWS 99 - Homer Marina<br />
Leuckartiara sp.<br />
PWS 99 - Sadie Cove, Katchemak Bay<br />
Melicertum octocostatum<br />
PWS 99 - Homer Marina<br />
Mitrocoma cellularia<br />
PWS 99 - Homer Marina<br />
Obelia sp. hydroids<br />
PWS 99 - Homer Marina<br />
Proboscidactyla flavicirrata hydroids PWS 99 - Homer Marina<br />
Sarsia/Coryne sp. hydroids PWS 99 - Homer Marina<br />
Sarsia spp. medusae<br />
PWS 99 - Homer Marina, Sadie Cove<br />
SCYPHOZOA<br />
Cyanea capillata<br />
Unidentified scyphistomae<br />
(probably Aurelia sp.)<br />
PWS 99 - Homer Marina, Sadie Cove<br />
PWS 99 - Homer Marina<br />
CTENOPHORA<br />
(none)<br />
MOLLUSCA<br />
Clione limacina PWS 99 - Homer marina<br />
Melibe leonina<br />
PWS 99 - not collected but told <strong>of</strong> site at Jakal<strong>of</strong> Bay<br />
by Carmen Field<br />
* PWS 99 refers to specimens collected by Ruiz et al., August 1999 in Cook Inlet.
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 7<br />
Table 9C2.4<br />
PRINCE WILLIAM SOUND ANNOTATED SPECIES LIST<br />
(combines both Cook Inlet and Prince William Sound locations)<br />
HYDROZOA<br />
Aequorea aequorea var. albida<br />
Distribution. Most <strong>of</strong> the Aequorea medusae that we saw were beached. Such specimens were seen at the<br />
Homer Marina, Lowell Point in Seward, the Whittier Marina, and in Cordova and Valdez. A few were<br />
seen in the water while underway south <strong>of</strong> Esther Island, along with Cyanea capillata.<br />
Remarks. This name was applied by Bigelow (1913) to Aequorea specimens measuring 120 mm and<br />
165 mm bell diameter, collected in Dutch Harbor. Such very-large Aequorea occur throughout southern<br />
Alaska, and are accompanied in some places by smaller specimens that seem very similar to Aequorea<br />
victoria at Friday Harbor (called Aequorea aequorea var. aequroea by Bigelow, 1913). Whether they<br />
are different sizes <strong>of</strong> the same species or 2 different species has still not been resolved (even the modern<br />
use <strong>of</strong> "A. victoria" as species name for Friday Harbor medusae is controversial). Only large-sized<br />
specimens (most 120-160 mm diameter) were seen in Prince William Sound in August 1999. Similar<br />
large Aequoreas in Prince William Sound were called A. victoria by Purcell (1998).<br />
Aequorea victoria or Aequorea aequorea var. aequorea<br />
Remarks. We did not collect any smaller specimens <strong>of</strong> Aequorea, but I am told by Dr. Jennifer Purcell,<br />
who has been doing a recent plankton study in Prince William Sound that small Aequoreas that look like<br />
those at Friday Harbor are also present. Bigelow (1913) calls these A. aequorea var. aequorea. Arai<br />
and Brinckmann-Voss later applied the name A. victoria to the same animals. It is not clear to me that<br />
A. victoria is not a junior synonym to A. aequorea.<br />
Aglantha digitale<br />
Distribution. Several Aglantha digitale medusae were collected at the head <strong>of</strong> Sadie Cove,<br />
Katchemak Bay, on August 8, 1999, by dipping from a small boat in about 8 feet <strong>of</strong> water. Most were<br />
within a layer <strong>of</strong> fresher water that occupied the upper 15" <strong>of</strong> the water column and were dead and<br />
decomposing. Many others were seen, but not collected.<br />
Remarks. There is no question about the identification <strong>of</strong> this material, although why these medusae<br />
were in the layer <strong>of</strong> low salinity water is not clear. This circumpolar species is well known in the<br />
North Pacific, North Atlantic and Arctic Oceans, including the Bering Sea.<br />
Bougainvillia superciliaris<br />
Distribution. Five Bougainvillia superciliaris medusae were collected at the head <strong>of</strong> Sadie Cove,<br />
Katchemak Bay, on August 8, 1999, by dipping from a small boat in about 8 feet <strong>of</strong> water. All were<br />
below a layer <strong>of</strong> fresher water that occupied the upper 15" <strong>of</strong> the water column. Several others were<br />
seen, but not collected.<br />
Remarks. These 6-12 mm high specimens best correspond with the description <strong>of</strong> Bougainvillia<br />
superciliaris, having its characteristic prominent peduncle above the manubrium. The Sadie Cove<br />
specimens had 36-40 tentacles on each <strong>of</strong> the four marginal bulbs, which is quite a bit higher than the
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 8<br />
10-22 tentacles described for B. superciliaris in Kramp (1961). Bigelow (1913) found a single specimen<br />
<strong>of</strong> B. superciliaris <strong>of</strong> the same size <strong>of</strong>f Attu Island in the Aleutians, but also with less than 20 tentacles<br />
in each group. This tentacle number discrepancy leads to the question about species identification for the<br />
Sadie Cove material.<br />
Catablema multicirrata<br />
Distribution. This species (2 medusae) was collected by Bigelow (1913) <strong>of</strong>f Orca in Prince William<br />
Sound on July 19, 1906.<br />
Remarks. We did not find it in August 1999, but did not sample at that location.<br />
Clytia gregaria (=Phialidium gregarium)<br />
Distribution. Many individuals <strong>of</strong> this species were collected in Sadie Cove, Katchemak Bay and in<br />
Fairmount Bay, Tatitlek, <strong>of</strong>f Busby Island and in the Cordova Marina.<br />
Remarks. Most <strong>of</strong> the specimens correspond well to the description <strong>of</strong> Clytia gregaria (as Phialidium<br />
gregarium) in Kramp (1961), with about 40 tentacles and a few rudimentary bulbs alternating with<br />
marginal vesicles in 12 mm diameter medusae; the gonads were on the distal 1/2 <strong>of</strong> the radial canals.<br />
Some smaller medusae (7 mm diameter), with a few less tentacles and shorter gonads may be C.<br />
gregaria, or could be C. lomae, looking very similar to specimens collected in September 1998 in Puget<br />
Sound by Claudia Mills and Erik Thuesen. Seasonal morphological variation with changes in<br />
zooplankton prey availability have not been described, so it is difficult to be positive about the species<br />
name in some cases. The genus name Clytia has typically been applied only to the hydroid form, but it is<br />
an older genus name than Phialidium, and should be applied to both phases <strong>of</strong> the life cycle.<br />
Eperetmus typus<br />
Distribution. One young Eperetmus typus medusa was collected at the head <strong>of</strong> Sadie Cove,<br />
Katchemak Bay, on August 8, 1999, by dipping from a small boat in about 8 feet <strong>of</strong> water. It was<br />
below a layer <strong>of</strong> fresher water that occupied the upper 15" <strong>of</strong> the water column. Three more Eperetmus<br />
typus medusae were collected in Fairmount Bay, Prince William Sound, in vertical plankton tows<br />
taken <strong>of</strong>f the side <strong>of</strong> the Kristina with Jeff Cordell's 130 µm mesh plankton net in about 70- 90 feet <strong>of</strong><br />
water.<br />
Remarks. These 8-12 mm diameter immature specimens were very lively swimmers, that sank rapidly<br />
when they were not swimming. Both locations were protected coves and in both cases the animals may<br />
have been fairly near the bottom, but not enough is known to verify whether this species is indeed<br />
typical <strong>of</strong> protected coves and associated with the bottom in the same way as Gonionemus or<br />
Polyorchis.<br />
Euphysa sp.<br />
Distribution. Three small Euphysa sp. medusa were collected in Fairmount Bay, Prince William<br />
Sound, in vertical plankton tows taken <strong>of</strong>f the side <strong>of</strong> the Kristina with Jeff Cordell's 130 µm<br />
mesh plankton net in about 70- 90 feet <strong>of</strong> water on August 10, 1999.<br />
Remarks. These 1.5–3.5 mm high medusae were found in the lab by Jeff Cordell in his plankton tow<br />
material. The two smaller specimens clearly had 3 larger tentacles and either one small tentacle or one
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 9<br />
bare bulb. The larger specimen had 4 tentacles. There is insufficient information to assign them to<br />
species.<br />
Eutonina indicans<br />
Distribution. Six Eutonina indicans medusae were collected in the Homer Marina, Katchemak Bay,<br />
Cook Inlet on August 8, 1999.<br />
Remarks. I am surprised that we did not find more <strong>of</strong> this species.<br />
Gonionemus vertens<br />
Distribution. More than 30 Gonionemus vertens medusae were seen in a dense eelgrass bed at low tide<br />
in front <strong>of</strong> Tatitlek, near the small town marina, on August 12, 1999.<br />
Remarks. These medusae emerged from within blades <strong>of</strong> eelgrass in the low intertidal as the tide came<br />
in. They were abundant. The pigmentation was less colorful and more brownish that specimens in the<br />
San Juan Islands. This species was previously not known north <strong>of</strong> Sitka (where it is mentioned by<br />
Ricketts and Calvin, 1939), except that probably the same species (as G. agassizii) was collected early<br />
this century by Trevor Kincaid in a salt lake on Unalaska Island, in the Aleutians (Murbach and<br />
Shearer, 1903). The same or a very similar species also occurs in Japan and the Russian Far East.<br />
Halitholus sp.<br />
Distribution. Three Halitholus sp. medusae were collected at the Tatitlek commercial (ferry) dock on<br />
August 11, 1999, in vertical plankton tows taken <strong>of</strong>f the side <strong>of</strong> the Kristina with Jeff Cordell's 130 µm<br />
mesh plankton net in about 50 feet <strong>of</strong> water. Two more <strong>of</strong> these medusae were seen using a flashlight,<br />
but not collected, later the same evening from the small Tatitlek town marina at about midnight.<br />
Remarks. These 7-8 mm high specimens cannot be referred to any described species <strong>of</strong> Halitholus.<br />
They are very similar to both Halitholus sp. I and Halitholus sp. II <strong>of</strong> Arai and Brinckmann-Voss (1980,<br />
pp. 48-52), which were previously known from British Columbia and Washington State.<br />
Leuckartiara sp.<br />
Distribution. Four Leuckartiara sp. medusae were collected at the head <strong>of</strong> Sadie Cove,<br />
Katchemak Bay, on August 8, 1999, by dipping from a small boat in about 8 feet <strong>of</strong> water. All<br />
were below a layer <strong>of</strong> fresher water that occupied the upper 15" <strong>of</strong> the water column. Several<br />
others were seen, but not collected.<br />
Remarks. These approximately 15 mm-high specimens cannot be referred to any described<br />
species <strong>of</strong> Leuckartiara. They bear some resemblance to Leuckartiara foersteri <strong>of</strong> Arai and<br />
Brinckmann-Voss (1980, pp. 52-53), which is known from British Columbia and Washington<br />
State. They had 8 large tentacles and 12 small tentacles, with no additional rudimentary marginal<br />
bulbs.<br />
Melicertum octocostatum<br />
Distribution. About ten Melicertum octocostatum medusae were found in the Homer Marina,<br />
Katchemak Bay, Cook Inlet, Fairmount Bay and the Cordova Marina.
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 10<br />
Remarks. This species is well known elsewhere in Alaska as well as in the North Pacific and Atlantic; it<br />
probably occurs throughout Prince William Sound. These specimens were relatively small, all being<br />
under 12 mm in bell height.<br />
Mitrocoma cellularia<br />
Distribution. Homer Marina, Katchemak Bay, Cook Inlet on August 8, 1999.<br />
Remarks. Only a single small (15 mm diameter) specimen was collected. This specimen was only<br />
provisionally identified as M. cellularia until it was compared with a comparable-sized living M.<br />
cellularia in Friday Harbor. The small and large tentacles on the margin are the same, confirming the<br />
species identification.<br />
Obelia longissima (Pallas, 1766)<br />
Distribution. Various sites in Prince William Sound, summer 1998.<br />
Remarks. John Chapman sent all <strong>of</strong> the hydroids he collected in 1998 to Claudia Mills, who passed<br />
them on to Dr. Wim Vervoort <strong>of</strong> the Natural History Museum at Leiden, the Netherlands. Dr. Vervoort<br />
identified all <strong>of</strong> the hydroids that he saw as Obelia longissima. John Chapman has both the hydroids and<br />
their specific collection information.<br />
Obelia spp. hydroids<br />
Distribution. These hydroids were collected at least at the Seward Marina and at Lowell Point on<br />
August 10 and 11, 1999.<br />
Remarks. In my inexpert opinion, the blackened portions <strong>of</strong> some <strong>of</strong> the stems imply that these were<br />
probably Obelia longissima. I originally guessed that they might be Garveia franciscana, but they are<br />
not. They should be inspected by a hydroid specialist.<br />
Proboscidactyla flavicirrata<br />
Distribution. The hydroid <strong>of</strong> P. flavicirrata was found at the distal tips <strong>of</strong> several, 6 cm-long sabellid<br />
worm tubes in the Homer Marina, Katchemak Bay, Cook Inlet on August 8, 1999. This hydroid was<br />
actively producing medusa buds although no medusae were seen in this marina. Several P. flavicirrata<br />
medusae were collected in Fairmount Bay, Tatitlek and the Cordova Marina.<br />
Remarks. These medusae are very small; they probably occur throughout Prince William Sound.<br />
Sarsia/Coryne sp. hydroids<br />
Distribution. One or more clumps <strong>of</strong> hydroids that looked like Sarsia were collected by Jeff Goddard in<br />
the Homer Marina, Katchemak Bay, Cook Inlet on August 8, 1999.<br />
Remarks. I did not look carefully at this material. If it was reproductive and making medusa buds, it can<br />
be assigned to Sarsia; if it was reproductive and bearing fixed gonophores, it could be assigned to<br />
Coryne. Sarsia hydroids cannot usually be identified to species without their mature medusae.<br />
Sarsia spp. medusae
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 11<br />
Distribution. About ten Sarsia medusae were collected from Sadie Cove and the Homer Marina,<br />
(Katchemak Bay, Cook Inlet) and Fairmount Bay and Busby Island.<br />
Remarks. Sarsia is a typical north-boreal hydrozoan genus. Many conspecific Sarsias are known from<br />
the Puget Sound / Strait <strong>of</strong> Georgia region and the entire life cycle - both hydroid and mature medusa - is<br />
usually needed for identification to species. Most <strong>of</strong> the medusae collected had the apical canal above<br />
the manubrium that is seen in Sarsia princeps and looked quite a bit like those pictured as S. princeps by<br />
Bigelow (1920), although they were rather small for that species. A second species seemed to also be<br />
present.<br />
Staurophora mertensii<br />
Distribution. This species (5 medusae) was collected by Bigelow (1913) in Prince William Sound - no<br />
further site description or date given.<br />
Remarks. We did not find it in August 1999.<br />
Tiaropsis multicirrata<br />
Distribution. Several Tiaropsis multicirrata were collected at station PWS 98-21 on June 24, 1998. John<br />
Chapman identified this site as Green Island, near Montague Island, on the south side <strong>of</strong> Prince William<br />
Sound.<br />
Remarks. Specimens sent to Claudia Mills for identification, summer 1998.<br />
SCYPHOZOA<br />
Aurelia labiata<br />
Distribution. Only two Aurelia labiata were seen, in the Cordova Marina, on August 13, 1999.<br />
Remarks. I would have expected to see this species or its northern congener Aurelia limbata (similar, but<br />
with a brown rim and tentacles) in many more locations. Aurelia tends to occur in dense aggregations at<br />
the surface - such "swarms" were described to me by resident kayakers as present in Sheep Bay near<br />
Cordova and at Long Bay <strong>of</strong>f Culross Passage near Whittier, but we did not see them. See Wrobel and<br />
Mills (1998) for a discussion <strong>of</strong> the differences between A. aurita and A. labiata. Purcell (1998) refers to<br />
the Prince William Sound species as A. aurita, but probably without knowing about the recently<br />
rediscovered A. labiata name.<br />
Cyanea capillata<br />
Distribution. Cyanea capillata was probably the most common medusa in Prince William Sound<br />
in August 1999. We saw it in the Homer Marina and Sadie Cove in Cook Inlet, as well as at the<br />
Whittier Marina, en route at the south end <strong>of</strong> Esther Island and south <strong>of</strong> Eaglek Bay, in<br />
Fairmount Bay, at Tatitlek and in the Cordova Marina. A.J. Paul told me that it is common in<br />
Resurrection Bay, but further out than the town <strong>of</strong> Seward.<br />
Remarks. In Prince William Sound this species comes in a range <strong>of</strong> colors, from red to pink or<br />
lilac, to yellowish, to a colorless "white". The size ranged from a couple <strong>of</strong> cm to about 30-40 cm<br />
in bell diameter. It was abundant in open water.
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 12<br />
Unidentified scyphozoan polyps<br />
Distribution. Jeff Goddard observed scyphistomae on the docks at both Homer and Whittier.<br />
Remarks. Most scyphistomae on docks on the west coast have proven to be those <strong>of</strong> Aurelia spp.<br />
Other species <strong>of</strong> scyphozoan scyphistomae have not been observed in the field and I assume that<br />
they select other types <strong>of</strong> habitats. If these were Aurelia, they may have been either Aurelia<br />
labiata or perhaps Aurelia limbata, which is likely to also occur in the region.<br />
CTENOPHORA<br />
Bolinopsis infundibulum<br />
Distribution. Only one Bolinopsis specimen was seen on the PWS 1999 trip. It was collected on<br />
August 11, 1999, at the Tatitlek commercial (ferry) dock, in vertical plankton tows taken <strong>of</strong>f the<br />
side <strong>of</strong> the Kristina with Jeff Cordell's 130 µm mesh plankton net in about 50 feet <strong>of</strong> water.<br />
Remarks. I do not hesitate to call this specimen Bolinopsis infundibulum, which I have also<br />
collected at Dutch Harbor.<br />
Pleurobrachia bachei<br />
Distribution. Only one Pleurobrachia specimen was seen on the PWS 1999 trip. It was collected on<br />
August 11, 1999, at the Tatitlek commercial (ferry) dock, in vertical plankton tows taken <strong>of</strong>f the side <strong>of</strong><br />
the Kristina with Jeff Cordell's 130 µm mesh plankton net in about 50 feet <strong>of</strong> water.<br />
Remarks. Examination <strong>of</strong> this preserved ctenophore left some question about its species identity.<br />
This animal was fairly contracted in its preserved state, at which point the funnel canal appeared<br />
to be shorter than the pharynx, which is indicative <strong>of</strong> Pleurobrachia pileus (see Bigelow, 1912).<br />
This species name should not be applied lightly, however, to a North Pacific specimen, since all<br />
those collected from British Columbia to California have been identified as Pleurobrachia<br />
bachei. Comparison with 3 living P. bachei <strong>of</strong> the same size (about 7 mm) from Friday Harbor,<br />
revealed that the pharynx/canal ratios in that species to be similar to the preserved specimen<br />
from PWS, so that name is applied here.<br />
MOLLUSCA<br />
Clione limacina<br />
Distribution. A young pteropod collected in the Homer Marina (Cook Inlet) on August 8, 1999 was<br />
probably Clione limacina.<br />
Remarks. The identification was not confirmed by careful microscopic examination. This species is<br />
found in boreal and temperate regions worldwide.<br />
Melibe leonina<br />
Distribution. Several Melibe leonina was observed either swimming in the water column or<br />
attached to kelp at each <strong>of</strong>: Fairmount Bay, Tatitlek and Cordova. In addition, Carmen Field
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 13<br />
informed me that this species is common in Jakal<strong>of</strong> Bay within Katchemak Bay, although we did<br />
not confirm that location.<br />
Remarks. This species was seen swimming well up in the water column at Fairmount Bay and over<br />
eelgrass at Tatitlek. It was attached to laminarian kelp in the marina at Cordova.
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 14<br />
NONINDIGENOUS CNIDARIA, (MOST) ALREADY PRESENT IN SOME<br />
WEST COAST ESTUARIES<br />
that might be positioned to ultimately invade locations in Alaska.<br />
CNIDARIA<br />
HYDROZOA<br />
Bougainvillia muscus (Allman, 1863). Hydroid known on the west coast only from Friday<br />
Harbor, Washington (Mills, 1981, as B. ramosa); possibly the same species <strong>of</strong> Bougainvillia that<br />
is a pest/contaminant in some aquariums in California. Temperature tolerances not known. This<br />
may actually be a complex <strong>of</strong> cryptic species rather than one species (Calder, 1988).<br />
Blackfordia virginica Mayer, 1910. Hydroids and medusae on the west coast known from north<br />
San Francisco Bay and Coos Bay (J. T. Carlton, personal communication); has a wide salinity<br />
and temperature tolerance. Also reported from the Chesapeake Bay and several European and<br />
Asian harbors, and its apparent point <strong>of</strong> origin, the Black Sea. Full temperature tolerances not<br />
known, but most <strong>of</strong> Alaska is north <strong>of</strong> its known distribution.<br />
Cladonema radiatum Dujardin, 1843. Hydroids and tiny medusae abundant in eelgrass community<br />
in Padilla Bay, Washington, not far from Anacortes and Cherry Point oil terminals. Temperature<br />
tolerances not known, but this species is found in numerous locations worldwide, including northern<br />
and Mediterranean Europe, its putative natural range.<br />
Cordylophora caspia (Pallas, 1771). Hydroid known from the mouth <strong>of</strong> the Samish River in<br />
Samish Bay, not far south <strong>of</strong> the Cherry Point oil terminals. Is also found in very low salinity<br />
tributaries to north San Francisco Bay and elsewhere on the west coast. Requires very low<br />
salinity, temperature tolerances not known; assumed to be <strong>of</strong> Ponto-Caspian origin.<br />
Ectopleura crocea (L. Agassiz, 1862). This Atlantic hydroid is probably established in at least<br />
California and British Columbia (see photo attributed to this species in Harbo (1999, p. 32).<br />
Species <strong>of</strong> Tubularia/Ectopleura cannot be positively identified without examining the<br />
reproductive medusoids, which is rarely done by non-specialists.<br />
Maeotias inexspectata Ostroum<strong>of</strong>f, 1896. Medusae on the west coast known from low<br />
salinity tributaries to north San Francisco Bay, seemingly always in salinities less than 15 psu,<br />
maybe to as low as 1-2 psu (Mills and Sommer, 1995). Also known intermittently from the<br />
Chesapeake Bay and several European estuaries, and its apparent point <strong>of</strong> origin, the Black<br />
Sea and Sea <strong>of</strong> Azov. Temperature tolerances not known, but most <strong>of</strong> Alaska is north <strong>of</strong> its<br />
known distribution. This species was newly collected in the Baltic Sea in Estonia in August<br />
1999 (Risto Vainola, pers. comm.).<br />
Moerisia spp. Several species in this genus have been described from a variety <strong>of</strong> widely<br />
separated locations worldwide, including some rivers emptying into north San Francisco Bay<br />
(J.T. Rees, personal communication), in which both polyps and medusae <strong>of</strong> this genus have been<br />
found. Also known from the Chesapeake Bay. It is not clear how many species are involved
Chapt 9C2. Planktonic Cnidaria, Ctenophora, and Pelagic Mollusca, page 9C2- 15<br />
worldwide. Some populations are known to be single-sexed, inplying a single introduction.<br />
Temperature and salinity tolerances not known.<br />
SCYPHOZOA<br />
Aurelia aurita (Linnaeus, 1758). The most commonly-reported species <strong>of</strong> Aurelia worldwide. With<br />
its seemingly highly-transportable sessile polyp, there is some reason to assume that this species was<br />
carried early to many additional locations, although its home range is so-far not defined - genetic<br />
studies are currently underway by several researchers. All Aurelia that I have inspected carefully in<br />
Alaska appear to be Aurelia labiata Chamisso and Eysenhardt, 1821 or Aurelia limbata Brandt, 1835<br />
(the latter species known from the Aleutians and the Bering Sea). A. labiata was originally described<br />
from central California, but seems to range all the way up the North American Pacific coast. It<br />
would not be too surprising to find A. aurita also living on the west coast. A report <strong>of</strong> a geneticallydifferent<br />
population <strong>of</strong> Aurelia (Greenberg et al., 1996) in San Francisco Bay is likely to be such.<br />
Because it is more amenable to culture, most public aquariums on the west coast have Aurelia aurita<br />
on display, providing a possible source <strong>of</strong> introduction. The name Aurelia aurita has been rather<br />
indiscriminately applied in the literature, including on the west coast <strong>of</strong> North America, without<br />
careful morphological inspection.<br />
ANTHOZOA<br />
Diadumene lineata Merrill, 1870. Cryptogenic sea anemone known from several locations in<br />
Washington and California, as well as worldwide. We searched in seemingly appropriate habitat for<br />
this species in several locations including the intertidal at Lowell Point, Seward, but none were found.<br />
Temperature tolerances are not known and might be an issue in Alaska.<br />
Nematostella vectensis Stephenson, 1935. Cryptogenic sea anemone known from quiet, lowsalinity<br />
lagoon habitats in most coastal American states, as well as numerous locations<br />
worldwide. This species was not found during the 1999 Prince William Sound Expedition, but<br />
we may not have encountered the right kind <strong>of</strong> habitat. Temperature tolerances are not known<br />
and might be an issue in Alaska.<br />
CTENOPHORA<br />
Mnemiopsis leidyi A. Agassiz, 1865. This is apparently the only species <strong>of</strong> ctenophore known to have<br />
invaded a marine habitat outside <strong>of</strong> its home range (the Black Sea). This genus, whose 3 putative<br />
species are not entirely resolved taxonomically, is native to the eastern coast <strong>of</strong> North America,<br />
extending from New England well into Argentina. It has not been found yet in the Pacific Ocean.<br />
NOTE: Many ctenophores are assumed to have very broad global distributions, and it is not known at<br />
this time to what extent, if any, their ranges have been extended artificially by man.
Chapt 9C3. Polychaete Worms, page 9C3- 1<br />
Chapter 9C3. Focal Taxonomic Collections: Polychaete Worms<br />
Jerry Kudenov, Department <strong>of</strong> Biology, University <strong>of</strong> Alaska, Anchorage<br />
Summary<br />
Nearly all <strong>of</strong> the Prince William Sound samples <strong>of</strong> polychaetes collected during the 1998<br />
Expedition have been examined. Certain taxa, such as the Spirorbinae, have not yet been<br />
identified. Excluding the latter subfamily, 61 species have been tentatively identified and<br />
partially described. In essence, there appear to be no clearly defined species that could be listed<br />
as a NIS. However, at least five species may be new to science (species <strong>of</strong> Eumida, Scoloplos,<br />
Exogone, Nephtys and Glycera). At least three new range extensions may be noted for<br />
Phyllodoce medipapillata, Chaetozone senticosa and Rhynchospio glutaea. Finally, six species<br />
that have widespread distributions in the northern hemisphere are represented in the present<br />
material, including: Pholoe minuta, Eteone longa, Barantolla americana, Harmothoe imbricata,<br />
Capitella capitata and Amphitrite cirrata. The systematics <strong>of</strong> each <strong>of</strong> these species is terribly<br />
confused and precise identifications are impossible to render presently. For example, Eteone<br />
longa was originally described from Greenland (1780), and has since been reported from<br />
numerous localities in the Arctic, Atlantic and Pacific Oceans where it appears to be<br />
phenotypically “identical” wherever it occurs. Of course, this is likely not true since the<br />
distributions <strong>of</strong> most species are restricted spatially and temporally. Resolving such dilemmas<br />
falls outside the scope <strong>of</strong> this study, and these six species are therefore identified as above,<br />
pending future revisions. Although all identifications are reasonably precise and non-indigenous<br />
species are represented in these samples, all results are based on literature descriptions and are<br />
preliminary; present materials must be more carefully compared to known reference specimens.<br />
References<br />
Agassiz, A. 1863. On alternate generations in the Annelida and the embryology <strong>of</strong> Autolytus<br />
cornutus. Journal <strong>of</strong> the Boston Society <strong>of</strong> Natural History series 3, 7:384-409.<br />
Annenkova, N.P. 1934. Kurze Übersicht der Polychaeten der Litoralzone der Bering-Insel<br />
(Kommandor-Inseln), nebst Beschreibung neuer arten. Zoologischer Anzeiger 106: 322-331.<br />
Audouin, J.V and H. Milne-Edwards. 1834. Classification des Annelides, et description de celles<br />
qui habitent les cotes de la France. Ann. Sci. Nat. Paris 28: 187-247.<br />
Berkeley, E. 1927. Polychaetous annelids from the Nanaimo District. 3. Leodicidae to Spionidae.<br />
Contr. Canad. Biol. Ottawa, n.s. 3: 405-422.<br />
Berkeley, E. & C. Berkeley. 1938. Notes on Polychaeta from the coast <strong>of</strong> western Canada. 2.<br />
Syllidae. Ann. Mag. Nat. Hist. London, ser. 11, 1:33-49.<br />
Berkeley, E. & C. Berkeley. 1942. North Pacific Polychaeta, chiefly from the west coast <strong>of</strong><br />
Vancouver Island, Alaska and Bering Sea. Can. J. Res. Ottawa 20: 183-208.<br />
Blake, J.A. 1996. Chapter 8. Family Cirratulidae Ryckholdt, 1851, Including a revision <strong>of</strong> the<br />
genera and species from the eastern North Pacific. In, J.A. Blake , B. Hilbig and P. H. Scott (eds.),
Chapt 9C3. Polychaete Worms, page 9C3- 2<br />
Taxonomic Atlas <strong>of</strong> the Benthic Fauna <strong>of</strong> the Santa Maria Basin and Western Santa Barbara<br />
Channel. Santa Barbara Museum <strong>of</strong> Natural History. Volume 6 (The Annelida Part III): 263-384.<br />
Blake, J.A. & K.H. Woodwick. 1971. New species <strong>of</strong> Polydora (Polychaeta: Spionidae) from the<br />
coast <strong>of</strong> California. Bull. So. Calif. Acad. Sci., 70:72-79.<br />
Bush. K.J. 1904. Tubicolous annelid <strong>of</strong> the tribes Sabellides and Serpulides from the Pacific<br />
Ocean. Harriman Alaska Exped., N.Y., 12:169-355.<br />
Chiaje, S. delle 1841. Descrizione e notomie degli animali invertibrati della Sicilia ateriore osservati<br />
vivi neglii anni 1822-1830.<br />
Ehlers, E. 1887. Report on the annelids <strong>of</strong> the dredging expedition <strong>of</strong> the U.S. coast survey steamer<br />
Blake. Memoirs <strong>of</strong> the Museum <strong>of</strong> Comparative Zoology, Harvard 15:1-335.<br />
Grube, A.E. 1840. Actinien, Echinodermen und Würmen des Atriatischen und Mittelmeers. J.H.<br />
Bon. Königsberg, pp. 61-88.<br />
Grube, A.E. 1863. Beschreibung neuer oder wenig bekannter Anneliden. Arch. Naturg. Berlin 29:<br />
37-69.<br />
Fabricius, O. 1780. Fauna Groenlandica, systematice sistens, Animalia Groenlandiae occidentalis<br />
hactenus indagata, quoad nomen specificum, triviale, vernaculumque; synonym auctorum plurium,<br />
descriptionem, locum, victum, generationem. mores, usum, capturamque singuli; prout detegendi<br />
occasio fuit, maximaque parti secundum proprias observationes. Hafniae et Lipsiae. xvi and 452 pp.<br />
Harrington, N.R. 1897. On nereids commensal with hermit crabs. Transactions <strong>of</strong> the New York<br />
Academy <strong>of</strong> Sciences 16:214-221.<br />
Hartman, O. 1938. Review <strong>of</strong> the annelid worms <strong>of</strong> the family Nepthyidae from the northeast<br />
Pacific, with descriptions <strong>of</strong> five new species. Proc. U.S. Nat. Mus., 85: 143-158.<br />
Hartman, O. 1961. Polychaetous annelids from Californa. Allan Hancock Pac. Exp., 25: 1-226.<br />
Hartman, O. 1963. Submarine canyons <strong>of</strong> southern California. Pt. 3. Systematics: Polychaetes.<br />
Allan Hancock Pac. Expeds., 27(3):1-93.<br />
Healy, E.A. and G.P. Wells. 1959. Three new lugworms (Arenicolidae, Polychaeta) from the north<br />
Pacific area. Proceedings <strong>of</strong> the Zoological Society <strong>of</strong> London 133:315-355.<br />
Hobson, K.D. and K. Banse. 1981. Sedentariate and archiannelid polychaetes <strong>of</strong> British Columbia<br />
and Washington. Canadian Bulletin <strong>of</strong> Fisheries and <strong>Aquatic</strong> Sciences 209:1-144.<br />
Jacobi, R. 1883. Anatomisch-histogische Untersuchung der Polydoren der Keiler Bucht. Inaugural<br />
Dissertation, Keil:1-35.
Chapt 9C3. Polychaete Worms, page 9C3- 3<br />
Johnson, H.P. 1897. A preliminary account <strong>of</strong> the marine annelids <strong>of</strong> the Pacific coast, with<br />
descriptions <strong>of</strong> new species. Euphrosynidae, Amphinomidae, Palmyridae, Polynoidae and<br />
Sigalionidae. Proc. Calif. Acad. Sci., 1:153-190.<br />
Johnson, H.P. 1901. The Polychaeta <strong>of</strong> the Puget Sound region. Proc. Boston Soc. Nat. Hist.,<br />
29:381-437.<br />
Johnston, G. 1840. British Annelids. Ann. Mag. Nat. Hist. London, ser. 1, 4: 368-375.<br />
Kinberg, J.G.H. 1867. Annulata nova. Öfversight af Kungliga Vetenskaps-Akademiens<br />
Förhandlingar, Stockholm 23: 337-357.<br />
Kudenov, J.D. and L. H. Harris. 1995. Chapter 1. Family Syllidae Grube, 1850. In, J.A. Blake and<br />
B. Hilbig (eds.), Taxonomic Atlas <strong>of</strong> the Benthic Fauna <strong>of</strong> the Santa Maria Basin and Western<br />
Santa Barbara Channel. Santa Barbara Museum <strong>of</strong> Natural History. Volume 5 (The Annelida Part<br />
II): 1-97.<br />
Linnaeus, C. 1767. Systema naturae. 12th ed.<br />
Malmgren, A.J. 1866. Nordiska Hafs-Annulater. Öfversight af Kungliga Vetenskaps-Akademiens<br />
Förhandlingar, Stockholm 22: 355-410.<br />
Malmgren, A.J. 1867. Annulata Polychaeta Spetsbergiae, Groenlandiae, Islandiae et Scandinaviae<br />
hactenus cognita. Öfversight af Kungliga Vetenskaps-Akademiens Förhandlingar, Stockholm 24:<br />
127-235.<br />
Mesnil, F. 1896. Études de morphologie externe chez les annélides. Les Spionidiens des cotes de la<br />
Manche. Bulletin Scientifique de la France et de la Belgique 29: 110-287.<br />
Moore, J.P. 1908. Some polychaetous annelids <strong>of</strong> the northern Pacific coast <strong>of</strong> North America. Proc.<br />
Acad. Nat. Sci. Phila., 60:321-364.<br />
Moore, J.P. 1909. Polychaetous annelids from Monterey Bay and San Diego, California. Proc.<br />
Acad. nat. Sci. Phila., 61: 235-295.<br />
Moore, J.P. 1911. The polychaetous annelids dredged by the U.S.S. Albatross <strong>of</strong>f the coast <strong>of</strong><br />
southern California in 1904. Euphrosynidae (sic) to Goniadidae. Proc. Acad. Nat. Sci. Phila., 63:<br />
234-318.<br />
Müller, O.F. 1771. Von Würmern des sussen und salzigen Wassers. Copenhagen, Heinich Mumme<br />
und Faber. 200 pp.<br />
Müller, O.F. 1776. Zoologica Danicae Prodromus seu Annimalium Daniae et Norvegiae<br />
indigenarum characters, nomine et synonyma imprimis populaium. Havniae. 274 pp.
Chapt 9C3. Polychaete Worms, page 9C3- 4<br />
Oersted, A.S. 1843. Groenlandiae Annulata dosibranchiata. K. Danske Videns. naturw. math-Afh.<br />
Copenhagen, 10: 153-216.<br />
Pettibone, M.H. 1957. North American genera <strong>of</strong> the family Orbiniidae. Journal <strong>of</strong> the Washington<br />
Academy <strong>of</strong> Sciences 47: 159-167.<br />
Rathke, H. 1843. Beitrage zur Fauna Norwegens. Nova Acta Acad. Leop. Carol. Nat. Cur. Halle,<br />
20: 1-264.<br />
Schmarda, L.K. 1861. Neue wirbellose Thiere beobachet und gesammelt auf einer Reise um die<br />
Erde 1853 bis 1857. a. Turbellarien, Rotatorien und Annelidien. Part 2: 1-164.<br />
Zachs, I.G. 1933. Polychaeta <strong>of</strong> the North-Japanese Sea. Explorations <strong>of</strong> the Seas <strong>of</strong> the<br />
USSR 19:125-137. (In Russian).<br />
Table 9C3.1 POLYCHAETA, PRINCE WILLIAM SOUND, SUMMER 1998<br />
SIGALIONIDAE<br />
Pholoe minuta Fabricius, 1780<br />
PHYLLODOCIDAE<br />
Eteone longa (Fabricius, 1780)<br />
Eulalia bilineata (Johnston, 1840)<br />
Eumida species A (new species)<br />
Phyllodoce medipapillata Moore, 1909<br />
Phyllodoce species<br />
NEREIDIDAE<br />
Chelonereis cyclurus Harrington, 1897<br />
Platynereis species (bicanaliculata)<br />
CAPITELLIDAE<br />
Barantolla americana Hartman, 1963<br />
GONIADIAE<br />
Glycinde picta Berkeley 1927<br />
GLYCERIDAE<br />
Glycinde armigera Moore, 1911
Chapt 9C3. Polychaete Worms, page 9C3- 5<br />
ORBINIIDAE<br />
Scoloplos species A New Species<br />
POLYNOIDAE<br />
Harmothoe imbricata (Linnaeus, 1767)<br />
Harmothoe extenuata (Grube, 1840)<br />
Harmothoinae<br />
Lepidonotinae<br />
CHRYSOPELATIDAE<br />
Chrysopetalum occidentale Johnson, 1897<br />
SYLLIDAE<br />
Exogone cf. dwisula Kudenov & Harris, 1995<br />
Sphaerosyllis cf. californiensis Hartman, 1961<br />
Trypanosyllis gemmipara Johnson, 1901<br />
Eudontosyllis species A<br />
Tyosyllis alternata (Moore, 1908)<br />
Typosyllis hyalina Grube 1863<br />
Typosyllis pulchra Berkeley & Berkeley, 1938<br />
Typosyllis stewarti Berkeley & Berkeley, 1942<br />
Autoylus (Procerea) cornutus Agassiz, 1863<br />
LUMBRINDERIDAE<br />
Lumbrineris latrielli Audouin & Milne-Edwards, 1834<br />
ORBINIIDAE<br />
Leitoscoloplos pugettensis (Pettibone, 1957)<br />
Naineris dendritica (Kinberg, 1867)<br />
SPIONIDAE<br />
Spio filicornis (Müller, 1776)<br />
OPHELIIDAE<br />
Armandia brevis Hartman, 1938<br />
Ophelia limacina (Rathke, 1843)<br />
NEREIDIDAE<br />
Nereididae (postmetamorphic juvenile)<br />
SPIONIDAE<br />
Prionospio steenstrupi Malmgren, 1867
Chapt 9C3. Polychaete Worms, page 9C3- 6<br />
CIRRATULIDAE<br />
Cirratulus cingulatus Johnson, 1901<br />
Chaetozone senticosa Blake, 1996<br />
CAPITELLIDAE<br />
Capitella capitata (Fabricius, 1780)<br />
NEPHTYIDAE<br />
Nephtys species A (N. ciliata)<br />
Nephtys species A (juvenile)<br />
Nephtys species B (juvenile)<br />
ARENICOLIDAE<br />
Abarenicola pacifica Healy & Wells 1959<br />
OWENIIDAE<br />
Owenia fusiformis della Chiaje, 1841<br />
GLYCERIDAE<br />
Glycera cf. nana Johnson, 1901<br />
SPIONIDAE<br />
Rhynchospio glutaea (Ehlers, 1887)<br />
Prionospio sp.<br />
Dipolydora cf. socialis (Schmarda, 1861)<br />
Dipolydora sp. A (near bidentata) Zachs, 1933<br />
Dipolydora sp. B<br />
Dipolydora sp. C (near giardi (Mesnil, 1896))<br />
Polydora limicola Annenkova, 1934<br />
Diplydora quadrilobata (Jacobi, 1883)<br />
Polydora sp.<br />
MALDANIDAE<br />
Nicomache personata Johnson 1901<br />
PECTINARIIDAE<br />
Pectinaria granulata Johnson 1901<br />
AMPHARETIDAE<br />
Ampharete species A<br />
TEREBELLIDAE<br />
Amphitrite cirrata Müller. 1771<br />
Polycirrus species III Hobson & Banse, 1981
Chapt 9C3. Polychaete Worms, page 9C3- 7<br />
SABELLIDAE<br />
Laonome cf. kroyeri (Malmgren, 1866)<br />
Schizobranchia insignis Bush 1904<br />
SERPULIDAE<br />
Crucigera zygophora (Johnson, 1901)<br />
Serpula vermicularis Linnaeus, 1767<br />
Spirorbis species<br />
Table 3. Polychaeta Collected in 1998 PWS Expedition<br />
Note: Materials listed below as PWS NIS 1998 include Stations followed by the number <strong>of</strong><br />
specimens in parentheses.<br />
Pholoe minuta Fabricius, 1780<br />
PWS NIS 1998: Sta 1(1 specimen); Sta 5(1); 6 (fragments); Sta 7 (2); Sta (3); Sta 10(4); Sta<br />
11(1); Sta 41(3).<br />
Based on specimens, these are not Pholoe minuta sensu Fabricius. Original taxon described as<br />
having papillae disbursed over entire ventral and parapodial surfaces. Specimens all have small,<br />
close-set, short papillae over entire ventral surface; parapodia with conspicuous digitiform<br />
papillae. Whatever “Pholoe minuta” represents, it must be a polyphyletic species at the very<br />
least. It has a recorded distribution in both the Arctic and south Atlantic Oceans. This species is<br />
correctly identified to a single taxon. However its actual identity is questionable in view <strong>of</strong> its<br />
widespread distribution.<br />
Eteone longa (Fabricius, 1780)<br />
PWS NIS 1998: Sta 3(1 specimen ); Sta 15(2); Sta 16(3); Sta 17(8); Sta 18(9); Sta 36(1); Sta<br />
41(1).<br />
Technically, the Eteone longa/flava group is in severe disarray and is undoubtedly represents a<br />
complex taxonomic assemblage <strong>of</strong> closely related species. Whatever taxon is represented by<br />
PWS specimens <strong>of</strong> “E. longa” must remain obscure until a definitive study is published.<br />
Eulalia bilineata (Johnson, 1840)<br />
PWS NIS 1998: Sta 21(1 specimen).<br />
Keys out according to Blake (1996) and Pleijel (1991). Only one specimen, which is poorly<br />
preserved used for this identification; therefore it is considered to be tentative.<br />
Eumida species A (new species)<br />
PWS NIS 1998: Sta 21(2 specimens).<br />
Genus identification correct. Prostomium small, with 4 distal and one median unpaired antenna.<br />
Dorsal cirri triangular, pointed, lanceolate. Ventral cirri subtriangular, subtly pointed. Segment<br />
1 highly reduced, not fused to segment 2; in largest specimen it actually extends onto<br />
prostomium (although this may be artifactual); tentacular cirri lateral to prostomium. Segment 2
Chapt 9C3. Polychaete Worms, page 9C3- 8<br />
with two pairs <strong>of</strong> tentacular cirri, ventral pair shortest; lacking setae. Segment 3 with one pair<br />
tentacular cirri, with fascicle <strong>of</strong> setae in neuropodium.<br />
Parapodia all distally rounded, without hint <strong>of</strong> dorsal lobe; all about the same length throughout<br />
the body. Ventral cirri asymmetrical, subquadrangular, longest anteriorly and gradually<br />
decreasing in length, size posteriorly.<br />
Pygidium lacking appendages (lost).<br />
Phyllodoce medipapillata Moore, 1909<br />
PWS NIS 1998: Sta 21(1); 36(1).<br />
Two beautiful specimens, complete, well preserved with proboscides everted. Key out according<br />
to Blake (1994). Present record is a range extension, and potentially also an introduced species<br />
to PWS, assuming there are no other records <strong>of</strong> it between here and central to southern California<br />
(0-300 m) where it seems to be restricted.<br />
Phyllodoce species<br />
PWS NIS 1998: Sta 15(2 specimens)<br />
Juvenile individuals, one <strong>of</strong> which has proboscis everted. Both extremely small, unidentifiable.<br />
Chelonereis cyclurus Harrington, 1897<br />
PWS NIS 1998: Sta 3(3 specimens); Sta 5(4); Sta 6(3); Sta 9(1); Sta 10(1); Sta 11(32); Sta<br />
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);<br />
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<br />
40(1); Sta 41(2); Sta 44(2).<br />
Characteristic species. Notosetae homogomph spinigers. Neurosetae heterogomph spinigers and<br />
falcigers. Largest specimen lacks homogomph falcigers in neuropodia.<br />
Platynereis species (bicanaliculata (Baird, 1863))<br />
PWS NIS 1998: Sta 12(1 specimen).<br />
Juvenile lacking tentacular cirri. This identification is highly tentative, based on a single<br />
specimen!<br />
Nereididae (postmetamorphic juvenile)<br />
PWS NIS 1998: Sta 6(1 specimen).<br />
Identified as “Platynereis” but the specimen is a postmetamorphic juvenile that is not<br />
identifiable to genus.<br />
Glycinde picta Berkeley 1927<br />
PWS NIS 1998: Sta 4(1 specimen ); Sta 7(1); Sta 15(1); Sta 16(2); Sta 41(1).<br />
All with ventral arc <strong>of</strong> micrognaths, characteristic <strong>of</strong> Glycinde picta along Pacific coast <strong>of</strong> North<br />
America.<br />
Glycinde armigera Moore, 1911<br />
PWS NIS 1998: Sta 13(1 specimen ); Sta 23(1).
Chapt 9C3. Polychaete Worms, page 9C3- 9<br />
Identification tentative in light <strong>of</strong> a dissection performed previously and prior to the present<br />
examination that damaged the critical region <strong>of</strong> macro- and micrognaths.<br />
Harmothoe imbricata (Linnaeus, 1776)<br />
PWS NIS 1998: Sta5(2 juveniles); Sta6(2 juveniles); Sta 10(3 specimens); Sta 11(10); Sta<br />
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);<br />
Sta 35(17); Sta 36(9).<br />
Another widespread species that must be re-examined critically. Tentatively assigned to<br />
Harmothoe imbricata.<br />
Harmothoe extenuata (Grube, 1840)<br />
PWS NIS 1998: Sta 35(3 specimens).<br />
Seems to key out well. Some <strong>of</strong> the specimens included as H. imbricata likely identical to this<br />
taxon.<br />
Harmothoinae<br />
PWS NIS 1998: Sta 19(1 specimen); Sta 24N(2).<br />
These are most likely “H. imbricata “ juveniles, and should be referred to above as “”<br />
Lepidonotinae<br />
PWS NIS 1998: Sta 23(1 specimen ); Sta 24N(1); 34(2).<br />
Note that Station 34(2 specimens) contains the best specimens, which seem to key out to<br />
Parhalosydna, which seems somewhat <strong>of</strong> a stretch. Most <strong>of</strong> the specimens are not well<br />
preserved, nearly all lack elytra, and identification can not be made positively.<br />
Chrysopetalum occidentale Johnson, 1897<br />
PWS NIS 1998: Sta 23(1 specimen); 24(1).<br />
Highly characteristic species.<br />
Exogone cf. dwisula Kudenov & Harris, 1995<br />
PWS NIS 1998: Sta 19(1 specimen); Sta 21(1); Sta 23(10); Sta 32(1).<br />
This species is very closely allied to E. dwisula, and also to E. gemmifera. Antennae closely set,<br />
laterals about .67-.75x length <strong>of</strong> median. Pharynx extends through 1.5-2 segments, with anterior<br />
unpaired middorsal tooth. Proventriculus extends through 2 segments, with 15 rows muscle<br />
cells. Peristomial antennae small, inconspicuous, not visible in dorsal view. Setae number 4-5<br />
per parapodium, <strong>of</strong> 3 kinds: a) falcigers with deeply incised blades, confined to anterior setigers;<br />
b) stout awl-shaped spinigers, numbering 1-2 per anterior parapodium, 1 per median and<br />
posterior parapodia; c) dorsal and ventral simple seta, the former present in all setigers, the latter<br />
in the last few setigers. Aciculae numbering 1 per parapodium, all terminating in distally<br />
enlarged heads (blunt or beaked)<br />
Sphaerosyllis cf. californiensis Hartman, 1961<br />
PWS NIS 1998: Sta 10(1 specimen); Sta 19(1); Sta 19(11); Sta 21(2); Sta 21(2); Sta 23(65);<br />
Sta 23(5).<br />
One difference between these specimens and those examined by Kudenov & Harris (1995) is the<br />
presence <strong>of</strong> an additional pair <strong>of</strong> conspicuous papillae on distal parapodial surfaces; one is
Chapt 9C3. Polychaete Worms, page 9C3- 10<br />
anterior, the other posterior. A most unusual aspect to the setal morphology is the fact that the<br />
cutting teeth on blades <strong>of</strong> compound falcigers are set in 2 rows, members <strong>of</strong> one row alternating<br />
with those <strong>of</strong> the other row.<br />
This has not been reported for S. californiensis. Then again, I don’t believe anyone has ever<br />
looked closely enough! These specimens will represent a new species if S. californiensis lacks<br />
these alternating rows <strong>of</strong> teeth on blade cutting surfaces.<br />
Trypanosyllis gemmipara Johnson, 1901<br />
PWS NIS 1998: Sta 23(1 specimen).<br />
One small specimen, 61 segments. Bidentate falcigers. Trepan with 10 teeth; middorsal tooth<br />
absent.<br />
Eudontosyllis species A<br />
PWS NIS 1998: Sta 19(1 specimen); 23(1).<br />
Only 2 specimens, both anterior fragments. Specimen <strong>of</strong> Sta 19 in 2 pieces, with dorsal cirri; Sta<br />
23 in 1 piece, lacking dorsal cirri. Specimens key out to Eudontosyllis Knox 1960, which<br />
according to Fauchald (1977) is represented by a single species.<br />
Essential descriptive elements include: Prostomium reduced, with 2 pairs <strong>of</strong> large lenticulate;<br />
eyes Palps reduced, fused only basally. Paired occipital nuchal organs extending over setiger 1,<br />
not fused to dorsum; Antennae very long, smooth basally, terminating in a few distal moniliform<br />
elements, each long and cylindrical; Peristomial tentacles long, number 1 pair; Notoacicula<br />
present, each conspicuous, with distally bent tips; Notosetae as multispinose capillaries in small<br />
inconspicuous tufts. Neurosetal fascicles with bidentate compound falcigers.<br />
The one specimen with notosetal fascicles may be epitokous (Sta 23). Need to check out the<br />
specimen from Sta 19 for comparison.<br />
Tyosyllis alternata (Moore, 1908)<br />
PWS NIS 1998: Sta 5(1 specimen); 9(1); Sta10(1, juvenile).<br />
Typosyllis hyalina Grube 1863<br />
PWS NIS 1998: Sta 21(1 specimen).<br />
Typosyllis pulchra Berkeley & Berkeley, 1938<br />
PWS NIS 1998: Sta 21(1 specimen); 24(1).<br />
Typosyllis stewarti Berkeley & Berkeley, 1942<br />
PWS NIS 1998: Sta 21(5 specimens); 21(1); 23(1).<br />
Characteristic increase in thickness <strong>of</strong> falcigers in posterior segments. Many <strong>of</strong> these have lost<br />
their blades.<br />
Autoylus (Procerea) cornutus Agassiz, 1863<br />
A.cornutus Okada, 1933:645-647, figs. 3,4; Pettibone, 1963:144, fig. 37e.<br />
A.cornatus Hartman, 1944:338, pl. 13, fig. 5.
Chapt 9C3. Polychaete Worms, page 9C3- 11<br />
A. (Regulatus) cornutus, Imajima, 1966:49-51, Text-fig. 13a-i.<br />
PWS NIS 1998: Sta 19(1 specimen); Sta 20(2); Sta 25(1).<br />
Only two specimens include in these samples. Specimen (Sta. 19) is small, lacking tentacular<br />
and dorsal cirri. Dorsal cirri <strong>of</strong> setiger 1 longest; those from setiger 2 all shorter and about the<br />
same size. Both asexual forms exhibiting stolons between segments 13-14. Trepan with 18<br />
teeth: 9 larger and 9 smaller. Dorsal simple setae thick, distally truncate and serrated. Nuchal<br />
organs restricted to posterolateral regions <strong>of</strong> prostomium; not extended to posterior margin <strong>of</strong><br />
setiger 1.<br />
The species was originally reported from Atlantic habitats (Labrador to Chesapeake Bay;<br />
Plymouth) and has also been reported from Japan to British Columbia-Washington.<br />
Specimens (Sta. 20) are sexual forms for which only the genus is a certain identification.<br />
Swarming or sexually swimming stages have not been related to asexual phases, unfortunately,<br />
along the Pacific coast <strong>of</strong> North America (or most other places, except see Gidholm 1965, 1966).<br />
Nephtys species A (N. ciliata)<br />
NIW NIS 1998: Sta 10(1 specimen); Sta11(1); Sta 15(3); Sta 16(1); Sta 16(4); Sta 36(5).<br />
This appears to be a new species. It does not key out to anything in Banse & Hobson (1974)<br />
where, in the key, this species drops out <strong>of</strong> the key at couplet 6 (page 73). The couplet provides<br />
a choice between large, postsetal notopodial lobes without a middorsal proboscideal papilla<br />
versus medium-sized postsetal notopodial lobes with out without a dorsal median proboscideal<br />
papilla.<br />
To couplet 9, the next choice is interramal cirri, proboscis with unpaired dorsal papilla, which<br />
leads to a choice between N. ciliata or N. caecoides.<br />
It is not N. caecoides. Key leads to N. ciliata which lacks a dorsal pigment pattern. Notopodial<br />
postsetal lobe partly covered by acicular lobe, which, in N. caeca is large, but is relatively small,<br />
compared to the postsetal notopodial lobe. Proboscis proximally with small warts.<br />
In all, this appears to be N. ciliata. One principal difference appears to be the size <strong>of</strong> the<br />
postsetal notopodial lobes.<br />
Specimen 11(1) poorly preserved; assignment tentative.<br />
Nephyts species A (juvenile)<br />
NIW NIS 1998: Sta 15(7 specimens); Sta 17(1); Sta 17(1).<br />
All specimens are postmetamorphic or young juveniles and are unidentifiable to species. Note<br />
that all have conical acicular lobes and poorly defined postsetal lamellae. Specimen (17(1))<br />
obviously a newly postmetamorphic juvenile; assigned to this taxon for convenience.<br />
Nephyts species B (juvenile)<br />
NIW NIS 1998: Sta 11(3 specimens).
Chapt 9C3. Polychaete Worms, page 9C3- 12<br />
Interramal cirri short, almost straight except for distally curved tip, hanging almost straight<br />
down. Interramal cirri beginning from setiger 5. Acicular lobes generally rounded although<br />
notopodial lobe slightly bilobed; neuropodial lobe more evenly rounded.<br />
Glycera cf. nana Johnson, 1901<br />
NIW NIS 1998: Sta 10(1 specimen); Sta 11(1); Sta 16(1); Sta 24(3); Sta 36(1).<br />
These specimens are mighty peculiar! Postsetal lobes are rounded, with biramous parapodia as<br />
per Glycera. Ailerons winged as per Glycera. Proboscis with 3 kinds <strong>of</strong> papillae including long,<br />
slender and shorter tapering forms plus spherical papillae. Inferior presetal lobe pointed,<br />
appearing rather different from that for Glycera nana.<br />
Hilbig (1994) describes Glycera nana in terms that, compared to the present materials, intimates<br />
that the PWS specimens are sufficiently different to represent a new species.<br />
Lumbrineris latrielli Audouin & Milne-Edwards, 1834<br />
PWS NIS 1998: Sta 5(2 specimens); Sta 11(1).<br />
Keys out according to both Banse & Hobson (1974) and also Ruff (1995), particularly in view <strong>of</strong><br />
the latter’s comments. Specimen (Sta. 11) with dental formula: 1+1, 5+4, 2+2, 1+1. Yellow<br />
aciculae. Compound falcigers in anterior segments. Posterior pre- and postsetal lobes not<br />
elongate. No obvious pigmentation patterns in preserved specimens.<br />
Scoloplos species A New Species<br />
As Scoloplos armiger PWS NIS 1998: Sta 4(2 specimens); Sta 10(4); Sta 15(2); Sta 36(1); Sta<br />
41(22).<br />
As Scoloplos species PWS NIS 1998: Sta 7(1 specimen); Sta 10(3).<br />
As Orbiniidae PWS NIS 1998: Sta 10(4 specimens); Sta 41(1).<br />
All <strong>of</strong> these individuals represent a new taxon. There are no subpodial lobes present whatsoever.<br />
Number <strong>of</strong> thoracic segments numbering 14-15. Branchiae from posterior thoracic segments.<br />
Thoracic neurosetae with distally smooth, transparent hoods. Abdominal neurosetae include<br />
both capillaries and delicate spines. Neuropodial lobes in larger specimens digitiform; smaller<br />
specimens notched. These lobes are clearly different from those <strong>of</strong> both S. armiger and S.<br />
acmeceps.<br />
Leitoscoloplos pugettensis (Pettibone, 1957)<br />
PWS NIS 1998: Sta 5(1 specimen); Sta 15(2); Sta 17(14).<br />
Specimens correctly identified to species. No abdominal subpodial lobes present.<br />
Naineris dendritica (Kinberg, 1867)<br />
PWS NIS 1998: Sta 17(36 specimens).<br />
Agrees well with descriptions. Originally identified as N. quadricuspida on label, however, only<br />
one record can be assigned to this species, and even then, Hartman (1961) noted distinct and<br />
significant differences between her material compared to those described by Fabricius. In other<br />
words, N. quaricuspida does not occur on this coast!
Chapt 9C3. Polychaete Worms, page 9C3- 13<br />
Spio filicornis (Müller, 1776)<br />
PWS NIS 1998: Sta 5(2 specimens); Sta 10(1); Sta 15(12); Sta 17(1).<br />
Keys out according to Blake (1996).<br />
Armandia brevis Hartman, 1938<br />
PWS NIS 1998: Sta 5(3 specimens); Sta 7(3); Sta 10(5); Sta 11(12); Sta 11(1*); Sta 15(4).<br />
Everything keys out extremely well to this taxon following Hartman (1969). Need to check out<br />
the validity <strong>of</strong> this genus based on Colin Herman’s comments a few years ago. Specimen (Sta.<br />
11(1*)) is poorly preserved, has a pair <strong>of</strong> prostomial eyespots, and hints <strong>of</strong> lateral eyespots, and<br />
is taken here to represent Armandia brevis.<br />
Ophelia limacina (Rathke, 1843)<br />
PWS NIS 1998: Sta 5(1 specimen).<br />
This is a supposedly cosmopolitan species. It has 37 setigers compared to 39 originally<br />
described. First 10 setigers abranchiate. Ventral groove present from around setiger 10-11.<br />
Prionospio steenstrupi Malmgren, 1867<br />
PWS NIS 1998: Sta 6(1 specimen); Sta 9(3); Sta 11(10).<br />
Specimens poorly preserved, and trashed in most cases. Gills not well intact, and in a few<br />
specimens (Sta. 11) they are <strong>of</strong> variable lengths. The neuropodial lamella <strong>of</strong> setiger 2 with the<br />
characteristic ventral protuberance; those <strong>of</strong> setiger 3 squarish to ventrally pointed also.<br />
Identification tentative pending additional specimens.<br />
Cirratulus cingulatus Johnson, 1901<br />
PWS NIS 1998: Sta 6(1 specimen); Sta 11(1); Sta 16(2); Sta 17(8) + Sta 17(3); Sta 24(3); Sta<br />
24N(1).<br />
Specimens agree fairly well with description provided by Blake (1996:350-351). Neurosetal<br />
spines begin from setigers 23-26; notosetal spines from setigers 35-37. Specimen from Sta 24N<br />
may be a juvenile, with neurosetal spines from setiger 9. and notosetal spines from setiger 16.<br />
Eyes present in all specimens as line <strong>of</strong> 4-6 individual eyespots. The three specimens from Sta<br />
17 are very large and show size-dependent morphology concerning transverse band <strong>of</strong><br />
tentacles/cirri.<br />
Chaetozone senticosa Blake, 1996<br />
PWS NIS 1998: Sta 15(9 specimens); Sta 17(57).<br />
Keys out according to Blake (1996), although the final identification needs to be confirmed<br />
based on methyl green. Specimens with around 60-70 setigers, hooks beginning from around<br />
setiger 35-40. Prostomium short, triangular, with a single achaetous annulus.<br />
Originally reported from Central and Northern California. This may be a range extension,<br />
assuming the identification is valid.<br />
Barantolla americana Hartman, 1963<br />
PWS NIS 1998: Sta 3(1 specimen); Sta 4(1); Sta 9(2); Sta 15(3); Sta 19(1); Sta 41(1).
Chapt 9C3. Polychaete Worms, page 9C3- 14<br />
This is a dubious taxon. Originally described has having capillary setae only in notosetiger 6<br />
and neurosetiger 7; mixed capillaries in notosetiger 7 and neurosetiger 8; hooks only in<br />
notosetigers 8-11 and neurosetigers 9-11. In contrast, Fauchald (1977) lists Barantolla as<br />
having 6 setigers with capillaries followed by 1 mixed capillaries and hooks, and then 4 more<br />
with hooks only.<br />
The present specimens are at odds with the above descrepancies. Specimen 3(1) with mixed<br />
notosetae on setiger 5; 7(2) with capillaries only in notosetigers 1-5 and neurosetigers 1-6,<br />
mixed setae in notosetiger 6 and neurosetiger 7, and hooks only thereafter in notosetigers 7-11<br />
and neurosetigers 8-11; specimen 41(1) with capillaries only in both noto- and neurosetigers 1-6,<br />
and hooks only in both noto- and neurosetigers 7-11 (setiger with mixed setae apparently<br />
absent).<br />
Capitella capitata (Fabricius, 1780)<br />
NIW NIS 1998: Sta 7(3 specimens); Sta 9(1); Sta 15(9); Sta 41(17).<br />
Whatever Capitella capitata is, these specimens can be assigned to the stem species. But this is<br />
one the “cosmopolitan” species that can be almost anything.<br />
Abarenicola pacifica Healy & Wells 1959<br />
NIW NIS 1998: Sta 7(1 specimen); Sta 17(3); Sta 18(9).<br />
One large specimen (Sta. 7), intact; all others are juveniles. No question concerning identity.<br />
Owenia fusiformis della Chiaje, 1841<br />
NIW NIS 1998: Sta 7(1 specimen); Sta 41(1).<br />
Only two specimens. Have a collar as per Owenia collaris. Setae appear to have configuration<br />
found in Owenia fusiformis. Refer to recent paper on the family from IP4 (Paris).<br />
Rhynchospio glutaea (Ehlers, 1887)<br />
PWS NIS 1998: Sta 10(1 specimen); Sta 10(1).<br />
New record for Alaska. Not particularly surprising.<br />
Prionospio sp.<br />
PWS NIS 1998: Sta 18(12 specimens).<br />
Recheck. One specimen appeared to have gills on middle body segments. These are not<br />
polydorids as noted on label.<br />
Dipolydora cf. socialis (Schmarda, 1861)<br />
PWS NIS 1998: Sta 17(1 specimen).<br />
Tentative identification, but not Dipolydora socialis. Falcate spines <strong>of</strong> setiger 5 strongly falcate,<br />
with hint <strong>of</strong> flange; bristle absent. Setiger 1 postsetal lamellae poorly developed. Gizzard-like<br />
structure present around setigers 17-18, but not as portrayed by Blake (1996).<br />
Dipolydora sp. A (near bidentata) Zachs, 1933<br />
PWS NIS 1998: Sta 17(18 specimens); Sta 19(1); Sta 23(2).<br />
This taxon may be a shell borer. Spines <strong>of</strong> setiger 5 without distal bristles, with flange-tooth on<br />
lateral surface (not on convex surface as fas as I can see). Caruncle to posterior setiger 4.
Chapt 9C3. Polychaete Worms, page 9C3- 15<br />
Notosetae present on setiger 1. Prostomium deeply incised, bifurcate. Posterior notosetae<br />
capillaries; spinous packets, modified setae absent. Neurosetae without manubrium, from setiger<br />
7, bidentate to end <strong>of</strong> body.<br />
Dipolydora. sp B<br />
PWS NIS 1998: Sta 17(1 specimen).<br />
Prostomium incised, strongly bilobed. 4 pairs <strong>of</strong> eyes. Notosetae setiger 1 present, lobe reduced<br />
to papillar lobe. Caruncle to posterior setiger 3. Setiger 5 strongly modified; heavy spines<br />
distally falcate, heavy triangular tooth on concave surface, bristles present in notch between<br />
tooth and tip <strong>of</strong> spine. Bidentate neurosetae without manubria, from setiger 7. Branchiae from<br />
setiger 7.<br />
Does not key out using Blake 1996…falls out at couplet 10 (10B where choice is presence <strong>of</strong> 2<br />
accessory teeth) since this specimen has only 1 visible accessory tooth, and appears to lack a<br />
cowling.<br />
Dipolydora sp. C (near giardi (Mesnil, 1896))<br />
PWS NIS 1998: Sta 21(>100 specimens); Sta30(1).<br />
Numerous specimens. Prostomium incised, bilobed. No eyes. Setiger 1 complete, notosetae<br />
present, notopodium reduced to digitiform lobe. Caruncle to setiger posterior margin setiger 3.<br />
Setiger 5 modified; spines with accessory tooth on concave surface, with partial cowling on<br />
opposite side <strong>of</strong> concave surface extending to convex surface; bristles absent.<br />
Branchiae from setiger 9. Neurosetal hooks without manubria, from setiger 7.<br />
Dipolydora quadrilobata (Jacobi, 1883)<br />
PWS NIS 1998: Sta 15(1 specimen).<br />
Incomplete specimen, and identification is tentative.<br />
Polydora limicola Annenkova, 1934<br />
PWS NIS 1998: Sta 10(1 specimen); Sta 16(1); Sta 23(1).<br />
Incomplete specimen, and identification is tentative.<br />
Polydora sp.<br />
PWS NIS 1998: Sta 17(1 specimen).<br />
This is a juvenile specimen. It is small and slender. Prostomium entire. Eyes numbering 8.<br />
Setiger 5 is not modified. Pygidium 4-lobed. Probably not assignable to any known taxon.<br />
Nicomache personata Johnson 1901<br />
PWS NIS 1998: Sta 11(1 specimen).<br />
Identification certain to this species, although specimen is incomplete. Pigmentation pattern<br />
characteristic <strong>of</strong> species.
Chapt 9C3. Polychaete Worms, page 9C3- 16<br />
Pectinaria granulata Johnson 1901<br />
PWS NIS 1998: Sta 11(5 specimens); Sta 16(3); Sta 17(1); Sta 36(22).<br />
Correct identification.<br />
Laonome cf. kroyeri (Malmgren, 1866)<br />
PWS NIS 1998: Sta 17(14 specimens).<br />
Avicular uncini with short bases; companion setae absent. Both capillary and spatulate setae<br />
present. Radioles lacking external stylodes; collar bilobed. Need to reconfirm identity.<br />
Schizobranchia insignis Bush 1904<br />
PWS NIS 1998: Sta 19(4 specimen); Sta 23(2) + Sta 23(>20).<br />
Identification correct. Smallest specimen placed into shell vial (Sta. 23) is perhaps a juvenile,<br />
with all the setal features consistent. Note that large vial (Sta. 23) mislabeled as Terebellidae<br />
(instead <strong>of</strong> Sabellidae).<br />
Amphitrite cirrata Müller, 1771<br />
PWS NIS 1998: Sta 19(2 specimens); Sta 23(3); Sta 24(2); Sta35(1); Sta 36(1).<br />
This is another “cosmopolitan” species…at least in the northern hemisphere. A taxonomic black<br />
hole similar to Phole minuta and Eteone longa!! Specimen (Sta. 35) is juvenile terebellid,<br />
probably Amphitrite cirrata; it is not Ampharetidae)!<br />
Polycirrus species III Hobson & Banse, 1981<br />
PWS NIS 1998: Sta 36(1 specimen).<br />
This keys out as per Hobson & Banse (1981). However, as a general comment, the key is<br />
extremely poor and relies on imprecise terminology that overlaps features used to describe<br />
notosetae! In any case, it would seem appropriate to attach names instead <strong>of</strong> roman numerals.<br />
Ampharete species A<br />
PWS NIS 1998: Sta 41(1 specimen).<br />
The generic identification is correct. Single specimen lacks a tail, and cannot be identified to<br />
species. It is not Ampharete labrops, which has eyespots on upper buccal lip; present specimen<br />
lacks eyespots in corresponding region.<br />
Crucigera zygophora (Johnson, 1901)<br />
PWS NIS 1998: Sta 19(6 specimens); Sta 23(>20); Sta 24(1).<br />
Identification correct. Present specimens are textbook examples.<br />
Serpula vermicularis Linnaeus, 1767<br />
PWS NIS 1998: Sta 23(3 specimens).<br />
Identification correct. Tentacles solid red or banded red and white, at least in preservative.<br />
Spirorbis species<br />
PWS NIS 1998: Sta 17(>50 specimens).<br />
Dextrally spiraled tubes, 3 thoracic setigers. Species identifications pending examination <strong>of</strong><br />
remaining materials.
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 1<br />
Chapter 9C4. Focal Taxonomic Collections: Peracaridan Crustaceans<br />
John W. Chapman, Department <strong>of</strong> Fisheries & Wildlife, Hatfield Marine Science Center, Oregon<br />
State University<br />
Summary<br />
No clear peracaridan NIS were discovered among the scores <strong>of</strong> species collected in three<br />
surveys <strong>of</strong> 72 sites in 21 general areas <strong>of</strong> Prince William Sound and south central Alaska<br />
between 1997 and 1999. No NIS were found in UAF samples from the area collected<br />
previously. Two peracaridan species previously considered to be introduced are likely to be<br />
misidentified. Five species <strong>of</strong> NIS gammaridean amphipods were found in ballast water <strong>of</strong><br />
tankers travelling to Prince William Sound, indicating that this is an active mechanism <strong>of</strong> NIS<br />
transport to Alaska, even though they do not appear to have invaded or become established there.<br />
<strong>Invasions</strong> <strong>of</strong> Alaskan estuaries and marine waters by a broad diversity <strong>of</strong> peracaridan species<br />
have not occurred. The diversities <strong>of</strong> peracaridan NIS invasions in the northern hemisphere vary<br />
with climate, as do invasions by other taxa noted previously. Most marine and estuarine<br />
peracaridan NIS thus appear to be incapable <strong>of</strong> invading Alaska from lower latitudes due to the<br />
extreme climate. The risk <strong>of</strong> invasions by high diversities <strong>of</strong> NIS <strong>of</strong> pericaridans thus appears to<br />
be extremely low.<br />
These findings do not indicate whether a few NIS could be present at ecologically<br />
catastrophic abundances, however. Eight peracaridans that are prominent members <strong>of</strong> either<br />
fouling or benthic communities sampled in the survey, have unclear origins or cannot yet be<br />
clearly distinguished from species that are nonindigenous to the northeast Pacific. They are<br />
therefore classified as cryptogenic. These cryptogenic peracaridan species occur in the same<br />
areas as the s<strong>of</strong>t shell clam, Mya arenaria Linnaeus, 1758, which is one <strong>of</strong> the most clearly<br />
documented NIS in south central Alaska. If proven to be NIS, these cryptogenic peracaridan<br />
species, would be evidence that even a few NIS capable <strong>of</strong> invading Alaskan estuaries can<br />
increase to ecologically catastrophic densities. They would indicate that surveys <strong>of</strong> peracaridan<br />
NIS diversity, such as this one, are an insufficient basis for estimates <strong>of</strong> risk. Whether these<br />
peracaridan crustaceans are, in fact, native to the region therefore should be tested by analyses <strong>of</strong><br />
morphological variation, molecular genetics and by crossbreeding viability tests with their<br />
presumed original populations.<br />
Introduction<br />
A major objective <strong>of</strong> the south central Alaskan NIS survey was to determine whether<br />
introductions <strong>of</strong> marine or estuarine species have already occurred. An ultimate objective <strong>of</strong> the<br />
overall risk analysis is to predict whether Alaskan waters are vulnerable to NIS invasions. The<br />
survey results and comparisons <strong>of</strong> climate effects on peracaridan NIS diversity over the northern<br />
hemisphere provide a basis for this prediction.<br />
Predicting which nonindigenous species (NIS) can be introduced, where, and the factors<br />
that control their survival are major objectives <strong>of</strong> invasion ecology. These predictions require<br />
knowledge <strong>of</strong> the interactions between dispersal and processes that determine NIS survival. The
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 2<br />
mechanisms <strong>of</strong> NIS dispersal among estuaries are becoming well known (e.g., Cangelosi 1999,<br />
Cohen 1998, Frey et al.1999, Draheim and Olson 1999, Miller and Chapman 2000, Moy 1999,<br />
Ruiz et al. 1999, Thresher 1999), while the processes limiting NIS survival and production<br />
among estuaries remain poorly known. The distributions <strong>of</strong> NIS reveal how survival varies as<br />
dispersal occurs and thus indicate the interactions <strong>of</strong> dispersal, ecology, and survival.<br />
Interpreting the geography NIS distributions is thus a necessary part <strong>of</strong> the search for factors<br />
controlling NIS invasions.<br />
NIS are particularly diverse and abundant in estuaries <strong>of</strong> the northeastern Pacific,<br />
including San Francisco Bay, California (Carlton and Geller, 1993, Cohen and Carlton 1995,<br />
1997, Ruiz et al. 1997a, 1997b) and in Europe (Leppakoski, 1994, Leppakoski and Olenin, 1999,<br />
Eno, et al. 1997). The majority <strong>of</strong> these NIS have origins from western ocean coasts (Cohen and<br />
Carlton 1995, Leppakoski and Olenin, 1999) and progressively fewer NIS are known with<br />
increasing latitudes (Carlton 1979, Cohen et al. 1998, Mills et al. 2000). These geographical<br />
patterns do not appear to result entirely from the mechanisms <strong>of</strong> dispersal or patterns <strong>of</strong> endemic<br />
species diversity. Other processes controlling NIS distributions warrant consideration.<br />
Potential climate effects on east and west and north and south patterns <strong>of</strong> NIS diversity<br />
among estuaries and coastal waters <strong>of</strong> the northern hemisphere are <strong>of</strong> paramount concern in any<br />
NIS risk analysis for Alaska. The cold temperate climate <strong>of</strong> Alaska is at the extreme northern<br />
range <strong>of</strong> many northeast Pacific intertidal species (O’Clair 1977, O’Clair and O’Claire 1998).<br />
Climate effects are therefore considered in this section. Salinity and temperature variations in<br />
estuaries are dominant processes <strong>of</strong> climate that limit NIS survival. Most estuarine species<br />
survive within narrow temperature and salinity ranges. Most chemical, biological, and<br />
hydrological processes that also limit the abundances and distributions <strong>of</strong> estuarine organisms are<br />
also controlled by, or closely correlated with, salinity and temperature (e.g., Southward 1969,<br />
Green 1971, Ebbesmeyer et al. 1991, Cohen and Carlton 1995 1999, Chapman 1998, Thompson<br />
1998). NIS distributions are therefore influenced by salinity and temperature within local<br />
estuaries. In turn, salinity and temperature are affected by climate (Ebbesmeyer et al. 1991,<br />
Cayan 1993). Precipitation and air temperature variations (Ebbesmeyer et al. 1991, Cayan 1993)<br />
can be interpreted to infer salinity and temperature variations in local estuaries even though<br />
direct, long-term measures <strong>of</strong> these parameters are lacking in most cases.<br />
Amphipods recovered from 34 ballast water samples taken from tankers during 1998<br />
were examined to determine whether amphipods are transported by ballast water traffic from<br />
west coast ports and harbors.<br />
Methods<br />
Rapid assessment survey<br />
Prince William Sound, Seward and Homer, Alaska (60 o 00' - 61 o 00' N) survey results are<br />
compared to results <strong>of</strong> NIS surveys <strong>of</strong> the native, cryptogenic and introduced peracaridans from<br />
Puget Sound, Washington (47 o 10' - 49 o 00' N) and San Francisco Bay, California (37 o 30' - 38 o<br />
10' N) to resolve how peracaridan NIS invasions are distributed over a broad range <strong>of</strong> latitudes.
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 3<br />
Collections were made from three sites in Port Valdez in early spring <strong>of</strong> 1997, 46 sites<br />
throughout Prince William Sound in June 1998, and from 23 sites in Prince William Sound,<br />
Seward and Homer in 1999 (Ruiz and Hines 1997, Hines and Ruiz 1998) (Figure 9C4.1).<br />
Twenty six sites were surveyed in Puget Sound, Washington in September 1998 (Cohen et al.<br />
1998). San Francisco Bay was surveyed in early fall, late spring or summer <strong>of</strong> 1993, 1994, 1996<br />
and 1997 at 25 regular sites plus several irregular sites (Cohen 1998, Cohen and Carlton 1995,<br />
1997, 1998). The three systems are excellent for comparison because they have all received and<br />
have been interconnected by significant aquaculture and shipping activities that are vectors <strong>of</strong><br />
NIS dispersal in at least the last century.<br />
Figure 9C4.1. Prince William Sound, Alaska and south central Alaska rapid assessment sites with open boxes<br />
indicating all 46 sampling sites <strong>of</strong> 1998, inset indicating the five port areas sampled in 1999 and the shaded boxes<br />
indicating the eleven general sampling areas <strong>of</strong> the sound in 1999. (Latitudes longitudes and site descriptions are in<br />
appendix table 9C4.5).<br />
Each area was surveyed by the author in the same fashion. Survey samples were<br />
collected by hand, scrapings, cores, or dredge as necessary to remove biological communities or<br />
substratum from floats, intertidal pilings rocks and intertidal or shallow subtidal mudflats<br />
accessible at each collection site. These samples were washed on an 0.5 mm mesh sieve directly<br />
or decanted onto an 0.5 mm mesh sieve and washed following vigorous sloshing in buckets <strong>of</strong>
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 4<br />
seawater, to suspend organisms from the removed substratum. Harbor float, rock and piling<br />
substratums were emphasized in all three survey areas but other available habitats were sampled<br />
extensively as available. Organisms were picked directly from substratums during sample<br />
collection or from the sieves after washing or from voucher samples <strong>of</strong> substratums and<br />
examined under a stereomicroscope. All collected organisms were fixed in 10% formalin before<br />
transfer to 70% ETOH for long-term preservation. All specimens were identified to lowest<br />
possible taxonomic category.<br />
Voucher specimens will be deposited in the Los Angeles County Museum, the California<br />
Academy <strong>of</strong> Sciences and the Smithsonian National Museum <strong>of</strong> Natural History. The precise<br />
locality records and notes for each collection site are available from the author. Temperature and<br />
salinity was measured at each collection site. Surface salinties ranged between 0 and 33 o / oo in<br />
all three survey areas. Surface water temperatures ranged between 8 and 20 o C in Prince William<br />
Sound, between 10 and 21 o C in Puget Sound and between 12 and 30 o C in San Francisco Bay.<br />
San Francisco Bay is a well mixed estuary. Low surface salinities and clear stratification occur<br />
in both sounds in summer and were apparent during the surveys.<br />
All species from the three surveys were collected and examined directly by the author<br />
and thus are assumed to be a more standardized sample than would be likely from comparisons<br />
<strong>of</strong> different surveys and sampling methods employed by different investigators. The indigenous<br />
origins <strong>of</strong> species are inferred from previously published records or herein using the criteria <strong>of</strong><br />
Carlton (1979) and Chapman and Carlton (1991, 1994). The criteria used for cyptogenic species<br />
(species that are not clearly native or introduced) are adopted from Carlton (1996a). Only<br />
populations <strong>of</strong> species that have been moved by human activities to new locations, that are<br />
reproductive there, and that satisfy the criteria for nonindigenous species are considered here to<br />
be NIS.<br />
Alaskan Climate and Peracaridan NIS Invasion Risks<br />
Sources <strong>of</strong> NIS to Alaskan estuaries are available from a global population. The<br />
peracaridan Crustacea <strong>of</strong> the northern hemisphere considered here are as a sample <strong>of</strong> that<br />
population. Crustacea comprise approximately 25% <strong>of</strong> the 250 NIS reported from San Francisco<br />
Bay (Cohen and Carlton 1995, J. T. Carlton, personal communication), where they are the most<br />
diverse NIS taxon. The majority <strong>of</strong> these crustacean NIS are peracaridans. The Peracarida<br />
consist <strong>of</strong> relatively small, short-lived species that are primarily mysids, amphipods, isopods,<br />
tanaidaceans and cumaceans. The Peracarida are prominent in most North Pacific and North<br />
Atlantic marine and estuarine NIS communities (Bowman et al. 1981, Chapman 1988, 1999,<br />
Chapman and Carlton 1991, 1994, Mees and Fockedey 1993, Leppakoski 1994, Cohen and<br />
Carlton 1995, 1997, Eno et al. 1997, T<strong>of</strong>t et al. 1999). Both native and nonindigenous<br />
Peracarida are diverse, taxonomically well known, and ubiquitous in aquatic environments (e.g.,<br />
Barnard and Barnard 1983, Barnard and Karaman 1991, Chapman 1988, Cohen and Carlton<br />
1995, Chapman 2000). Peracarida develop directly, without larval dispersal stages or unique life<br />
history traits that complicate identifications and interpretations <strong>of</strong> their geographical<br />
distributions. Peracaridans may thus provide clear indications <strong>of</strong> the patterns <strong>of</strong> diversity within<br />
and among broad geographical regions.
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 5<br />
The species selected for the east to west geographical analysis <strong>of</strong> northern hemisphere<br />
peracaridan NIS are common or abundant where they occur and documented either in the<br />
literature or by personal observations. Species that are poorly documented, not examined<br />
directly, cryptogenic, or that are not introduced across the North Atlantic or North Pacific, were<br />
not included. For instance, the amphipod Chelicorophium curvispinum (Sars, 1895), which<br />
spread from the Black and Caspian Sea to northern Europe (Eno et. al. 1997), and the introduced<br />
mysid Acanthomysis bowmani Modlin and Orsi, 1997 in San Francisco Bay, which has unknown<br />
origins, and many northern NIS that are native to the southern hemisphere are not in the scope <strong>of</strong><br />
this study and are therefore excluded from the analysis.<br />
Long-term climate conditions in the northeast Pacific, including San Francisco Bay,<br />
Puget Sound and Prince William Sound are inferred from monthly average climate time series<br />
data for the Pacific Ocean and western Americas (Cayan et al. 1991). These data extend over<br />
approximately 100 years up to 1986. Global records <strong>of</strong> sea surface temperature and precipitation<br />
minus evaporation (http://www.cdc.noaa.gov/ 1998) are used for comparisons <strong>of</strong> temperature<br />
among ocean regions. The term “western ocean” is used in reference to the Pacific Ocean<br />
bordering the east Asian coast and the Atlantic Ocean bordering the eastern North American<br />
coast. The term “eastern ocean” refers to the ocean areas bordering the west coasts <strong>of</strong> Europe<br />
and North Africa and the west coast <strong>of</strong> North America.<br />
Amphipods in Ballast <strong>Water</strong><br />
Ballast water samples were collected in vertical plankton tows from “dedicated” ballast<br />
tanks, which are not contaminated by oil. Amphipods retained in the 0.25 mm mesh plankton<br />
nets preserved in 5% formalin, subsequently transferred to 70% ethanol for final sorting and<br />
identification. Samples were initially sorted under stereo microscopes at the Smithsonian<br />
Environmental Research Center or at the field <strong>of</strong>fice in Port Valdez and final amphipod<br />
identifications were performed at HMSC, Oregon State University.<br />
The origins <strong>of</strong> the ballast water sampled (Table 9C4.3) were the Los Angeles-Long<br />
Beach area, the San Francisco Bay area, Puget Sound (Anacortes) and the open ocean. One ship,<br />
from the San Francisco Bay area, exchanged the ballast water at sea during transit. The potential<br />
for dispersal <strong>of</strong> nonindigenous species is assessed from the presence <strong>of</strong> nonindigenous species in<br />
samples.<br />
Results<br />
North - South Climate<br />
The maximum, minimum, mean and range <strong>of</strong> monthly sea surface temperatures <strong>of</strong> the<br />
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<br />
maximum average monthly temperatures <strong>of</strong> Seward, Kodiak and Sitka, Alaska (between 58 and<br />
60 o N) overlap the Neah Bay, Washington maximum surface temperatures at 48 o N and are<br />
similar to the 13 o C average sea surface temperatures adjacent to San Francisco at 38 o N (Figure<br />
2). Nonindigenous species expected to reach south central Alaska might include those that can<br />
reproduce within Puget Sound in summer or in average San Francisco temperatures (Figure<br />
9C4.2).
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 6<br />
Figure 9C4.2. Sea surface temperature monthly average minimum, mean, maximum and range <strong>of</strong> northeast Pacific<br />
coastal waters estimate over an approximately 100 years up to 1986 (Cayan 1991).<br />
North - South Biodiversity<br />
Of the 106 peracaridan crustacean species identified from the surveys <strong>of</strong> Prince William<br />
Sound, Puget Sound and San Francisco Bay, 54 are native, 14 are cryptogenic and 38 are<br />
introduced (Table 9C4.1). Seven peracaridan crustaceans that prominent members <strong>of</strong> benthic or<br />
fouling communities that were recovered in the survey are cryptogenic (Table 9C4.1). They are<br />
the tanaidacean Leptochelia dubia (Kroyer, 1842) a cosmopolitan species (Miller, 1975); the<br />
cumacean Cumella vulgaris Hart, 1930 which occurs in Asia (Lomakina 1958) as well as the<br />
eastern Pacific; the amphipod Monocorophium carlottensis Bousfield and Hoover, 1997 is not<br />
clearly distinguished from the nonindigenous amphipods Monocorophium acherusicum and<br />
Monocorophium insidiosum (Ruiz and Hines 1997); the amphipod Hyale plumulosa (Stimpson,<br />
1857) is reported also from the western Atlantic (Bousfield 1973); the amphipod Jassa staudei<br />
Conlan 1990 is extemely similar to the cosmopolitan Jassa marmorata Holmes, 1903; the<br />
amphipod Pontogeneia rostrata Gurjanova, 1938 is reported from the eastern and western<br />
Pacific (Gurjanova 1938, 1951, Barnard 1962, 1964); the caprellid Caprella depranochir Mayer,<br />
1880 is reported from the eastern and western Pacific (Arimoto 1976, Kozl<strong>of</strong>f and Price1997).
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 7<br />
Table 9C4.1. The 106 peracaridan crustaceans identified as nonindigenous, cryptogenic or native, and the records<br />
per species collected from San Francisco Bay, California, (w/o “*”), south central Alaska and Prince William Sound<br />
(with “*”), or Puget Sound, Washington (underlined). Gnorimosphaeroma lutea was collected from San Francisco<br />
Bay and Prince William Sound only. No species were collected only in Puget Sound.<br />
Nonindigenous Cryptogenic Native<br />
Records Records Records<br />
Mysidacea Tanaidacea Mysidacea<br />
Accanthomysis aspera 1 Leptochelia dubia 3 Mysis littoralis* 1<br />
Tanaidacea Cumacea Isopoda<br />
Tanais stanfordi 2 Cumella vulgaris 3 Dynamenella glabra* 2<br />
Isopoda Gammaridea Gnorimosphaeroma lutea 2<br />
Asellus sp. 1 Ampithoe lacertosa 3 Gnorimosphaeroma oregonense 3<br />
Dynoides dentisinus 1 Dulichia sp. 1 Ianiropsis kincaidi* 2<br />
Euylana arcuata 1 Hyale plumulosa 3 Idotea montereyensis 3<br />
Ianiropsis serricadus 1 Ischyrocerus sp. 2 Idotea obscura* 1<br />
Limnoria quadripunctata 1 Jassa staudei 3 Idotea resecata 2<br />
Limnoria tripunctata 2 Monocorophium carlottensis* 2 Idotea wosnsenskii 3<br />
Munna ubiquita 1 Pontogeneia rostrata 3 Ligia pallasi* 1<br />
Paranthura sp. 1 Caprellidea Limnoria lignorum 2<br />
Sphaeroma quoyanum 1 Caprella depranochir* 2 Cumacea<br />
Synidotea laevidorsalis 1 Caprella laeviuscula 3 Diastylis alaskensis* 1<br />
Cumacea Caprella penantus 1 Diastylis sp.* 1<br />
Nippoleucon hinumensis 2 Caprella verrucosa 2 Lamprops beringi* 1<br />
Gammaridea Tritella sp. 1 Lamprops quadriplicata* 2<br />
Ampelisca abdita 1 Gammaridea<br />
Ampithoe valida 2 Allorchestes angusta 3<br />
Crangonyx sp. 1 Americorophium brevis* 2<br />
Eochelidium sp. 2 Americorophium salmonis* 2<br />
Gammarus daiberi 1 Americorophium spinicorne 2<br />
Grandidierella japonica 2 Ampithoe dalli* 2<br />
Hyalella azteca 1 Ampithoe kussakini* 1<br />
Jassa marmorata 2 Ampithoe sectimanus* 1<br />
Laticorophium baconi 2 Ampithoe simulans* 2<br />
Leucothoe alata 1 Anisogammarus pugettensis 3<br />
Melita nitida 2 Aoroides columbiae 3<br />
Melita sp. 1 Aoroides intermedius* 2<br />
Monocorophium<br />
2 Calliopius carinatus* 2<br />
acherusicum<br />
Monocorophium insidiosum 2 Calliopius pacificus* 2<br />
Monocorophium<br />
1 Eogammarus confervicolus 3<br />
oaklandense<br />
Monocorophium uenoi 1 Eogammarus oclairi* 1<br />
Trasorchestia enigmatica 1 Gammaridae n. gen. n. sp.* 1<br />
Paradexamine sp. 1 Gnathopleustes pugettensis 2<br />
Parapleustes derzhavini 1 Hyale frequens 3<br />
Senothoe valida 2 Lagunogammarus setosus* 2<br />
Sinocorophium<br />
1 Locustogammarus locustoides* 1<br />
heteroceratum<br />
Caprellidea Megamoera subtener* 2<br />
Caprella acanthogaster 2 Microdeutopus schmitti 1<br />
Caprella bidentata 1 Najna n. sp.* 1<br />
Caprella californica 2 Paracalliopiella pratti* 2<br />
Caprella equilibra 2 Parallorchestes ochotensis 3<br />
Table 9C4.1. Continued
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 8<br />
Paramoera bousfieldi* 2<br />
Paramoera mohri* 1<br />
Parampithoe humeralis.* 2<br />
Parampithoe mea* 1<br />
Photis brevis 2<br />
Pontogeneia inermis 3<br />
Pontogeneia ivanovi* 1<br />
Pontoporeia femorata* 2<br />
Spinulogammarus subcarinatus* 1<br />
Trasorchestia traskiana 3<br />
Caprellidea<br />
Caprella alaskana* 1<br />
Caprella gracilior* 1<br />
Caprella irregularis* 2<br />
Metacaprella kennerlyi* 2<br />
Total Species 38 14 54<br />
Records/Species 1.4 2.1 1.8<br />
The diversity <strong>of</strong> species collected is nearly nearly constant among sites (66 to 60 and 59,<br />
respectively, between San Francisco, Puget Sound and Prince William Sound). Of the 54 native<br />
species collected in Alaska, 52 (96%) were collected also in San Francisco Bay or Puget Sound<br />
(Table 9C4.1). The common pool <strong>of</strong> native species and the similar species diversities collected<br />
among the three areas both indicate that the habitat selection, collection and sample processing<br />
methods <strong>of</strong> the rapid assessment surveys were consistent among the three areas. Little variation<br />
in NIS diversity among these three sample sets was therefore likely to result from sample biases.<br />
From San Francisco Bay north to Puget Sound and Prince William Sound, introduced<br />
peracaridan species declined from 38 to 15 to 0 (Figure 9C4.3, X 2 > 27.01; p < 0.0001; df = 2),<br />
while the frequencies <strong>of</strong> cryptogenic peracaridan species were nearly constant at 11, 10 and 8,<br />
respectively (X 2 = 2.0; p > 0.73; df = 2). In contrast, the frequencies <strong>of</strong> native species increased<br />
to the north from 17 to 35 and 52 (Figure 9C4.3, X 2 > 27.01; p < 0.0001; df = 2 ). All<br />
peracaridan NIS and all but two <strong>of</strong> the cryptogenic peracaridan species at any site also occurred<br />
in San Francisco Bay while only 17 <strong>of</strong> the 54 native species were recovered from San Francisco<br />
Bay (Table 9C4.1, Figure 9C4.3). The peracaridan NIS that managed to invade northeast Pacific<br />
estuaries thus have adaptations to lower latitude climates than do the native species. The<br />
affinities <strong>of</strong> peracaridan NIS for low latitude climates closely corresponds to a pattern that would<br />
be expected if the peracaridan NIS are poorly adapted to cold water conditions or other factors <strong>of</strong><br />
climate that vary with latitude.
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 9<br />
Native Cryptogenic Nonindigenous<br />
60<br />
50<br />
Species<br />
40<br />
30<br />
20<br />
10<br />
0<br />
SFB<br />
PS<br />
PWS<br />
Figure 9C4.3. Native, cryptogenic, nonindigenous and total species in samples from San Francisco Bay, California,<br />
Puget Sound, Washington and Prince William Sound, Alaska (SFB, PS and PWS, respectively).<br />
Some NIS may be among the 8 cryptogenic peracaridan species <strong>of</strong> Prince William Sound<br />
(Figure 9C4.3). Six <strong>of</strong> these cryptogenic Alaskan peracaridan species have broad thermal<br />
tolerance ranges and occur also in Puget Sound and San Francisco Bay (Table 9C4.1). The<br />
nearly complete faunal peracaridan overlap between Alaska and the two more southern sites<br />
indicate that the low diversity <strong>of</strong> Alaskan NIS peracaridans are not likely to result from unique<br />
types <strong>of</strong> NIS peracaridans that were not collected in the survey. The nearly uniform pool <strong>of</strong><br />
native peracaridan species and uniform pool <strong>of</strong> NIS peracaridans among the areas require few<br />
qualifications or assumptions to arrive at conclusions <strong>of</strong> the patterns <strong>of</strong> diversity. The possibility<br />
<strong>of</strong> overlooking populations <strong>of</strong> nonindigenous peracaridan crustaceans in Alaska that occur at<br />
similar diversities and abundances as in Puget sound or San Francisco Bay, by these rapid<br />
assessment methods, is remote. The presence <strong>of</strong> peracaridan NIS that are not among the<br />
cryptogenic species <strong>of</strong> peracardans in Alaskan estuaries and marine waters is not significantly<br />
different from zero.<br />
These results are not surprising. The extreme climate <strong>of</strong> Alaska was previously assumed<br />
to limit the survival <strong>of</strong> nearly all NIS peracaridans. However, these data include only a single<br />
coast and three major areas <strong>of</strong> reference. NIS pericarican diversity was therefore compared<br />
between other geographical regions with climates that vary similarly to the variation that occurs<br />
between San Francisco and south central Alaska as a further test <strong>of</strong> the climate pattern.<br />
East -West Climate<br />
Sea surface temperatures are relatively constant between latitudes <strong>of</strong> 25 o and 50 o N in<br />
eastern oceans compared to western ocean areas (Figures 9C4.4 and 9C4.5). Eastern ocean<br />
species from 32 o to 50 o N evolved in temperature ranges that span only latitudes 40 to 42 o N in
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 10<br />
western oceans (Figure 9C4.6). Eastern ocean species thus have not evolved in conditions that<br />
would create broad temperature tolerance ranges. In contrast, most western ocean organisms<br />
survive temperature ranges exceeding the total temperature range <strong>of</strong> broad eastern ocean areas.<br />
Eastern oceans below 50 o N, are thus broad thermal targets for western ocean species over space<br />
and time while western ocean coasts are narrow thermal targets for eastern ocean species.<br />
N.W. Pacific<br />
N.E. Pacific<br />
30<br />
30<br />
25<br />
25<br />
SST (Deg. C)<br />
20<br />
15<br />
10<br />
5<br />
‘<br />
SST (Deg. C)<br />
20<br />
15<br />
10<br />
5<br />
0<br />
-5<br />
JF MA MJ JA SO ND<br />
60<br />
50<br />
30<br />
40<br />
Lat. N<br />
0<br />
-5<br />
JF MA MJ JA SO ND<br />
60<br />
50<br />
30<br />
40<br />
Lat. N<br />
Months<br />
Months<br />
N.W. Pacific<br />
N.E. Pacific<br />
5<br />
5<br />
4<br />
4<br />
Precip. (mm / day)<br />
3<br />
2<br />
1<br />
0<br />
-1<br />
-2<br />
-3<br />
JF MA MJ JA SO ND<br />
30<br />
60<br />
50<br />
40<br />
Lat. N<br />
Precip. (mm / day)<br />
3<br />
2<br />
1<br />
0<br />
-1<br />
-2<br />
-3<br />
JF MA MJ JA SO ND<br />
30<br />
60<br />
50<br />
40<br />
Lat. N<br />
Months<br />
Months<br />
Figure 9C4.4. Northwest and northeast Pacific sea surface temperature in degrees Celsius and bimonthly<br />
precipitation minus evaporation in mm -d at ten degree latitude intervals for the bimonthly periods <strong>of</strong> Jan-Feb, Mar-<br />
Apr, May-Jun, Jul-Aug, Sep-Oct and Nov-Dec. (Note: axes <strong>of</strong> latitude are reversed between graphs <strong>of</strong> temperature<br />
and
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 11<br />
N.W. Atlantic<br />
N.E. Atlantic<br />
30<br />
30<br />
25<br />
25<br />
SST (DEG. C)<br />
20<br />
15<br />
10<br />
SST (DEG. C)<br />
20<br />
15<br />
10<br />
5<br />
0<br />
JF MA MJ<br />
JA SO ND<br />
60<br />
30<br />
40<br />
50<br />
Lat. N.<br />
5<br />
0<br />
JF MA MJ JA SO ND<br />
60<br />
30<br />
40<br />
50<br />
Lat. N.<br />
Months<br />
Months<br />
N. W. Atlantic<br />
N.E. Atlantic<br />
4<br />
4<br />
3<br />
3<br />
Precip. (mm / day)<br />
2<br />
1<br />
0<br />
-1<br />
-2<br />
-3<br />
JF MA MJ<br />
JA<br />
Months<br />
SO<br />
ND<br />
30<br />
60<br />
50<br />
40<br />
Lat. N<br />
Precip. (mm/day)<br />
2<br />
1<br />
0<br />
-1<br />
-2<br />
-3<br />
JF<br />
MA<br />
MJ<br />
JA<br />
Months<br />
SO<br />
ND<br />
30<br />
60<br />
50<br />
40<br />
Lat. N<br />
Figure 9C4.5. Northwest and northeast Atlantic sea surface temperature in degrees Celsius and bimonthly<br />
precipitation minus evaporation in mm -d at ten degree latitude intervals for the bimonthly periods <strong>of</strong> Jan-Feb, Mar-<br />
Apr, May-Jun, Jul-Aug, Sep-Oct and Nov-Dec. (Note: axes <strong>of</strong> latitude are reversed between graphs <strong>of</strong> temperature<br />
and precipitation.)
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 12<br />
Degrees Celsius<br />
30<br />
25<br />
20<br />
15<br />
10<br />
5<br />
0<br />
25 30 35 40 45 50 55 60<br />
North Latitude<br />
6<br />
Precip - Evap / Day<br />
4<br />
2<br />
0<br />
-2<br />
-4<br />
25 30 35 40 45 50 55 60<br />
North Latitude<br />
NWA NEA NWP NEP<br />
Figure 9C4.6. Maximum and minimum monthly sea surface temperature in degrees Celsius (A) and monthly<br />
precipitation minus evaporation in mm d -1 (B) at ten degree latitude intervals for the northwest (dashed lines) and<br />
northeast (solid lines) Atlantic (thin lines) and Pacific (thick lines). Northeast Pacific, northwest Pacific, northeast<br />
Atlantic and northwest Atlantic are NEP, NWP, NEA and NWA, respectively.<br />
Precipitation, and thus salinity in estuaries, also vary from east to west in patterns that<br />
resemble the south to north pattern from San Francisco to Alaska. The broadest ranges <strong>of</strong><br />
precipitation occur in the eastern Pacific north <strong>of</strong> 35 o N. Lat. (Figure 9C4.4). The narrowest<br />
ranges <strong>of</strong> precipitation and negative net precipitation (desert conditions) occur in the eastern<br />
Pacific and Atlantic south <strong>of</strong> 35 o N. Lat. (Figure 9C4.4). Desert conditions do not occur at low<br />
latitudes in the northwest Pacific and occur in the northwest Atlantic only below 30 o N (Figure
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 13<br />
9C4.4). The latitudinal range and areal extent <strong>of</strong> low salinity estuaries is therefore less in eastern<br />
oceans than in western oceans and the climates are more uniform.<br />
The seasonal patterns <strong>of</strong> precipitation (Figures 9C4.4 and 9C4.5) also differ consistently<br />
between eastern and western oceans. More precipitation occurs in western oceans during<br />
summer when temperatures are maximum while most precipitation occurs in eastern oceans in<br />
winter when temperature are low (Figures 9C4.4 and 9C4.5). Where snow-melt is not important,<br />
and in the absence <strong>of</strong> major water impoundments, the salinity-temperature pattens <strong>of</strong> eastern and<br />
western ocean estuaries are out <strong>of</strong> synchrony. In high latitude regions, such as Alaska, run<strong>of</strong>f<br />
varies most with snow-melt and salinity is lower in warm seasons in correspondence with<br />
western ocean climates.<br />
East and West Biodiversity<br />
Peracaridan NIS diversity varied among the four northern hemisphere ocean coasts ( X 2 =<br />
17.27, p = 0.001 df = 3), with five times as many introductions to eastern ocean coasts (Table<br />
9C4.2; X 2 = 16.05, p = 0.001 df = 1). Except for the gammaridean amphipods Orchestia<br />
gammarella (Pallas, 1766) and Corophium volutator (Pallas, 1766) in the tidal mudflats and<br />
marshes <strong>of</strong> the Bay <strong>of</strong> Fundy, peracaridan NIS abundance and diversity in western Atlantic<br />
estuaries appeared to be low. Similarly, <strong>of</strong> the 28 peracaridan species included in the east to<br />
west analysis, only 2 - 3 NIS were in the northwest Pacific, compared to 20-23 in the<br />
northeastern Pacific and 10-12 in the northeast Atlantic (Table 9C4.2). None <strong>of</strong> the common<br />
northeast Pacific peracaridans were introduced to other areas <strong>of</strong> the world. Only 2 <strong>of</strong> the<br />
northeast Atlantic species were clearly introduced to other regions compared to13 from the<br />
northwest Pacific and 14 from the northwest Atlantic (Table 9C4.2). Remarkably, 5 - 9 <strong>of</strong> the 28<br />
NIS (Table 9C4.2) have been reported on two coasts and 4 <strong>of</strong> these species have been discovered<br />
on 3 coasts.<br />
Climate and NIS Diversity<br />
The ranks <strong>of</strong> climates from least to most similar based on overall temperature variation<br />
and seasonal precipitation (Figures 9C4.2, 9C4.3 and 9C4.4) were, northeast Pacific, northeast<br />
Atlantic, northwest Pacific and northwest Atlantic (NEP, NEA, NWP and NWA, respectively).<br />
The ranks <strong>of</strong> NIS invaders <strong>of</strong> these regions (Imports, Table 1) were from highest to lowest: NEP,<br />
NEA, NWP and NWA. The ranks <strong>of</strong> native species that have been introduced to other regions<br />
(Exports, Table 1) were from lowest to highest: NEP, NEA, and NWP and NWA. The east and<br />
west variations in NIS imports and exports were thus correlated with climate variation (Kendall<br />
coefficient <strong>of</strong> concordance W = 1.0; X 2 = 8.2; p < 0.02; df = 2) (Siegal 1956) in a similar pattern<br />
to the south to north pattern <strong>of</strong> NIS between San Francisco Bay and Prince William Sound.
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 14<br />
Table 9C4.2. The east and west destinations and sources <strong>of</strong> common introduced peracaridan crustaceans <strong>of</strong> the<br />
Northwest Pacific Northeast Pacific, Northwest Atlantic and Northeast Atlantic (NWP, NEP NWA and NEA,<br />
respectively), their native, introduced, or probable introduced status (N, I, and I, respectively) and the numbered<br />
reference sources.<br />
Common Introduced Nonindigenous Estuarine Peracaridan<br />
Crustaceans <strong>of</strong> the Northern Hemisphere<br />
NWP NEP NWA NEA Sources<br />
Mysidacea<br />
Acanthomysis aspera N I 1,2<br />
Cumacea<br />
Nippoleucon hinumensis N I 1,3<br />
Isopoda<br />
Asellus communis I N I 4<br />
Caecidotea racovitzai I N 7<br />
Dynoides dentisinus N I 1<br />
Ianiropsis serricatus N I 1,8<br />
Paranthura sp. N I 1<br />
Synidotea laevidorsalis N I I I 1,5<br />
Amphipoda<br />
Ampelisca abdita I N 1,6,10<br />
Ampithoe valida I N 1,9,10<br />
Apocorophium lacustre N I 13,14<br />
Caprella acanthogaster N I I 1,10,11<br />
Corophium sp. N I 15,16<br />
Corophium volutator I N 13,14,17<br />
Crangonyx floridanus I N 7<br />
Crangonyx pseudogracilis N I 18<br />
Gammarus daiberi I N 1,7<br />
Gammarus tigrinus N I 13,14,18,19<br />
Grandidierella japonica N I I 10,20,21<br />
Jassa marmorata I I N I 10,22,23<br />
Leucothoe alata N I 1,24<br />
Melita nitida I I N 1,6,10<br />
Microdeutopus gryllotalpa I N I 13,25<br />
Moncorophium uenoi N I 9,10<br />
Monocorophium acherusicum I I N I 1,9,10<br />
Monocorophium insidiosum I I N I 1,9,10<br />
Orchestia gammarella I N 13,14,19<br />
Parapleustes derzhavini N I 1,6,10<br />
Sinocorophium heteroceratum N I 1,12<br />
Total Natives (Exports) 13 0 14 2<br />
Total NIS (Imports) 1 - 3 21 - 23 2 - 3 10 - 12<br />
N = native; I = introduced; = distribution not completely resolved but probable<br />
1) Cohen and Carlton 1995; 2) Modlin and Orsi 1997; 3) Watling 1991; 4) Williams 1972; 5) Chapman and Carlton 1991, 1994; 6)<br />
Chapman 1988; 7) T<strong>of</strong>t et al. 1999; 8) Kussakin 1988; 9) Barnard 1975; 10) Carlton 1979; 11) Platvoet et al. 1995; 10) Carlton 1979;<br />
12) Chapman and Cole In Prep.; 13) Bousfield 1973; 14) Lincoln 1979; 15) Hirayama 1984, 1986; 16) Janta 1995; 17) Chapman and<br />
Smith In prep.; 18) Costello 1993; 19) Watling 1979; 20) Chapman and Dorman 1975; 21) Smith et al. 1999; 22) Conlan 1990; 23)<br />
Mills et al. 1999; 24) Nagata 1965a-d; 25) Chapman and Miller In Prep.
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 15<br />
Table 9C4.3. Ballast and Port Valdez amphipos zooplankton samples subdivided into major source areas (Port<br />
Valdez, Benicia water exchanged at sea, San Francisco Bay area and southern California, respectively).
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 16<br />
Amphipods in Ballast <strong>Water</strong><br />
The 125 specimens recovered include one hyperiid species and fourteen species and ten<br />
families <strong>of</strong> gammaridean amphipods. Five <strong>of</strong> the gammaridean amphipods are NIS in the<br />
northeast Pacific and were present in ballast tanks discharged into waters <strong>of</strong> Port Valdez. These<br />
preliminary data indicate that ballast water traffic is a potential mechanism for transporting<br />
amphipods among harbors and coastal U. S. waters. The amphipod diversity in these samples<br />
(Table 9C4.4) is high given the small number <strong>of</strong> specimens involved. Except for the Ocean<br />
exchanged water and the water from Anacortes, Puget Sound, the heterogeneity among<br />
zooplankton sources is almost complete with a significant difference among the four species<br />
represented by more than 8 specimens (O 2 ; p < 0.001, df = 12). Only Ampelisca abdita and<br />
Gammarus daiberi occurred in more than one zooplankton source. Descriptions <strong>of</strong> the<br />
amphipods in these records are given in Appendix Table 94C. 8. The occurrence <strong>of</strong> amphipod<br />
species as a function <strong>of</strong> temperature and salinity <strong>of</strong> ballast water is shown in Figures 9C4.7 and<br />
9C4.8, respectively.<br />
Table 9C4.4. The average salinities, teperatures, subtotals, cryptogenic, introduced and native origins and total<br />
numbers <strong>of</strong> amphipod Crustacea collected in Port Valdez waters and from dedicated ballast tanks containing water<br />
exchanged at sea, or entrained from Puget Sound, San Francisco Bay or southern California.<br />
Discussion<br />
Most estuarine peracaridan NIS <strong>of</strong> the northeast Pacific are from the western sides <strong>of</strong> the<br />
Pacific or the Atlantic oceans. Peracaridan crustacean NIS diversity coincided with particular<br />
climates between 25 and 60 o N. Lat. Annual sea surface temperatures at latitudes below 50 o N<br />
vary less along northeast Pacific and Atlantic coasts than along western ocean coasts and, also in<br />
contrast to western ocean coasts, low salinity conditions occur in winter months rather than the<br />
summer months.
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 17<br />
Figure 9C4.7. Native, introduced and cryptogenic amphipod numbers with temperature from 34 ballast water<br />
samples.<br />
Figure 9C4.8. Native, introduced and cryptogenic amphipod numbers with salinity from 34 ballast water samples.<br />
The great diversity <strong>of</strong> invading species in northeast Pacific estuaries may thus result, in<br />
part, from the great diversity <strong>of</strong> climates that invading species are adapted to relative to the<br />
narrow range <strong>of</strong> climates in the region. The low diversity <strong>of</strong> native northeast Pacific species that<br />
invade other areas may also result, in part, from the relatively broad range <strong>of</strong> climate variations<br />
they must endure to survive elsewhere. The decline <strong>of</strong> northeast Pacific NIS diversity from<br />
south to north coincides with fewer introductions occurring where greater annual variations in<br />
temperature occur and where low salinity conditions occur during warm water periods.<br />
All <strong>of</strong> the peracaridan NIS known from Puget Sound occurred also in San Francisco Bay.<br />
The absence <strong>of</strong> cold water NIS in Southern Alaska is consistent with a pattern <strong>of</strong> introductions
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 18<br />
occurring less in continental climates <strong>of</strong> high temperature variations and low summer salinities.<br />
The overall pattern <strong>of</strong> NIS peracaridan diversity in the northern hemisphere strongly suggests<br />
that northeast Pacific estuarine peracaridan NIS <strong>of</strong> San Francisco Bay and north are<br />
predominantly from lower latitudes. Few <strong>of</strong> the presently known recognized northeast Pacific<br />
NIS peracaridans are thus likely to become established in southern Alaskan waters, even though<br />
Alaskan weather variations resemble western ocean climates from where most NIS originate.<br />
<strong>Aquatic</strong> species with nearly any life history and from nearly any taxon can be introduced<br />
(e.g. Carlton 1985, Cohen and Carlton 1995, Eno et al. 1997, Smith et al. 1999, Hewitt et al.<br />
1999). The many vectors, directions, distributions, routes <strong>of</strong> introduction (e.g., Carlton 1979,<br />
1985, 1987, 1999, Carlton and Geller 1993, Ruiz et al. 1997) and taxa available for introductions<br />
over the last 500 years (e.g. Carlton 1992, Carlton and Hodder 1995, Ruiz et al. 1997) have<br />
moved broad diversities <strong>of</strong> species to many suitable and unsuitable areas. The present<br />
distributions <strong>of</strong> NIS are a mosaic <strong>of</strong> surviving populations composed <strong>of</strong> a broad diversity <strong>of</strong> taxa<br />
and life histories. These surviving nonindigenous populations are surrounded by unsuitable<br />
areas in which they were also introduced but failed to survive. The patterns <strong>of</strong> transport<br />
mechanisms superimposed on this mosaic reveal where introductions fail. Resolution <strong>of</strong> this<br />
global pattern can reveal sources, destinations, targets and vectors <strong>of</strong> NIS and which ecosystems<br />
are most vulnerable.<br />
The probability <strong>of</strong> particular species introductions from one region to another cannot be<br />
determined from these data. NIS invasions in northern hemisphere estuaries, even confined to<br />
peracaridan crustaceans, are more complex than Tables 9C4.1 and 9C4.2 indicate. Additionally<br />
many evolutionary and ecological processes that are likely to contribute to the patterns <strong>of</strong> NIS<br />
invasions are not addressed. Western ocean estuaries may be older, with more diverse biotas and<br />
could be less intensely altered by human activities, for instance, than eastern ocean estuaries.<br />
When and how climates control NIS distributions are complicated by other processes including<br />
human and natural disturbances, and the timing, geography and magnitude <strong>of</strong> transport vectors.<br />
Also, the complexity <strong>of</strong> climate variations are drastically simplified in Figure 9C4.2, and<br />
Figures 9C4.4 - 9C4.6 on the assumption that large populations distributed over broad<br />
geographic areas are more likely to reflect average conditions over extended periods. However,<br />
other time intervals for integration could be better than the one and two month averages selected<br />
here for climate analyses. These simplifications may reduce the fit between biogeographical<br />
boundaries and climate and thus obscure interactions between NIS distributions and climate.<br />
These correlations between west ocean to east ocean NIS peracaridan invasions and<br />
climate nevertheless deserve close inspection. Source and destination climates for peracaridan<br />
NIS may be easier to identify than particular species that are likely to be introduced by particular<br />
dispersal vectors. Moreover, peracaridans appear to be a sufficient taxon to sample NIS<br />
biogeography. Similar east to west patterns <strong>of</strong> other NIS taxa have also been noted (Cohen and<br />
Carlton 1995, Leppakoski and Olin 2000, Miller and Chapman 2000). Moreover, the patterns <strong>of</strong><br />
NIS invasions must correspond to climate patterns if climate is an important ecological and<br />
evolutionary mechanism controlling the geography species.
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 19<br />
The correlations between marine and estuarine peracaridan NIS invasions with<br />
temperature and salinity conditions may reveal where, and perhaps how, climates control NIS<br />
distributions. NIS peracaridans are unlikely to be as suited to local climates as native species<br />
that have had more evolutionary time to adapt. From a maximum possible <strong>of</strong> 3, the average<br />
occurrence <strong>of</strong> NIS between San Francisco Bay, Puget Sound and Prince William Sound is 1.4<br />
compared to 1.8 records per native species (Table 9C4.2). This restriction <strong>of</strong> NIS peracaridan<br />
distributions in the northeast Pacific relative to native peracaridan species may result from their<br />
different adaptations to climate. NIS peracaridan dispersal vectors are sufficient to move NIS<br />
peracaridans to Prince William Sound. Port Valdez is the third largest ballast water port in the<br />
U.S. and the associated nonindigenous ballast water species that it receives are diverse and<br />
abundant (Hines et al. 1999) but none <strong>of</strong> these peracaridan species have been discovered in<br />
Prince William Sound. At the same time, the same nonindigenous ballast water species are<br />
invading San Francisco Bay at an accelerating rate (Cohen and Carlton 1998). The greater<br />
diversity <strong>of</strong> NIS in San Francisco Bay (Figure 9C4.1), the complete overlap between Puget<br />
Sound NIS and San Francisco Bay NIS and the absence <strong>of</strong> NIS in the Prince William Sound<br />
collections (Table 9C4.1), indicate that peracaridans have warm water origins and survive poorly<br />
in the cold-water areas <strong>of</strong> the northeast Pacific.<br />
The introductions <strong>of</strong> Corophium volutator and Orchestia gammarellus (Table 9C4.2)<br />
from Europe to eastern North America are exceptions to the western ocean to eastern ocean<br />
pattern <strong>of</strong> introductions listed here. Corophium volutator is confined in North America to the<br />
Bay <strong>of</strong> Fundy. Orchestia gammarellus is confined in North America to the Bay <strong>of</strong> Fundy and<br />
the outer coasts north to Newfoundland (Bousfield 1973, Watling 1979). The climates <strong>of</strong> these<br />
areas are isolated from the Gulf Current and have narrower temperature ranges than areas either<br />
to the north or south (Bousfield 1973). The successes <strong>of</strong> these two species, and other European<br />
species such as the green crab Carcinus maenus and the European littorine snail Littorina littorea<br />
in this region, and the Bay <strong>of</strong> Fundy in particular, may result from the closer match between the<br />
climate <strong>of</strong> this region and the climate <strong>of</strong> northern Europe.<br />
The occurrence <strong>of</strong> 5 NIS species <strong>of</strong> amphipods in segregated ballast water <strong>of</strong> tankers<br />
discharging into Port Valdez indicates that this is an active mechanism <strong>of</strong> transport introduding<br />
NIS arriving to Prince William Sound, even though none <strong>of</strong> these species appears to have<br />
become established yet.<br />
Conclusions<br />
These results reveal that climate and evolution interact to prevent estuarine peracaridan<br />
NIS from invading south central Alaska. Western ocean introductions <strong>of</strong> peracaridans to eastern<br />
ocean estuaries are common while few eastern ocean peracaridan species spread in the opposite<br />
direction. Unless Arctic Ocean passages <strong>of</strong> ballast water from the north Atlantic become<br />
possible, or global climate changes significantly, or massive shipping from high latitudes <strong>of</strong> the<br />
southern hemisphere occurs, ballast water sources NIS peracaridans from climates similar to<br />
south central Alaska are too remote to pose a significant risk.
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 20<br />
Like a lock and key, the adaptations <strong>of</strong> successful invaders must be sufficient to survive<br />
in climates and types <strong>of</strong> disturbances that occur in the new invaded areas. Thermal adaptations<br />
<strong>of</strong> western ocean species may thus fit eastern ocean climates while the thermal adaptations <strong>of</strong><br />
most eastern ocean species may be insufficient for western ocean climates. By the same<br />
mechanism, seasonal salinity disturbances may confine western ocean NIS peracaridans in<br />
eastern ocean estuaries to areas where the most stable salinity conditions occur and control which<br />
taxonomic groups <strong>of</strong> NIS peracaridans predominate in particular estuaries. This pattern <strong>of</strong><br />
invasion appears to be consistent among many taxa even though the particular interacting<br />
mechanisms <strong>of</strong> climate and adaptation creating the pattern remain unresolved and the particular<br />
species that invade remain unpredictable. However, predicting where species can survive<br />
provides a means to identify, particular mechanisms <strong>of</strong> introduction, source regions for NIS <strong>of</strong><br />
greatest concern, the likely secondary dispersal routes <strong>of</strong> newly introduced species and the<br />
possible role <strong>of</strong> climate changes on further invasions.<br />
Seasonal temperature variations are difficult for shallow water organisms to avoid. The<br />
extreme high and low temperature ranges <strong>of</strong> western ocean climates (Figures 9C4.4 and 9C4.5)<br />
create adaptations that span most eastern ocean temperature ranges. The reverse is less likely<br />
and must prevent survival <strong>of</strong> many western ocean introductions <strong>of</strong> eastern ocean species. The<br />
entire sea surface temperature range <strong>of</strong> the northeast Pacific between 35 and 50 o N is overlapped<br />
by the temperature ranges <strong>of</strong> the western ocean coasts between 37 and 42 o N (Figure 9C4.6).<br />
Thermal limits for the NIS peracaridans at 60 o N (Prince William Sound) are apparent<br />
also for the Pacific oyster, Crassostrea gigas (Thunberg, 1795), native to the western Pacific and<br />
cultured commercially in Asia as far north as Hokkaido, Japan (Quayle, 1969) at about 44 o N.<br />
Crassostrea grows but does not spawn in the low temperatures <strong>of</strong> Prince William Sound or south<br />
central Alaska (Foster 1991, Hines and Ruiz 1997). The minimum monthly temperatures <strong>of</strong><br />
Prince William Sound match western Atlantic and western Pacific minimum temperatures as far<br />
south as 44 o N (Figure 9C4.6). Thus, lethal low temperatures might not be encountered in the<br />
sound by western ocean NIS peracaridans from 44 o N or even slightly farther south. However,<br />
maximum northeast Pacific sea surface temperatures at 60 o N match western Atlantic and<br />
western Pacific maximums only as far south as 48 o N (Figure 9C4.6) the inability <strong>of</strong> C. gigas to<br />
spawn in Alaska is therefore not surprising.<br />
The poor match <strong>of</strong> seasonal precipitation across oceans (Figures 9C4.4 and 9C4.5) may<br />
also affect the patterns <strong>of</strong> introduction. The range <strong>of</strong> coinciding temperature and salinity<br />
tolerances that peracaridan species require to survive and reproduced in south central Alaska<br />
may prohibit NIS peracaridans that also survive in Long Beach, San Francisco or Puget Sound.<br />
The low summer salinities in Prince William Sound (Hines and Ruiz 1997) due to snow melt,<br />
more closely matches a western ocean climate (Figures 9C4.4 and 9C4.5). Only the largest<br />
eastern ocean estuaries or estuaries with impounded freshwater sources, such as San Francisco<br />
Bay, are likely to have stable haloclines in summer that match those <strong>of</strong> western ocean estuaries.<br />
The high diversity and predominance <strong>of</strong> benthic peracaridan NIS in the northeast Pacific<br />
(Table 9C4.1) may result in part from their superior adaptive responses to large salinity ranges
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 21<br />
(Figure 9C4.6). All peracaridan life stages are highly mobile, and their short reproductive cycles<br />
allow dispersal away from unsuitable conditions and rapid recruitment when conditions improve<br />
(e.g., Watkin 1941). Peracaridans can avoid rapid changes in salinity (Figures 9C4.4 and 9C4.5)<br />
by migrating short vertical distances or by swimming into water masses in which transport them<br />
to higher or lower salinity areas.<br />
San Francisco Bay may be particularly suited to NIS peracaridans that cannot avoid or<br />
quickly adapt to changing salinities. The particular predominance and high diversity <strong>of</strong> large,<br />
long-lived, suspension feeding NIS in San Francisco Bay (Carlton 1979, Nichols et al. 1990,<br />
Thompson 1998) may result from human water impoundments that limit major freshwater run<strong>of</strong>f<br />
events. San Francisco Bay is the largest estuary <strong>of</strong> the eastern Pacific. Massive water diversions<br />
and impoundments in the San Francisco Bay watershed, aided by its large size, create a stable<br />
salinity structure more typical <strong>of</strong> western ocean estuaries. Sedentary, long-lived species,<br />
including molluscs (Nicholls et al. 1990), burrowing decapods (Posey et al. 1991, Grosholz and<br />
Ruiz 1995, Cohen and Carlton 1997) and sedentariate polychaetes (Pearson and Rosenberg<br />
1978) predominate in benthic communities in the absence <strong>of</strong> major salinity disturbances. These<br />
taxa can control the trophic dynamics in estuaries (Nichols et al. 1986, Kimmerer et al. 1994,<br />
Barber 1997, Thayer et al. 1997, Thompson 1998) but are slow to repopulate areas following<br />
disturbances.<br />
The vulnerability <strong>of</strong> estuaries to invasion and the potentials <strong>of</strong> particular taxa and life<br />
history types to become invasive may be increasing globally and increase the risk <strong>of</strong> NIS<br />
invasions <strong>of</strong> Alaskan waters in the future. <strong>Water</strong> diversions and impoundments and land use<br />
practices on western coasts combined with global temperature increases are reducing the<br />
differences between climates <strong>of</strong> eastern and western ocean estuaries. The convergence <strong>of</strong><br />
climates will increase the potential for biological exchanges.<br />
The predominantly western ocean to eastern ocean direction <strong>of</strong> peracaridan invasions and<br />
south to north gradient in peracaridan invasions support a lock and key hypothesis in which<br />
peracaridan NIS cannot be introduced outside <strong>of</strong> their tolerance ranges. Obvious predictions <strong>of</strong><br />
this hypothesis that should be tested are: 1) sources <strong>of</strong> eastern ocean invaders are from a narrow<br />
range <strong>of</strong> western ocean latitudes; 2) eastern ocean invaders <strong>of</strong> western oceans have restricted<br />
geographical ranges; 3) specific physiological tolerances and life history adaptations <strong>of</strong> most NIS<br />
exceed the stresses experienced in the climates invaded; 4) southern hemisphere NIS invasions<br />
superimpose on and are superimposed upon by northern hemisphere NIS when their origins are<br />
from similar climates; 5) climates affect the life histories and taxonomic composition <strong>of</strong> invaders<br />
and; 6) invaders <strong>of</strong> western to western or eastern to eastern oceans have broader ecological and<br />
geographical ranges than mixed climate invaders.<br />
The analysis <strong>of</strong> climates and the geography <strong>of</strong> introductions must include more species<br />
and taxa in more detail than is possible here. However, failure to determine the origins <strong>of</strong> the<br />
cryptogenic peracaridan species (Table 9C4.1) are a more critical short-coming <strong>of</strong> this risk<br />
analysis. The eight cryptogenic peracaridans are abundant over broad ecological distributions<br />
within the south central Alaska. Even though none <strong>of</strong> the cryptogenic peracaridan species
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 22<br />
appear to be associated with ballast water as a means <strong>of</strong> introduction, these species are abundant<br />
and wide spread. If introduced, they are pro<strong>of</strong> that even a few NIS invaders <strong>of</strong> Alaskan estuaries<br />
can increase to ecologically catastrophic densities. Moreover surveys <strong>of</strong> NIS diversity, such as<br />
this one, are an insufficient for estimating ecological risks if these species are introduced. Due to<br />
the potential for a single species to produce massive impacts, conclusions <strong>of</strong> risk from global<br />
patterns <strong>of</strong> diversity <strong>of</strong> diversity and climate therefore could conflict with conclusions <strong>of</strong> risk<br />
based on presence <strong>of</strong> even a single NIS in a system. The occurrence <strong>of</strong> 5 NIS species <strong>of</strong><br />
amphipods in segregated ballast water <strong>of</strong> tankers discharging into Port Valdez indicates that this<br />
is an active mechanism <strong>of</strong> transport introduding NIS arriving to Prince William Sound, even<br />
though none <strong>of</strong> these species appears to have become established yet.<br />
Acknowledgments<br />
James Carlton, Williams College, Maritime Studies Program provided many inspirations<br />
leading to this project, pointing out problems and providing solutions. Todd Miller read an early<br />
draft <strong>of</strong> this manuscript. Robert Malouf and Jan Auyong, NOAA Oregon State University Sea<br />
Grant College Program; and Leon Cammen, NOAA National Sea Grant College Program and<br />
the Oregon State legislature for assistance with in kind and direct financial support from Grants<br />
NA0061RD09 and NA0061RD19; Patrick Clinton and Bryan Coleman, OAO, and Walter<br />
Nelson, US EPA, <strong>Coastal</strong> Ecology Branch, Newport, Oregon for technical assistance and<br />
facilities use. The Prince William Sound survey was greatly assisted by Joel Kopp and Robert<br />
Benda, Regional Citizens’ Advisory Council <strong>of</strong> Prince William Sound (RCAC), Nora Foster,<br />
Howard Feder and Mike Stekoll, University <strong>of</strong> Alaska; Anson Hines, Greg Ruiz and George<br />
Smith, Smithsonian Environmental Research Center; Gary Sonnevil, US Fish and Wildlife<br />
Service; the personnel <strong>of</strong> the Alyeska Pipeline Service Company and Valdez Marine; Todd<br />
Miller and Gayle Hansen, Oregon State University, Milos Falta, owner, and Capt. Richard<br />
Vodica <strong>of</strong> the “Kristina”, Steve Totemov and Gary Kompk<strong>of</strong>f, <strong>of</strong> Tatitlek and Gail Irvine <strong>of</strong> the<br />
USGS, Anchorage. The Puget Sound survey was funded by the United States Department <strong>of</strong><br />
Natural Resources, the Washington State Department <strong>of</strong> Natural Resources through Helen Berry<br />
and Betty Bookheim and facilities were provided by the University <strong>of</strong> Washington (UW),<br />
Western Washington State University (WWU), organized by Claudia Mills, (UW) and Andrew<br />
Cohen, San Francisco Estuary Institute, participants included Brian Bingham,, James Carlton,<br />
Leslie Harris, L.A. County Museum, Alan Kohn, UW, Terry Klinger, Charles and Gretchen<br />
Lambert, Kevin Li, David Secord, WWU, Jason T<strong>of</strong>t, UW and Marjorie Wonham, UW. The<br />
San Francisco Bay survey was funded through grants from the USFWS, Oregon Sea Grant and<br />
PEW travel funds to James Carlton and organized by Andrew Cohen and James Carlton.<br />
Participants included Claudia Mills, Leslie Harris, John Holliman, Gretchen Lambert, Jan<br />
Thompson, USGS, Menlo Park, Terry Gosliner, California Academy <strong>of</strong> Sciences, Jeff Crooks,<br />
U.C. San Diego.<br />
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Bull. Zool. Mus. Univ. Amsterdam, 15(1):1-4.<br />
Posey, M. H., B. R. Dumbauld and D. A. Armstrong 1991. Effects <strong>of</strong> burrowing mud shrimp<br />
Upogebia pugettensis (Dana), on abundances <strong>of</strong> macr<strong>of</strong>auna, J. Exp. Mar. Biol. Ecol. 148:283-<br />
294.<br />
Quayle, D. B. 1969. Pacific oyster culture in British Columbia, Fish. Res. Bd. Canada, Bull.,<br />
169:1-192<br />
Ruiz, G. M., J. T. Carlton, E. D. Grosholz and A. H. Hines 1997a. Global invasions <strong>of</strong> marine<br />
and estuarine habitats by non-indigenous species: mechanisms, extent, and consequences,<br />
American Zoologist 37:621-632.<br />
Ruiz, G. M., P. F. F<strong>of</strong>on<strong>of</strong>f, A. H. Hines and J. T. Carlton 1997b. Analysis <strong>of</strong> nonindigenous<br />
species invasions <strong>of</strong> the Chesapeake Bay (USA) Part 1. Report submitted to the U.S. Fish and<br />
Wildlife Service, Washington.<br />
Ruiz, G. M. and A. H. Hines 1997. The risk <strong>of</strong> nonindigenous species invasion in Prince<br />
William Sound associated with oil tanker traffic and ballast water management: Pilot study, Final<br />
Report to Regional Citizens’s Advisory Council <strong>of</strong> Prince William Sound, 46 pp. plus<br />
appendices.
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 30<br />
Siegel, S. 1956. Nonparametric statistics for the behavioral sciences, McGraw-Hill, New York,<br />
312 pp.<br />
Smith, L. D., M. J. Wonham, L. D. McCann, G. M. Ruiz, A. H. Hines and J. T. Carlton 1999.<br />
Invasion pressure to a ballast-flooded estuary and an assessment <strong>of</strong> inoculant survival, Biol.<br />
Invas., 1(1):67-87.<br />
Smith, P., J. Perrett, P. Garwood and P. G. Moore 1999. Two additions to the UK marine fauna:<br />
Desdemona ornata Banse, 1957 (Polychaeta, Sabellidae) and Grandidierella japonica<br />
Stephensen, 1938 (Amphipoda, Gammaridea), Newslet. Porcupine Mar. Nat. Hist. Soc., (2):8-11<br />
Southward, A. J. 1969. Life on the seashore, Harvard University Press, Cambridge, Maryland<br />
153 pp.<br />
Thayer, S. A., R. C. Haas, R. D. Hunter, R. H. Kushler 1997. Zebra mussel Dreissena<br />
polymorpha effects on sediment, other zoobenthos, and the diet and growth <strong>of</strong> adult yellow perch<br />
(Perca flavescens) in pond enclosures. Canadian Journal <strong>of</strong> Fisheries and <strong>Aquatic</strong> Science,<br />
54(8): 1903-1915.<br />
Thompson, B. 1998. Benthic macr<strong>of</strong>aunal assemblages <strong>of</strong> San Francisco Bay and Delta,<br />
Interagency Ecological Program for the Sacramento-San Joaquin Estuary Newsletter, 11(2):26-<br />
32.<br />
Thompson, J. K. 1998. Trophic effects <strong>of</strong> Potamocorbula amurensis in San Francisco Bay,<br />
California, Proc. Eighth Int. Zebra Mussel Conf., 1(Pembroke, Ontario, Canada):171-172.<br />
Thresher, R. E. 1999. Marine bio-invasions: Take-home from ten years <strong>of</strong> managing the problem<br />
in Australia, Abstract, In J. Pederson (ed.) National Conference on Marine Bioinvasions, 24-27<br />
Jan. 1999, Massachutsets Institute <strong>of</strong> Technology, Cambridge, Massachusetts.<br />
T<strong>of</strong>t, J., J. Cordell and C. Simenstad 1999. More non-indigenous species First records <strong>of</strong> one<br />
amphipod and two isopods in the delta, IEP Newsletter, 12(4):35-37.<br />
Watkin, E. E. 1941. Observations on the night tidal migrant Crustacea <strong>of</strong> Kames Bay, Journal <strong>of</strong><br />
the Marine <strong>Biological</strong> Association, U.K. 25:81-96.<br />
Watling, L. 1979. Zoogeographic affinities <strong>of</strong> northeaster North American gammaridean<br />
Amphipoda, pp. 256-282 In A. B. Williams (ed.), “Symposium on the composition and evolution<br />
<strong>of</strong> crustaceans in the cold and temperate waters <strong>of</strong> the world ocean” Bulletin <strong>of</strong> the <strong>Biological</strong><br />
Society <strong>of</strong> Washington, No. 3.<br />
Watling, L. 1991. Revision <strong>of</strong> the cumacean family Leuconidae, Journal <strong>of</strong> Crustacean Biology,<br />
11(4):569-582
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 31<br />
Williams, W. D. 1972. Occurrence in Britain <strong>of</strong> Asellus communis Say, 1818, a North American<br />
freshwater isopod, Crustaceana, Supplement 3:134-138.
Appendix Table 9C4.5. Site Descriptions<br />
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 32
Appendix Table 9C4.5. Continued<br />
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 33
Appendix Table 9C4.6. Peracardian Crustacea August 1999<br />
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 34
Appendix Table 9C4.7. UAF unidentified Amphipoda<br />
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 35
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 36<br />
APPENDIX TABLE 9C4.8 Descriptions <strong>of</strong> Amphipod Species Identified in Ballast <strong>Water</strong><br />
Tanks.<br />
Hyperidae<br />
Hyperidae<br />
Hyperia cf. medusarum (Bowman 1973:6-10, figs. 2-6 and references therein; Brusca<br />
1981:21, fig. 9e; Vinogradov et al. 1996:323-327, fig. 131 and references therein). Hyperiaa<br />
medusarum is a morphologically variable bipolar pelagic species <strong>of</strong> cold and moderately cold<br />
water marine regions <strong>of</strong> both hemispheres, occurring in the Bering Sea, the Gulf <strong>of</strong> Alaska and<br />
the coastal waters <strong>of</strong> Canada and the western U. S. (Vinogradov et al. 1996). Specimens <strong>of</strong> this<br />
study are all less than 4 mm in length, and none are mature. Gnathopods, mandibles and<br />
pereopods closely resemble H. medusarum but the identifications are tentative. The sutures<br />
between pereonites 1 and 2 and between coxal plates and pereionites are faint. Their appearance<br />
only in the Anacortes samples and in the open ocean exchanged ballast water samples (Table 1)<br />
are consistent with the life history <strong>of</strong> typical hyperiids and this species is very likely native to the<br />
region.<br />
Gammaridea<br />
Ampeliscidae<br />
Ampelisca abdita Mills, 1964a; Northeast Pacific records <strong>of</strong> Ampelisca milleri are: Jones<br />
1961:253-254; Filice 1959a:183; Filice 1959b:10; Chapman and Dorman 1975:107, 106;<br />
Chapman 1988:365-368, fig. 2 (and references therein); Dickensen 1982:15-17, fig. 9; (not<br />
Barnard 1954b:9-11). The range <strong>of</strong> A. abdita on the east coast <strong>of</strong> the U.S.extends from Maine to<br />
the eastern Gulf <strong>of</strong> Mexico (Mills, 1967, Bousfield 1973, Chapman 1988).<br />
Ampelisca abdita from San Francisco Bay, Bolinas Lagoon and Tomales Bay was<br />
confused with Ampelisca milleri Barnard 1954 for 40 years and was probably introduced into<br />
San Francisco Bay from the eastern U. S. with shipments <strong>of</strong> the eastern oyster Crassostrea<br />
virginica(Chapman 1988). The species dominates s<strong>of</strong>t, subtidal sedimentst’s <strong>of</strong> San Francisco<br />
Bay at salinities between 10 and 25 PSU and must occasionally occur as zooplankton in massive<br />
numbers as it seasonally exits and repopulates shallow subtidal an intertidal mudflats (Mills<br />
1967).<br />
Arigissidae<br />
Argissa hamatipes (Norman 1869); Syrrhoe hamatipes Norman 1869:279; Boeck<br />
1871:125; Argissa stebbingi Bonnier 1896:626-630, pl.36, fig.4; Argissa typica Sars<br />
1895:141-142, pl.48; Chevreux & Fage 1925:90, figs.81-82; Ruffo 1982:159-161, figs. 106-107;<br />
Argissa hamatipes Walker 1904:246; Stebbing 1906:277; Shoemaker 1930:37-40, figs.15-16;<br />
Stephensen 1935:140; Stephensen 1940:41; Stephensen 1944:52; Gurjanova 1951:327-328,<br />
fig.193; Gurjanova 1962:392-393; J.L. Barnard 1962c:151; J.L. Barnard 1964a:218-219; Nagata<br />
1965:154-155, fig.7; J.L. Barnard 1966a:61; J.L. Barnard 1967a:14-15, fig.1d-i; J.L. Barnard<br />
1969b:159, fig. 65; J.L. Barnard 1971b:9; Bousfield 1973:121-122, pl.XX; Griffiths 1975:;<br />
Lincoln 1979:334, fig.157; Ledoyer 1982:144-146, fig. 50; Hirayama 1983:147-149, figs.38-41;<br />
Barnard & Barnard 1983:607-608; Thomas & McCann 1997:22, fig.2.1.<br />
Argissa hamatipes is an entirely marine, nearly cosmopolitan species with an extensive<br />
bathymetric distribution that ranges throughout coastal north Pacific shelf regions from southern<br />
California to southern Japan and the north Atlantic from North Carolina Greenland and Iceland,
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 37<br />
throughout northern Europe and the western Mediterranean, Madagascar and South Africa.<br />
Many <strong>of</strong> the the synonymies are unclear. This extremely dispersed species is likely to be a<br />
species complex and the identity <strong>of</strong> the North Pacific population is probably incorrect.<br />
Nevertheless, the very broad open ocean distribution in the northeast Pacific population strongly<br />
indicates that it is a native to the region.<br />
Corophiidae<br />
Monocorophium acherusicum Podacerus cylindricus Lucas 1842:232; Corophium<br />
cylindricum Smith 1873:566; Paulmier 1905:167, fig. 37; Holmes 1905:521-522, fig. ; C.<br />
cylindricus Stebbing 1914:372-373; Kunkel 1918:171-173, fig. 52; Corophium contractum<br />
Thompson 1881:220-221, fig. 9; Corophium bonnellii K. H. Barnard 1932:244 (in Crawford<br />
1937); Corophium acherusicum 1853:178; Costa 1857:232, Fig. 1827; Bate 1862:282; Heller<br />
1867:51-52, pl 4 fig. 14; DeElla Valle 1893:364-367, pl. I, Fig. II, Pl. 8, figs. 17, 18, 20-41;<br />
Sowinsky 1897:9; Sowinsky 1898:455; Chevreux 1900a:109; Graeffe 1902:20; Holmes<br />
1905:521-522, fig. ; Stebbing 1906:692-740; Chevreux 1911:271; K. H. Barnard 1916:272-274;<br />
Stebbing 1917a:448; Ussing and Stephensen 1924:78-79; Chevreux 1925c:271; Chevreux and<br />
Fage 1925:368, fig. 376; Chevreux 1926:392; Cecchini 1928e:8, pl. 1, fig. 6a; Cecchini<br />
1928b:309-312, fig. 1; Schellenberg 1928:672;Schijfsma 1931a:22-25; Monod 1931a:499; Fage<br />
1933:224; Candeias 1934:3; Shoemaker 1934c:24-25; Cecchini-Parenzan 1935:227-229, fig. 52;<br />
Shoemaker 1935c:250; Crawford 1936:104; Schellenberg 1936c:21; Schijsfma 1936:122-123;<br />
Crawford 1937:617-620, 650, fig. 2; Monod 1937:13; Miloslavskaya 1939:148-149; K. H.<br />
Barnard 1940:482; Bassindale 1941:174; Stephensen 1944a:134; Shoemaker 1947:53, figs. 2, 3;<br />
Shoemaker 1949a:76; Soika 1949:210-211; Gurjanova 1951:977-978, fig. 680; Reid 1951:269;<br />
Stock and Bloklader 1952:4-5; J.L. Barnard 1954a:36; Hurley 1954e:442-445, figs. 35-39; J. L.<br />
Barnard 1955a:37; Irie 1957:5-6, fig. 6; Irie 1958c:145; Irie 1959: tab. 4; J.L.Barnard 1959:38;<br />
Nayar 1959:43-44, pl. XV, figs. 14-20; Irie In Okada and Ochida et al. 1960:122, pl. 61, fig. 12;<br />
Nagata 1960:177; J. L. Barnard 1961:182; Irie and Nagata 1962:20; Nagata 1964:10; Barnard<br />
1964a:111, chart 5; Nagata 1965c:317; Nagata 1966:334; Kikuchi 1966:, tab 21; Kikuchi<br />
1968:179; Reish and J. L. Barnard 1967:16; Ledoyer 1968:214; Fearn-Wannan 1968b:134-135;<br />
Mordhukai-Boltovskoi 1969:485, pl. 25, fig. 2; Sivaprakasam 1969d:156, fig. 14; Bellan-Santini<br />
1971:260-261; J. L. Barnard 1971a:59; J. L. Barnard 1972b:48; Bousfield 1973:201, Pl. LXII.2;<br />
Griffiths 1974:181-182;Griffiths 1974b:228; Griffiths 1974c:281; Griffiths 1975:109; Fox and<br />
Bynum 1975:225; Hirayama 1984:13, fig. 50; Azuma 1986:77; Sudo et al. 1987:1570; Inaba<br />
1988:141; Monocorophium acherusicum Bousfield and Hoover 1997:111-114, figs. 26-27.<br />
Monocorophium acherusicum could be the most widely distributed and widely<br />
introduced estuary invertebrate in the world, occurring in fouling and benthic mud communities<br />
<strong>of</strong> shallow and intertidal areas on all continents except Antarctica. C. acherusicum has been<br />
reported at all latitudes between 60 o North and South. However, many <strong>of</strong> the records <strong>of</strong> this<br />
species at latitudes greater than 60 o , including all records from Alaska are doubtful (Hines et al.<br />
1999) due likely confusion with other species. M. acherusicum was not found in the present<br />
surveys <strong>of</strong> south central Alaska, including Port Valdez and Prince William Sound.<br />
Parthenogenic populations <strong>of</strong> an extremely similar species, that closely resembles<br />
Monocorophium carlottensis Bousfield and Hoover, 1997, occurs in nearly all fouling<br />
communities <strong>of</strong> these areas (Chapman 1999, Peracarida and Decapoda). All Alaskan records <strong>of</strong>
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 38<br />
the nonindigenous species, M. insidiosum and M. acherusicum are probably referrable to M.<br />
carlottensis.<br />
Sinocorophium heteroceratum (Yu, 1938) Corophium heteroceratum Yu 1938:93-101,<br />
figs. 7-11; Sinocorophium heteroceratum Bousfield and Hoover 1997:75, 78.<br />
Sinocorophium heteroceratum was introduced into San Francisco Bay and Los Angeles<br />
Harbor from Asia in the mid 1980s where it presumed to have been transported with ballast<br />
water traffic (Cohen and Carlton 1995). This species occurs predominantly in s<strong>of</strong>t sediments in<br />
high salinity, sub-tidal areas <strong>of</strong> San Francisco Bay and Los Angeles Harbor (Chapman and Cole,<br />
MS in preparation). Sinocorophium heteroceratum are also unlikely to have been introduced<br />
with aquaculture industries, or ship fouling in San Francisco Bay or Los Angeles Harbor because<br />
its lack <strong>of</strong> association with fouling communities.<br />
Grandidierella japonica Stephensen, 1938:179-184; Nagata 1960:179; Nagata<br />
1965c:320-321; Chapman and Dorman 1975:104-108, 4 figs., Nagata 1984:15, fig. 53-56; Muir<br />
1997; Smith et al. 1999:8-9, fig. 3.<br />
Grandidierella japonica occurs in salinities from 5-40 PSU predominantly in warm<br />
intertidal areas <strong>of</strong> nearly all estuaries from Puget Sound to San Diego. Dense populations are<br />
especially common in low-salinity tidepools and seepage areas; mixed sediment. The species has<br />
been introduced to Hawaii (Muir 1997) England (Smith et al. 1999), and estuaries <strong>of</strong> the<br />
northeast Pacific between Puget Sound and San Diego (Chapman and Dorman 1975, Staude<br />
1997, Bay et al. 1988). Grandidierella was most likely introduced to the northeast Pacific with<br />
Pacific oysters from Japan between the 1930s and 1950s (Chapman and Dorman 1975). Its<br />
recent arrival in Europe (Smith et al. 1999) is more likely to be associated with ballast water. It<br />
was misidentified in Hawaii for 25 years as Neogamphopus cabinae (Muir 1997) where the<br />
mechanism <strong>of</strong> its introduction is unclear. The massive ballast water traffic from San Francisco<br />
Bay to Hawaii is a possible mechanism for its introduction there.<br />
Cyphocharidae<br />
Cyphocharis challengeri Stebbing 1888: ;Birstein and Vinogradov 1955:212 (with<br />
references); J. L. Barnard 1961b:31; Bowman and McCain 1967:1-14, figs. 1-9 (with<br />
references).<br />
The most common epipelagic gammaridean amphipod in sub-Arctic <strong>of</strong>fshore and coastal<br />
waters <strong>of</strong> the North Pacific is Cyphocharis challengeri (Bowman and McCain 1967). Bousfield<br />
and McCain (1967) demonstrated that most <strong>of</strong> the variation in the anterior protrusion <strong>of</strong><br />
pereonite 1 varies with size. The species is endemic to the North Pacific but has been reported<br />
from a broad range <strong>of</strong> latitudes in the Atlantic and Indian Ocean. .<br />
Eusiridae<br />
Pontogeneia rostrata Gurjanova, 1938; Gurjanova, 1938:330,398, fig.39; Gurjanova,<br />
1951:719, fig.500; Nagata 1960:171-173, pl.14; J.L.Barnard, 1962b:81; J.L.Barnard,<br />
1964b:114-116, fig.20; Nagata 1965b:185, fig. 26; Nagata 1965e:563; Nagata 1966:334;<br />
J.L.Barnard, 1969a:111,112,114; Itoh 1970:29; Mukai 1971:178; Itoh, Honma and Kakimoto<br />
1972:25; Honma and Kitami 1978:40; Azuma 1980:28; Barnard 1979a:49, figs. 25-27 (part);<br />
Imada et al. 1981:127; Itoh 1981:24; Itoh, Honma and Kitami 1982:41; Hirayama 1985a:28;<br />
Azuma et al. 1985:4; Azuma 1986:74; Sudo et al. 1987:1570; Inaba 1988:146; Ariyama<br />
1988:129, fig. 13f; Barnard and Karaman 1991:334; Ishimaru 1994:45.
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 39<br />
A widely reported species in the North Pacific from Bahia de San Quintin, Mexico<br />
(Barnard 1964) to Alaska and down the Asian coast to southern Japan (Ishimaru, 1994).<br />
Pontogeneia rostrata was recovered in the nonindigenous species surveys <strong>of</strong> Prince William<br />
Sound (Hines 1999). However, P. rostrata can easily be confused with species <strong>of</strong> Accedemorea<br />
its extremely broad range in the Northeast Pacific and uncertain taxonomic status prevents clear<br />
resolution <strong>of</strong> whether it is endemic or introduced. This species is cryptogenic (Carlton 1996).<br />
Gammaridae<br />
Gammarus daiberi Bousfield, 1969; Bousfield 1969:10(1)4-8, figs. 1& 4; Bousfield<br />
1973:52, pl. IV.2; T<strong>of</strong>t et al. 1999:36.<br />
A probable 1980s ballast water introduction into San Francisco Bay from the eastern<br />
Unites States. Gammarus daiberi is an estuarine species most abundant in the low salinity<br />
ranges between 1.5 and 15 PSU and is largely pelagic. Except for this report, this species is<br />
unknown in the northeast Pacific outside <strong>of</strong> San Francisco Bay. Its range in the eastern U.S.<br />
extends from Delaware Bay to South Carolina (Bousfield 1973). Gammarus daiberi is a<br />
probable ballast water introduction into San Francisco Bay that arrived in the 1980s.<br />
Eogammarus confervicolus Mara confervicola Stimpson 1856:90; Gammarus<br />
confervicolus Stimpson 1857:520-521; Bate 1862:218, pl. 38, fig. 9; Holmes 1904:239; Melita<br />
confervicola Stebbing 1906:428; Anisogammarus confervicolus Saunders 1933:248 (in part); J.L.<br />
Barnard 1954a:9-12, pls.9-10; Shoemaker 1964:423, figs. 14-15; Pamamat 1968:211; Bousfield<br />
and Hubbard 1968:3; Anisogammarus (Eogammarus) confervicolus Schellenberg 1937a:274;<br />
Tzvetkova 1975:145-147, fig. 57; Bousfield 1958a:86, fig. 10; Eogammarus confervicolus<br />
Bousfield 1979:317-319, fig.4.<br />
Eogammarus confervicolus is the only native species recovered from ballast water that<br />
was also collected in the Prince William Sound nonindigenous species survey (Chapman 1999).<br />
Eogammarus confervicolus is extremely euryhaline and is occasionally pelagic. It occurs mainly<br />
in estuaries and protected coastal shores. Eogammarus confervicolus is the most common and<br />
widely distributed gammaroidean amphipod <strong>of</strong> the North American Pacific coast (Bousfield<br />
1979).<br />
Megaluropidae<br />
Gibberosus longimerus Hoek; Megaluropus longimerus; J.L.Barnard 1962b:103,<br />
figs.20-21; J.L.Barnard, 1964a:224; J.L. Barnard 1966b:19; J.L.Barnard, 1969a:126; J.L.<br />
Barnard 1971b:15; Gibberosus myersi (McKinney 1980); Megaluropus myersi McKinney 1980;<br />
Megaluropus longimerus <strong>of</strong> Cadien et al. NEP (in part); (not Schellenberg 1925:151-153,<br />
fig.14).<br />
The genus is poorly studied north <strong>of</strong> northern California (Cadien et al. 1997).<br />
Schellenberg (1925) figured only two appendages from his specimens from Lagos, Nigeria and<br />
the type specimens have not been compared to California material. The possibility <strong>of</strong> these<br />
populations comprising a single species seems remote. The common occurrence <strong>of</strong> this species<br />
over a broad range <strong>of</strong> coastal and nearshore waters (Barnard 1962) indicates that it is endemic.<br />
Melphidipiidae<br />
Melphisana bola Barnard 1962b:81-82, fig.7; Thomas & McCann 1997:42, fig.2.20.<br />
This native species is limited to depths no greater than 130 m on the southern California<br />
coastal shelf (Barnard 1962b, Thomas and McCann 1997). Appendages are usually missing this<br />
species on recovery from benthic samples (Barnard 1962b, Thomas and McCann 1997) greatly
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 40<br />
complicating identifications. These specimens are in excellent condition is in sharp contrast ot<br />
previous material on which the species is described.<br />
Oedicerotidae<br />
Hartmanodes hartmanae Monoculodes hartmanae J.L. Barnard 1962:363, figs. 6-7;<br />
Barnard & Karaman 1991:560; Hartmanodes hartmannae Bousfield & Chevrier 1996:92-93, fig.<br />
10. Bousfield and Chevrier (1996) did not find this species in their 200 samples from the shelf<br />
benthos and coastal waters <strong>of</strong> Canada and it is not listed in by Staude (1997). This species is<br />
native to southern California marine waters where it occurs at less than 40 m depths (Barnard<br />
1962). Hartmanodes hartmannae is the most abundant gammaridean amphipod found in the<br />
samples.<br />
Westwoodilla caecula Halimedon caecula Bate 1857:140; Halimedon Molleri Boeck<br />
1871:169-170; Halimedon Mulleri Sars 1895:327-329, pl.115; Halimedon acutifrons Sars<br />
1895:329-330, pl.116, fig.1; Westwoodilla caecula Enequist 1950:333-338, figs.40-56;<br />
Gurjanova 1951:541-543, pl. 357; Mills 1962:5-9, fig.1; J.L. Barnard 1962e:370; J.L. Barnard<br />
1964a:235; J.L. Barnard 1966a:80 (forma acutifrons); J.L. Barnard 1966b:27; J.L. Barnard<br />
1971b:51; Lincoln 1979:354, fig.167; Thomas & McCann 1997:58, figs. 2.37, 2.38; Beare, D. J.<br />
and P. G. Moore 1998.<br />
This species may not be part <strong>of</strong> a complex <strong>of</strong> similar species distributed around the Arctic<br />
Ocean, the Japan Sea, the north east Pacific from British Columbia to southern California and the<br />
North Atlantic from Greenland to the Gulf <strong>of</strong> St. Lawrence and northern Europe. This extremely<br />
widespread, common, species in <strong>of</strong>fshore marine s<strong>of</strong>t-sediment environments is most likely<br />
endemic to cold water areas <strong>of</strong> the northeast Pacific. Its occurrence in Port Valdez zooplankton<br />
samples is not surprising.<br />
Synopiidae<br />
Tiron sp.<br />
The short dactyls, spineless telson, tiny mandibular palp, smooth dorsal urosome <strong>of</strong> this<br />
single specimen do not agree with either <strong>of</strong> the local species Tiron tropakis J. L. Barnard, 1972<br />
or Tiron biocellata J. L. Barnard, 1962. Pelagic dispersal <strong>of</strong> benthic peracaridans usually occurs<br />
as adults and <strong>of</strong>ten is preceded by slight morphological changes that are adaptive for swimming.<br />
These changes are poorly understood. The low morphological correspondence <strong>of</strong> this single<br />
specimen with a known species is therefore not surprising. The specimen is therefore considered<br />
more likely to be a member <strong>of</strong> one <strong>of</strong> the above native species than an introduced species.<br />
References for Appendix Table 9C4.8 (Descriptions <strong>of</strong> Amphipod Species Identified in<br />
Ballast <strong>Water</strong> Tanks:<br />
Alldredge, A. L. and J. M. King 1980. Effects <strong>of</strong> moonlight on the vertical migration patterns <strong>of</strong><br />
demersal zooplankton, J. Exp. Mar. Biol. Ecol., 44:133-156<br />
Barnard, J. L. 1952b. Some Amphipoda from central California, Wasmann J. Biol., 10:9-36, 9<br />
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years 1873-1876, Zoology, 29, Eyrie and Spottiswoodie, London, xxiv + 1737 pp.<br />
Stebbing, T. R. R., 1906., Amphipoda. I. Gammaridea, Das Tierreich, 21, Oxford University<br />
Press, Berlin, 806 pp.<br />
Stephensen, A. 1938. The Amphipoda <strong>of</strong> northern Norway and Spitzbergen with adjacent waters,<br />
Tromso Museum Skr., 3:141-278<br />
Stephensen, K. 1944. Some Japanese Amphipods, Vidensk. Medd. Dansk Naturh. Foren.,<br />
108(4):25-88
Chapt 9C4. Peracaridan Crustaceans, page 9C4- 47<br />
Stimpson, W. 1856. On some Californian Crustacea, Proc. Calif. Acad. Sci., 1:87-90<br />
Stimpson, W. 1857. On the Crustacea and Echinodermata <strong>of</strong> the Pacific shores <strong>of</strong> North<br />
America, Boston J. Nat. Hist., 6:444-532<br />
Thomas, J. D. and L. D. McCann 1997. The families Argissidae, Dexaminidae, Eursiridae,<br />
Gammaridae, Leucothoidae, Melphidippidae, Oedicerotidae, Pardaliscidae, Phoxocephalidae,<br />
Podoceridae, Stegocephalidae, Stenothoidae, Stilipedidae, Synopiidae, and Urothoidae, pp.<br />
21-136, In J. A. Blake, L. Watling and P. H. Scott (eds.) "Taxonomic Atlas <strong>of</strong> the Benthic<br />
Fauna <strong>of</strong> the Santa Maria Basin and Western Santa Barbara Channel" Santa Barbara Museum<br />
<strong>of</strong> Natural History, Santa Barbara, California<br />
T<strong>of</strong>t, J., J. Cordell and C. Simenstad 1999. More non-indigenous species First records <strong>of</strong> one<br />
amphipod and two isopods in the delta, IEP Newslet., 12(4):35-37<br />
Tzvetkova, N. L. 1975a. Pribezhnye gammaridy severny kh I dal’nevostochnykh morei SSSR I<br />
sopredel’nykh vod, Akad. Nauk SSSR, Zool. Inst. Izdatel. "Nauka", Leningrad, :1-256<br />
Vinogradov, M. E., A. F. Volkov and T. N. Semenova, 1996., Hyperiid amphipods (Amphipoda,<br />
Hyperiidea) <strong>of</strong> the world oceans,,, Science Publishers, Inc., 10 <strong>Water</strong> Street, #310, Lebanon,<br />
NH 03766, xxvii + 632 pp. pp.<br />
Williams, A. B. and K. H. Bynum 1972. A ten-year study <strong>of</strong> meroplankton in North Carolina<br />
estuaries: Amphipods, Chesapeake Sci., 13:175-192<br />
Williams, R. J., F. B. Griffiths, E. J. Van der Wal and J. Kelly 1988. Cargo vessel ballast water<br />
as a vector for the transport <strong>of</strong> nonindigenous marine species, Estuar. Coast. Shelf. Sci.,<br />
26:409-420<br />
Yu, S. C. 1938. Descriptions <strong>of</strong> two new amphipod Crustacea from Tankgu, Bulletin <strong>of</strong> the Fan<br />
Memorial Institute <strong>of</strong> Zoology, 8:83-1
Chapt 9C5. Copepod Crustaceans, page 9C5- 1<br />
Chapter 9C5. Focal Taxonomic Collections: Copepod Crustaceans<br />
Jeffery R. Cordell, Wetland <strong>Ecosystems</strong> Team, Fishery Sciences, University <strong>of</strong> Washington,<br />
Seattle<br />
Methods<br />
Copepods were identified from three types <strong>of</strong> samples. The first method consisted <strong>of</strong><br />
sweeps that were made through algal and fouling assemblages on the underside <strong>of</strong> docks, using a<br />
small hand-held net consisting <strong>of</strong> 130 µm mesh material attached to a stainless steel hoop <strong>of</strong><br />
approximately 15 cm diameter. An effort was made to disturb algal and bivalve holdfasts in<br />
order to capture copepods from those microhabitats. The second type <strong>of</strong> samples taken were<br />
vertical water column plankton hauls made either <strong>of</strong>f a dock or from a small boat in the harbor<br />
with a 0.25 m diameter 250 µm mesh plankton net. The net was lowered to the bottom, and after<br />
waiting approximately 1 minute for disturbance to dissipate, the net was slowly pulled to the<br />
surface. The third type <strong>of</strong> samples were taken from settling plates used for collecting fouling<br />
macro-invertebrates by briefly soaking plates in 5% formaldehyde solution in a plastic tub and<br />
washing the residue through 130 µm mesh.<br />
In the laboratory, each sample was examined under a dissecting microscope, and several<br />
representatives <strong>of</strong> each species <strong>of</strong> copepod were removed. Each species was then further<br />
examined under a compound microscope. Identification was made as far as possible without<br />
dissection <strong>of</strong> individuals (with the exception <strong>of</strong> occasional removal <strong>of</strong> the abdomen to facilitate<br />
viewing the fifth leg).<br />
Results and Discussion<br />
We identified 68 species <strong>of</strong> harpacticoid copepods, six calanoids, four cyclopoids, and<br />
several unidentified poecilostomatoid species (Table 9C5.1). Of these, none is a confirmed<br />
introduced species.<br />
For harpacticoids, our results were similar to those <strong>of</strong> Kask et al. (1982) for the Nanaimo<br />
estuary in southern British Columbia in both the number <strong>of</strong> species found (75 in B.C.) and in that<br />
most <strong>of</strong> the species that were identified were either northern Pacific, broadly distributed boreal<br />
species (i.e., Pacific and Atlantic records), or probably undescribed species. Kask et al. (1982)<br />
speculated that the presence <strong>of</strong> many <strong>of</strong> the species that they found might have been the result <strong>of</strong><br />
introductions from ship fouling communities. Harpacticoid copepods may be particularly likely<br />
to be transported and introduced because as a group they have successfully occupied almost all<br />
benthic and epibenthic habitats. Also, many species have multiple life history modes (e.g.,<br />
resting and planktonic stages) that may also increase their chance <strong>of</strong> being transported and<br />
introduced. However, these same factors may also explain wide natural distributions. The<br />
paucity <strong>of</strong> studies <strong>of</strong> harpacticoid taxonomy in the northeastern Pacific makes it nearly<br />
impossible to determine whether or not a given species has been introduced without extensive<br />
distribution or genetic studies. Many <strong>of</strong> the species described in Lang’s monograph on the<br />
harpacticoids <strong>of</strong> central California (Lang, 1965) occur in Puget Sound (J. Cordell, unpublished<br />
data) and southern British Columbia (Kask et. al., 1982), and Lang’s species that also occur in<br />
Prince William Sound (Table 9C5.2) probably have continuous distributions. Other species that<br />
we encountered have Arctic and circumboreal distributions. An example <strong>of</strong> this is the important
Chapt 9C5. Copepod Crustaceans, page 9C5- 2<br />
Table 9C5.1. Copepoda<br />
Order Harpacticoida<br />
Fam. Ameiridae<br />
Ameira longipes Boeck, 1865<br />
Ameira sp. 1<br />
Ameira sp. 2<br />
Ameiridae, unid. sp. 1<br />
Fam. Ancorabolidae<br />
Arthropsyllus serratus Sars, 1909<br />
Fam. Canthocamptidae<br />
Mesochra pygmaea (Claus, 1863)<br />
Mesochra sp. 1<br />
Fam. Canthocamptidae, incertae sedis<br />
Leimia vaga Willey, 1923<br />
Fam. Cletodidae<br />
Acrenhydrosoma sp.<br />
Fam. Danielsseniidae<br />
Danielssenia typica<br />
Fam Diosaccidae<br />
Diosaccus spinatus Lang, 1965<br />
Diosaccus sp. 1<br />
Amphiascopsis cinctus<br />
Amphiascus minutus<br />
Amphiascus sp. 1<br />
Amphiascoides cf. debilis<br />
Amphiascoides sp. 1<br />
Amonardia perturbata Lang, 1965<br />
Amonardia normani<br />
Robertsonia sp.<br />
Stenhelia peniculata Lang, 1965<br />
Fam. Ectinosomatidae<br />
Ectinosoma sp.<br />
Halectinosoma sp. 1<br />
Halectinosoma sp. 2<br />
Halectinosoma sp. 3<br />
Microsetella norvegica<br />
Fam. Tachidiidae<br />
Microarthridion littorale (Poppe, 1881)
Chapt 9C5. Copepod Crustaceans, page 9C5- 3<br />
Table 9C5.1. (continued) Copepoda<br />
Fam. Harpacticidae<br />
Harpacticus uniremis Kröyer, 1842<br />
Harpacticus septentrionalis Klie, 1941<br />
Harpacticus compressus Frost, 1967<br />
Harpacticus sp.- uniremis group 1<br />
Harpacticus sp.- obscurus group 1<br />
Harpacticus sp.- obscurus group 2<br />
Zaus sp.<br />
Fam. Laophontidae<br />
Echinolaophonte sp.<br />
Heterolaophonte discophora (Willey, 1929)<br />
Heterolaophonte longisetigera Klie, 1950<br />
Heterolaophonte variabilis Lang, 1965<br />
Heterolaophonte sp. 1<br />
Laophonte elongata Boeck, 1872<br />
Laophonte applanata<br />
Laophonte sp. 1<br />
Pseudonychocamptus spinifer Lang, 1965<br />
Paralaophonte cf. congenera (Sars, 1908)<br />
Paralaophonte pacifica Lang, 1965<br />
Paralaophonte perplexa (T. Scott, 1898)<br />
Paralaophonte hyperborea (Sars, 1909)<br />
Paralaophonte sp. 1<br />
Laophontidae, unid.<br />
Fam. Longipediidae<br />
Longipedia sp.<br />
Fam. Parastenheliidae<br />
Parastenhelia sp. 1<br />
Parastenhelia sp. 2<br />
Fam. Tegastidae<br />
Tegastes sp. 1<br />
Tegastes sp. 2<br />
Fam. Thalestridae<br />
Idomene sp.<br />
Diarthrodes sp. 2<br />
Parathalestris sp. 1<br />
Parathalestris sp. 2<br />
Parathalestris sp. 3<br />
Dactylopusia vulgaris Sars, 1905<br />
Dactylopusia glacialis Sars, 1909<br />
Dactylopusia cf. glacialis<br />
Dactylopusia paratisboides Lang, 1965<br />
Dactylopusia sp. 1<br />
Paradactylopodia sp. 1
Chapt 9C5. Copepod Crustaceans, page 9C5- 4<br />
Table 9C5.1. (continued) Copepoda<br />
Fam. Tisbidae<br />
Tisbe cf. furcata (Baird, 1837)<br />
Tisbe spp.<br />
Scutellidium arthuri Poppe, 1884<br />
Order Cyclopoida<br />
Fam. Cyclopinidae<br />
Fam. Cyclopidae<br />
Euryte sp.<br />
Halicyclops sp.<br />
Fam. Oithonidae<br />
Oithona similis Claus, 1863<br />
O. spinirostris Claus, 1863<br />
Order Poecilostomatoida<br />
Unidentified spp.<br />
Order Calanoida<br />
Fam. Acartiidae<br />
Acartia cf. clausi Giesbrecht, 1889<br />
Acartia longiremis (Lilljeborg, 1853)<br />
Fam. Centropagidae<br />
Centropages abdominalis Sato, 1913<br />
Fam. Paracalanidae<br />
Paracalanus sp.<br />
Fam. Pseudocalanidae<br />
Pseudocalanus spp.<br />
Fam. Temoridae<br />
Eurytemora herdmani Thompson and Scott, 1897
Chapt 9C5. Copepod Crustaceans, page 9C5- 5<br />
Table 9C5.2. Copepods Collected at 6 Locations in Southcentral Alaska in August 1999.<br />
1999 Copepoda Homer Whittier Cordova Valdez Seward Shotgun<br />
Cove<br />
Order Harpacticoida<br />
Fam. Ameiridae<br />
Ameira longipes<br />
x<br />
Ameira sp. 1 x x x x<br />
Ameira sp. 2 x x<br />
Family Ancorabolidae<br />
Arthropsyllus serratus<br />
x<br />
Fam. Canthocamptidae<br />
Mesochra pygmaea x x<br />
Mesochra sp. 1<br />
x<br />
Fam. Canthocamptidae, incertae sedis<br />
Leimia vaga<br />
x<br />
Fam. Danielsseniidae<br />
Danielssenia typica<br />
x<br />
Fam Diosaccidae<br />
Diosaccus spinatus x x<br />
Amphiascopsis cinctus x x x<br />
Amphiascus minutus x x x x<br />
Amphiascus sp. 1 x x<br />
Amphiascoides sp. 1<br />
x<br />
Amonardia perturbata x x<br />
Fam. Ectinosomatidae<br />
Ectinosoma sp.<br />
Fam. Harpacticidae<br />
Harpacticus uniremis x x x x x<br />
H. septentrionalis x<br />
H. compressus x<br />
H. unidentified sp. x<br />
Harpacticus sp. A- uniremis group<br />
x<br />
Harpacticus sp.- obscurus group 1 x x x<br />
Harpacticus sp.- obscurus group 2<br />
x<br />
Zaus sp.<br />
Fam. Laophontidae<br />
Unidentified sp.<br />
x<br />
Heterolaophonte discophora x x<br />
Heterolaophonte longisetigera x x x<br />
Laophonte elongata<br />
x<br />
Pseudonychocamptus spinifer x x<br />
Paralaophonte cf. congenera<br />
x<br />
Paralaophonte pacifica<br />
x<br />
Paralaophonte perplexa x x x<br />
Paralaophonte sp. 1<br />
x<br />
Family Parastenheliidae<br />
Parastenhelia sp. 1<br />
x<br />
Parastenhelia sp. 2<br />
x
Chapt 9C5. Copepod Crustaceans, page 9C5- 6<br />
Table 9C5.2. (continued) Copepods Collected at 6 Locations in Southcentral Alaska in<br />
August 1999.<br />
1999 Copepoda Homer Whittier Cordova Valdez Seward Shotgun<br />
Cove<br />
Fam. Tegastidae<br />
x<br />
Tegastes sp. 1 x x<br />
Tegastes sp. 2<br />
Fam. Thalestriade<br />
Diarthrodes sp. 1<br />
x<br />
Diarthrodes sp. 2<br />
x<br />
Parathalestris sp. 1<br />
x<br />
Parathalestris sp. 2<br />
x<br />
Parathalestris sp. 3<br />
x<br />
Dactylopusia vulgaris x x x x<br />
Dactylopusia cf. glacialis<br />
x<br />
Paradactylopodia sp.<br />
x<br />
Fam. Tisbidae<br />
Tisbe cf. furcata x x<br />
Tisbe sp. x x x x<br />
Scutellidium arthuri x x<br />
Order Cyclopoida<br />
Family Cyclopinidae<br />
x<br />
Family Cyclopidae<br />
Euryte sp. x x x<br />
Halicyclops sp. x x<br />
Order Poecilostomatoida<br />
x<br />
Total Number <strong>of</strong> Taxa 28 11 20 1 3 23
Chapt 9C5. Copepod Crustaceans, page 9C5- 7<br />
juvenile salmon prey species Harpacticus uniremis, which occurs in the arctic and as far south as<br />
the English Channel in the Atlantic (Kask et al. 1982) and La Jolla, California in the Pacific<br />
(Gunnill, 1982).<br />
Settling plates appear to be a good way to sample the diversity <strong>of</strong> harpacticoid and other<br />
epibenthic/epiphytic copepods. In each case where we had dock sweep samples to compare with<br />
settling plate samples, more species were collected from the settling plates. Only one species,<br />
the algal blade dwelling Scutellidium arthuri was found only in the dock sweep samples.<br />
Reduced numbers <strong>of</strong> copepod taxa in the dock sweep samples was probably due to low and/or<br />
highly fluctuating surface salinities and/or temperatures. This assertion is supported by the fact<br />
that dock sweep samples taken in harbors with high freshwater input (e.g., Valdez, Seward) had<br />
extremely low taxa numbers, and those without much freshwater (Homer, Shotgun Cove) had the<br />
highest number <strong>of</strong> taxa.<br />
One species <strong>of</strong> harpacticoid copepod that we found in dock sweep collections, Leimia<br />
vaga, may be regarded as “probably introduced”. This species, which was described from Nova<br />
Scotia, is also abundant in many estuaries in Oregon and Washington, where it is restricted to<br />
brackish reaches (J. Cordell, unpublished data); however, this species was not reported from the<br />
Nanaimo estuary by Kask (1982). The fact that L. vaga has restricted habitat requirements and<br />
apparently disjunct populations on the Pacific coast may indicate that it has been introduced.<br />
Although a number <strong>of</strong> Asian planktonic copepods have become established in California,<br />
Oregon, and Washington estuaries (e.g., Cordell and Morrison, 1996, Orsi and Ohtsuka, 1999),<br />
we found no introduced species in the vertical haul samples. In fact, overall planktonic copepod<br />
diversity was quite low, and almost all <strong>of</strong> the copepod numbers were made up <strong>of</strong> only three taxa:<br />
Acartia longiremis, Pseudocalanus spp., and Oithona similis. Also, we did not encounter several<br />
taxa that have been previously reported from ballast water arriving to Prince William Sound<br />
(Ruiz and Hines, 1997; Hines et al., 1998; Chapt. 3 <strong>Biological</strong> Characteristics <strong>of</strong> Ballast <strong>Water</strong>).<br />
As with dock-associated harpacticoids, our shallow sampling depths that were probably subject<br />
to large fluctuations in salinity and temperature may have decreased diversity <strong>of</strong> planktonic<br />
copepods in our samples.<br />
References<br />
Gunnill, F.C. 1982. Effects <strong>of</strong> plant size and distribution on the numbers <strong>of</strong> invertebrate species<br />
and individuals inhabiting the brown alga Pelvetia fastigiata. Mar. Biol., 69: 263-280.<br />
Kask, B.A., J.R. Sibert, and B. Windecker. 1982. A check list <strong>of</strong> marine and brackish water<br />
harpacticoid copepods from the Nanaimo estuary, southwestern British Columbia. Syesis, 15:<br />
25-38.<br />
Lang, K. 1965. Copepoda Harpacticoidea from the Californian Pacific coast. Kungliga Svenska<br />
Vetenskapsakademiens Handlingar, (4)10(2), 1-560.<br />
Cordell, J.R. and S.M. Morrison. 1996. The invasive Asian copepod Pseudodiaptomus inopinus<br />
in Oregon, Washington, and British Columbia estuaries. Estuaries, 19 (3): 629-638.
Chapt 9C5. Copepod Crustaceans, page 9C5- 8<br />
Orsi, J.J., and S. Ohtsuka. 1999. Introduction <strong>of</strong> the Asian copepods Acartiella sinensis,<br />
Tortanus dextrilobatus (Copepoda: Calanoida), and Limnoithona tetraspina (Copepoda:<br />
Cyclopoida) to the San Francisco estuary, California, USA. Plank. Biol. Ecol., 46 (2): 128-131.
Chapt 9C6 Decapod Crustaceans, page 9C6- 1<br />
Chapter 9C6. Focal Taxonomic Collections: Decapod Crustaceans<br />
Anson Hines, Smithsonian Environmental Research Center<br />
Lise Schickel, University <strong>of</strong> California, Santa Barbara<br />
Nora Foster, University <strong>of</strong> Alaska Museum<br />
Results<br />
We collected a total <strong>of</strong> 21 species <strong>of</strong> decapods, including 12 species <strong>of</strong> brachyuran crabs,<br />
3 species <strong>of</strong> lithodid crabs, 3 species <strong>of</strong> hermit crabs, and 5 species <strong>of</strong> caridean shrimp (Table<br />
9C6.1). All <strong>of</strong> these species were collected by hand or dip net in the intertidal and shallow<br />
subtidal zones, or on float fouling communities. None <strong>of</strong> these species is a new record for the<br />
region (Jensen, 1995).<br />
We also noted parasitic castrators (rhizocephalan cirripedes, entoniscid isopods) <strong>of</strong><br />
decapods in the collections during August 1999. Several <strong>of</strong> the samples <strong>of</strong> the hermit crab<br />
Pagurus hirsutiusculus exhibited infections by rhizocephalan parasites. The crab<br />
Lophopanopeus bellus also has a high prevalence <strong>of</strong> infection by the rhizocephalan<br />
“Loxothylacus panopaei” at Tatitlek. (Note: The name given to this rhizocephalan in the<br />
literature by Boschma (1955) for the west coast <strong>of</strong> North America is undoubtedly incorrect and<br />
should be renamed, as this is not the same species that is found in xanthid crabs <strong>of</strong> the eastern<br />
and gulf coasts.) The population <strong>of</strong> the shore crab Hemigrapsus oregonensis also had high<br />
prevalence <strong>of</strong> the entoniscid isopod Portunion conformis.<br />
References<br />
Boschma, H. 1955. The described species <strong>of</strong> the family Sacculinidae. Zool. Verhandel. 27: 48-<br />
76.<br />
Jensen, G.C. 1995. Pacific Coast Crabs and Shrimps. Sea Challengers, Monterey, California. 87<br />
pp.
Chapt 9C6 Decapod Crustaceans, page 9C6- 2<br />
Table 9C6.1. Decapod Crustaceans in Field Surveys<br />
Decapod Crustaceans<br />
1997; 1998; 1999<br />
Homer Sew<br />
-<br />
ard<br />
Whittier<br />
Port Saw-<br />
Valdez Mill<br />
Bay<br />
Rocky<br />
Pt.,<br />
Gallen<br />
a Bay<br />
Busby<br />
Is.<br />
Tatitlek<br />
Cordovcier<br />
Gla-<br />
Is.<br />
Green<br />
Island<br />
Consta-<br />
Tine<br />
Harbor<br />
Port<br />
Chalmers<br />
Brachura<br />
Cancer magister X X X X X<br />
Cancer oregonensis X X X X X X X X X<br />
Cancer productus X X<br />
Cancer gracilis<br />
X<br />
Chionoecetes bairdi<br />
X<br />
(molts)<br />
Chorilla longipes<br />
X<br />
Hemigrapsus oregonensis X X X X X X,9 X,4 X,7 X X<br />
Lophopanopeus bellus X X X,5 X X<br />
Oregonia gracilis X X X X<br />
Pugettia gracilis X X X<br />
Scyra acutifrons<br />
X<br />
Telmessus cheiragonus X X X X X X X X X<br />
Anomura<br />
Cryptolithodes typicus<br />
X<br />
Hapalogaster grebnitzii X X X X<br />
Phyllolithodes papillosus<br />
X<br />
Pagurus beringanus X X<br />
Pagurus granosimanus X X X X<br />
Pagurus hirsutiusculus X,1 X,2 X X,3 X X X X,6 X,8 X X X X<br />
Caridea<br />
Eualus b9iunguis<br />
X<br />
Hyppolyte clarki X X<br />
Heptacarpus stimpsoni X X X<br />
Spirontocaris ochotensis X X X X<br />
Spirontocaris prionota X X<br />
Parasite Notes:<br />
Aug 1999:<br />
1 = 7/9 with bopyrid isopod<br />
2 = 1/32 with rhizocephalan Peltogaster paguri; 1/32 with rhizocephalan<br />
Peltogasterella gracilis<br />
3 = 0/9 infected<br />
4 = 30/45 with entoniscid isopod Portunion<br />
conformis<br />
5 = 19/44 with rhizocephalan Loxothylacus<br />
panopei<br />
6 = 1/6 with bopyrid isopod; 2/6 with rhizocephalan Peltogaster paguri; 1/6 with isopod<br />
and P. paguri<br />
7 = 0/13 with Portunion conformis<br />
8 = 3/30 with Peltogaster paguri;<br />
1/30 with Peltogasterella gracilis<br />
June 1998: 9 = 2/93 with<br />
rhizocephalan
Chapt 9C7. Shelled Molluscs, page 9C7- 1<br />
Chapter 9C7. Focal Taxonomic Collections: Shelled Molluscs<br />
Nora Foster, <strong>Aquatic</strong> Collection, University <strong>of</strong> Alaska Museum<br />
Methods<br />
Molluscs were collected and identified by N. Foster at sites in Prince William Sound<br />
during June 20-28, 1998 and August 8-14, 1999. In 1999 the field survey was expanded to<br />
include sites in Homer, Seward, and Hinchinbrook Entrance (see Fig. 9A.2 above). Also, J.<br />
Goddard and L. Schickel participated in the second year’s field collecting and identification,<br />
providing expertise in opisthobranch taxonomy (see Chapt 9C8. Opisthobranch Molluscs).<br />
Site information. Site information was collected by J. Chapman & T. Miller (see Chapt 9B).<br />
Sampling and processing. Presence and relative abundance (recorded as abundant, common, or<br />
rare) <strong>of</strong> the easily identified and abundant species were recorded in the field. Field notes were<br />
then transferred to spreadsheets maintained by J. Chapman. Voucher specimens <strong>of</strong> small or rare,<br />
mollusks were collected and or were preserved in 10% formalin. Identifications were made by<br />
N. Foster at the University <strong>of</strong> Alaska Museum <strong>Aquatic</strong> Collection.<br />
Identifications and distributions. No comprehensive reference is available for Alaskan marine<br />
mollusca. Major references used for both identifications and distribution records include Foster<br />
(1991) , Coan and Scott (unpublished draft ), Baxter (1987), Turgeon (1998), Kozl<strong>of</strong>f (1996),<br />
and Harbo (1997). Voucher collections form this study will be accessioned into the UAM<br />
<strong>Aquatic</strong> Collection as accession 1998-003 and 1999-001.<br />
Results<br />
Seventy-nine mollusc species were collected or identified from 27 sites in 1998 (Table<br />
9C7.1). Sixty- six three species were collected or identified from 16 sites in 1999 (Table 9C7.2).<br />
Mollusks were collected from three general types <strong>of</strong> habitats:<br />
1. Human-made structures, including oyster lantern nets, and associated floats and buoys, and<br />
docks. Characteristic mollusks from fouling communities are Mytilus trossulus, Lacuna<br />
vincta, Hiatella arctica. This represented the richest habitat for nudibranchs, including 12<br />
new records. Small mytilids were examined, to look for Musculista stenhousia.<br />
2. Mudflats. Upper intertidal areas <strong>of</strong> mudflats, near Cordova and Valdez and Homer were<br />
dominated by Mya arenaria. A search was made for Nuttalia obscurata, Batillaria cumingi,<br />
Ilyanassa obsoleta, and Nassa fratercula, but these potenial NIS were not found at any site <strong>of</strong><br />
our surveys.<br />
3. Rocky intertidal zones <strong>of</strong> beaches, sheltered bays with eelgrass. Fauna <strong>of</strong> rocky intertidal<br />
areas, both in bays and headlands is fairly well documented. It is the habitat that is richest in<br />
species, but has fewest NIS candidates or new collecting records.<br />
In combination for the two years, 115 species <strong>of</strong> molluscs were sampled (Table 9C7.3).<br />
Of these species, the s<strong>of</strong>t-shelled clam Mya arenaria is an NIS (Strasser 1999) that was widely<br />
distributed as a self-sustaining population in intertidal s<strong>of</strong>t sediments <strong>of</strong> protected bays<br />
throughout Prince William Sound. Recently, Foster examined archaeological evidence for
Chapt 9C7. Shelled Molluscs, page 9C7- 2<br />
Table 9C7.1. Shelled<br />
Molluscs 1998 Survey<br />
Harbors<br />
Valdez<br />
Cordova<br />
Whittier<br />
Mud bays<br />
Valdez<br />
Bligh Island<br />
Valdez Arm<br />
Growler Island<br />
Cordova<br />
Headlands &<br />
Reefs<br />
Valdez Arm<br />
Busby Island<br />
Green Island<br />
Rocky<br />
Bays<br />
Gastropoda<br />
x<br />
Acmaea mitra x x x<br />
Acteocina harpa<br />
x<br />
Aglaja ocelligera<br />
Alia gauspata x x x x<br />
Alvania sp. x x x x x<br />
Archidoris montereyensis x x x x<br />
Barleeia sp.<br />
x<br />
Buccinum baeri x x x<br />
Cerithiidae<br />
x<br />
Cerithiopsis sp. x x<br />
Crepidula dorsata<br />
x<br />
Crepidula perforans<br />
x<br />
Crepidula sp.<br />
x<br />
Cryptobranchia concentrica<br />
x<br />
Cryptonatica affinis x x<br />
Dendronotus<br />
Northwest Bay<br />
Fouling<br />
Valdez<br />
Cordova<br />
Whittier<br />
Bligh Island<br />
Eagilik Bay<br />
Evans Island<br />
Knight Island<br />
Lake Bay<br />
Main bay<br />
Sheep Bay<br />
Shotgun Cove<br />
Squaw Bay<br />
Port Valdez<br />
Valdez Arm<br />
Windy Bay<br />
Dendronotus albopunctatus<br />
x<br />
Dendronotus frondosus x x<br />
Diplodonta impolita<br />
x<br />
Dorid nudibranch x x x<br />
Doridella steinbergi x x x<br />
Eubranchus olivaceous x x<br />
Fusitriton oregonensis<br />
x<br />
Granulina margaritula<br />
x<br />
Haminoea virescens x x x<br />
Hermissenda crassicornis x x x x x<br />
Lacuna sp. x x x x x x x<br />
Lacuna marmorata x x x x x<br />
Lacuna vincta x x x x x x x<br />
Lepeta caeca<br />
x
Chapt 9C7. Shelled Molluscs, page 9C7- 3<br />
Table 9C7.1.<br />
(Continued) Shelled<br />
Molluscs 1998 Survey<br />
Harbors<br />
Valdez<br />
Cordova<br />
Whittier<br />
Mud bays<br />
Valdez<br />
Bligh Island<br />
Valdez Arm<br />
Growler Island<br />
Cordova<br />
Headlands &<br />
Reefs<br />
Valdez Arm<br />
Busby Island<br />
Green Island<br />
Rocky<br />
Bays<br />
Northwest Bay<br />
Valdez<br />
Cordova<br />
Whittier<br />
Bligh Island<br />
Eagilik Bay<br />
Evans Island<br />
Knight Island<br />
Lake Bay<br />
Main bay<br />
Sheep Bay<br />
Shotgun Cove<br />
Squaw Bay<br />
Port Valdez<br />
Valdez Arm<br />
Lepetidae<br />
x<br />
Lirularia lirulata<br />
x<br />
Littorina scutulata x x x x x x x<br />
Littorina sitkana x x x x x x x x x x x<br />
Lottia pelta x x x x x x x x x x x x x<br />
Lottia sp. x x x x x x x x x x<br />
Margarites berhigensis x x<br />
Margarites helicinus x x<br />
Margarites pupilliis x x x x x x x x<br />
Margarites sp.<br />
Nassarius mendicus x x<br />
Neptunea lyrata<br />
x<br />
Nucella lamellosa x x x x x<br />
Nucella lima x x x<br />
Ocenebra interfossa<br />
x<br />
Odostomia spp.<br />
x<br />
Olivella baetica x x x<br />
Onepota spp. x x<br />
Scabrotrophon maltzani<br />
x<br />
Searlesia dira x x x<br />
Tectura fenestrata x x<br />
Tectura persona x x x x x x x x<br />
Tectura scutum x x x x x x x<br />
Trichotropis cancellata x x x<br />
Trichotropis insignis x x<br />
unidentified Turridae<br />
x<br />
Velutina rubra<br />
x<br />
Bivalvia<br />
Axinopsida sp.<br />
x<br />
Bankia setacea x x<br />
Chlamys rubida x x x x<br />
Clinocardium nuttalli x x x x x x<br />
Crassostrea gigas<br />
x<br />
Fouling<br />
Windy Bay
Chapt 9C7. Shelled Molluscs, page 9C7- 4<br />
Table 9C7.1.<br />
(Continued) Shelled<br />
Molluscs 1998 Survey<br />
Harbors<br />
Valdez<br />
Cordova<br />
Whittier<br />
Mud bays<br />
Valdez<br />
Bligh Island<br />
Valdez Arm<br />
Growler<br />
Island<br />
Cordova<br />
Headlands &<br />
Reefs<br />
Valdez Arm<br />
Busby Island<br />
Green Island<br />
Rocky<br />
Bays<br />
Northwest<br />
Bay<br />
Fouling<br />
Valdez<br />
Cordova<br />
Whittier<br />
Bligh Island<br />
Eagilik Bay<br />
Evans Island<br />
Knight Island<br />
Lake Bay<br />
Main bay<br />
Sheep Bay<br />
Shotgun Cove<br />
Squaw Bay<br />
Port Valdez<br />
Valdez Arm<br />
Gari californica<br />
x<br />
Hiatella arctica x x x x x x x x x x x x x x x x x x x<br />
Macoma balthica x x x x x x x x<br />
Macoma inquinata x x x x x x x<br />
Macoma nasuta x x<br />
Macoma sp. x x<br />
Modiolus modiolus x x<br />
Montacutidae x x<br />
Musculus vernicosus x x x x x<br />
Mya arenaria x x x x<br />
Mya pseudoarenaria<br />
x<br />
Mya sp. x x x x<br />
Mya truncata x x x x<br />
Mysella tumida x x x<br />
Mytilus trossulus x x x x x x x x x x x x x x x x x x x x x<br />
Pododesmus macroschisma x x x x x x x<br />
Protothaca staminea x x x x x x x<br />
Pseudopythinia compressa x x<br />
Saxidomus giganteus x x x x x x x x x x<br />
Serripes groenlandicus x x x x<br />
Serripes laperousii<br />
x<br />
Tresus capax x x<br />
Turtonia minuta x x<br />
Yoldia hyperborea<br />
x<br />
Polyplacophora<br />
Cryptochiton stelleri<br />
x<br />
Katherina tunicata<br />
x<br />
Lepidozona interstinctus<br />
x<br />
Mopalia ciliata x x x<br />
Mopalia lignosa x x x x x<br />
Mopalia sp. x x<br />
Schizoplax brandtii x x<br />
Tonicella insignis x x<br />
Tonicella lineata x x x<br />
Windy Bay
Chapt 9C7. Shelled Molluscs, page 9C7- 5<br />
Table 9C7.2. Shelled<br />
Molluscs 1999 Survey<br />
Homer<br />
Seward<br />
Lowell Pt<br />
Whittier<br />
Shot-gun<br />
Cove<br />
Fairmont<br />
Bay<br />
Valdez<br />
Cloudman<br />
Bay<br />
Busby Is.<br />
Cordov<br />
a<br />
Windy<br />
Bay<br />
Constantine<br />
Harbor<br />
Tatitlek<br />
a = abundant<br />
r = rare<br />
c = common<br />
p = present<br />
floats<br />
benthic<br />
homer<br />
spit<br />
benthic<br />
floats<br />
float<br />
fouling<br />
benthic<br />
grab<br />
benthic<br />
benthic<br />
floats<br />
floats<br />
benthic<br />
Gastropoda<br />
Alia gauspata<br />
r<br />
Archidoris montereyensis<br />
r<br />
Boreotrophon<br />
r<br />
Buccinum baeri<br />
c<br />
Cerithiidae<br />
r<br />
Collisella digitalis<br />
c<br />
Collisellla strigatellla c c<br />
Crepipatella dorsata<br />
r<br />
Cryptobranchia alba r r r r<br />
Cryptonatica <strong>of</strong>finis r r<br />
dorid nudibranch c c<br />
Granulina margaritula<br />
Haminoea vesicula c r r c<br />
Hermissenda crassicornis<br />
c<br />
Lacuna vincta c c c c c c<br />
Littorina sitkana a c c a c r c<br />
Llttoria scutulata c c<br />
Lottia pelta a c c c c<br />
Lottia strigatella<br />
r<br />
Margarites pupillis r r<br />
Melibe leonina<br />
c<br />
Nassarius mendicus<br />
r<br />
Neptunea lyrata<br />
p<br />
Nucella lamellosa c r<br />
Nucella lima c c r<br />
Ocinebrina interfossa<br />
r<br />
Odostomia sp. r r r<br />
Oenopota sp<br />
r<br />
Olivella baetica r r<br />
Onchidoris bilammelata<br />
c<br />
Polycera zosterae<br />
c<br />
Tectura fenestrata<br />
c<br />
Tectura persona c c<br />
Tectura scutum c c c<br />
Trichotropis cancellata c r<br />
Velutina plicatilis<br />
r<br />
Polyplacophora<br />
Mopalia cf M. imporcata r<br />
Mopallia cf. M.spectabilis<br />
r<br />
Moplalia ciliata r r<br />
Schizoplax brandtii<br />
r<br />
Tonicella lineata r r<br />
Tonicella rubra<br />
r<br />
Bivalvia<br />
Bankia setacea<br />
p<br />
Chlamys rubida r r<br />
Clinocardium californiense r<br />
Clinocardium nuttalli c c c c<br />
Clinocardium sp. c p<br />
Crassicardia crassidens<br />
r<br />
Cryptomya californica<br />
r<br />
r
Chapt 9C7. Shelled Molluscs, page 9C7- 6<br />
Table 9C7.2.<br />
(Continued) Shelled<br />
Molluscs 1999 Survey<br />
a = abundant<br />
r = rare<br />
c = common<br />
p = present<br />
Homer Valdez Cordov<br />
a<br />
floats<br />
benthic<br />
homer<br />
spit<br />
Seward<br />
Lowell Pt<br />
benthic<br />
Whittier<br />
floats<br />
Shot-gun<br />
Cove<br />
float<br />
Fairmont<br />
Bay<br />
fouling<br />
benthic<br />
grab<br />
Cloudman<br />
Bay<br />
benthic<br />
Busby Is.<br />
benthic<br />
floats<br />
Windy<br />
Bay<br />
Constantine<br />
Harbor<br />
floats<br />
benthic<br />
Tatitlek<br />
Cyclocardia sp. r r<br />
Hiatella acrtica c c c c c p c c c<br />
Humilaria kennerleyi<br />
r<br />
Macoma balthica a c a c a c<br />
Macoma calcarea<br />
p<br />
Macoma inquinata c c a c c c<br />
Macoma lama<br />
r<br />
Macoma obliqua r r<br />
Macoma sp. p p<br />
Modiolus modiolus r r<br />
Mya arenaria c a c<br />
Mya pseudoarenaria c c<br />
Mya sp.<br />
p<br />
Mya truncata c c r<br />
Mytilus trossulus a a a a c a a a a a a<br />
Neaeromya compressa<br />
r<br />
Pododesmus macroschisma<br />
r<br />
Protothaca staminea c c p c<br />
Saxidomus giganteus c c p c<br />
Serripes groenlandicus p r<br />
Serripes laperousii<br />
r<br />
Siliqua patula p p<br />
Spisula polynyma c c c<br />
Tresus capax c c<br />
Vilasina vernicosa c c c<br />
Yoldia myalis<br />
p
Chapt 9C7. Shelled Molluscs, page 9C7- 7<br />
Table 9C7.3. All Mollusks from Both 1998 and 1999<br />
Species Distribution NIS status<br />
Acmaea mitra<br />
NE Pac<br />
Acteocina harpa<br />
NE Pac<br />
Aglaja ocelligera NE Pac new record<br />
Astyris gauspata<br />
NE Pac<br />
Alvania sp.<br />
Archidoris montereyensis NE Pac<br />
Axinopsida sp.<br />
Bankia setacea<br />
NE Pac<br />
Barleeia sp. <br />
Boreotrophon<br />
Buccinum baeri<br />
NE Pac<br />
Cerithiidae<br />
Cerithiopsis<br />
Chlamys rubida<br />
NE Pac<br />
Clinocardium californiense NE Pac<br />
Clinocardium nuttallii<br />
NEW Pac<br />
Clinocardium sp.<br />
Lottia digitalis<br />
NE Pac<br />
Lottia strigatellla<br />
NE Pac<br />
Cyclocardia crassidens NEW Pac<br />
Crassostrea gigas<br />
introduced<br />
Crepidula sp.<br />
Crepipatella dorsata<br />
NE Pac<br />
Cryptobranchia alba<br />
NE Pac<br />
Cryptobranchia concentrica NEW Pac<br />
Cryptomya californica<br />
NEW Pac<br />
Cryptonatica affinis<br />
arctic circumboreal<br />
Cyclocardia sp.<br />
Dendronotus frondosus northern hemisphere<br />
Dendronotus sp.<br />
Diplodonta impolita<br />
NE Pac<br />
dorid nudibranch<br />
Acanthodoris<br />
Doridella steinbergi<br />
NE Pac<br />
Eubranchus olivaceous NE Pac new record<br />
Fusitriton oregonensis<br />
NE Pac<br />
Gari californica<br />
NE Pac<br />
Granulina margaritula<br />
NE Pac<br />
Haminoea vesicula<br />
NE Pac<br />
Haminoea virescens<br />
NE Pac<br />
Hermissenda crassicornis NE Pac<br />
Hiatella arctica<br />
northern hemisphere<br />
Humilaria kennerleyi<br />
NE Pac<br />
Lacuna marmorata<br />
NE Pac<br />
Lacuna sp.<br />
Lacuna vincta<br />
amphiboreal<br />
Lepidozona interstinica<br />
NE Pac<br />
Lepetidae<br />
Lirularia lirulata<br />
NE Pac
Chapt 9C7. Shelled Molluscs, page 9C7- 8<br />
Table 9C7.3. (Continued) All Mollusks from Both 1998 and 1999<br />
Species Distribution NIS status<br />
Littorina scutulata<br />
NE Pac<br />
Littorina sitkana<br />
NE Pac<br />
Lottia pelta<br />
NE Pac<br />
Lottia sp<br />
Lottia strigatella<br />
NE Pac<br />
Macoma balthica circumboreal poss. cryptogenic<br />
Macoma calcarea<br />
ampiboreal<br />
Macoma inquinata<br />
NE Pac<br />
Macoma lama<br />
NEW pac<br />
Macoma nasuta<br />
NE Pac<br />
Macoma obliqua<br />
NE Pac<br />
Macoma sp.<br />
Margarites beringensis<br />
AK<br />
Margarites helicinus<br />
circumboreal<br />
Margarites pupillus<br />
NE Pac<br />
Margarites sp.<br />
Melebe leonina<br />
NE Pac<br />
Modiolus modiolus<br />
circumboreal<br />
Montacutidae<br />
Mopalia cf M. imporcata NE Pac<br />
Mopalia ciliata<br />
NE Pac<br />
Mopalia lignosa<br />
NE Pac<br />
Moplalia cf. M.spectabilis NE Pac<br />
Mya arenaria amphiboreal introduced<br />
Mya pseudoarenaria<br />
NE Pac<br />
Mya sp.<br />
Mya truncata<br />
amphiboreal<br />
Mysella tumida<br />
NE Pac<br />
Mytilus trossulus<br />
NE Pac<br />
Nassarius mendicus<br />
NE Pac<br />
Neaeromya compressa NE Pac<br />
Neptunea lyrata<br />
NE Pac<br />
Nucella lamellosa<br />
NE Pac<br />
Nucella lima<br />
NE Pac<br />
Ocinebrina interfossa<br />
NE Pac<br />
Odostomia spp.<br />
Oenopota sp.<br />
Olivella baetica<br />
NE Pac<br />
Onchidoris bilamellata<br />
amphiboreal<br />
Onchidoris sp.<br />
Pododesmus macroschisma NE Pac<br />
Polycera zosterae<br />
NE Pac<br />
Protothaca staminea<br />
NE Pac<br />
Saxidomus giganteus<br />
NE Pac<br />
Scabrotrophon maltzani NE Pac<br />
Schizoplax brandtii<br />
NEW Pac<br />
Searlesia dira<br />
NE Pac<br />
Serripes groenlandicus circumboreal<br />
Serripes laperousii<br />
circumboreal<br />
Siliqua patula<br />
NE Pac
Chapt 9C7. Shelled Molluscs, page 9C7- 9<br />
Table 9C7.3. (Continued) All Mollusks from Both 1998 and 1999<br />
Species Distribution NIS status<br />
Spisula polynyma<br />
amphiboreal<br />
Tectura fenestrata<br />
NE Pac<br />
Tectura persona<br />
NE Pac<br />
Tectura scutum<br />
NE Pac<br />
Tonicella insignis<br />
AK<br />
Tonicella lineata<br />
NE Pac<br />
Tonicella rubra<br />
circumboreal<br />
Tresus capax<br />
NE Pac<br />
Trichotropis cancellata<br />
NE Pac<br />
Trichotropis insignis<br />
NEW Pac<br />
Turtonia minuta<br />
amphiboreal<br />
Velutina plicatilis<br />
arctic circumboreal<br />
Velutina rubra<br />
AK<br />
Vilasina vernicosa<br />
NEW Pac<br />
Yoldia hyperborea<br />
circumboreal<br />
Yoldia myalis<br />
amphiboreal
Chapt 9C7. Shelled Molluscs, page 9C7- 10<br />
occurrence <strong>of</strong> Mya arenaria in Alaskan waters before the 1890s. No M. arenaria were found in<br />
samples from shell middens from Sitka (1070-470 yr BP) (Foster, unpublished data) and S.<br />
Hawkins Island (500-200 yr BP and 2150-1380 yr BP) (Yarborough, unpublished data), and<br />
faunal lists from two other Prince William Sound sites (ca. 400-200 yr BP) (Yarborough,<br />
unpublished data), and from Resurrection Bay and Aialik Bay (1700s –1800s AD) (Yarborough,<br />
unpublished data). These negative findings are consistent with M. arenaria being an invasive<br />
species that was introduced in the late 1880s. In addition, the Asian oyster Crassostrea gigas<br />
was also widely distributed in protected bays <strong>of</strong> Prince William Sound and Katchemak Bay,<br />
where it is cultured abundantly in nets suspended vertically in the water column. However, it is<br />
sustained as an aquaculture species by importation <strong>of</strong> laboratory produced spat, and is not a selfsustaining<br />
population, probably because water temperatures are too cold for gonad development<br />
and spawning. The common clam Macoma balthica may be viewed as a cryptogenic species that<br />
may be introduced from the north Atlantic; but because <strong>of</strong> its circumboreal distribution and<br />
variation in shell morphology, it is difficult to reconstruct the history <strong>of</strong> this species’<br />
biogeography.<br />
References:<br />
Baxter, R. 1987. Mollusks <strong>of</strong> Alaska. Shells and Sea Life. Bayside, California, 163pp.<br />
Behrens, D.W. 1991. Pacific coast nudibranchs. Second edition. Monterey California. 207 pp.<br />
Coan, E.V. and P.V. Scott. 1999. Personal Communication. (unpublished draft <strong>of</strong> guide to<br />
Pacific coast bivalves.)<br />
Feder, H.M. and G.E.M. Matheke. 1980. Distribution, abundance, community structure and<br />
tropic structure <strong>of</strong> the infauna <strong>of</strong> the northeast Gulf <strong>of</strong> Alaska. Inst. Mar. Sci. Rept. R78-8, Univ.<br />
<strong>of</strong> Alaska, Fairbanks. 209 pp.<br />
Fisher, W.K. 1930. Asteroidea <strong>of</strong> the North Pacific and adjacent waters. Part 3. Forcipulata<br />
(concluded). Smithsonian Institution. U. S. Nat. Mus. Bull 76. 245 pp.<br />
Foster, N.R. 1981. A synopsis <strong>of</strong> the marine prosobranch and bivalve mollusks in Alaskan<br />
waters. Univ. Alaska, Institute <strong>of</strong> Marine Science Technical Rept IMS 81-3, 479 p.<br />
Foster, N. R. 1991. Intertidal Bivalves: A Guide to the Common Marine Bivalves <strong>of</strong> Alaska.<br />
University <strong>of</strong> Alaska Press, Fairbanks, Alaska. 152 pp.<br />
Harbo, R. M. 1997. Shells and Shellfish <strong>of</strong> the Pacific Northwest A field guide. Harbour<br />
Publishing, Madiera Park, BC, Canada. 270 pp.<br />
Kozl<strong>of</strong>f, E. N. 1996. Marine Invertebrates <strong>of</strong> the Pacific Northwest. University <strong>of</strong> Washington<br />
Press. 539 pp.<br />
Strasser, M. 1999. Mya arenaria- an ancient invader <strong>of</strong> the North Sea Coast. Helgolander<br />
Meeresuntersuchungen 52: 309-324.
Chapt 9C7. Shelled Molluscs, page 9C7- 11<br />
Turgeon , D.A. (ed.) 1998. Common and Scientific Names <strong>of</strong> <strong>Aquatic</strong> Invertebrates from the<br />
United States and Canada: Mollusks, Second Edition. American Fisheries Society Special<br />
Publication 26. 526 pp.
Chapt 9C8. Opisthobranch Molluscs, page 9C8- 1<br />
Chapter 9C8. Focal Taxonomic Collections: Opisthobranch Molluscs<br />
Jeffrey H. R. Goddard, Marine Science Institute, University <strong>of</strong> California, Santa Barbara<br />
Introduction<br />
The Smithsonian Environmental Research Center, in conjunction with the University <strong>of</strong><br />
Alaska, U.S. Fish and Wildlife Service and Prince William Sound Advisory Council, conducted<br />
in August 1999 a survey <strong>of</strong> Prince William Sound for non-indigenous marine species. Owing to<br />
the abundance <strong>of</strong> opisthobranch molluscs observed in a similar survey the previous summer, I<br />
was invited to help identify members <strong>of</strong> this taxon in the 1999 survey. The following report<br />
summarizes the 1999 collection <strong>of</strong> opisthobranch molluscs and attempts to identify nonindigenous<br />
and cyptogenic members <strong>of</strong> the fauna.<br />
Methods<br />
Specimens were collected by hand from floating docks, intertidal mudflats and rocky<br />
shores, jetties, and from settling panels deployed earlier in the year. Observations were made <strong>of</strong><br />
living animals, either with a dissecting microscope in the laboratory, or with a hand lens while<br />
traveling to the next collecting station. Voucher specimens were fixed in either 5 to 10 %<br />
formalin or 70 % ethanol. Specimens not identified during the expedition were examined,<br />
dissected and identified in the laboratory at UCSB.<br />
Results<br />
Twenty-eight species were found, consisting primarily <strong>of</strong> dorid nudibranchs. These are<br />
listed below with notes on their classification, habitats, and prey; an asterisk marks range<br />
extensions. Distributions and abundance in our field collections are summarized in Table 9C8.1<br />
following the list.<br />
Cephalaspidea<br />
Aglaja ocelligera (Bergh, 1894)<br />
Numerous specimens were found on the intertidal mudflats west <strong>of</strong> the Cordova marina.<br />
Haminoea sp. (either vesicula or virescens)<br />
Egg masses <strong>of</strong> this species were abundant on Zostera in the Cordova marina, where Nora Foster<br />
collected a single specimen.<br />
Melanochlamys diomedea (Bergh, 1894)<br />
Adults and egg masses <strong>of</strong> this species were abundant on mudflats just west <strong>of</strong> the Cordova<br />
marina.<br />
Sacoglossa<br />
*Alderia modesta (Loven, 1844)<br />
Adults and their egg masses were abundant on Vaucheria sp. on the high intertidal mudflats<br />
immediately west <strong>of</strong> the Cordova marina. Range extension from Vancouver Island, British<br />
Columbia (Millen, 1980).
Chapt 9C8. Opisthobranch Molluscs, page 9C8- 2<br />
*Olea hansineensis Agersborg, 1923<br />
One specimen <strong>of</strong> this opisthobranch egg eating sacoglossan was found on the egg masses <strong>of</strong><br />
Melanochlamys diomedea on the mudflat west <strong>of</strong> the Cordova marina. Range extension from<br />
Sechelt Inlet, British Columbia (Millen, 1980).<br />
Nudibranchia, Doridacea<br />
Acanthodoris nanaimoensis O’Donoghue, 1921<br />
A specimen about 35 mm long was found in the low intertidal on the jetty at the Cordova marina.<br />
This species was recently collected from Little Green Island, Prince William Sound (Nora<br />
Foster, personal communication, 1999).<br />
Acanthodoris pilosa (Abildgaard, 1789)<br />
Three specimens, 2.2 to 5 mm long, were found on drift pieces <strong>of</strong> a large, unidentified,<br />
fleshy, orange-brown colored ctenostome bryozoan on the mudflats just west <strong>of</strong> the Cordova<br />
marina. Note: unidentified species <strong>of</strong> cheilostome bryozoans were growing on the above<br />
ctenostome and may have been the prey <strong>of</strong> these specimens <strong>of</strong> A. pilosa. Two additional<br />
specimens, 5.5 and 7 mm long, were found on settling panels in the Cordova marina.<br />
*Adalaria jannae Millen, 1987<br />
This species was abundant, along with its ribbon shaped egg masses, on Membranipora sp.<br />
growing on Laminaria on the docks at Whittier and on the moored buoy at Shotgun Cove. A.<br />
jannae closely resembles Onchidoris muricata, but lacks the medial radular teeth found in the<br />
latter; A. jannae also has 4 to 6 small lateral teeth on each half row <strong>of</strong> the radula, as well as a<br />
ribbon shaped egg masses (Millen, 1987). The radular formula from an 8 mm long (preserved)<br />
specimen from Shotgun Cove was 30 (4.1.0.1.4.). Range extension from Sointula, British<br />
Columbia (Millen, 1987).<br />
Adalaria proxima (Alder & Hancock, 1854)<br />
Two specimens, 3 mm long (alive) and 10 mm long (preserved) were found on the unidentified<br />
ctenostome bryozoan mentioned above for Acanthodoris pilosa and on the jetty, respectively, at<br />
Cordova (the above note for the prey <strong>of</strong> Acanthodoris pilosa also applies to this species). The<br />
radular formula for these two specimens was 30 (7.1.1.1.7) and 36 (9.1.1.1.9), respectively, and<br />
the changes in shape <strong>of</strong> the teeth with increasing body size match that described for this species<br />
by Thompson & Brown (1984). Note: previous Alaskan records <strong>of</strong> this species can be found<br />
under the names Adalaria albopapillosa, A. pacifica, and A. virescens (see Lee & Foster,<br />
1985:444), all three <strong>of</strong> which Millen (1987) considered junior synonyms <strong>of</strong> A. proxima.<br />
*Adalaria sp. 1 <strong>of</strong> Behrens (1991) and Millen (1987:2701)<br />
One specimen, 3.3 mm long (preserved) <strong>of</strong> this distinctive species was found on the low<br />
intertidal rocky shore at Tatitlek. Range extension from Ketchikan, Alaska (Millen, 1989).<br />
Adalaria sp.<br />
Two specimens, 2 and 3 mm long, were found on the same fleshy ctenostome bryozoan that<br />
Acanthodoris pilosa was found at Cordova (and the same note on prey also applies here). An
Chapt 9C8. Opisthobranch Molluscs, page 9C8- 3<br />
additional specimen, 3 mm long, was found on a settlement panel at Valdez. The radular<br />
formula (24 x 4.1.0.1.4 in a 3 mm specimen), tooth shape, and bipinnate gills inserted in separate<br />
pits place this onchidoridid in the genus Adalaria, but the presence <strong>of</strong> long slender dorsal<br />
papillae lacking spicules does not appear to match any described species.<br />
*Ancula pacifica MacFarland, 1905<br />
A single specimen, lacking orange lines on the body, was found on the floating docks in the<br />
Cordova marina. This species (or the color form <strong>of</strong> A. pacifica lacking orange lines on the body)<br />
may be a junior synonym <strong>of</strong> Ancula gibbosa (Risso, 1818). Range extension from Grant Island,<br />
Ketchikan, Alaska (Millen, 1989).<br />
Archidoris montereyensis (Cooper, 1863)<br />
Two specimens were found eating the sponge Halichondria panicea growing on oysters in<br />
Fairmont Bay, and two specimens were found in the marina at Cordova.<br />
Doridella steinbergae (Lance, 1962)<br />
This species was found on its prey, Membranipora sp., on Laminaria on the floating docks at<br />
Whittier, and on drift Laminaria on the mudflats at Cordova. The range <strong>of</strong> this species was<br />
extended by Foster (1987) northward to Prince William Sound and more recently westward to<br />
Mink Island, Katmai National Park (Nora Foster, personal communication, 1999).<br />
*Geitodoris heathi (MacFarland, 1905)<br />
Four specimens were found on the low intertidal rocky shore at Tatitlek. Range extension from<br />
Ketchikan, Alaska (Millen, 1989).<br />
Onchidoris bilamellata (Linnaeus, 1767)<br />
This circumboreal species feeds on Balanus spp. and was abundant, along with its egg masses, in<br />
the Homer marina, on the settling panels at Fairmont Bay and Tatitlek, and on rocks in and<br />
around the Cordova marina.<br />
Onchidoris muricata (Müller, 1776)<br />
Atotal <strong>of</strong> five small specimens were found, two on settling panels at Valdez, and three on settling<br />
panels at Cordova. Note: previous Alaskan records <strong>of</strong> this species can be found under the names<br />
O. hystricina and O. varians (see Lee & Foster, 1985:444), which Millen (1985) synonomized<br />
with O. muricata.<br />
*Palio zosterae (O’Donoghue, 1924)<br />
Adults and egg masses were abundant on the bryozoan Membranipora sp. growing on Laminaria<br />
on the floating docks at Whittier; a few specimens were also found on the buoy at Shotgun Cove<br />
and on the docks at Cordova. P. zosterae may be a junior synonym <strong>of</strong> P. pallida Bergh, 1880; if<br />
not these specimens represent a small westward range extension from Hawkins Island, Prince<br />
William Sound (Nora Foster, personal communication, 1999).<br />
Triopha catalinae (Cooper, 1863)<br />
Three specimens were found under cobbles in the low intertidal at Tatitlek.
Chapt 9C8. Opisthobranch Molluscs, page 9C8- 4<br />
Nudibranchia, Dendronotacea<br />
Dendronotus frondosus (Ascanius, 1774)<br />
Specimens were found on Obelia-like hydroids on settling panels at Seward, Fairmont Bay,<br />
Potato Point in Valdez narrows, and Tatitlek. Adults and egg masses were also common on the<br />
docks at Cordova.<br />
Melibe leonina (Gould, 1852)<br />
Specimens were found on Zostera in Fairmont Bay, on a settling panel at Tatitlek, and on the<br />
docks at Cordova.<br />
Nudibranchia, Arminacea<br />
Dirona albolineata MacFarland in Cockerell & Eliot, 1905<br />
Four small specimens were found on the floating docks in the Cordova marina, an additional<br />
juvenile specimen was found on a settling panel at Tatitlek.<br />
*Janolus fuscus (O’Donoghue, 1924)<br />
Two specimens, 60 to 70 mm long, were found with their egg masses on Bugula sp. on docks in<br />
the Homer marina. Range extension from Klu Bay, Revillagigedo Island, Alaska (Robilliard &<br />
Barr, 1978).<br />
Nudibranchia, Aeolidacea<br />
Aeolidia papillosa (Linnaeus, 1761)<br />
One 70 mm long specimen was found on the docks at Homer.<br />
*Cuthona albocrusta (MacFarland, 1966)<br />
A single specimen <strong>of</strong> this distinctive species was found on the docks at Cordova. Range<br />
extension from White Rock, southern British Columbia (Millen, 1983).<br />
*Cuthona pustulata (Alder & Hancock, 1864)<br />
Four specimens, 4 to 5 mm long, were found feeding on the hydroid Sarsia sp. on a dock in the<br />
marina at Homer. These specimens resembled Gosliner & Millen’s (1984) description <strong>of</strong><br />
Cuthona pustulata from British Columbia, especially with regard to overall shape <strong>of</strong> the body,<br />
cerata, and head tentacles. Our specimens differed however by lacking large white spots on the<br />
cerata (they did have smaller opaque white flecks) and by having slightly fewer rows <strong>of</strong> cerata<br />
with fewer cerata per row. The radula and shape <strong>of</strong> the radular teeth <strong>of</strong> our specimens are<br />
virtually identical to that described by Gosliner & Millen (1984) but differed in having 4 to 5<br />
lateral denticles, compared to 5 to 9. Range extension from Galiano Island, British Columbia<br />
(Gosliner & Millen, 1984).<br />
*Eubranchus olivaceus (O’Donoghue, 1922)<br />
This distinctive aeolid was found with its egg masses on Obelia-like hydroids on the docks at<br />
Homer and at Whittier. Range extension from Prince William Sound (Nora Foster, personal<br />
communication, 1999).
Chapt 9C8. Opisthobranch Molluscs, page 9C8- 5<br />
Hermissenda crassicornis (Eschscholtz, 1831)<br />
Hermissenda was found at Whittier, Shotgun Cove, Fairmont Bay, Tatitlek, and in the Cordova<br />
marina. Specimens were small at all sites except Cordova, where they were up to 35 mm long.<br />
Discussion<br />
Most <strong>of</strong> the opisthobranchs collected during this survey are cold-temperate species endemic to<br />
the northern Pacific Ocean, especially the northeastern Pacific. The remainder are circumboreal<br />
species, a few <strong>of</strong> which (e.g., Dendronotus frondosus and Onchidoris bilamellata) penetrate the<br />
Arctic Ocean (see information on distributions in Marcus, 1961; McDonald, 1983; Thompson &<br />
Brown, 1984; Behrens, 1991). To my knowledge, none <strong>of</strong> the species collected in our survey are<br />
non-indigenous in the Prince William Sound region. While native/non-native status can not be<br />
assigned with certainty to the unidentified species <strong>of</strong> Adalaria or the tentatively identified<br />
specimens <strong>of</strong> Cuthona pustulata, the radiation <strong>of</strong> both <strong>of</strong> these genera in boreal waters suggests<br />
that both <strong>of</strong> these species are probably also indigenous to Prince William Sound.<br />
References<br />
Behrens, D.W. 1991. Pacific coast nudibranchs. Sea Challengers: Monterey, California. 107 pp.<br />
Foster, N.R. 1987. Range extension for Doridella steinbergae (Lance, 1962) to Prince William<br />
Sound, Alaska. The Veliger 30: 97-98.<br />
Gosliner, T.M. and S.V. Millen, 1984. Records <strong>of</strong> Cuthona pustulata (Alder & Hancock, 1854)<br />
from the Canadian Pacific. The Veliger 26: 183-187.<br />
Lee, R.S. and N. R. Foster. 1985. A distributional list with range extensions <strong>of</strong> the opisthobranch<br />
gastropods <strong>of</strong> Alaska. The Veliger 27: 440-448.<br />
Marcus, E. 1961. Opisthobranch mollusks from California. The Veliger 3(Supplement): 1-85.<br />
McDonald, G.R. 1983. A review <strong>of</strong> the nudibranchs <strong>of</strong> the California coast. Malacologia 24:<br />
114-276.<br />
Millen, S.V. 1980. Range extensions, new distribution sites, and notes on the biology <strong>of</strong><br />
sacoglossan opisthobranchs (Mollusca: Gastropoda) in British Columbia. Canadian Journal <strong>of</strong><br />
Zoology 58: 1207-1209.<br />
Millen, S.V. 1983. Range extensions <strong>of</strong> opisthobranchs in the northeastern Pacific. The Veliger<br />
25: 383-386.<br />
Millen, S.V. 1985. The nudibranch genera Onchidoris and Diaphorodoris (Mollusca,<br />
Opisthobranchia) in the northeastern Pacific. The Veliger 28: 80-93.<br />
Millen, S.V. 1987. The nudibranch genus Adalaria, with a description <strong>of</strong> a new species from the<br />
northeastern Pacific. Canadian Journal <strong>of</strong> Zoology 65: 2696-2702.
Chapt 9C8. Opisthobranch Molluscs, page 9C8- 6<br />
Millen, S.V. 1989. Opisthobranch range extensions in Alaska with the first records <strong>of</strong> Cuthona<br />
viridis (Forbes, 1840) from the Pacific. The Veliger 32: 64-68.<br />
Robilliard, G.A. and L. Barr. 1978. Range extensions <strong>of</strong> some nudibranch molluscs in Alaskan<br />
waters. Canadian Journal <strong>of</strong> Zoology 56: 152-153.<br />
Thompson, T.E. and G.H. Brown. 1984. Biology <strong>of</strong> opisthobranch molluscs, Vol. II. Ray<br />
Society: London. 229 pp.
Chapt 9C8. Opisthobranch Molluscs, page 9C8- 7<br />
Table 9C8. 1. Numbers <strong>of</strong> opisthobranch molluscs collected at 8 sites on the Kenai Peninsula and in Prince William Sound, Alaska,<br />
August 1999. A number followed by (S) means those specimens were found on fouling plates only.<br />
Table 9C8.1 Opisthobranch Homer Seward Whittier Shotgun Fairmont Tatitlek Cordova<br />
Molluscs 1999<br />
Species marina mudflat marina Lowell Pt. marina Cove Bay Valdez rocky shore mudflats marina<br />
Aeolidia papillosa 1<br />
Eubranchus olivaceus >10 1<br />
Cuthona pustulata 4<br />
Janolus fuscus 2<br />
Onchidoris bilamellata >10 >10(S) 3(S) >10* >10<br />
Melanochlamys diomedea 1 >100<br />
Dendronotus frondosus 7(S) >10 2(S) >1(S) >10<br />
Palio zosterae >100 >1 >1<br />
Adalaria jannae >100 >10<br />
Hermissenda crassicornis 5 >10 >10 >1 >10<br />
Doridella steinbergae 2 >10*<br />
Archidoris montereyensis 1 2<br />
Melibe leonina 1 >1(S) 1<br />
Onchidoris muricata 2(S) 3(S)<br />
Adalaria sp. 1(S) 2*<br />
Adalaria sp. 1 <strong>of</strong><br />
Behrens (1991) 1<br />
Geitodoris heathi 4<br />
Triopha catalinae 3<br />
Aglaja ocelligera >10<br />
Acanthodoris pilosa 3* 2(S)<br />
Adalaria proxima 1* 1<br />
Alderia modesta >10<br />
Olea hansineensis 1<br />
Haminoea sp. 1<br />
Acanthodoris nanaimoensis 1<br />
Dirona albolineta 1(S) 4<br />
Ancula pacifica 1<br />
Cuthona albocrusta 1<br />
Number <strong>of</strong> species per site 5 1 1 0 5 3 5 3 8 9 14<br />
* specimens found on cobbles, drift kelp, or drift bryozoans lying on the mudflat.
Chapt 9C9. Echinoderms, page 9C9- 1<br />
Chapter 9C9. Focal Taxonomic Collections: Echinoderms<br />
Nora Foster, University <strong>of</strong> Alaska Museum<br />
Anson H. Hines, Smithsonian Environmental Research Center<br />
Results<br />
Echinoderm collections over the two years <strong>of</strong> field sampling included 4 species <strong>of</strong><br />
holothuroid, 1 species <strong>of</strong> ophiuroid, 6 species <strong>of</strong> asteroid, and 1 species ophiurioid (Table<br />
9C9.1). No crinoid species were collected. All <strong>of</strong> these species are reported as native to Alaska.<br />
However, Asterias amurensis, a northwest Pacific seastar, was found at Homer spit <strong>of</strong> Kachemak<br />
Bay, <strong>of</strong>f Cook Inlet. This is a range extension form the Shumagin Islands (Fisher 1930) and<br />
from the Kodiak Shelf and Izhut Bay, Kodiak Island, and Bering and Chukchi Seas, where it was<br />
found by Jewett and Feder (1981). Specimens <strong>of</strong> A. amurenis are not represented in the<br />
University <strong>of</strong> Alaska Museum’s collections for trawl surveys in either the Gulf <strong>of</strong> Alaska (Foster,<br />
1999 pers. comm.) or Cook Inlet (Feder and Paul 1981, Feder et al. 1981). The survey <strong>of</strong> Cook<br />
Inlet also included scuba surveys. It would have been very unusual for such collections to miss<br />
a large conspicuous sea star (30 cm from ray tip to ray tip). Prior experienced naturalists<br />
working in Kachemak Bay reported the sudden recent appearance <strong>of</strong> this species. This species is<br />
an invasive species in Tasmania, where it was probablyintroduced by ballast water (it has a longlived<br />
planktotrophic larva). In Tasmania it is having adverse impacts on native bivalves and is<br />
considered to be a serious pest. We consider this species to be cryptogenic in Cook Inlet, with<br />
characteristics that are very suspicious <strong>of</strong> an NIS. Because it is a voracious predator, it could<br />
have a major impact on benthic communities.<br />
References<br />
Feder, H.M. and A.J. Paul. 1981. Distribution and abundance <strong>of</strong> some epibenthic invertebrates<br />
<strong>of</strong> Cook Inlet, Alaska. Univ. Alaska Fairbanks, Institute <strong>of</strong> Marine Science Tech. Rept. R80-3,<br />
154 p.<br />
Feder, H.M., A.J. Paul, M. Hoberg, and S. Jewett. 1981. Distribution, abundance, community<br />
structure and trophic relationships <strong>of</strong> the nearshore benthos <strong>of</strong> Cook Inlet. In: Environmental<br />
assessment <strong>of</strong> the Alaskan Continental Shelf. Final Reports, <strong>Biological</strong> Studies 14:45-676.<br />
Fisher, W.K. 1930. Asteroidea <strong>of</strong> the North Pacific and Adjacent <strong>Water</strong>s. Part 3. Forcipulata<br />
(concluded). Smithsonian Institution. U. S. Nat. Mus. Bull 76. 245 pp.<br />
Jewett, S.C. and H. M Feder. 1981. Epifaunal invertebrates <strong>of</strong> the continental shelf <strong>of</strong> the Bering<br />
and Chukchi Seas. Pp. 1131-1153 in Hood, D.W. and J. Clader (ed.), The Eastern Bering Sea<br />
Shelf: Oceanography and Resources. Vol. II, U.S. Dept <strong>of</strong> Commerce. Distributed by the<br />
University <strong>of</strong> Washington Press, Seattle. 1339 p.
Chapt 9C9. Echinoderms, page 9C9- 2<br />
Table 9C9.1. Echinoderms 1998, 1999<br />
Homer Seward Whittier Growler Sawmill<br />
Island Bay<br />
Port<br />
Valdez<br />
Rockey<br />
Point<br />
Busby<br />
Island<br />
Tatitlek Cloudman<br />
Bay<br />
Cordova Constantine<br />
Harbor<br />
Port<br />
Chalmers<br />
Green<br />
Island<br />
Chenega Northwest<br />
Bay<br />
Main<br />
Bay<br />
Eupentacta X<br />
pseudoquesemita<br />
Cucumaria<br />
X<br />
frondosa japonica<br />
Psolus chitonoides X X<br />
Cucumaria vegae<br />
X<br />
Ophiopholis<br />
X X X<br />
aculeata<br />
Pycnopodia X X X X X X X X X X X X X<br />
helianthoides<br />
Evasterias<br />
X X X X X X<br />
troschelii<br />
Pisaster ochraceus X X X<br />
Asterias amurensis X<br />
Dermasterias<br />
X X X X<br />
imbricata<br />
Leptasterias sp. X X X<br />
Strongylocentrotus<br />
droebachiensis<br />
X X X
Chapt 9C10. Ascidians, page 9C10- 1<br />
Chapter 9C10. Focal Taxonomic Collections: Ascidians<br />
Gretchen Lambert, Friday Harbor Laboratories, University <strong>of</strong> Washington<br />
Results<br />
A total <strong>of</strong> 12 species <strong>of</strong> ascidians were collected during this expedition. The highest<br />
number <strong>of</strong> species and <strong>of</strong> individuals was at Homer, Tatitlek and Cordova, the three locations<br />
with the highest salinity. No ascidians were recorded from Seward or Whittier, or the floats at<br />
Valdez, where the salinity was only 10-12 parts per thousand, although fouling panels at Valdez,<br />
submerged at about 7m, did have some ascidians. Ascidians require at least 25-27 parts per<br />
thousand and prefer a salinity <strong>of</strong> 30 or higher. Three <strong>of</strong> the 12 species are colonial, the other 9<br />
are solitary forms. Although there were not a large number <strong>of</strong> species collected, they do<br />
represent a good diversity. Both orders and all three suborders <strong>of</strong> the class Ascidiacea are<br />
represented and include a total <strong>of</strong> 6 families. Seven <strong>of</strong> the 12 species are known to brood their<br />
embryos and release swimming tadpoles; all 7 <strong>of</strong> these did harbor mature embryos. At the end <strong>of</strong><br />
this report is a list <strong>of</strong> the ascidian voucher specimens collected in 1998; there were no additional<br />
species to the 1999 collections.<br />
The collection includes an undescribed species <strong>of</strong> Distaplia, a colonial form in the<br />
suborder Aplousobranchia. It is surprising that this species is very abundant at both Homer and<br />
Cordova marinas, yet has gone unrecognized for nearly a century. It is possible that if Ritter<br />
(1901) or Huntsman (1912) collected it, they may have considered it merely a variant <strong>of</strong><br />
Distaplia occidentalis. On the other hand, this species may have been much rarer during Ritter’s<br />
and Huntsman’s time. Its preferred habitat is apparently sheltered surfaces, shallow but never<br />
exposed at low tide and away from very much light. It is thus “pre-adapted” for the huge surface<br />
area and specialized environment provided by marina floats and the many submerged ropes up to<br />
3m. in length that are <strong>of</strong>ten suspended from floats. Before the building <strong>of</strong> large marinas, this<br />
particular type <strong>of</strong> habitat was not abundant. The marina environment is also somewhat unstable,<br />
subject to changing seasonal conditions. Many ascidians are well adapted to take advantage <strong>of</strong><br />
this instability. Distaplia spp., like most aplousobranchs, are very fast growing, reach sexual<br />
maturity in a few weeks, reproduce and then die back to a dedifferentiated basal portion than can<br />
survive until environmental conditions are again suitable for rapid growth. This new species is<br />
somewhat similar to a cold-water form from the Kamchatka Peninsula <strong>of</strong> Russia, and an analysis<br />
and description <strong>of</strong> it is being prepared in collaboration with Dr. Karen Sanamyan <strong>of</strong> the<br />
Kamchatka Institute <strong>of</strong> Science.<br />
Two species <strong>of</strong> the genus Ascidia were collected and present a taxonomic problem. One<br />
corresponds to Ascidia adhaerens Ritter, 1901, the other to Ascidiopsis columbiana Huntsman,<br />
1912. (The genus Ascidiopsis is no longer valid; it was synonymized under the genus Ascidia.)<br />
Van Name (1945) synonymized these two NE Pacific species under Ascidia callosa, probably<br />
incorrectly; A. callosa was described in 1852 by Stimpson from U.S. east coast specimens from<br />
Massachusetts. The recent genetic analysis <strong>of</strong> many Atlantic and Pacific species <strong>of</strong> various taxa<br />
that closely resemble each morphologically and were originally lumped into the same species has<br />
resulted in many cases in their separation into different species (R. Strathmann pers. comm.).<br />
However, a recent re-examination <strong>of</strong> Stimpson’s syntypes <strong>of</strong> Ascidia callosa and Ritter’s<br />
paratypes <strong>of</strong> Ascidia adhaerens, borrowed from the Smithsonian National Museum <strong>of</strong> Natural
Chapt 9C10. Ascidians, page 9C10- 2<br />
History, has confirmed that these are the same species. Thus, Ascidia adhaerens remains a<br />
synonym under A. callosa. The type specimen <strong>of</strong> Huntsman’s Ascidiopsis columbiana was<br />
borrowed from the Royal Ontario Museum in Toronto. I was able to confirm that this species is<br />
indeed distinct from Ascidia callosa and thus should be resurrected as a valid species, which will<br />
be done in a publication now in preparation.<br />
Botrylloides violaceus is not native to Alaska, or to any part <strong>of</strong> the NE Pacific. It is a<br />
Japanese species that appeared on the U.S. Pacific coast about 20 years ago (J. Carlton pers.<br />
comm.). Since that time it has spread and become extremely abundant from southern California<br />
to British Columbia (Lambert and Lambert, 1998). This is the first report <strong>of</strong> its presence in any<br />
part <strong>of</strong> Alaska, however. Although large colonies were not observed, the fouling panels at<br />
Tatitlek contained numerous newly settled zooids most <strong>of</strong> which appeared healthy. Thus<br />
somewhere close to the panels there had to be mature colonies which were supplying the shortlived<br />
tadpoles that settled on the panels. B. violaceus incubates its embryos; the tadpoles that are<br />
released are huge, complex, and usually swim for just a very brief period, perhaps only a few<br />
minutes, before settling. The unusual tadpole morphology allows for easy recognition <strong>of</strong> this<br />
invasive species (Saito et al., 1981).<br />
Molgula retortiformis and Ascidia callosa are apparently the only species collected on<br />
this expedition that occur in both the North Pacific and North Atlantic (Van Name, 1945).<br />
Chelyosoma productum, Corella inflata and C. willmeriana, Distaplia occidentalis, Pyura<br />
haustor and Styela truncata have been recorded only from the NE Pacific. Halocynthia<br />
hilgendorfi is known from both the NW and NE Pacific.<br />
Sixty-five species <strong>of</strong> ascidians are known to occur in Alaskan waters. Most <strong>of</strong> these were<br />
obtained by dredging many decades ago and may be restricted to the Bering Sea or the Arctic<br />
Ocean. The ascidian fauna <strong>of</strong> Alaska is still mostly unexplored, and probably there exist a<br />
number <strong>of</strong> undescribed species.<br />
Publication <strong>of</strong> the present work, including a description <strong>of</strong> the new Distaplia species and<br />
redescription <strong>of</strong> the two Ascidia species, is in preparation with Dr. Karen Sanamyan <strong>of</strong> the<br />
Kamchatka Institute <strong>of</strong> Ecology.<br />
References<br />
Huntsman, A.G. 1912. Ascidians from the coasts <strong>of</strong> Canada. Trans. Canad. Inst. 9: 111-148.<br />
Kott, P. 1985. The Australian Ascidiacea. Mem. Qd. Mus. 23: 1-440.<br />
Lambert, C.C. & G. Lambert 1998. Non-indigenous ascidians in southern California harbors and<br />
marinas. Mar. Biol. 130: 675-688.<br />
Lambert, G., C.C. Lambert & D.P. Abbott 1981. Corella species in the American Pacific<br />
Northwest: distinction <strong>of</strong> C. inflata Huntsman, 1912 from C. willmeriana Herdman, 1898<br />
(Ascidiacea, Phlebobranchia). Can. J. Zool. 59: 1493-1504.<br />
Nishikawa, T. 1991. The ascidians <strong>of</strong> the Japan Sea. II. Publ. Seto Mar. Biol. Lab. 35: 25-170.
Chapt 9C10. Ascidians, page 9C10- 3<br />
O’Clair, R.M. & C.E. O’Clair 1998. Southeast Alaska’s Rocky Shores. . Plant Press, Auke Bay,<br />
AK, 564 pp.<br />
Ritter, W.E. 1901. Papers from the Harriman Alaska Expedition. XXIII. The ascidians. Proc.<br />
Wash. Acad. Sci. 3: 225-266.<br />
Ritter, W.E. 1913. The simple ascidians from the northeastern Pacific in the collection <strong>of</strong> the<br />
United States National Museum. Proc. U.S. Natl. Mus. 45: 427-505.<br />
Saito, Y., H. Mukai & H. Watanabe 1981. Studies on Japanese compound styelid ascidians II. A<br />
new species <strong>of</strong> the genus Botrylloides and redescription <strong>of</strong> B. violaceus Oka. Publ. Seto Mar.<br />
Biol. Lab. 26: 357-368.<br />
Sanamyan, K. 1993. Ascidians from the north-western Pacific region. 2. Molgulidae. Ophelia<br />
38: 127-135.<br />
Sanamyan, K. 1996. Ascidians from the north-western Pacific region. 3. Pyuridae. Ophelia 45:<br />
199-210.<br />
Van Name, W.G. 1945. The North and South American Ascidians. Bull. Amer. Mus. Nat. Hist.<br />
84: 1-476.
Chapt 9C10. Ascidians, page 9C10- 4<br />
Table 9C10.1 Sites visited:<br />
1. Aug. 8 Homer Marina, Kachemak Bay, Cook Inlet<br />
Salinity 27 % o 1 ft. depth, 30 % o at 2m depth; Temp. 10 o C<br />
Most <strong>of</strong> the following records are from suspended ropes.<br />
Ascidia callosa--common esp. about halfway between the inner and outer ends <strong>of</strong> the marina.<br />
Some contain brooded larvae in atrial chamber. Two small specimens from fouling panel:<br />
one from E1 and one from OH--2-P2.<br />
Corella inflata--reported by John Chapman but not personally seen in spite <strong>of</strong> extensive<br />
sampling <strong>of</strong> floats and ropes. Samples lost; record not verified.<br />
Distaplia new sp. very abundant at most locations in the marina, esp. on ropes 1-2 m. deep. A<br />
few small colonies on fouling panel.<br />
Molgula retortiformis: 2 large and 8 small specimens on ropes. Two small specimens on fouling<br />
plate E1. The large animals contain brooded embryos in the atrial chamber.<br />
Styela truncata - 1 small individual.<br />
2. Aug. 9: Seward Marina.<br />
Salinity 11 % o about a foot down.<br />
Temperature 11.5 o C.<br />
No ascidians either at the marina or Lowell Pt. rocky intertidal (11 % o , 11.5 o C.)<br />
3. Aug. 10, 1999 Whittier marina.<br />
No sal. or temp. readings taken. Sal. very low; no ascidians on floats.<br />
4. Aug. 10, 1999 Fairmont Bay oyster farm, Prince Wm. Sound.<br />
Salinity 25 % o about a foot down.<br />
Temperature 14 o C.<br />
Ascidia columbiana--numerous small specimens on concrete blocks <strong>of</strong> the fouling panels and<br />
one large one on bottom <strong>of</strong> submerged 4 ft long oyster bag.<br />
5. Aug. 11, 1999 Valdez Marina<br />
No salinity or temp. readings taken<br />
No ascidians observed on marina floats or suspended ropes.<br />
6. Aug. 11-12, 1999 Tatitlek<br />
Salinity 27 % o about a foot down.<br />
Temperature 17 o C. surface<br />
a) Fouling panels and frames, 3m depth:<br />
Ascidia columbiana -- a few small specimens on the concrete blocks for the fouling panels and<br />
one on panel PL3-l.<br />
Botrylloides violaceus -- numerous very small zooids on panels. Largest one with 2 zooids in<br />
colony. Youngest appeared to be only hours post metamorphosis.<br />
Corella inflata -- common on both the panels and frames about 3m depth, with brooded larvae in<br />
the atrial chamber.
Chapt 9C10. Ascidians, page 9C10- 5<br />
Table 9C10.1 (Continued) Sites visited:<br />
b) Intertidal rocks, low tide, morning Aug. 12<br />
Ascidia columbiana -- one large specimen coll. by J. Goddard.<br />
Chelyosoma productum -- about 12 seen, 2 collected.<br />
Halocynthia igaboja -- one very small immature specimen coll. by J. Goddard.<br />
Pyura haustor -- 4 seen, one incomplete specimen collected. Difficult to collect; wedged tightly<br />
into rock crevices.<br />
c) oyster cages from across bay: one small Corella inflata.<br />
7. Aug. 13, 1999 Cordova Marina. Tide just beginning to come in.<br />
Salinity 27 % o about a foot down.<br />
Temperature 13 o C. surface<br />
Ascidia callosa-- numerous on floats and especially on suspended ropes. Some contain brooded<br />
larvae in atrial chamber.<br />
Corella inflata -- large, common, full <strong>of</strong> brooded larvae in atrial chamber.<br />
Distaplia new sp. --abundant colonies, with many mature larvae.<br />
Distaplia occidentalis -- common; several color morphs, large heads, with mature larvae.<br />
Styela truncata -- numerous. Some contain brooded larvae in atrial chamber.<br />
b) Fouling panels<br />
Ascidia callosa -- 12 small specimens from I dock, Pl. 3, 3 in a second vial.<br />
Corella inflata -- one small specimen from I dock, Pl. 3; two in a second vial.<br />
Corella willmeriana-- (specimens identified on site, not saved apparently.)<br />
Distaplia new sp. -- numerous small colonies from I dock Pl. 3, one in a second vial.<br />
Styela truncata -- 8 on fouling panels in voucher collection in one vial, 3 in another.<br />
8. Aug. 14, 1999 Valdez fouling panels, Berth 5<br />
Corella willmeriana -- 3 individuals, 1 large and 2 small. Specimens identified on site; not saved<br />
apparently.<br />
Ascidia columbiana -- one small specimen from fouling panel (or perhaps the concrete block<br />
anchoring the panels) at 20 ft. (sampled 8/14).
Chapt 9C10. Ascidians, page 9C10- 6<br />
Table 9C10.2 Ascidian vouchers from fouling panels 1998, identified Sept. 1999<br />
Data taken from jar labels.<br />
Homer 3/9/98 -- Styela truncata (1)<br />
Homer: 9/3/98 -- Ascidia callosa (1)<br />
Homer Oct. 1998 -- Distaplia new sp. collected by G. Sonnevil. Colonies without gonads or<br />
larvae.<br />
Homer 5/11/99 -- Distaplia new sp. collected by G. Sonnevil. Colonies small, immature.<br />
Chenega 7/9/98 -- Corella inflata (2) Port San Juan, Chenega sm. boat hbr<br />
Chenega 9/7/98 -- Ascidia callosa (1)<br />
Chenega no date -- Corella inflata (7) panel MI - 05 - (can’t read label)<br />
Valdez 9/8/98 -- Corella inflata (2) at Alyeska berths<br />
No location given -- Ascidia sp. (1) too small to ID to species. Panel WH-04-P3.
Chapt 9C10. Ascidians, page 9C10- 7<br />
Table 9C10.3 Systematics<br />
Class Ascidiacea<br />
OrderAplousobranchia<br />
Family Holozoidae<br />
Distaplia occidentalis Bancr<strong>of</strong>t, 1899<br />
Distaplia new species<br />
Order Phlebobranchia<br />
Family Corellidae<br />
Chelyosoma productum Stimpson, 1864<br />
Corella inflata Huntsman, 1912<br />
Corella willmeriana Herdman, 1898<br />
Family Ascidiidae<br />
Ascidia callosa Stimpson, 1852<br />
Ascidia columbiana (Huntsman, 1912)<br />
Order Stolidobranchia<br />
Family Styelidae<br />
Botrylloides violaceus Oka, 1927<br />
Styela truncata Ritter, 1901<br />
Family Pyuridae<br />
Halocynthia igaboja Oka, 1906<br />
Pyura haustor (Stimpson, 1864)<br />
Family Molgulidae<br />
Molgula retortiformis Verrill, 1871<br />
Table 9C10. 4 Ascidian References by Depth and Habitat<br />
Depth(m)<br />
Selected References<br />
intertidal, floats;to 10m Ritter ’01, ’13; VName 45; PWS Expedition 1999<br />
us. 0-60m<br />
Ritter’01,Huntsman’12,VName45,Abbott66,O’Clair98;PWS Exped.’99<br />
0-few m. Saito et al 81,Lambert & Lambert 98, PWS Exped. 1999<br />
intertidal - 50m Huntsman ’12,VName 45,O’Clair98; PWS Exped. ’99<br />
0-50m Huntsman ’12,VName 45; PWS Exped. ’99<br />
0-few m. Huntsman ’12,VName 45; PWS Exped. ’99<br />
float PWS Expedition 1999<br />
floats; intertidal-few m. VName 45, O’Clair 98, PWS Expedition 1999<br />
to 57m VName 45, Nishikawa 91,Sanamyan 96,O’Clair98, PWS Exped. ’99<br />
4-75m; floats Ritter 01,VName 45,Abbott66,Sanamyan 93, PWS Exped. ’99<br />
intertidal - 114m VName 45, Sanamyan 96,O’Clair98; PWS Exped. ’99<br />
ropes on floats;0-20m Ritter ’01, Van Name 45, PWS Exped. ’99
Chapt 9C10. Ascidians, page 9C10- 8<br />
Table 9C10.5 Ascidan Species Distribution in Alaska<br />
Genus Species Author/date Family Distribution in Alaska<br />
Ascidia callosa Stimpson, 1852 Ascidiidae Homer;PWS:Fairmont Bay<br />
oyster<br />
farm,Tatitlek,Cordova,Valdez,<br />
Chenega<br />
Ascidia columbiana (Huntsman,<br />
1912)<br />
Ascidiidae Arctic coast to Alask. Penin. &<br />
SE Alaska<br />
Botrylloides violaceus Oka, 1927 Styelidae PWS: Tatitlek<br />
Chelyosom<br />
a<br />
productum Stimpson, 1864 Corellidae PWS: Tatitlek; Sitka<br />
Sound:Passage Is.<br />
Corella willmeriana Herdman, 1898 Corellidae PWS: Cordova, Valdez<br />
Corella inflata Huntsman, 1912 Corellidae PWS: Tatitlek, Cordova,<br />
Valdez, Chenega<br />
Distaplia new sp. Holozoidae Cook Inlet: Homer Marina;<br />
PWS: Cordova Marina<br />
Distaplia occidentalis Bancr<strong>of</strong>t, 1899 Holozoidae PWS: Cordova Marina;<br />
Chichag<strong>of</strong> Is.<br />
Halocynthia igaboja Oka, 1906 Pyuridae Gulf <strong>of</strong> Alaska: Kodiak<br />
Is.;Chichag<strong>of</strong> Is.; PWS:<br />
Tatitlek; Prince Rupert BC<br />
Molgula retortiformis Verrill, 1871 Molgulidae SE Bering Sea to Sitka; Canoe<br />
Bay; SE Chukchi Sea; Homer<br />
Marina<br />
Pyura haustor (Stimpson, 1864) Pyuridae Alaska Gulf:Sanak Is.,<br />
Shumagin Is.; Sitka<br />
Sound:Alice Is.; PWS: Tatitlek<br />
Styela truncata Ritter, 1901 Styelidae Yakutat Bay; Cook Inlet:<br />
Homer Marina; PWS: Cordova<br />
Marina
Chapt 9D. Fouling Community Surveys, page 9D- 1<br />
9D. Fouling Community Surveys<br />
Anson H. Hines, Smithsonian Environmental Research Center<br />
Gregory M. Ruiz, Smithsonian Environmental Research Center<br />
9D1. Purpose<br />
Fouling communities in many bays, harbors and estuaries are frequently invaded by NIS<br />
associated with shipping traffic (Cohen & Carlton 1995, Hewitt 1993, Coles et al. 1999).<br />
Fouling communities have major impact on ships and floating structures, ranging from buoys<br />
and floats to aquaculture pens and nets, as well as surfaces <strong>of</strong> oysters and mussels themselves.<br />
Consequently, fouling communities are well-studied in many parts <strong>of</strong> the world, but they have<br />
received little study in Alaskan waters. We conducted surveys <strong>of</strong> fouling communities in Prince<br />
William Sound, Seward and Homer using experimental fouling plates. The use <strong>of</strong> fouling plates<br />
provides a replicated standardized assay for NIS in a community that is prone to invasions, but<br />
which has received little prior ecological analysis in Alaska. We also surveyed fouling<br />
communities on floats, pilings, and buoys to compare these substrates with our experimental<br />
plates.<br />
9D2. Methods<br />
We conducted two surveys that focused on fouling communities that settled on natural<br />
surfaces and experimental plates, which we deployed in earlier spring months at an array <strong>of</strong> sites<br />
in Homer, Seward and Prince William Sound. We sampled these fouling plates in 7-17<br />
September 1998 and in 8-16 August 1999.<br />
The team for the fouling community surveys consisted <strong>of</strong>:<br />
• Greg Ruiz (SERC), molluscs, parasites, fouling communities;<br />
• Anson Hines (SERC), barnacles and decapod crustaceans;<br />
• James Carlton (Mystic Seaport), marine/estuarine invertebrates and global NIS<br />
• Melissa Frey (SERC); technical and field assistance<br />
• George Smith (SERC); technical and field assistance<br />
• Lea Ann Henry (University <strong>of</strong> Ontario), hydroid identifications;<br />
• Judy Winston (Virginia Museum <strong>of</strong> Natural History), bryozoan identifications.<br />
We deployed arrays <strong>of</strong> fouling plates at 12 locations, including 2 on the Kenai Peninsula<br />
(Homer, Seward) and 10 in Prince William Sound (Table 9D.1). At each site we deployed 5<br />
arrays suspended at depths <strong>of</strong> either 1 meter (3 arrays) or 3 meters (2 arrays) below mean low<br />
water level. Each array consisted <strong>of</strong> settling plates attached to a frame made <strong>of</strong> 2 crossed pieces<br />
<strong>of</strong> pvc pipe (20 mm diameter, 50 cm long), which was suspended in a horizontal position by a<br />
line attached to a dock or float and weighted by a concrete block. Settling plates made <strong>of</strong> 14 cm<br />
x 14 cm x 7 mm thick pvc (3 plates) or plywood (1 plate) were attached to each <strong>of</strong> the 4 ends <strong>of</strong><br />
the cross in a horizontal position. The horizontal orientation assured that sediment would not<br />
accumulate on the underside, providing a clean surface addition to several irregular surfaces <strong>of</strong><br />
the frame and top surfaces for settlement. To maximize the chance <strong>of</strong> detecting possible NIS<br />
within each site, the 5 arrays were dispersed as widely as possible to sample the range <strong>of</strong><br />
microhabitats present. We deployed the plates during April-May <strong>of</strong> each year and retrieved them<br />
in August-September, providing a “soak time” <strong>of</strong> about 4 months.
Chapt 9D. Fouling Community Surveys, page 9D- 2<br />
Table 9D.1. Sampling Locations for Fouling Community Surveys.<br />
Site 1998 1999<br />
Kenai Sites<br />
Homer X X<br />
Seward X X<br />
Prince William Sound<br />
Port Valdez<br />
Marine Terminal X X<br />
Valdez Area X X<br />
Growler Island<br />
X<br />
Whittier<br />
X<br />
Chenega<br />
X<br />
Port Chalmers<br />
X<br />
Fairmont Bay<br />
X<br />
Tatitlek<br />
Oyster Culture<br />
X<br />
Docks<br />
X<br />
Cordova<br />
X<br />
At the time <strong>of</strong> retrieving the arrays, the fouling plates were removed from their frame and<br />
placed individually into ziplock plastic bags for transport to a laboratory. Each plate was<br />
examined under a dissecting microscope and sessile and motile species were scored as present or<br />
absent. Voucher specimens were preserved and sent to taxonomic experts for authoritative<br />
identification. At the present stage <strong>of</strong> this project, we have used the fouling plates to develop an<br />
inventory <strong>of</strong> species. However, using this technique, the occurrence and abundance <strong>of</strong> species<br />
can be quantified and compared statistically for frequency <strong>of</strong> plates with species present. This<br />
type <strong>of</strong> comparison will proceed in new work during 2000 and 2001 as we sample other locations<br />
in Alaska and the major ports <strong>of</strong> western North America along the lower 48 states.<br />
9D3. Results<br />
During the soak time <strong>of</strong> ca. 4 months, the fouling plates accumulated dense community<br />
assemblages with large biomasses <strong>of</strong> mussels, barnacles, ascidians, hydroids, and bryozoans<br />
(Table 9.D.2). The fouling communities were rich in species, particularly for bryozoans,<br />
hydroids, nudibranchs, and ascidians Combining the 2 years <strong>of</strong> study across the12 sampling<br />
locations, we recorded more than 107 taxa/species on the fouling plates (Table 9D.2), including:<br />
8 families <strong>of</strong> polychaete worms, 13 species <strong>of</strong> acidians, 34 species <strong>of</strong> bryozoans, 1 species <strong>of</strong> sea<br />
anemone, 3 species <strong>of</strong> barnacle and 13 taxa <strong>of</strong> motile crustacea, 2 species <strong>of</strong> echinoderms, 12<br />
species <strong>of</strong> hydroids, 29+ taxa <strong>of</strong> molluscs, and 1 species <strong>of</strong> protozoan. Diversity <strong>of</strong> hydroids was<br />
particularly high at Homer (11 species). Byozoans were most diverse at Chenega (10 species),<br />
Homer (13 species), Port Valdez (14 species), and Tatilek (13 species). Acidians were most<br />
diverse at Homer (6 species) and Cordova (6 species).
Chapt 9D. Fouling Community Surveys, page 9D- 3<br />
Table 9D.2. Taxa Recovered on Fouling Plates. Asterisk denotes NIS. x denotes presence in 1998 and or 1999.<br />
Site Key: AL=Alyeska terminal; CH=Chenega Bay; GR=Growler Isl.; HO=Homer, MI=Montague Isl.;<br />
SE=Seward;VA=Port Valdez; WH=Whittier; CORD=Cordova; FRMNT=Fairmount Bay; PTOPT=Potato Pt.<br />
TAT=Tatitlek<br />
Site<br />
AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT<br />
Protozoan Foliculina x x x x x x x x x<br />
AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT<br />
Cnidaria Metridium senile x<br />
Metridium sp. x x x<br />
AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT<br />
Hydroids Calycella syringa x<br />
Clytia hemispherca<br />
x<br />
Clytia kincaidi<br />
x<br />
Companulina rugosa<br />
x<br />
* Garveia franciscana x<br />
Gonoththyraea clarki x x x x<br />
Obelia longissima x x x x x x x x<br />
Obelia sp. x x x x x x x x<br />
Opercularella lacerata<br />
x<br />
Sarsia eximia<br />
x<br />
Sarsia tubulosa<br />
x<br />
Sertularia robusta<br />
x<br />
AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT<br />
Annelida Capitellidae x x x x<br />
Cirratulidae x x<br />
Nereidae x x x x x x x x x<br />
Polynoidae x x x x x x x x x<br />
Sabellidae x x x<br />
Serpulidae x x x x x x x x x x<br />
Spirorbidae x x x x x x x x x x x<br />
Syllidae x x x x x x<br />
Terebellidae<br />
x<br />
AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT<br />
Mollusca Acanthodoris sp. x<br />
Acmaeidae<br />
x<br />
Aelolididae<br />
x<br />
Alvania sp. x x x x x<br />
Chlamys sp. x x x<br />
Clinocardium sp.<br />
x<br />
Dendronotus frondosus x x<br />
Dendronotus sp x x<br />
Fusitron oregonensis<br />
x<br />
Haminoea sp. x x x<br />
Hermissenda sp. x x x x x x x x x x<br />
Hiatella arcitca x x x x x x x x x x x<br />
Hinnites sp.<br />
x<br />
Lacuna sp. x x x x<br />
Lacuna vincta<br />
x<br />
Limacina helicina<br />
x
Chapt 9D. Fouling Community Surveys, page 9D- 4<br />
Table 9D.2. Continued<br />
AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT<br />
Mollusca Lottia sp. x<br />
cont. Margarites helinicus x<br />
Margarites pupillis x x<br />
Mytilus trossulus x x<br />
Mytilus sp. x x x x x x x x x x x x<br />
Odostomia sp. x x x x<br />
Onchidoris sp. x x x x<br />
Onchidoris murricata x x<br />
Onchidoris sp. x x x x x x<br />
Pododesma macroshisma<br />
x<br />
Pododesmus sp. x x<br />
Sacoglossan sp. x x<br />
Tritonia sp.<br />
x<br />
Vilisina vernicosa<br />
x<br />
AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT<br />
Crustacea Ampithoe sp. x<br />
Balanus cariosus x x x x x x x<br />
Balanus crenatus x x x x x x x x x x x x<br />
Semibalanus carriosus x x x<br />
Cancer magister x x<br />
Caprellidae x x x x x x<br />
Corophium sp. x x x x<br />
Eogammarus sp.<br />
x<br />
Gnorimosphaeroma oregonense x x x x x<br />
Harpacticus uniramus<br />
x<br />
Jassa sp. x x<br />
Munna sp.<br />
x<br />
Oregonia gracilis<br />
x<br />
Scyra acutifems<br />
x<br />
Spirontocaris sp.<br />
x<br />
AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT<br />
Bryozoa Alcyonidium sp. x x x x x x<br />
Alcyonidium hirsutum x x<br />
Bugula californica x x<br />
Bugula pacifica x x x<br />
Bugula sp. x x x x x x x x<br />
Calloporella craticula x x x<br />
Calloporella "lineata"<br />
x<br />
Cauloramphus pseudospinifer<br />
x<br />
Celleporella hyalina x x x x x x<br />
Cribrilina sp. x x<br />
Cribrilina corbicula x x x x x x<br />
Cribrilina sp. x x x x x x x<br />
Crisia serrulata<br />
x<br />
Cryptosula okadai<br />
x<br />
Electra sp. x x x<br />
Fenestruloides eopacifica x x<br />
Filicrisia franciscana x x x<br />
Harmeria scutulata<br />
x<br />
Hippothoa sp. x x x<br />
Lichenopora sp.<br />
x
Chapt 9D. Fouling Community Surveys, page 9D- 5<br />
Table 9D.2. continued<br />
AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT<br />
Bryozoa Membranipora menbrenacea x<br />
cont. Membranipora sp. x x<br />
Microporella germana<br />
x<br />
Parasmittina trispinosa<br />
x<br />
Phidolopora pacifica x x<br />
Porella major x x<br />
Rhamphostomella gigantea<br />
x<br />
Rhamphostomella hincksi<br />
x<br />
* Schizoporella "unicornis" x<br />
Tegella armifera<br />
x<br />
Tegella aquilirostris<br />
x<br />
Tubulipora tuba x x x<br />
Tubulipora sp.<br />
x<br />
AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT<br />
Echino- Psolus sp. x<br />
dermata Strongylocentrotus sp. x x<br />
AL CH GR HO MI SE VA WH CORD FRMNT PTOPT TAT<br />
Ascidia Ascidia sp. x<br />
Ascidia callosa x x x<br />
Ascidia columbiana x x x<br />
* Botrylloides violaceus x<br />
Corella sp. x x x<br />
Corella inflata x x x x<br />
Corella willmariana x x<br />
Distaplia sp. (new) x x<br />
Distaplia alaskensis x x<br />
Molgula retortiformis<br />
x<br />
Molgula sp. x x<br />
Ritterella sp. x x x<br />
Styela truncata<br />
x<br />
Among these species, there were 3 NIS: the ascidean Botrylloides violaceus, the bryozoan<br />
Schizoporella unicornis, and the hydroid Garvieia franciscana. These are the first records for<br />
two <strong>of</strong> these species (B. violaceus and G. franciscana) in Alaska. Botrylloides violaceus<br />
(=Botryllus aurantius is a colonial tunicate that is native to the northwest Pacific (Japan), and<br />
may have been first found on the West Coast in 1973, in San Francisco Bay (Cohen and Carlton<br />
1995). It is now widespread, from southern California to British Columbia (Cohen et al. 1998;<br />
Lambert and Lambert 1998). B. violaceus was abundant on fouling plates in Prince William<br />
Sound in 1999 (G. Lambert 1999 pers. comm.). Schizoporella unicornis is a bryozoan found in<br />
the northwest Pacific but was first collected in the Eastern Pacific in 1927, in Puget Sound<br />
(Carlton 1979; Cohen and Carlton 1995). Its first Alaska collection was made between 1944 and<br />
1949, in Kodiak (U. S. Navy 1951; Carlton, pers. comm.). Schizoporella unicornis may have<br />
been introduced in ship fouling or with plantings <strong>of</strong> Pacific Oysters (Cohen and Carlton 1995).<br />
In 1999, it was found in Tatilek. This form, while definitely introduced to the Pacific coast, may<br />
actually be a complex <strong>of</strong> several species (Winston 1999 pers. comm.). Garveia franciscana<br />
(Rope Grass Hydroid) has been found in many estuaries around the world, but its origin is<br />
uncertain. The Indo-Pacific and the Black--Caspian Sea basin have been suggested as possible<br />
native regions (Cohen and Carlton 1995; Calder 1997 pers. comm.) It was first described from<br />
San Francisco Bay in 1902, which was its only known location on the west coast <strong>of</strong> North<br />
America (Cohen and Carlton 1995), until we found it near Homer in 1999 (Lea-Anne Henry
Chapt 9D. Fouling Community Surveys, page 9D- 6<br />
pers. comm. 1999). In other regions <strong>of</strong> the world, this hydroid has been an economically<br />
important fouling organism, adversely affecting ships, power plants and fishing gear (Simkina<br />
1963; Andrews 1973; McLean 1972). In addition, fouling plates at Homer and Cordova had<br />
large biomasses <strong>of</strong> the new/undescribed ascidian, Distaplia sp. nov, which has many suspicious<br />
characteristics <strong>of</strong> NIS. Diastaplia sp. nov. is a new, undescribed species <strong>of</strong> tunicate, which is<br />
very abundant in fouling communities on floats and man-made substrates in marinas at Homer<br />
and Cordova. It was first collected in 1998 in Homer and was found in both Homer and Cordova<br />
in 1999. It was not found at other sites within Prince William Sound where other, native species<br />
<strong>of</strong> tunicate were common in fouling communities but lack similar shipping/boating traffic (e.g.,<br />
Tatitlek, Chenega, Port Chalmers). Its appearance is also suspicious, because it was not found in<br />
1901 when tunicates were collected in the region at nearby sites. This tunicate could be a<br />
formerly rare native species that has taken advantage <strong>of</strong> the newly created marina habitat, or a<br />
recent introduction (G. Lambert 1999 pers. comm.).<br />
Fouling communities on the plates suspended at 1m depth were greatly diminished at<br />
Seward, Whittier and Port Valdez during the summer, when snow and glacial melt produced<br />
markedly low salinities in the surface waters and also thick sediment deposits that covered the<br />
plates. Fouling plate arrays at 3 m depth in these locations experienced higher salinities, but still<br />
suffered from heavy sediment deposits. It was evident, however, that these plates had developed<br />
rich fouling communities prior to the summer season <strong>of</strong> greatest freshwater run<strong>of</strong>f and siltation.<br />
Species composition <strong>of</strong> fouling communities on the experimental plates were generally<br />
quite similar to the composition on surrounding surfaces (floats, pilings, lines, oysters, etc) at<br />
comparable tidal levels. Since the arrays were suspended below mean low water, the main<br />
groups <strong>of</strong> specie s that the plates did not sample adequately were species on pilings in the<br />
intertidal zone, such as Balanus glandula, Semibalanus balanoides, and various species <strong>of</strong><br />
acmaeid limpets.
Chapt 9E. Museum, Reference and Voucher Specimens, page 9E- 1<br />
Chapter 9E. Re-Examination <strong>of</strong> Museum, Reference, and Voucher Specimens<br />
Nora Foster, University <strong>of</strong> Alaska Museum, Fairbanks<br />
9E1. Purpose<br />
As a result <strong>of</strong> environmental assessment work in Port Valdez during the 1970’s for<br />
construction <strong>of</strong> the Valdez Marine Terminal, and the Exxon Valdez oil spill (EVOS), a large<br />
ecological database exists for Prince William Sound. Through a careful scrutiny <strong>of</strong> both the<br />
species lists and specimens archived in museum, reference and voucher collections for the past<br />
work, this part <strong>of</strong> our study was intended to test the idea that this extensive prior work in Prince<br />
William Sound should have detected NIS. However, in spite <strong>of</strong> the amount <strong>of</strong> sampling done in<br />
Prince William Sound, some species lists were developed without input from taxonomic experts,<br />
so that the biota is still poorly known. Few prior surveys have focused on biogeography or<br />
taxonomy, and until this study, none has specifically looked for NIS.<br />
9E2. Methods<br />
Literature sources for Prince William Sound invertebrates include:<br />
• Reports on marine fauna <strong>of</strong> the northeastern Gulf <strong>of</strong> Alaska, compiled as a result <strong>of</strong> the Outer<br />
Continental Shelf Environmental Assessment Project (OCSEAP);(Feder and Matheke, 1980;<br />
Feder and Jewett, 1988);<br />
• Thesis by M. Hoberg (1986);<br />
• Project reports on the Port Valdez environment (Cooney and Coyle, 1988; Feder et al., 1976;<br />
Feder and Bryson-Schwafel, 1988; Feder and Keiser, 1980; Jewett and Feder, 1977);<br />
• Eyerdam’s (1924) species lists;<br />
• Haven (1971); and<br />
• Several taxonomic references (Butler, 1980; Hart, 1982; Lambert, 1981; 1991; Behrens, 1991<br />
and others) for additional distribution records in Alaskan waters.<br />
A vast array <strong>of</strong> preserved biological samples from Prince William Sound and adjacent<br />
Gulf <strong>of</strong> Alaska are housed in the <strong>Aquatic</strong> Collection <strong>of</strong> the University <strong>of</strong> Alaska Museum<br />
(UAM). The major part <strong>of</strong> these are collections made as part <strong>of</strong> the OCSEAP projects in the mid-<br />
1970’s, environmental monitoring <strong>of</strong> Port Valdez, and Max Hoberg’s thesis material from three<br />
bays in the outer part <strong>of</strong> the Sound. Intertidal and benthic biota from the s<strong>of</strong>t bottom<br />
communities <strong>of</strong> Prince William Sound are well represented in the collection. Species lists for<br />
this material were generated using the museum accession records.<br />
EVOS Specimen Archives<br />
The specimens collected as part environmental surveys after the 1989 Exxon Valdez oil<br />
spill supplement the UAM collection. Subtidal invertebrates in 1,154 lots have been transferred<br />
to the UAM from warehouse storage for examination. These represent sampling conducted<br />
during 1990, 1991, 1993, and 1995, from 15 selected unoiled sites. The collections provide<br />
samples <strong>of</strong> shallow subtidal fauna from silled fjords, eelgrass beds, and habitats dominated by<br />
Laminaria. With limited resources, each specimen from this large number <strong>of</strong> samples cannot be<br />
examined individually, so it has been necessary to select subset <strong>of</strong> specimens for careful reexamination.
Chapt 9E. Museum, Reference and Voucher Specimens, page 9E- 2<br />
The subset <strong>of</strong> reference and voucher specimens examined as part <strong>of</strong> this study was<br />
selected by first compiling a list <strong>of</strong> 88 invertebrate taxa by comparing the lists <strong>of</strong> known taxa<br />
from Prince William Sound (discussed above) with lists <strong>of</strong> known NIS from west coast source<br />
ports: Puget Sound, San Francisco Bay, southern California, and other localities (see Ruiz &<br />
Hines, 1997). This compilation <strong>of</strong> 88 species includes:<br />
• species that may be confused with known NIS in Alaska;<br />
• NIS known from western North America (especially those extending to higher latitudes);<br />
• taxonomically difficult species and species complexes;<br />
• biogeographic outliers; and<br />
• range extensions.<br />
For example, polychaete specimens <strong>of</strong> most Boccardia and Polydora, were removed for reexamination<br />
because for their resemblance to Polydora ligni. Similarly, capitellid and many<br />
syllid polychaetes were selected to check for the presence <strong>of</strong> Barontolla americana, Capitella<br />
capitata, and Decamastus spp. The gastropod slipper shell Crepidula was partitioned from the<br />
collections to check for Crepidula fornicata and C. plana. Specimens <strong>of</strong> the amphipod<br />
Corophuim were selected to check for several NIS Corophium reported from Puget Sound. This<br />
original list was considered a working document and has been refined as the project has<br />
progressed (Table 9.E.1).<br />
Table 9.E.1. Museum and Reference Samples Selected for Re-Examination<br />
Family Genus Species Specimen Source<br />
Amphitoidae Amphitoe sp. Jewett samples<br />
Amphitoidae Amphitoe simulans Jewett samples<br />
Corophidae Corophium sp. Jewett samples<br />
Corophidae Corophium brevis Jewett samples<br />
Gammaridae Melita sp. UAM uncataloged<br />
Gammaridae Gammarus sp. Jewett samples<br />
Gammaridae Jassa sp. Jewett samples<br />
Gammaridae Melita dentata UAM uncataloged<br />
Cardiidae Clinocardium fucanum Jewett samples<br />
Glycymeridae Glycymeris septentrionalis Jewett samples<br />
Kelliidae Pseudopythina* compressa Jewett samples<br />
Kelliidae Pseudopythina* rugifera* Jewett samples<br />
Mytilidae Dacrydium vitreum* Jewett samples<br />
Mytilidae Dacrydium pacificum* UAM cataloged<br />
Ungulinidae Diplodonta orbella Port Valdez uncataloged<br />
Ungulinidae Diplodonta impolita UAM cataloged<br />
Adontorhina sp. Jewett samples<br />
Cryptosula okaidai Jewett samples<br />
Atyidae Haminoea virescens Little Green Island<br />
Atyidae Haminoea vesicula PWS 98 samples<br />
Atyidae Haminoea sp. Jewett samples<br />
Calyptraeidae Crepidula dorsata* Jewett samples<br />
Calyptraeidae Crepidula sp. Jewett samples<br />
Calyptraeidae Crepidula grandis Jewett samples<br />
Calyptraeidae Crepidula sp. Jewett samples<br />
Diaphanidae Diaphana minuta* Jewett samples
Chapt 9E. Museum, Reference and Voucher Specimens, page 9E- 3<br />
Table 9.E.1. Continued<br />
Diaphanidae Diaphana brunnea Jewett samples<br />
Diaphanidae Diaphana sp. UAM cataloged<br />
Fissurellidae Puncturella cooperi* Jewett samples<br />
Lacunidae Lacuna sp. UAM cataloged<br />
Lacunidae Lacuna vincta* Jewett samples<br />
Lacunidae Lacuna variegata* UAM cataloged<br />
Lacunidae Lacuna marmorata* UAM cataloged<br />
Lamellariidae Velutina sp. UAM cataloged<br />
Rissoidae Barleeia acuta* Jewett samples<br />
Scaphandridae Cylichna alba UAM cataloged<br />
Scaphandridae Cylichna occulta UAM cataloged<br />
Scaphandridae Cylichna sp. UAM cataloged<br />
Scaphandridae Cylichna attonsa Jewett samples<br />
Scaphandridae Cylichnella culcitella Jewett samples<br />
Scaphandridae Cylichnella harpa Jewett samples<br />
Scaphandridae Cylichnella sp. Jewett samples<br />
Turridae Kurtziella plumbea Jewett samples<br />
Turridae Taranis strongi Jewett samples<br />
Limnoriidae Limnoria lignorum Jewett samples<br />
Limnoriidae Limnoria Jewett samples<br />
Capitellidae Barantolla americana Jewett samples<br />
Capitellidae Barantolla sp. Jewett samples<br />
Capitellidae Capitella sp. Jewett samples<br />
Capitellidae Capitella capitata Jewett samples<br />
Capitellidae Decamastus sp. Jewett samples<br />
Capitellidae Decamastus sp. Jewett samples<br />
Capitellidae Heteromastus sp. Jewett samples<br />
Capitellidae Heteromastus filliformis Jewett samples<br />
Capitellidae Mediomastus sp. Jewett samples<br />
Capitellidae Notomastus sp. Jewett samples<br />
Nereidae Platynereis bicanaliculata Jewett samples<br />
Polynoidae Harmothoe imbricata Jewett samples<br />
Spionidae Malacoceros sp. Jewett samples<br />
Spionidae Nerine cirratulus Jewett samples<br />
Spionidae Polydora sp. Jewett samples<br />
Spionidae Polydora cf P.bracycephalata Jewett samples<br />
Spionidae Polydora socialis Jewett samples<br />
Spionidae Prionospio cirrifera Jewett samples<br />
Spionidae Prionospio sp. Jewett samples<br />
Spionidae Prionospio steenstrupi Jewett samples<br />
Spionidae Prionospoi malmgreni Jewett samples<br />
Spionidae Pygospio sp. Jewett samples<br />
Spionidae Pygospio elegans Jewett samples<br />
Spionidae Rhynchospio glutaeus Jewett samples<br />
Spionidae Scolelepis sp. Jewett samples<br />
Spionidae Scolelepis squamata Jewett samples<br />
Spionidae Spio cirrifera Jewett samples<br />
Spionidae Spio fillicornis Jewett samples<br />
Spionidae Spio sp. Jewett samples<br />
Spionidae Spiophanes berkeleyorum Jewett samples<br />
Spionidae Spiophanes bombyx Jewett samples<br />
Spionidae Spiophanes sp. Jewett samples
Chapt 9E. Museum, Reference and Voucher Specimens, page 9E- 4<br />
Table 9.E.1. Continued<br />
Spionidae Sternapsis scutata Jewett samples<br />
Syllidae Typosyllis armillaris UAM uncataloged<br />
Syllidae Typosyllis fasciata Jewett samples<br />
Syllidae Typosyllis harti UAM uncataoged<br />
Syllidae Typosyllis sp. UAM uncataoged<br />
Jewett samples<br />
<strong>Biological</strong> Technician Max Hoberg (UAF) used the suspect list to remove 1,154 specimens for<br />
screening: 154 crustaceans, 1081 polychaetes, 307 molluscs, 30 bryozoans. From these,<br />
specimens in the best condition were removed for further study. Thirty-five small crustacea<br />
were loaned to focal taxonomic expert J. Chapman (OSU), and similarly polychaete specimens<br />
were loaned to focal taxonomic expert J. Kudenov (UA Anchorage). Analyses <strong>of</strong> these<br />
specimens (presently on-going) will be reported in the 2000 final report for the University <strong>of</strong><br />
Alaska Sea Grant.<br />
9E3. Results<br />
Mollusc Taxa, including One Known NIS<br />
Most east Pacific records <strong>of</strong> the s<strong>of</strong>t shell clam, Mya arenaria, are the result <strong>of</strong> its<br />
accidental introduction along with Atlantic oysters, starting in San Francisco Bay in 1869, and<br />
Puget Sound in 1888 or 1889. We found that the clam was abundant in mud sediments in<br />
Cordova. We also found it in Port Valdez, Tatitlek, Constantine Harbor, Homer and Seward.<br />
Mya arenaria is also documented in Alaska from Nunivak Island, Norton Sound, and Kodiak<br />
Island (UAM collection records) as well as southeastern Alaska.<br />
Nora Foster examined mollusc specimens from both the EVOS specimens and cataloged<br />
and uncataloged specimens the UA Museum, selecting the following taxa for re-examination:<br />
• small venerids and Turtonia, were examined as possibly misidentified specimens <strong>of</strong> potential<br />
NIS Protamcorbicula amurensis, Venerupsis philipinarium, or Nuttalia obscurata;<br />
• small individuals <strong>of</strong> Musuculus spp. were reexamined as possibly misidentified specimens <strong>of</strong><br />
the potential NIS Musculista stenhousei;<br />
• Cerithiidae were re-examined were reexamined as possibly misidentified specimens <strong>of</strong> the<br />
potential NIS Battilaria;<br />
• Ocenibrina were reexamined as possibly misidentified specimens <strong>of</strong> the potential NIS<br />
Urosalpinx cineria and Ocenebra inornata, two introduced oyster drills; and<br />
• Crepidula specimens were reexamined as possibly misidentified specimens <strong>of</strong> the potential<br />
NIS Crepidula fornicata.<br />
None <strong>of</strong> these potential NIS taxa were found. However, to date it has been possible to examine<br />
carefully only a small subsample <strong>of</strong> the collections, and we are faced with the situation <strong>of</strong> there<br />
being too many samples and too little time.<br />
University <strong>of</strong> Alaska Sea Grant funding for this analysis <strong>of</strong> existing museum and<br />
reference collections continues until June 30, 2000. The goals for this remaining time frame are<br />
to:<br />
• Incorporate information from additional taxonomic experts and ecologists.
Chapt 9E. Museum, Reference and Voucher Specimens, page 9E- 5<br />
• Check database against additional grey literature sources (e.g., O’Clair & Zimerman,1987).<br />
• Add to the database a biogeographical descriptor, indicating whether the PWS record<br />
represents a range extension, and a reference source for the animal’s distribution.<br />
• Publish with J. Goddard an analysis <strong>of</strong> our 1999 collections <strong>of</strong> opisthobranch molluscs <strong>of</strong><br />
Prince William Sound (see also focal taxonomic subsection 9C8 Opisthobranch Molluscs,<br />
above).<br />
• Incorporate <strong>of</strong> additional specimens collected as part <strong>of</strong> this assessment <strong>of</strong> EVOS collections<br />
into the UAM <strong>Aquatic</strong> Collection.<br />
References<br />
Anonymous. 1999. Draft long-term monitoring plan to be presented to trustees. Exxon Valdez<br />
Oil Spill Trustees Council Restoration Update 6 (4): 7.<br />
Austin, W.C. 1985. An annotated checklist <strong>of</strong> marine invertebrates in the <strong>Cold</strong> Temperate<br />
Northeast Pacific. Khoyatan Marine Laboratory, Cowichan Bay, British Columbia. Vols. 1-3.<br />
682 pp.<br />
Banse, K. and K.D. Hobson. 1968. Benthic errantiate polychaetes <strong>of</strong> British Columbia and<br />
Washington. Bull. Fish. Res. Bd. Can. 185: 111 pp.<br />
Barnard, J.L. 1969. The families and genera <strong>of</strong> Marine Gammaridean Amphipoda. Smithsonian<br />
Institution (USNM Bull. 271), Washington. D.C. 535 pp.<br />
Baxter, R. 1987. Mollusks <strong>of</strong> Alaska. Shells and Sea Life. Bayside, California. 163 pp.<br />
Behrens, 1991. Pacific coast nudibranchs; A guide to the Opisthobranchs Alaska to Baja<br />
California. Sea Challengers, Monterey California. 107 pp.<br />
Berkeley, E. and C. Berkeley. 1948. Annelida. Polycheata Errantia. Canadian Pacific Fauna,<br />
9b(1). 111pp.<br />
Berkeley, E. and C. Berkeley. 1952. Annelida. Polycheata Sedentaria. Canadian Pacific Fauna,<br />
9b(1), 139.<br />
Bernard, F.R. 1979. Bivalve mollusks <strong>of</strong> the Western Beaufort Sea. Contrib. Sci.: Natur. Hist.<br />
Mus. Los Angeles County. Los Angeles, California. 313: 1-80.<br />
Blanchard, A. and H.M. Feder. 1997. Reproductive timing and nutritional storage cycles <strong>of</strong><br />
Mytilus trossulus Gould, 1850, in Port Valdez, Alaska, Site <strong>of</strong> a Marine Oil Terminal. Veliger<br />
40(2): 121-130.<br />
Butler, T.H. 1980. Shrimps <strong>of</strong> the Pacific Coast <strong>of</strong> Canada. Can. Bull. Fish. Aquat. Sci., Bull.<br />
202, 280p.<br />
Coan, E.C. Unpublished manuscript.
Chapt 9E. Museum, Reference and Voucher Specimens, page 9E- 6<br />
Cooney, R.T. and K.O. Coyle. 1988. <strong>Water</strong> Column Production. In Environmental Studies in<br />
Port Valdez, Alaska: A Basis for Management. Pages 93-110. D.G. Shaw and M.J. Hameedi<br />
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Dick, M.H. and J.R.P. Ross. 1988. Intertidal Bryozoa (Cheilostomata) <strong>of</strong> the Kodiak Vicinity,<br />
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Eyerdam, W. 1924. Marine shells <strong>of</strong> Drier Bay, Knight Island, Prince William Sound, Alaska.<br />
The Nautilus. 38(1):22-28.<br />
Feder, H.M., L.M. Cheek, P. Flannagan, S.C. Jewett, M.H. Johnston, A.S. Naidu, S.A. Norrell,<br />
A.J. Paul, A. Scarborough and D. Shaw. 1976. The sediment environment <strong>of</strong> Port Valdez,<br />
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Foster, N.R. 1991. Intertidal Bivalves: A Guide to the Common Marine Bivalves <strong>of</strong> Alaska.<br />
University <strong>of</strong> Alaska Press, Fairbanks. 152p.<br />
Hart, J.F.L. 1968. Crab-like Anomura and Brachyura (Crustacea: Decapoda) from southeastern<br />
Alaska and Prince William Sound. Nat. Mus. Can. Nat Hist. Papers. 38:6.<br />
Hart, J.F.L. 1982. Crabs and their relatives <strong>of</strong> British Columbia. British Columbia Provincial<br />
Museum, Victoria. 40: 267p.<br />
Haven, S. B. 1971. Effect <strong>of</strong> land-level changes on intertidal invertebrates, with discussion <strong>of</strong><br />
post-earthquake ecological succession. The great Alaska earthquake <strong>of</strong> 1964: Biology. National<br />
Academy <strong>of</strong> Sciences. pp82-125.<br />
Hoberg, M.K. 1986. A numerical analysis <strong>of</strong> the benthic infauna <strong>of</strong> three bays in Prince William<br />
Sound, Alaska. Dept. Biology. Humboldt State University, Arcata, California. 153p.<br />
Hobson, K.D. and K. Banse. 1981. Sedentariate and Archiannelid polychaetes <strong>of</strong> British<br />
Columbia and Washington. Can. Bull. Fish. Aquat. Sci. 209: 144p.<br />
Jewett, S.C. and H.M. Feder. 1977. Biology <strong>of</strong> the Harpacticoid copepod, Harpacticus uniremis<br />
Kroyer on Dayville Flats, Port Valdez, Alaska. Ophelia. 16(1):111-119.<br />
Kluge, G.A. 1962. Bryozoa <strong>of</strong> the northern seas <strong>of</strong> the USSR. Smithsonian Institution and the<br />
National Science Foundation (transl., 1975), TT 72-52010. Amerind Publishing Co. Pvt. Ltd.,<br />
New Delhi. 711p.<br />
Kozl<strong>of</strong>f, E.N. 1987. Marine Invertebrates <strong>of</strong> the Pacific Northwest. University <strong>of</strong> Washington<br />
Press, Seattle. 511p.<br />
Lambert, P. 1981. The Sea Stars <strong>of</strong> British Columbia. British Columbia Provincial, Victoria. 39:<br />
152p.<br />
Lambert, P. 1997. Sea Cucumbers <strong>of</strong> British Columbia, Southeast Alaska and Puget Sound.<br />
UBC Press. Vancouver, British Columbia. 166p.<br />
Lee, R.S. and N.R. Foster. 1985. A distributional list with range extensions <strong>of</strong> the opisthobranch<br />
gastropods <strong>of</strong> Alaska. Veliger. 27(4):440-448.<br />
Millen, S. 1985. The nudibranch genera Onchidoris and Diaphorodoris (Mollusca,<br />
Opisthobranchia) in the northeastern Pacific. Veliger. 28(1):80-93.<br />
Millen, S. 1987. The nudibranch genus Adalaria with a description <strong>of</strong> a new species from the<br />
northeastern Pacific. Can. J. Zool. 65:2696-2702.
Chapt 9E. Museum, Reference and Voucher Specimens, page 9E- 8<br />
O’Clair, C. E. , and S. T. Zimmerman. 1987, Biogeography and ecology <strong>of</strong> intertidal and subtidal<br />
communities. Chapter 11, pp. 305-337. In The Gulf <strong>of</strong> Alaska: Physical environment and<br />
biological resources. OCS study MMS-96-0095.<br />
Pavlovskii, E.N. 1955. Atlas <strong>of</strong> the Invertebrates <strong>of</strong> the Far Eastern Seas <strong>of</strong> the USSR. Acad. Sci.<br />
USSR, Zool. Inst. Israel Program Scient. Transl. (1966), Jerusalem. 457p.<br />
Paul, A.J. and H.M. Feder. 1973. Growth, recruitment and distribution <strong>of</strong> the Littleneck clam,<br />
Protothaca staminea, in Galena Bay, Prince William Sound, Alaska. Fishery Bull. 71(3):665-<br />
677.<br />
Scheel, D., N.R. Foster and K.R. Hough. 1997. Habitat and <strong>Biological</strong> Assessment Shepard Point<br />
Road and Project. Final Rept. Prince William Sound Science Center. Cordova, Alaska.<br />
www.pwssc.gen.ak/~shepard.<br />
Ruiz, G.M. and A.H. Hines. 1997. The risk <strong>of</strong> nonindigenous species invasion in Prince William<br />
Sound associated with oil tanker traffic and ballast water management: Pilot study. Technical<br />
Report , Regional Citizens’ Advisory Council <strong>of</strong> Prince William Sound, 80 p.<br />
Schultz, G.A. 1969. The Marine Isopod Crustaceans. Wm. C. Brown Company. Dubuque, Iowa.<br />
359p.<br />
Schuster, R.O. and A.A. Grigarick. 1965. Taridigrada from Western North America: with<br />
emphasis on the fauna <strong>of</strong> California. Univ. Calif. Publ. Zool., Berkeley. 76: 67p.<br />
Squires, J.J. and A.F.G. Figueria. 1974. Shrimps and shrimp-like Anomura (Crustacea:<br />
Decapoda) from southeastern Alaska and Prince William Sound. Publ. Biol. Oceanogr. Nat.<br />
Mus. Can. 6:23.<br />
Stoker, S.W. 1978. Benthic invertebrate macr<strong>of</strong>auna <strong>of</strong> the eastern continental shelf <strong>of</strong> the<br />
Bering/Chukchi Seas. Ph.D. Disseration. Inst. Mar. Sci., Univ. Alaska, Fairbanks, 259p.<br />
Ushakov, P.V. 1955. Polychaeta <strong>of</strong> the Far Eastern Seas <strong>of</strong> the USSR. Acad. Sci. USSR. Israel<br />
Program Scient Transl. (1965), Jerusalem. 419p.
Chapt 10. Biodiversity, page 10- 1<br />
Chapter 10. Biodiversity <strong>of</strong> Prince William Sound<br />
Nora Foster, University <strong>of</strong> Alaska Museum<br />
Howard M. Feder, Institute <strong>of</strong> Marine Sciences, University <strong>of</strong> Alaska Fairbanks<br />
10A. Summary<br />
Basic taxonomic, biogeographic, and habitat information for 1876 species <strong>of</strong> marine<br />
plants and animals <strong>of</strong> Prince William Sound were compiled into structured data sets. This species<br />
inventory is intended as a baseline from which biodiversity responses to future environmental<br />
changes can be assessed measured. The data sets include 39 possibly undescribed species, 89<br />
range extensions, and 17 nonindigenous species. Two hundred thirty-one marine plant species,<br />
mostly algae, are present. Twenty invertebrate phyla are represented, and Annelida, Mollusca,<br />
and Crustacea account for over 60% <strong>of</strong> the invertebrates. Vertebrates include 175 fish, 14<br />
mammal, and 113 bird species. The fauna and flora <strong>of</strong> Prince William Sound is a mixture <strong>of</strong><br />
species with biogeographic affinities that overlap the northeastern and northwestern Pacific as<br />
well as the Arctic and Atlantic regions.<br />
10B. Purpose<br />
A single inventory <strong>of</strong> Prince William Sound biodiversity is currently lacking. This gap in<br />
taxonomic and biogeographic knowledge is an impediment to efforts to monitor environmental<br />
change, to understand the diversity <strong>of</strong> plant and animal life and to clarify biogeographic<br />
relationships among the living organisms in the rich waters <strong>of</strong> Prince William Sound and the<br />
adjacent shelf <strong>of</strong> the northern Gulf <strong>of</strong> Alaska. (Hines et al. 2000) The purpose <strong>of</strong> this report is to<br />
compile a database for the plant and animal species <strong>of</strong> Prince William Sound. The project is one<br />
component <strong>of</strong> a research project on potential introductions <strong>of</strong> nonindigenous species into Prince<br />
William Sound, especially through the discharge <strong>of</strong> ballast water from oil tankers traveling into<br />
Port Valdez. Annotated species lists were developed for the project to help taxonomic experts<br />
establish a current baseline for the status <strong>of</strong> nonindigenous species in Prince William (Hines et<br />
al. 2000).<br />
Working versions <strong>of</strong> the data sets were compiled by M. Fry and expanded by N. Foster<br />
with assistance by M. Hoberg, and review by K. Coyle (zooplankton), J. Goddard<br />
(opisthobranch gastropods), J. Kudenov (polychaete annelids), G. Lambert (urochordates), P.<br />
Lambert (echinoderms), C. Mecklenberg (fishes), C. Mills (Cnidaria and Ctenophora), J.<br />
Norenburg (nemerteans) and others. Additional polychaete identifications were accomplished for<br />
this report by J. Kudenov. The draft versions <strong>of</strong> the data sets were included in the final project<br />
report (Hines et al. 2000).<br />
10C. Scope<br />
Taxonomic<br />
The data sets list free-living macrophytes and animals . Protists and most endoparasites<br />
are outside the scope <strong>of</strong> the project.<br />
Geographic<br />
Prince William Sound waters are well-defined We have used literature and collecting<br />
records from Port Bainbridge east to Orca Inlet, and south and east to the outer coasts <strong>of</strong>
Chapt 10. Biodiversity, page 10- 2<br />
Montague and Hinchinbrook islands. However, for some taxa, (e.g. Bryozoa), the fauna <strong>of</strong><br />
adjacent areas (northeastern Gulf <strong>of</strong> Alaska and Kodiak Island waters) are better documented and<br />
some records for those areas are included. Migratory birds and marine mammals that inhabit<br />
Prince William Sound waters seasonally are included in the data set. Planktonic Cnidaria and<br />
Crustacea from the northern Gulf <strong>of</strong> Alaska listed in the data sets occur in Prince William Sound<br />
rarely, with their presence dependent on oceanographic conditions.<br />
10D. Methods<br />
Literature sources<br />
A large number <strong>of</strong> species included in the data sets are based on their listing in the<br />
environmental surveys <strong>of</strong> the Gulf <strong>of</strong> Alaska, Prince William Sound, and Port Valdez in the late<br />
1970’s and early 80’s. We compiled species lists from Cooney et al. (1973) and Cooney and<br />
Coyle (1988):plankton, Feder et al. (1976):sediments, Feder and Bryson-Schwafel (1988):the<br />
intertidal zone, Feder and Jewett(1987):subtidal benthos, Feder and Keiser (1980):intertidal<br />
biology, Feder and Matheke (1980):benthic infauna, Feder et al. (1979):benthos <strong>of</strong> Resurrection<br />
and Aialik bays, Feder and Paul (1973):the intertidal zone, Feder and Paul (1980):Cook Inlet,<br />
Hoberg (1986):benthic infauna, Jewett and Feder (1977):Port Valdez, and Rogers et al.<br />
(1986):nearshore fishes.<br />
Species names from field guides and taxonomic revisions <strong>of</strong> characteristic northeastern<br />
Pacific taxa have been added, for example Banse and Hobson (1968) (benthic errantiate<br />
polychaetes), Barnard (1969) (gammaridean amphipods), Butler (1980) (shrimp), Coan et al.<br />
(2000) (bivalve mollusks), Dick and Ross (1988) (Bryozoa), Lambert (1981, 1997) (sea stars and<br />
sea cucumbers), and Schultz (1969) (isopods). For detailed information on the geographical<br />
ranges, we have relied on the same taxonomic revisions and regional publications. However, this<br />
information in a consistent format has been difficult to obtain for many species. When specific<br />
reference to a species’ occurrence in Prince William Sound was not available, we used Austin<br />
(1985). We have also deleted inaccurate records that have crept into the literature, and have<br />
made changes in nomenclature based on reviews and taxonomic revisions accomplished after<br />
the environmental surveys mentioned above.<br />
Biogeographic analyses presented here use conventions based on a global system<br />
developed by the World Conservation Union and described in Kelleher et al. (1995) (Figures<br />
10.1 and 10.2). The biogeographic summaries are based on numbers <strong>of</strong> species for which we<br />
have confident identifications, leaving out unidentified or undescribed species. Some <strong>of</strong> the<br />
analyses exclude birds, because the migratory habits <strong>of</strong> some species fit poorly into the<br />
bioregional classification used for algae, invertebrates and fishes.
Chapt 10. Biodiversity, page 10- 3<br />
Figure 10.1. Biogeographic areas <strong>of</strong> the North Pacific.<br />
Figure 10.2. Biogeographic areas <strong>of</strong> the North Atlantic.
Chapt 10. Biodiversity, page 10- 4<br />
Specimens and observations<br />
The second major source for names <strong>of</strong> Prince William Sound species is the collections <strong>of</strong><br />
the University <strong>of</strong> Alaska Museum (UAM). Names <strong>of</strong> algae, mollusks, bryozoans, and fishes were<br />
entered from specimen catalogues. Voucher specimens for many <strong>of</strong> the species listed in the<br />
environmental studies are in the UAM <strong>Aquatic</strong> Collection, but may not be catalogued. A large<br />
number <strong>of</strong> specimens was also collected in 1990-1995 by Dr. Stephen Jewett as part <strong>of</strong> damage<br />
assessment after the 1989 Exxon Valdez oil spill. While not accessioned into the UAM, these<br />
specimens were available for this study . Each species entry should be traceable to a specimen<br />
identified to the species level by an expert. Citing specimens from other museum collections,<br />
however, goes beyond the scope <strong>of</strong> the project.<br />
Rapid community assessments, focal taxonomic collections, and fouling plate surveys in<br />
Prince William Sound in 1998 and 1999 were intended to detect well-established nonindigenous<br />
species, especially in areas at risk for invasion (Hines and Ruiz 2000). Taxonomic experts who<br />
participated in the field work in Prince William Sound and the Kenai Peninsula contributed<br />
names and distribution information for marine plants (G. Hansen), Cnidaria and Ctenophora (C.<br />
Mills), opisthobranch gastropods (J. Goddard), polychaete annelids (J. Kudenov), Crustacea (J.<br />
Chapman and J. Cordell), Bryozoa (J. Winston), and urochordates (G. and C. Lambert).<br />
Results<br />
The data sets<br />
The data sets comprising the bulk <strong>of</strong> this report are in the form <strong>of</strong> separate Excel<br />
worksheets for algae and vascular plants, Cnidaria and Ctenophora, Annelida, Mollusca,<br />
Arthropoda , Bryozoa, Echinodermata, miscellaneous invertebrate taxa, fishes, birds, mammals<br />
(Tables 10.1 – 10.11). Data are presented in tabular form, one species per row.<br />
The following data fields, as columns, are common to all data sets:<br />
• Family, Genus, Species<br />
An attempt has been made to use the currently accepted name<br />
• Other Name<br />
The column contains family, genus or species names under which the taxon has<br />
appeared in the cited literature, museum catalogs, or specimens labels, which may not<br />
reflect revisions made in the past 20 years.<br />
• A Source for the name<br />
UAM specimen available in the University <strong>of</strong> Alaska Museum <strong>Aquatic</strong> Collection<br />
EVOS specimen archived as part <strong>of</strong> Exxon Valdez oil spill damage assessment studies<br />
Other sources are in Literature Cited section <strong>of</strong> this report.<br />
• Habitat<br />
For most invertebrates the following conventions are used:<br />
I intertidal Inf infauna P plankton<br />
ST subtidal Epi epifauna
Chapt 10. Biodiversity, page 10- 5<br />
Abbreviations used in the fish, bird and mammal sections will be given with the<br />
Summary Information for those taxa.<br />
• Bioregion<br />
An “x” marks the presence <strong>of</strong> a species in the large scale bioregions: Northwestern<br />
Pacific (NWP), Arctic (AR), and Northeastern Atlantic (NWA) (Figures 1, 2).<br />
Occurrence <strong>of</strong> each species with in the northeastern Pacific bioregion is further<br />
designated by Roman numerals denoting a small scale bioregion (Figure 3).<br />
• Origin<br />
For species that represent a new collecting record or nonindigenous species, “origin”<br />
indicates the area closest to Prince William Sound from which the species is likely to<br />
have spread.<br />
• NIS status<br />
nr<br />
NR<br />
C<br />
definite<br />
possible<br />
range extension within Alaska to Prince William Sound<br />
new record for Alaska<br />
cryptogenic<br />
definite nonindigenous species<br />
likely a nonindigenous species<br />
• Reference to Distribution<br />
Literature source for distributional information.<br />
Summary information for major taxa<br />
Marine Plants<br />
Red, brown, and green algae account for over 90% <strong>of</strong> the 231 marine plants listed in the<br />
data set (Table 10.1). Ten taxa could not be determined to the species level. The non-vascular<br />
plant species lists is based on G. Hansen’s (2000) report and cataloged UAM specimens. Records<br />
<strong>of</strong> Chrysophyta and Ascomycota are based on Feder and Keiser 1980, and Feder and Bryson-<br />
Schwafel 1988. Additional distribution information is derived from Scagel et al. 1986 and<br />
Lindstrom 1977. Ten vascular plants species are included. The surfgrasses and eelgrass are<br />
obvious vascular plants, but a few other species are included because <strong>of</strong> their presence in the<br />
upper intertidal and splash zones. Sources used are Scheel et al. 1997; Hulten 1968, and Hansen<br />
2000. Hansen (2000) found 17 new distributional records for marine plants in Prince William<br />
Sound. There is one undescribed species, a Coilodesme and six species are possibly introduced.
Chapt 10. Biodiversity, page 10- 6<br />
Table 10.1. Marine plants.<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE<br />
COMMUNI<br />
TY NEP NWP AR NWA ORIGIN<br />
NIS<br />
STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
PHYLUM FAMILY GENUS SPECIES OTHER NAMES<br />
Feder and Bryson-<br />
Ascomycota Verrucaricaea Verrucaria mucosa<br />
Schwafel 1988 I II III IV x x C Hansen 2000<br />
Ascomycota Verrucaricaea Verrucaria maura Hansen 2000 I II III IV x x C Hansen 2000<br />
Chlorophyta Acrosiphoniaceae Acrosiphonia saxitilis Spongomorpha Hansen 2000 I II III IV x Hansen 2000<br />
Chlorophyta Acrosiphoniaceae Acrosiphonia arcta Hansen 2000 I II III IV x x C Hansen 2000<br />
Chlorophyta Acrosiphoniaceae Acrosiphonia coalita Hansen 2000 I II III IV Hansen 2000<br />
Chlorophyta Acrosiphoniaceae Urospora penicilliformis Hansen 2000 I x x x x C Hansen 2000<br />
Chlorophyta Capsosiphonaceae Capsosiphon fulvescens Hansen 2000 I II III IV x x BC NR, C Hansen 2000<br />
Chlorophyta Chlorocystidaceae Halochlorococcum moorei Hansen 2000 I II III IV x x BC NR, C Hansen 2000<br />
Chlorophyta Cladophoraceae Chaetomorpha capillaris/ cannabina Hansen 2000 I II III IV x x Hansen 2000<br />
Chlorophyta Cladophoraceae Chaetomorpha recurva Hansen 2000 I II III IV WA NR Hansen 2000<br />
Chlorophyta Cladophoraceae Cladophora sericea C. gracilis Hansen 2000 I II III IV x C Hansen 2000<br />
Chlorophyta Cladophoraceae Cladophora albida Hansen 2000 I II III IV x x C Hansen 2000<br />
Chlorophyta Cladophoraceae Cladophora hutchinsiae Hansen 2000 I II III IV x x C Hansen 2000<br />
Chlorophyta Cladophoraceae Cladophora stimpsonii Hansen 2000 I II III IV Hansen 2000<br />
Chlorophyta Cladophoraceae Rhizoclonium implexum R. implexum Hansen 2000 I x x x x C Hansen 2000<br />
Chlorophyta Cladophoraceae Rhizoclonium riparium Hansen 2000 I x x x x C Hansen 2000<br />
Chlorophyta Cladophoraceae Rhizoclonium tortuosum Hansen 2000 I x x x x C Hansen 2000<br />
Chlorophyta Codiaceae Codium fragile subsp. fragile Hansen 2000 I II III IV V VI SE Alaska nr Hansen 2000<br />
Chlorophyta Codiaceae Codium<br />
fragile subsp.<br />
tomentosoides fragile Hansen 2000 I x x x x WA probable Hansen 2000<br />
Chlorophyta Collinsiellaceae Collinsiella tuberculata<br />
Feder and Bryson-<br />
Schwafel 1988 I II III IV Scagel et al. 1986<br />
Chlorophyta Gayraliaceae Gayralia oxyspermum Hansen 2000 I II III IV x x C Hansen 2000<br />
Chlorophyta Monostromataceae Monostroma fractum Hansen 2000 I II III IV WA NR Hansen 2000<br />
Chlorophyta Monostromataceae Monostroma grevillei/arcticum M. arcticum Hansen 2000 I II III IV x x C Hansen 2000<br />
Chlorophyta Ulotrichaceae Ulothrix flacca U. pseud<strong>of</strong>lacca<br />
Feder and Bryson-<br />
Schwafel 1988 I II III IV x x C Scagel et al. 1986<br />
Chlorophyta Ulotrichaceae Ulothrix implexa ( non flacca) Hansen 2000 I II III IV x x C Scagel et al. 1986<br />
Chlorophyta Ulvaceae Blidingia chadefauldi Hansen 2000 I x x x C Hansen 2000<br />
Chlorophyta Ulvaceae Blidingia marginata Hansen 2000 I x x x x BC NR, C Hansen 2000<br />
Chlorophyta Ulvaceae Blidingia minima Hansen 2000 I x x x x C Hansen 2000<br />
Chlorophyta Ulvaceae Blidingia subsalsa Hansen 2000 I II III IV x x C Hansen 2000<br />
Chlorophyta Ulvaceae Enteromorpha compressa<br />
Feder and Bryson-<br />
Schwafel 1988 I x x x x C Scagel et al. 1986<br />
Chlorophyta Ulvaceae Enteromorpha clathrata E. crinita Hansen 2000 I x x x x C Scagel et al. 1986<br />
Chlorophyta Ulvaceae Enteromorpha intestinalis Hansen 2000 I x x x x C Hansen 2000<br />
Chlorophyta Ulvaceae Enteromorpha linza Hansen 2000 I x x x x C Hansen 2000<br />
Chlorophyta Ulvaceae Enteromorpha prolifera/torta Hansen 2000 I x x x x C Hansen 2000<br />
Chlorophyta Ulvaceae Kornmannia leptoderma Hansen 2000 I/epilithic II III IV x x North Atlantic NR, C Hansen 2000
Chapt 10. Biodiversity, page 10- 7<br />
Table 10.1. Continued.<br />
BIOREGION<br />
PHYLUM FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE<br />
COMMUNI<br />
TY NEP NWP AR NWA ORIGIN<br />
NIS<br />
STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Chlorophyta Ulvaceae Kornmannia zostericola Hansen 2000 I/epiphytic II III IV x x C Hansen 2000<br />
Feder and Bryson-<br />
Chlorophyta Ulvaceae Ulva fenestrata U. lactuca Schwafel 1988 I II III IV x Scagel et al. 1986<br />
fenestrata/expansa/<br />
Chlorophyta Ulvaceae Ulva<br />
lactuca Hansen 2000 I x x x x C Hansen 2000<br />
Chlorophyta Ulvaceae Ulvaria obscura<br />
Monostroma<br />
fuscum Hansen 2000 I II III IV x x C Hansen 2000<br />
Chlorophyta Ulvellaceae Ulvella setchelli Hansen 2000<br />
I/epiphytic/e<br />
ndophytic II III IV V VI x C Hansen 2000<br />
Chlorophyta Cladophoraceae Percursaria percursa Hansen 2000 brack/I x x x x C Hansen 2000<br />
Cyanophyta Calothryx crustacea Hansen 2000 I x x x x C Hansen 2000<br />
Cyanophyta Rivularia atra Hansen 2000 I x x x x C Hansen 2000<br />
Phaeophyta Alariaceae Alaria prelonga/marginata Hansen 2000 I/ST II III IV V x Hansen 2000<br />
Phaeophyta Alariaceae Alaria<br />
taeniata/augusta/<br />
crispa Hansen 2000 I/ST II III IV x Hansen 2000<br />
Phaeophyta Alariaceae Alaria<br />
tenuifolia/pylaii/<br />
membranacea Hansen 2000 I/ST II III IV x Hansen 2000<br />
Phaeophyta Chordaceae Chorda filum Hansen 2000 I/ST II III IV x x Hansen 2000<br />
Phaeophyta Chordariacea Chordaria flagelliformis Hansen 2000 I/ST II III IV x x x C Hansen 2000<br />
Phaeophyta Chordariacea Chordaria gracilis Hansen 2000 I/ST II III IV x C Hansen 2000<br />
Phaeophyta Chordariaceae Eudesme virescens Hansen 2000 I II III IV x x Hansen 2000<br />
Phaeophyta Chordariaceae Saundersella simplex UAM I/epiphytic II III IV x Scagel et al. 1986<br />
Phaeophyta Coilodesmaceae Coilodesme bulligera C. polygnampta Hansen 2000 I II III IV x x C Hansen 2000<br />
Phaeophyta Coilodesmaceae Coilodesme californica Hansen 2000<br />
I/ST/<br />
epiphytic II III IV V VI Hansen 2000<br />
Phaeophyta Coilodesmaceae Coilodesme undescribed Hansen 2000 I II III x Hansen 2000<br />
Phaeophyta Cystoseiraceae Cystoceria germinata Hansen 2000 I/ST II III IV x Hansen 2000<br />
Phaeophyta Desmarestiaceae Desmarestia lingulata UAM I/ST II III IV V VI Scagel et al. 1986<br />
Phaeophyta Desmarestiaceae Desmarestia viridis D. media UAM I/ST II III IV V VI x x C Scagel et al. 1986<br />
Phaeophyta Desmarestiaceae Desmarestia viridis Hansen 2000 I/ST II III IV V VI x x Hansen 2000<br />
Phaeophyta Desmarestiaceae Desmarestia aculeata D. intermedia UAM I/ST II III IV x x C Hansen 2000<br />
Phaeophyta Dictyosiphonaceae Dictyosiphon foeniculaceus Hansen 2000 I II III IV x x C Hansen 2000<br />
Phaeophyta Dictyosiphonaceae Dictyosiphon sinicola UAM I II III IV Scagel et al. 1986<br />
Phaeophyta Ectocarpaceae Ectocarpus parvus Hansen 2000 I II III IV V VI Hansen 2000<br />
Phaeophyta Ectocarpaceae Ectocarpus siliculosus Hansen 2000 I x x x x C Hansen 2000<br />
Phaeophyta Ectocarpaceae Ectocarpus acutus Hansen 2000 I II III IV V BC nr Hansen 2000<br />
Phaeophyta Ectocarpaceae Ectocarpus dimorpha E. dimorphus Hansen 2000 I II III IV V BC nr Hansen 2000<br />
Phaeophyta Ectocarpaceae Ectocarpus sp. (Acinetospora) Hansen 2000 I II III IV V Hansen 2000<br />
Phaeophyta Ectocarpaceae Pilayella littoralis Hansen 2000 I x x x x Hansen 2000
Chapt 10. Biodiversity, page 10- 8<br />
Table 10.1. Continued.<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE<br />
COMMUNI<br />
TY NEP NWP AR NWA ORIGIN<br />
NIS<br />
STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
PHYLUM FAMILY GENUS SPECIES OTHER NAMES<br />
littoralis/<br />
Phaeophyta Ectocarpaceae Pilayella<br />
washingtoniensis Hansen 2000 I x x x x C Hansen 2000<br />
Phaeophyta Ectocarpaceae Pilayella unidentified Hansen 2000 I/epiphytic II Hansen 2000<br />
Phaeophyta Ectocarpiaceae Spongonema tomentosum Hansen 2000 I/epiphytic II III IV V VI x x x C Hansen 2000<br />
Phaeophyta Elachistaceae Elachista fucicola Hansen 2000 I/epiphytic II III IV V x x Hansen 2000<br />
Phaeophyta Elachistaceae Elachista lubrica Hansen 2000 I/epiphytic II III x x Hansen 2000<br />
gardneri/distichus/<br />
Phaeophyta Fucaceae Fucus<br />
evanensis Hansen 2000 I II III IV V C Hansen 2000<br />
Phaeophyta Fucaceae Fucus cottoni Hansen 2000 I II III IV x x North Atlantic probable Hansen 2000<br />
Phaeophyta Fucaceae Fucus spiralis UAM I II III IV C Hansen 2000<br />
Phaeophyta Heterochordariaceae Analipus japonicus UAM I II III IV V x Scagel et al. 1986<br />
Phaeophyta Laminariaceae Agarum clathratum (cribrosum) Hansen 2000 ST II III IV x x C Hansen 2000<br />
Phaeophyta Laminariaceae Costaria costata Hansen 2000 I II III IV x Hansen 2000<br />
Phaeophyta Laminariaceae Cymathere triplicata Hansen 2000 ST II III IV x Hansen 2000<br />
Phaeophyta Laminariaceae Hedophyllum sessile UAM I II III IV V Scagel et al. 1986<br />
Phaeophyta Laminariaceae Laminaria groenlandica<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST II III IV Scagel et al. 1986<br />
"groenlandica"/<br />
Phaeophyta Laminariaceae Laminaria<br />
bongardiana Hansen 2000 ST II III IV x x Hansen 2000<br />
Phaeophyta Laminariaceae Laminaria dentigera UAM ST II III IV x Scagel et al. 1986<br />
Phaeophyta Laminariaceae Laminaria longipes UAM I/ST II III IV x Scagel et al. 1986<br />
Phaeophyta Laminariaceae Laminaria yezoensis Hansen 2000 I II III IV x Hansen 2000<br />
Phaeophyta Laminariaceae Laminaria saccharina Hansen 2000 I II III IV x x C Hansen 2000<br />
Phaeophyta Laminariaceae Pleurophycus gardneri UAM ST II III IV V Scagel et al. 1986<br />
Phaeophyta Leathesiaceae Leathesia difformis<br />
Feder and Bryson-<br />
Schwafel 1988 I x x x x C Scagel et al. 1986<br />
Phaeophyta Leathesiaceae Leathesia nana Hansen 2000 I II III IV V VI Hansen 2000<br />
II III IV V VI<br />
Phaeophyta Lessoniaceae Macrocystis integrifolia UAM ST VII VIII IX SE Alaska definate Hansen 2000<br />
Phaeophyta Lessoniaceae Nereocystis leutkeana UAM ST II III IV V Scagel et al. 1986<br />
Phaeophyta Punctariaceae Delamarea attenuata Hansen 2000 I II III IV x x<br />
Commander<br />
Islands NR, C Hansen 2000<br />
Feder and Bryson-<br />
Phaeophyta Punctariaceae Myelophycus intestinalis Melanosiphon Schwafel 1988 I II III IV x x C Scagel et al. 1986<br />
latifolia<br />
Phaeophyta Punctariaceae Punctaria<br />
(Desmotrichium) Hansen 2000 ST II III IV x x SE Alaska nr Hansen 2000<br />
Phaeophyta Punctariaceae Punctaria lobata Hansen 2000<br />
E/ST/<br />
epiphytic II III IV Hansen 2000<br />
Phaeophyta Punctariaceae Punctaria plantagenea Hansen 2000 I II III IV x x Japan NR, C Hansen 2000
Chapt 10. Biodiversity, page 10- 9<br />
Table 10.1. Continued.<br />
BIOREGION<br />
PHYLUM FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE<br />
COMMUNI<br />
TY NEP NWP AR NWA ORIGIN<br />
NIS<br />
STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Phaeophyta Punctariaceae Punctaria tenuimissima Hansen 2000 I II III IV x x C Hansen 2000<br />
Phaeophyta Punctariaceae Soranthera ulvoidea Hansen 2000 I II III IV x Hansen 2000<br />
Phaeophyta Punctariaceae Soranthera ulvoidea f. difformis Hansen 2000 I/epiphytic II III IV x Hansen 2000<br />
Phaeophyta Ralfsiaceae Microspongium globosum Hansen 2000 I/epiphytic II III IV x x Japan probable Hansen 2000<br />
Phaeophyta Ralfsiaceae Ralfsia fungiformis Hansen 2000 I II III IV x x C Hansen 2000<br />
Phaeophyta Scytosiphonaceae Colpomenia peregrina Hansen 2000 I x x x x C Hansen 2000<br />
Phaeophyta Scytosiphonaceae Colpomenia bullosa Hansen 2000 I/epiphytic II III IV x Hansen 2000<br />
Phaeophyta Scytosiphonaceae Petalonia fascia Hansen 2000 I/ST x x x x Hansen 2000<br />
Phaeophyta Scytosiphonaceae Scytosiphon lomentaria<br />
Feder and Bryson-<br />
Schwafel 1988 I x x x x Scagel et al. 1986<br />
Phaeophyta Scytosiphonaceae Scytosiphon simplicissima Hansen 2000 I x x x x C Hansen 2000<br />
Phaeophyta Sphacelariaceae Sphacelaria racemosa Hansen 2000 I/ST II III IV x x C Hansen 2000<br />
Phaeophyta Sphacelariaceae Sphacelaria rigidula Hansen 2000 I x x x x C Hansen 2000<br />
Rhodophyta Acrochaetiaceae Acrochaetium unidentified UAM I C<br />
Rhodophyta Acrochaetiaceae Audouienlla membranaceum<br />
Rhodochorton<br />
membranaceum UAM I x x x x C Scagel et al. 1986<br />
Rhodophyta Acrochaetiaceae Audouinella purpurea Hansen 2000 I x x x x C Hansen 2000<br />
Rhodophyta Bangiaceae Bangia atropurpurea Hansen 2000 I x x x x C Hansen 2000<br />
Rhodophyta Bangiaceae Porphyra cuneiformis Hansen 2000 I/ST II III IV Hansen 2000<br />
Rhodophyta Bangiaceae Porphyra miniata Hansen 2000 I/ST II III IV x x<br />
Commander<br />
Islands NR Hansen 2000<br />
Rhodophyta Bangiaceae Porphyra mumfordii Hansen 2000 I/ST II III IV Hansen 2000<br />
Rhodophyta Bangiaceae Porphyra nereocystis Hansen 2000<br />
ST/<br />
epiphytic II III IV Hansen 2000<br />
Rhodophyta Bangiaceae Porphyra perforata Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Bangiaceae Porphyra purpureo-violacea Hansen 2000 I/ST II III IV x x North Atlantic NR, C Hansen 2000<br />
Rhodophyta Bangiaceae Porphyra rediviva Hansen 2000 I II III IV WA NR Hansen 2000<br />
Rhodophyta Bangiaceae Porphyra cf. P. thuretti<br />
Feder and Bryson-<br />
Schwafel 1988 I II III IV Scagel et al. 1986<br />
Rhodophyta Bangiaceae Porphyra torta/abbottae Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Ceramiaceae Antithamnion dendroideum UAM ST<br />
II III IV V VI<br />
VII VIII IX Scagel et al. 1986<br />
Rhodophyta Ceramiaceae Antithamnionella pacifica Hansen 2000<br />
ST/<br />
epiphytic II III IV Hansen 2000<br />
Rhodophyta Ceramiaceae Antithamnionella spiriographidis Hansen 2000 I II III IV x x x C Scagel et al. 1986<br />
Rhodophyta Ceramiaceae Callithamnion acutum Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Ceramiaceae Callithamnion pikeanum v. laxum Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Ceramiaceae Callithamnion pikeanum v. pikeaum Hansen 2000 I II III IV Hansen 2000
Chapt 10. Biodiversity, page 10- 10<br />
Table 10.1. Continued.<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE<br />
COMMUNI<br />
TY NEP NWP AR NWA ORIGIN<br />
NIS<br />
STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
PHYLUM FAMILY GENUS SPECIES OTHER NAMES<br />
Feder and Bryson-<br />
Rhodophyta Ceramiaceae Callophyllis unidentified<br />
Schwafel 1988 I II III IV<br />
Rhodophyta Ceramiaceae Ceramium gardneri Hansen 2000 I II III IV Hansen 2000<br />
pacificum/<br />
Rhodophyta Ceramiaceae Ceramium<br />
washingtoniensis Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Ceramiaceae Ceramium rubrum/kondoi Hansen 2000 I x x x x Hansen 2000<br />
Rhodophyta Ceramiaceae Ceramium cimbricum Hansen 2000 I II III IV x x Hansen 2000<br />
Rhodophyta Ceramiaceae Ceramium sinicola Hansen 2000 I II III IV CA probable Hansen 2000<br />
Rhodophyta Ceramiaceae Ceramium rubrum UAM I x x x x Scagel et al. 1986<br />
Rhodophyta Ceramiaceae Hollenbergia unidentified UAM I/ST Scagel et al. 1986<br />
Rhodophyta Ceramiaceae Hollenbergia subulata UAM I/ST II III IV V Scagel et al. 1986<br />
Rhodophyta Ceramiaceae Microcladia borealis Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Ceramiaceae Neoptilota asplenioides Hansen 2000 I/ST II III IV x Hansen 2000<br />
Rhodophyta Ceramiaceae Platythamnion pectinatum Hansen 2000 ST II III IV Hansen 2000<br />
Rhodophyta Ceramiaceae Plenosporium cf. P. vancouverianum Hansen 2000 ST II III IV x Hansen 2000<br />
Rhodophyta Ceramiaceae Ptilota serrata ( incl. pectinata) Hansen 2000 I/ST II III IV x x C Hansen 2000<br />
Rhodophyta Ceramiaceae Ptilota californica UAM I/ST II III IV x C Scagel et al. 1986<br />
Abbott and Hollenberg<br />
Rhodophyta Ceramiaceae Ptilota plumosa P. p v. filicina UAM I/ST II III IV V VI x C<br />
1976<br />
Rhodophyta Ceramiaceae Ptilota filicina P. tenuis Hansen 2000 I/ST II III IV Hansen 2000<br />
Rhodophyta Ceramiaceae Scagiella americana Hansen 2000 I II III IV x x C Hansen 2000<br />
Rhodophyta Choreocolacaceae Leachiella pacifica Hansen 2000 I II III IV V VI x Hansen 2000<br />
Rhodophyta Cladophoraceae Chroodactylon ramosus Hansen 2000 I II III IV x x CA probable Hansen 2000<br />
Rhodophyta Corallinaceae Bossiella chiloensis UAM I/ST II III IV Scagel et al. 1986<br />
Rhodophyta Corallinaceae Bossiella cretacea UAM I/ST II III IV x Scagel et al. 1986<br />
Rhodophyta Corallinaceae Bossiella plumosa UAM I II III IV Scagel et al. 1986<br />
Rhodophyta Corallinaceae Calliarthron unidentified UAM I Scagel et al. 1986<br />
Rhodophyta Corallinaceae Clathromorphum reclinatum I/epiphytic II III IV V VI x Scagel et al. 1986<br />
Rhodophyta Corallinaceae Corallina frondescens Hansen 2000 I II III IV Hansen 2000<br />
II III IV V VI<br />
Rhodophyta Corallinaceae Corallina <strong>of</strong>ficianalis v. chilensis Hansen 2000 I/ST VII VII VIII IX x Hansen 2000<br />
Rhodophyta Corallinaceae Corallina vancouverensis UAM I II III IV Scagel et al. 1986<br />
Rhodophyta Corallinaceae Lithophyllum dispar Hansen 2000 II III IV Hansen 2000<br />
Rhodophyta Corallinaceae Lithothamnion unidentified I Scagel et al. 1986<br />
Rhodophyta Corallinaceae Mesophyllum lamellatum Scagel 1986 I II III IV V VI Scagel et al. 1986<br />
Rhodophyta Delesseriacaea Delesseria decipiens Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Delesseriaceae Phycodrys riggii Hansen 2000 I/ST II III IV x Hansen 2000<br />
Rhodophyta Delesseriaceae Tokidadendron kurilensis T. bullata Hansen 2000 I II III IV x Hansen 2000
Chapt 10. Biodiversity, page 10- 11<br />
Table 10.1. Continued.<br />
BIOREGION<br />
PHYLUM FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE<br />
COMMUNI<br />
TY NEP NWP AR NWA ORIGIN<br />
NIS<br />
STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Rhodophyta Dumontiacaea Cryptosiphonia woodii Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Dumontiacaea Weeksia coccinea Hansen 2000 II III IV Hansen 2000<br />
Rhodophyta Dumontiaceae Constantinea subulifera Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Dumontiaceae Constantinea simplex<br />
Feder and Bryson-<br />
Schwafel 1988 I II III IV V Scagel et al. 1986<br />
Rhodophyta Dumontiaceae Dumontia simplex Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Dumontiaceae Dumontia contorta D. incrassata Hansen 2000 I II III IV x x C Hansen 2000<br />
Rhodophyta Dumontiaceae Farlowia mollis UAM I/ST II III IV V VI x Scagel et al. 1986<br />
Rhodophyta Dumontiaceae Neodilsea unidentified UAM I Hansen 2000<br />
Rhodophyta Endocladiaceae Endocladia muricata Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Endocladiaceae Gloiopeltis furcata Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Erythropeltidaceae Erythrotrichia carnea Hansen 2000 I x x x x Hansen 2000<br />
ST/<br />
Rhodophyta Erythropeltidiaceae Smithoria naiadum Hansen 2000 epiphytic II III IV C Hansen 2000<br />
Rhodophyta Gigartinaceae Gigartina unidentified UAM I/ST Hansen 2000<br />
Rhodophyta Gigartinaceae Iridaea cordata UAM I/ST II III IV x Scagel et al. 1986<br />
Rhodophyta Gigartinaceae Iridaea cornucopiae<br />
Feder and Bryson-<br />
Schwafel 1988 I II III IV x Scagel et al. 1986<br />
Rhodophyta Gigartinaceae Rhodoglossum latissimum UAM I II III IV Scagel et al. 1986<br />
Rhodophyta Halemeniaceae Cryptonemia obovata Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Halemeniaceae Cryptonemia borealis Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Helminthocladiaceae Nemalion helminthoides Hansen 2000 I x x x x C Hansen 2000<br />
Rhodophyta Hildenbrandiaceae Hildenbrandia rubra H. prototypus Hansen 2000 I x x x x C Scagel et al. 1986<br />
Rhodophyta Nemastomataceae Neodilsea borealis<br />
N. americana/<br />
Schizymenia UAM I II III IV Scagel et al. 1986<br />
Rhodophyta Palmariaceae Devaleraea ramentacea Hansen 2000 I II III IV x x C Hansen 2000<br />
Rhodophyta Palmariaceae Halosaccion glandiforme Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Palmariaceae Halosaccion firmum Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Palmeriaceae Palmaria mollis/palmata Hansen 2000 I II III IV x x C Hansen 2000<br />
Rhodophyta Palmeriaceae Palmaria<br />
calophylloides/<br />
stenogona Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Palmeriaceae Palmaria hecatensis Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Petrocelidaceae Mastocarpus cf. M. pacificus Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Petrocelidaceae Mastocarpus papillatus complex Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Petrocelidaceae Mastocarpus papillatus Gigartina cristata UAM I<br />
II III IV V VI<br />
VII VII VIII IX Scagel et al. 1986<br />
Rhodophyta Phyllophoraceae Ahnfeltia plicata UAM I II III IV x x Scagel et al. 1986<br />
Rhodophyta Phyllophoraceae Ahnfeltia fastigata Hansen 2000 I x x x x C Hansen 2000<br />
Rhodophyta Phyllophoraceae Ahnfeltiopsis gigartinoides Arnfeltia Hansen 2000 I II III IV Hansen 2000
Chapt 10. Biodiversity, page 10- 12<br />
Table 10.1. Continued.<br />
BIOREGION<br />
PHYLUM FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE<br />
COMMUNI<br />
TY NEP NWP AR NWA ORIGIN<br />
NIS<br />
STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Rhodophyta Rhodomelaceae Polysiphonia brodiaei Hansen 2000 I x x x x C Hansen 2000<br />
Rhodophyta Rhodomelaceae Polysiphonia eastwoodae Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Rhodomelaceae Polysiphonia<br />
hendryi v.<br />
deliquescens Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Rhodomelaceae Polysiphonia hendryi v. hendryi Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Rhodomelaceae Polysiphonia hendryi v. luxurians Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Rhodomelaceae Polysiphonia hendryi v. pacifica Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Rhodomelaceae Polysiphonia<br />
pacifica v.<br />
determinata Hansen 2000 I/ST II III IV Hansen 2000<br />
Rhodophyta Rhodomelaceae Polysiphonia pacifica v. pacifica Hansen 2000 I/ST II III IV Hansen 2000<br />
Rhodophyta Rhodomelaceae Polysiphonia pungens Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Rhodomelaceae Polysiphonia hendryi P. collinsii UAM I II III IV x Scagel et al. 1986<br />
Rhodophyta Rhodomelaceae Polysiphonia pacifica<br />
Feder and Bryson-<br />
Schwafel 1988 I II III IV Scagel et al. 1986<br />
Rhodophyta Rhodomelaceae Polysiphonia senticulosa Hansen 2000 I II III IV SE Alaska nr Hansen 2000<br />
Rhodophyta Rhodomelaceae Polysiphonia stricta (ureceolata) Hansen 2000 I II III IV x x Hansen 2000<br />
Rhodophyta Rhodomelaceae Polysiphonia ureceolata Hansen 2000 I x x x x Hansen 2000<br />
Rhodophyta Rhodomelaceae Pterosiphonia bipinnata Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Rhodomelaceae Rhodomela lycopodioides Hansen 2000 I II III IV x x C Hansen 2000<br />
Rhodophyta Rodomelaceae Neorhodomela larix Rhodomela Hansen 2000 I/ST II III IV x Hansen 2000<br />
Rhodophyta Rodomelaceae Neorhodomela aculaeta Hansen 2000 I/ST II III IV x Hansen 2000<br />
Rhodophyta Rodomelaceae Neorhodomela oregona Hansen 2000 I/ST II III IV x Hansen 2000<br />
Rhodophyta Rodomelaceae Odonthalia floccosa Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Rodomelaceae Odonthalia kamtschatica Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Rodomelaceae Odonthalia setacea Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Rhodymeniaceae Rhodymenia pertusa Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Solieriaceae Opuntiella californica Hansen 2000 ST II III IV Hansen 2000<br />
Rhodophyta Gigartinaceae Mazzaella heterocarpa Hansen 2000 I II III IV Hansen 2000<br />
Rhodophyta Gigartinaceae Mazzaella phyllocarpa Hansen 2000 I II III IV x Hansen 2000<br />
Rhodophyta Gigartinaceae Mazzaella splendescens Hansen 2000 I II III IV Hansen 2000<br />
Xanthophyta Vaucheria longicaulis () Hansen 2000 I II III IV x x BC NR Hansen 2000<br />
vascular plant Caryophyllaceae Honchneya peploides Scheel et al. 1997 I II III IV x Hulten 1968<br />
vascular plant Cyperaceae Carex unidentified Feder and Bryson-SchwI Hulten 1968<br />
vascular plant Gramineae Puccinellia nutkaensis Scheel et al. 1997 I II III IV Hulten 1968<br />
vascular plant Gramineae Puccinellia plumila Hulten 1968 I II III IV Hulten 1968<br />
vascular plant Primulaceae Glaux maritima Scheel et al. 1997 I II III IV x x Hulten 1968<br />
vascular plant Rosaceae Potentilla unidentified Feder and Bryson-SchwI Hulten 1968<br />
vascular plant Sparganiaceae Phyllospadix scouleri Hansen 2000 I/ST II III IV Hulten 1968<br />
vascular plant Sparganiaceae Phyllospadix serrulatus Hansen 2000 I/ST II III IV Hulten 1968<br />
vascular plant Sparganiaceae Zostera marina Hansen 2000 I/ST II III IV x x Hulten 1968
Chapt 10. Biodiversity, page 10- 13<br />
Cnidaria and Ctenophora<br />
One hundred two species <strong>of</strong> Cnidaria and Ctenophora are listed in this report (Table<br />
10.2). Twenty-seven <strong>of</strong> the taxa have been identified to the generic level, and not to species.<br />
Anthozoa are probably under-counted, because they are difficult to identify based on preserved<br />
specimens. The hydrozoans and scyphozoans include 34 species, four <strong>of</strong> which, the hydrozoans<br />
Tiaropsis multicirrata, Eperetmus typus, Gonionemus vertens and Proboscidactyla flavicirrata,<br />
are distributional range extensions (Mills 2000). Thirty species records are derived from T.<br />
Cooney‘s (1987) compilation <strong>of</strong> Gulf <strong>of</strong> Alaska plankton.<br />
Table 10.2. Cnidaria and Ctenophora.<br />
PHYLUM/<br />
SPECIMEN or<br />
CLASS FAMILY GENUS SPECIES OTHER NAMES SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
Cnidaria:<br />
Anthozoa Actiniidae Anthopleura artemesia<br />
Cnidaria:<br />
Anthozoa Actiniidae Cribirnopsis unidentified Rosenthal 1977 ST/Epi<br />
Cnidaria:<br />
Anthozoa Actiniidae Urticina crassicornis<br />
NIS<br />
STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Feder and Bryson-<br />
Schwafel 1988 I II III IV Austin 1985<br />
Feder and Jewett<br />
1987 I/ST/Epi II III IV x Austin 1985<br />
Cnidaria:<br />
Anthozoa Actinostolidae Stomphia unidentified Scheel et al. 1997 ST/Epi<br />
Cnidaria:<br />
Anthozoa Carophllidae Carophyllia alaskensis UAM ST II III IV Austin 1985<br />
Cnidaria:<br />
Anthozoa Cerianthidae Pachycerianthus unidentified UAM ST<br />
Cnidaria:<br />
Anthozoa Epizoanthidae Epizoanthus unidentified UAM ST<br />
Cnidaria:<br />
Anthozoa Halcalvidae Peachia unidentified<br />
Cnidaria:<br />
Anthozoa Metridiidae Metridium senile<br />
Feder and Matheke<br />
1980 ST<br />
Feder and Jewett<br />
1987 I/ST/Epi II x x x Austin 1985<br />
Cnidaria:<br />
Anthozoa Metridiidae Metridium unidentified NRF observation ST<br />
Cnidaria:<br />
Anthozoa Pennatulidae Ptilosarcus gurneyi Feder et al. 1979 ST II III IV Austin 1985<br />
Cnidaria:<br />
Anthozoa Primnoidae Callogorgia unidentified UAM ST<br />
Cnidaria:<br />
Anthozoa Primnoidae Primnoa unidentified UAM ST<br />
Cnidaria:<br />
Anthozoa Virgularidae Acanthoptilum ptile<br />
Feder and Matheke<br />
1980 ST II III<br />
Cnidaria:<br />
Anthozoa Virgularidae Stylatula elongata Rosenthal 1977 ST II III IV V<br />
Cnidaria:<br />
British<br />
columbia NR Austin 1985<br />
Hydrozoa Aeginidae Aegina citrea A. rosea Cooney 1987 P II III IV V VI x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Aequoreidae Aequorea<br />
aequorea v.<br />
albida Mills 2000 ST II III IV<br />
Wrobel and Mills<br />
1998<br />
Cnidaria:<br />
Hydrozoa Aequoreidae Aequorea unidentified Cooney:SEA ST<br />
Cnidaria:<br />
Hydrozoa Aequoreidae Aequorea<br />
victoria/ A.<br />
aequorea v.<br />
aequorea Mills 2000 ST II III IV<br />
Wrobel and Mills<br />
1998<br />
Cnidaria:<br />
Hydrozoa Agalmidae Agalma elegans Cooney 1987 P II III x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Agalmidae Nanomia unidentified Cooney 1987 P<br />
Cnidaria:<br />
Hydrozoa Bougainvilliidae Bougainvillia superciliaris Mills 2000 ST II III IV x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Bougainvilliidae Garveia franciscana Mills 2000 ST x x x x uncertain definate<br />
Cnidaria:<br />
Hydrozoa Calycopsidae Calycopsis simulans<br />
Cnidaria:<br />
Hydrozoa Campanulariidae Clytia gregaria<br />
Cnidaria:<br />
Hydrozoa Campanulariidae Clytia hemispherca<br />
Wrobel and Mills<br />
1998 P x<br />
Phialidium<br />
gregarium Mills 2000 I II III IV<br />
BIOREGION<br />
Hines and Ruiz<br />
2000<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Hines and Ruiz<br />
2000 ST/Epi II III IV V VI x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Campanulariidae Clytia kincaidi Mills 2000 ST x x x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Campanulariidae Gonothyraea clarki Mills 2000 I II III IV x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Campanulariidae Obelia longissima Mills 2000 I II x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Campanulariidae Obelia borealis O. dichotoma Cooney 1987 P II III IV V x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Campanulariidae Obelia unidentified Mills 2000 I<br />
Cnidaria:<br />
Hydrozoa Campanulariidae Obelia spp. (hydriods) Mills 2000 I<br />
Cnidaria:<br />
Hydrozoa Campanulariidae Phialidium gregarium P. languidum Cooney 1987 P II III IV V Austin 1985
Chapt 10. Biodiversity, page 10- 14<br />
Table 10.2 continued.<br />
PHYLUM/<br />
SPECIMEN or<br />
CLASS FAMILY GENUS SPECIES OTHER NAMES SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
Cnidaria:<br />
Hydrozoa Campanulariidae Verticillina verticillata<br />
Feder and Matheke<br />
1980 ST II III IV V x x<br />
NIS<br />
STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Cnidaria:<br />
Hydrozoa Clausophyidae Chuniphyes multidentata Cooney 1987 P II III IV V VI Austin 1985<br />
Cnidaria:<br />
Hydrozoa Corymorphidae Euphysa japonica Cooney 1987 P II III IV x x x<br />
Wrobel and Mills<br />
1998<br />
Cnidaria:<br />
Hydrozoa Corymorphidae Euphysa unidentified Mills 2000 I/ST/Epi<br />
Cnidaria:<br />
Hydrozoa Corynidae Sarsia spp. (medusae) Mills 2000 I/ST/Epi<br />
Cnidaria:<br />
Hydrozoa Corynidae Sarsia tubulosa Mills 2000 I/ST/Epi x x x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Corynidae Sarsia eximia Mills 2000 I/ST/Epi II x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Corynidae Sarsia princeps Cooney 1987 P II III IV x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Corynidae Sarsia rosaria Cooney 1987 P II III IV x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Corynidae Sarsia/Coryne sp. (hydroids) Mills 2000 I/ST/Epi<br />
Cnidaria:<br />
Hydrozoa Cuninidae Cunina globosa Cooney 1987 P II III IV Cooney 1987<br />
Cnidaria:<br />
Hydrozoa Diphyidae Dimophyes arctica Cooney 1987 P<br />
II III IV V VI<br />
VII VIII x x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Diphyidae Lensia concoidea Cooney 1987 P II III IV V VI x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Diphyidae Muggiaea atlantica Cooney 1987 P II III x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Eirenidae Eutonina indicans<br />
Wrobel and Mills<br />
1998 P II III IV x<br />
Wrobel and Mills<br />
1998<br />
Cnidaria:<br />
Hydrozoa Eutimidae Eutonima indicans Mills 2000 ST II III IV Austin 1985<br />
Cnidaria:<br />
Hydrozoa Haleciidae Grammaria immersa Rosenthal 1977 ST II III x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Halicreatidae Halicreas unidentified Cooney:SEA P<br />
Cnidaria:<br />
Hydrozoa Halimedusidae Halimedusa typus Cooney 1987 P II III IV Austin 1985<br />
Cnidaria:<br />
Hydrozoa Hippopodiidae Vogtia serrata V. pentacantha Cooney 1987 P II III x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Laodicidae Cuspidella unidentified Cooney:SEA P<br />
Cnidaria:<br />
Hydrozoa Laodicidae Staurophora mertensii Mills 2000 ST II x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Laodicidae Stegopoma plicatile<br />
BIOREGION<br />
Feder and Matheke<br />
1980 ST II x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Lovenellidae Calycella springe Mills 2000 I II x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Melicertidae Melicertum campanula Cooney:SEA ST II III IV x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Melicertidae Melicertum octocostatum Mills 2000 ST II x x<br />
Cnidaria:<br />
Hydrozoa Mitrocomidae Mitrocoma cellularia Halistaura Mills 2000 ST II III IV V<br />
Cnidaria:<br />
Hydrozoa Mitrocomidae Tiaropsidium unidentified Cooney 1987 P<br />
Cnidaria:<br />
Hydrozoa Mitrocomidae Tiaropsis multicirrata Mills 2000 I/ST/Epi II III IV x x SE Alaska nr<br />
Cnidaria:<br />
Hydrozoa Olindiasidae Eperetmus typus Mills 2000 ST II III IV SE Alaska nr<br />
Cnidaria:<br />
Hydrozoa Olindiasidae Gonionemus unidentified Cooney:SEA ST<br />
Cnidaria:<br />
Hydrozoa Olindiasidae Gonionemus vertens Mills 2000 ST II III IV SE Alaska nr<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998
Chapt 10. Biodiversity, page 10- 15<br />
Table 10.2 continued.<br />
PHYLUM/<br />
SPECIMEN or<br />
CLASS FAMILY GENUS SPECIES OTHER NAMES SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
NIS<br />
STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Cnidaria:<br />
Hydrozoa Pandeidae Catablema multicirrata Mills 2000 ST/Epi II III IV x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Pandeidae Halitholus unidentified Mills 2000 ST<br />
Cnidaria:<br />
Hydrozoa Pandeidae Leuckartiara breviconis Neoturris Cooney 1987 P II III IV Austin 1985<br />
Cnidaria:<br />
Hydrozoa Pandeidae Leuckartiara nobilis<br />
Catablema<br />
nodulosa Cooney 1987 P II III IV Austin 1985<br />
Cnidaria:<br />
Hydrozoa Pandeidae Leuckartiara octona Cooney 1987 P II III IV Austin 1985<br />
Cnidaria:<br />
Hydrozoa Pandeidae Leuckartiara unidentified Mills 2000 I/ST/Epi<br />
Cnidaria:<br />
Hydrozoa Pandeidae Neoturris breviconis<br />
Wrobel and Mills<br />
1998 P II II III IV V<br />
Cnidaria:<br />
Hydrozoa Pandeidae Perigonimus unidentified Cooney:SEA P<br />
Cnidaria:<br />
Hydrozoa Pandeidae Stomotoka atra<br />
Wrobel and Mills<br />
1998 P II III<br />
Cnidaria:<br />
Hydrozoa Periphyllidae Periphylla periphylla P. hyacinthina Cooney 1987 P II III IV V VI x<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Cnidaria:<br />
Hydrozoa Phialellidae Campanulina rugosa Mills 2000 ST II III IV Austin 1985<br />
Cnidaria:<br />
Hydrozoa Phialellidae Opercularella lacerata Mills 2000 ST II x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Polyorchidae Polyorchis penicillatus<br />
Wrobel and Mills<br />
1998 P II III IV V VI<br />
Wrobel and Mills<br />
1998<br />
Cnidaria:<br />
Hydrozoa Prayidae Nectopyramis diomedeae Cooney 1987 P II III IV V VI x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Prayidae Praya reticulata Nectodroma Cooney 1987 P<br />
II III IV V VI<br />
VII VIII Austin 1985<br />
Cnidaria:<br />
Hydrozoa Proboscidactylidae Proboscidactyla flavicirrata Mills 2000 I/ST/Epi II x x SE Alaska nr<br />
Cnidaria:<br />
Hydrozoa Protohydridae Protohydra unidentified<br />
Feder and Bryson-<br />
Schwafel 1988<br />
Wrobel and Mills<br />
1998<br />
Cnidaria:<br />
Hydrozoa Rathkeidae Rathkea jaschnowi Cooney:SEA P II x Bering Sea nr Naumov 1960<br />
Cnidaria:<br />
Hydrozoa Rathkeidae Rathkea octopunctata R. blumenbachii Cooney 1987 P II III IV x x<br />
Cnidaria:<br />
Hydrozoa Rathkeidae Rathkea unidentified Cooney:SEA P<br />
Cnidaria:<br />
Hydrozoa Rhopalonematidae Aglantha digitale<br />
Cooney and Coyle<br />
1988 P/ST II III IV x x<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Cnidaria:<br />
Hydrozoa Rhopalonematidae Pantachogon haekeli Cooney 1987 P II III IV x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Sertulariidae Sertularia robusta Mills 2000 ST II x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Tubularidae Hybocodon prolifer Cooney 1987 P II III x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Tubularidae Plotocnide borealis Cooney:SEA P II III IV x x Austin 1985<br />
Cnidaria:<br />
Hydrozoa Tubularidae Tubularia prolifera Cooney:SEA P II x x x Austin 1985<br />
Cnidaria:<br />
Scyphozoa Cyaneidae Cyanea capillata Mills 2000 ST II x x<br />
Cnidaria:<br />
Scyphozoa Pelagiidae Chrysaora melangaster Cooney 1987 P II III IV V VI x<br />
Cnidaria:<br />
Scyphozoa Ulmariidae Aurelia aurita Mills 2000 ST II x x<br />
Cnidaria:<br />
Scyphozoa Ulmariidae Aurelia labiata Mills 2000 ST II III IV x<br />
Cnidaria:<br />
Scyphozoa Ulmariidae Aurelia unidentified Mills 2000 P<br />
Cnidaria:<br />
Scyphozoa Ulmariidae Phacellophora camtschatica Catablema Cooney:SEA P II III IV V VI x x<br />
Ctenophora Beroidae Beroe unidentified Cooney 1987 P<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Ctenophora Bolinopsidae Bolinopsis infundibulum Mills 2000 ST II x x<br />
Wrobel and Mills<br />
1998<br />
Ctenophora Mertensiidae Mertensia sp. (ovum)<br />
Cooney and Coyle<br />
1988 P II x x Austin 1985<br />
Ctenophora Pleurobranchiidae Hormiphora cucumis<br />
Wrobel and Mills<br />
1998 P II III x<br />
Ctenophora Pleurobranchiidae Pleurobrachia bachei Mills 2000 ST II III IV<br />
BIOREGION<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998
Chapt 10. Biodiversity, page 10- 16<br />
Annelida<br />
The region’s polychaete annelid fauna is particularly rich, with 233 species in 46<br />
families. Kudenov (2000) has noted three range extensions, for Phyllodoce medipapillata,<br />
Chaetozone senticosa, and Rhynchospio glutaea. Some species <strong>of</strong> Eumida, Scoloplos, Exogone,<br />
Nephtys, Glycera, and the archaeannelid Polygordius are likely to be undescribed. Seven taxa,<br />
including members <strong>of</strong> the Spirorbidae have not been identified below the generic level. Nineteen<br />
new records from the UAM collection and Exxon Valdez oil spill specimens represent range<br />
extensions.<br />
Hirudinea and Oligochaeta are not included in the data sets (Table 10.3), because we lack<br />
reliable identifications for specimens found in Prince William Sound. At least one species <strong>of</strong><br />
marine leech is present in UAM samples from the adjacent Gulf <strong>of</strong> Alaska. Kozl<strong>of</strong>f (1987) lists<br />
14 leech species with distributions from Oregon north. Oligochaetes have been collected, but no<br />
effort has been made to identify them to family or lower taxon. Austin (1985) lists eight<br />
oligochaete species whose ranges include Alaska or southeast Alaska.<br />
Information on distribution, habitat, and nomenclatural changes have been drawn from<br />
Austin (1985), Banse and Hobson (1968), Berkeley and Berkeley (1948, 1952), Hobson and<br />
Banse (1981), Kudenov (2000) and Ushakov (1955).<br />
Table 10.3. Annelida.<br />
GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Ampharete acutifrons EVOS ST/Inf II III IV Austin 1985<br />
Ampharete finmarchia A. arctica<br />
Feder et al.<br />
1979 ST/Inf II III IV x x x Austin 1985<br />
Amphicteis<br />
gunneri<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x Ushakov 1955<br />
Amphicteis<br />
scaphobranchiata<br />
Feder et al.<br />
1979 ST/Inf II III IV Austin 1985<br />
Anobothrus<br />
gracilis<br />
Feder et al.<br />
1979 ST/Inf II III IV x Austin 1985<br />
Lysippe<br />
labiata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Melinna<br />
cf. M. cristata<br />
Feder et al.<br />
1979 ST/Inf II III IV x Austin 1985<br />
Melinna<br />
cristata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Melinna<br />
elisabethae<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Pseudosabellides lineata Asabellides EVOS ST/Inf II III IV x Austin 1985<br />
Pseudosabellides sibirica Asabellides littoralis UAM ST/Inf II III IV Austin 1985<br />
Sosanella unidentified UAM<br />
Aphrodita japonica UAM ST/Inf II III IV x Austin 1985<br />
Drilonereis<br />
falcata minor<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV<br />
BIOREGION<br />
Drilonereis longa UAM ST/Inf II x x<br />
British<br />
Columbia NR Austin 1985<br />
British<br />
Columbia NR Austin 1985<br />
Abarenicola<br />
pacifica<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV x Austin 1985<br />
Barantolla americana EVOS I/ST/Inf II III IV C Austin 1985<br />
Capitella<br />
capitata<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II x x x C Austin 1985<br />
Heteromastus filiformis EVOS ST/Inf II III IV V x x possible Austin 1985<br />
Mediomastus unidentified EVOS I/ST/Inf<br />
Mesochaetopterus taylori EVOS ST/Inf II III IV<br />
British<br />
Columbia NR Austin 1985<br />
Feder and<br />
Spiochaetopterus costarum<br />
Matheke 1980 ST/Inf x x x x Austin 1985
Chapt 10. Biodiversity, page 10- 17<br />
Table 10.3 continued.<br />
BIOREGION<br />
GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Chrysopetalium occidentale Kudenov 2000 II III IV V VI x Kudenov 2000<br />
Chaetozone senticosa Kudenov 2000 ST/Inf II IV V California NR Kudenov 2000<br />
Feder and<br />
Chaetozone<br />
setosa<br />
Matheke 1980 ST/Inf x x x x Austin 1985<br />
Cirratulus cingulatus Kudenov 2000 ST/Inf II III IV Kudenov 2000<br />
Cirratulus cirratus EVOS ST/Inf x x x x<br />
British<br />
Columbia NR Austin 1985<br />
Tharyx<br />
monilaris<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 ST/Inf II III IV Austin 1985<br />
Tharyx<br />
multifilis<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV Austin 1985<br />
Tharyx<br />
parvus<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Tharyx secundus EVOS ST/Inf II III IV<br />
British<br />
Columbia NR Austin 1985<br />
Cossura<br />
longocirrata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Dorvillea rudolphi EVOS ST/Inf II III IV<br />
Dorvillea<br />
pseudorubrovittata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Protodorvillea gracilis EVOS ST/Inf II III IV Austin 1985<br />
Schistomeringos unidentified UAM ST<br />
Eunice valens E. kobiensis UAM ST/Inf II III IV x Austin 1985<br />
Brada<br />
granulata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Flabelligera affinis EVOS ST/Inf II III IV x Austin 1985<br />
Pherusa plumosa EVOS I/ST/Inf II III IV x Austin 1985<br />
Pherusa<br />
papillata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Glycera sp. undescribed Kudenov 2000 Kudenov 2000<br />
Glycera cf. G. nana Kudenov 2000<br />
Glycera nana G. capitata<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 ST/Inf II III IV x Austin 1985<br />
Hemipodus borealis EVOS I/ST/Inf II III IV Austin 1985<br />
Glycinde armigera EVOS ST/Inf II III IV x Austin 1985<br />
Glycinde<br />
Goniada<br />
Goniada<br />
picta<br />
annulata<br />
maculata<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV Austin 1985<br />
Feder and<br />
Jewett 1988 ST/Inf II III IV Austin 1985<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Microphthalmus sczelkowii<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 ST/Inf II x x California NR Austin 1985<br />
Micropodarke dubia EVOS ST/Inf II III IV Austin 1985<br />
Feder and<br />
Podarkeopsis glabrus Gyptis brevipalpa<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Pelagobia longicirrata Cooney 1987 P II III IV V VI x x Austin 1985<br />
Feder et al.<br />
II III IV V VI<br />
Lumbrineris<br />
latreilli<br />
1979 ST/Inf VII VIII IX x Austin 1985<br />
Lumbrineris limicola UAM ST/Inf II III IV SE Alaska nr<br />
Lumbrineris<br />
Lumbrineris<br />
luti<br />
similabris<br />
Feder and<br />
Jewett 1988 ST/Inf II III IV<br />
British<br />
Columbia NR Austin 1985<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Lumbrineris<br />
zonata<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV Austin 1985<br />
Ninoe<br />
gemmea<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Ninoe<br />
simpla<br />
Feder et al.<br />
1979 ST/Inf II III IV Austin 1985<br />
Magelona berkeleyi EVOS ST/Inf II III IV<br />
British<br />
Columbia NR Austin 1985<br />
Magelona hobsonae M. pitekai<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV<br />
British<br />
Columbia NR Austin 1985
Chapt 10. Biodiversity, page 10- 18<br />
Table 10.3 continued.<br />
GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Metasychis disparidentata Asychis<br />
Feder et al.<br />
1979 ST/Inf II III IV x Austin 1985<br />
Nicomache personata UAM ST/Inf II III IV Austin 1985<br />
Notoproctus<br />
pacificus<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Petaloproctus borealis P. tenuis EVOS ST/Inf II III IV x Austin 1985<br />
Praxillella praetermissa EVOS ST/Inf II III IV x Austin 1985<br />
Praxillella<br />
gracilis<br />
Feder and<br />
Jewett 1988 ST/Inf II III IV Austin 1985<br />
Praxillella pacifica P. affinis<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Rhodine<br />
bitorquata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Aglophamus<br />
rubella anops<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Nephtys cornuta cornuta UAM ST/Inf II III IV Austin 1985<br />
Nephtys ferruginea EVOS ST/Inf II III IV Austin 1985<br />
Nephtys<br />
punctata<br />
Feder and<br />
Jewett 1988 ST/Inf II III IV x Austin 1985<br />
Nephtys<br />
caeca<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV V x x x Austin 1985<br />
Nephtys ciliata EVOS I/ST/Inf II x x Austin 1985<br />
Nephtys<br />
cornuta<br />
Feder and<br />
Matheke 1980 I/ST/Inf II III IV Austin 1985<br />
Nephtys<br />
cornuta franciscana<br />
Feder and<br />
Matheke 1980 I/ST/Inf II III IV Austin 1985<br />
Nephtys sp. undescribed Kudenov 2000<br />
Chelonereis cyclurus Kudenov 2000 ST/Inf II III IV V x Austin 1985<br />
Nereis grubei UAM I/ST/Inf II III IV x Austin 1985<br />
Nereis pelagica UAM I/ST/Inf II III IV Austin 1985<br />
Nereis<br />
procera<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV V VI Austin 1985<br />
Nereis<br />
vexillosa<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/inf II III IV V VI Austin 1985<br />
Nereis<br />
zonata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV V VI x Austin 1985<br />
Platynereis bicanaliculata P. agassizi EVOS I/ST/Inf II III IV Austin 1985<br />
Onuphis iridescens UAM ST/Inf II III IV x Austin 1985<br />
Onuphis<br />
conchylega<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x<br />
BIOREGION<br />
Banse and Hobson<br />
1968<br />
Onuphis<br />
geophiliformis<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Onuphis<br />
parva<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x<br />
Banse and Hobson<br />
1968<br />
Onuphis stigmatis ST/Inf II III IV Austin 1985<br />
Armandia brevis EVOS ST/Inf II III IV Austin 1985<br />
Ophelia<br />
limacina<br />
Ophelina acuminata Amnotrypane aulogaster<br />
Travisia<br />
forbesii<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf x x x x Austin 1985<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV V VI x Austin 1985<br />
Feder et al.<br />
1979 ST/Inf II x x x Austin 1985<br />
Leitoscoloplos<br />
panamensis<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV Austin 1985<br />
Leitoscoloplos pugettensis L. elongatus<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Naineris dendritica Kudenov 2000 ST/Inf II III IV V Austin 1985<br />
Naineris drenritica N. laevigata UAM ST/Inf II III IV Austin 1985<br />
Naineris uncinata EVOS ST/Inf II III IV Austin 1985<br />
Naineris<br />
unidentified<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Scoloplos acmeceps EVOS ST/Inf II III IV Austin 1985<br />
Scoloplos armiger EVOS ST/Inf II III IV x x Austin 1985<br />
Scoloplos sp. undescribed Kudenov 2000<br />
Feder and<br />
Myriochele oculata M. heeri<br />
Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Owenia<br />
fusiformis<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV x x Austin 1985<br />
Aricidea ramosa EVOS ST/Inf II III IV x Austin 1985
Chapt 10. Biodiversity, page 10- 19<br />
Table 10.3 continued.<br />
BIOREGION<br />
GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Aricidea<br />
lopezi<br />
Feder and<br />
Jewett 1988 ST/Inf II III IV x Austin 1985<br />
Aricidea<br />
neosuecica<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Aricidea nolani A. suecica UAM ST/Inf II III IV Austin 1985<br />
Cirrophoras lyra EVOS ST/Inf II III IV Austin 1985<br />
Cirrophoras branchiatus UAM ST/Inf II III IV Austin 1985<br />
Paraonis<br />
gracilis<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x x Austin 1985<br />
Amphictene moorei A. auricoma<br />
Feder and<br />
Matheke 1980 I/ST/Inf II III IV x Austin 1985<br />
Cistenides granulata C. brevicoma<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV x Austin 1985<br />
Pectinaria californiensis EVOS ST/Inf II III IV Austin 1985<br />
Pholoides asperus Polyodontidae:Peisidice<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Anaitides groenlandica Phyllodoce UAM ST/Inf II III IV x x Austin 1985<br />
Anaitides<br />
mucosa<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Anaitides<br />
maculata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Eteone<br />
longa<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV x C Austin 1985<br />
Eulalia viridis UAM ST/Inf II III IV x Austin 1985<br />
Eumida sp. undescribed Kudenov 2000 ST/Inf<br />
Hypoeulalia bilineata Eulalia UAM ST/Inf II III IV x x Austin 1985<br />
Mysta barbata UAM ST/Inf II x x Chukchi Sea nr Ushakov 1955<br />
Nereiphylla castanea Phyllodoce, Genetylus UAM ST/Inf II III IV x Austin 1985<br />
Notophyllum tectum UAM ST/Inf II III IV Austin 1985<br />
Phyllodoce medipapillata Kudenov 2000 ST/Inf II III IV V California NR Kudenov 2000<br />
Ancistrosyllis<br />
hamata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Polygordius sp. undescribed UAM ST/Inf Kudenov 2000<br />
Feder and<br />
Antinoella<br />
sarsi<br />
Matheke 1980 ST/Inf II III IV x x Ushakov 1955<br />
Arcteobia<br />
spinelytris<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Feder and<br />
Byglides macrolepidus Antinoella<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Eunoe depressa UAM ST/Inf II III IV x Austin 1985<br />
Eunoe oerstedi EVOS ST/Inf II III IV Austin 1985<br />
Eunoe senta UAM ST/Inf II III IV Austin 1985<br />
Gattyana<br />
ciliata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Gattyana<br />
cirrosa<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x x Austin 1985<br />
Gattyana<br />
treadwelli<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Halosydna brevisetosa UAM ST/Inf II III IV Austin 1985<br />
Harmothoe extenuata UAM ST/Inf II III IV x x Austin 1985<br />
Harmothoe imbricata EVOS I/ST/Inf II III IV x x C Austin 1985<br />
Hesperonoe<br />
complanata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Lepidonotus helotypus UAM ST/Inf II III IV Ushakov 1955<br />
Feder and<br />
Lepidonotus<br />
squamatus<br />
Matheke 1980 ST/Inf x x x x Austin 1985<br />
Nemida tamarae UAM ST/Inf II Bering Sea nr Ushakov 1955<br />
Feder et al.<br />
Polynoe<br />
canadensis<br />
1979 ST/Inf II III IV x Austin 1985<br />
Idanthyrsus<br />
armatus<br />
Feder and<br />
Matheke 1980 ST/Inf<br />
II III IV V VI<br />
VII VIII IX Austin 1985<br />
Idanthyrsus ornamentatus EVOS ST/Inf II III IV Austin 1985<br />
Chone magna EVOS ST/Inf II III IV Austin 1985<br />
Chone cincta UAM ST/Inf II III IV x Austin 1985<br />
Euchone hancocki EVOS ST/Inf II III IV Austin 1985<br />
Euchone longifissuirata UAM ST/Inf II III IV x x Ushakov 1955<br />
Euchone<br />
analis<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV x x Ushakov 1955<br />
Fabricia<br />
sabella<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II x x Austin 1985
Chapt 10. Biodiversity, page 10- 20<br />
Table 10.3 continued.<br />
BIOREGION<br />
GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
REFERENCE to<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS DISTRIBUTION<br />
Laonome<br />
kroyeri<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II x x<br />
Berkeley and<br />
Berkeley 1952<br />
Myxicola<br />
infundibulum<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Potentilla ocellata UAM ST/Inf II III IV Austin 1985<br />
Schizobranchia insignis Kudenov 2000 II III IV V Austin 1985<br />
Scalibregma<br />
inflatum<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Crucigera<br />
irregularis<br />
Feder and<br />
Matheke 1980 ST/Epi II III IV Austin 1985<br />
Crucigera zygophora UAM ST/Epi II III IV x Austin 1985<br />
Pseudochitinopoma occidentalis EVOS ST/Epi II III IV Austin 1985<br />
Serpula vermicularis Kudenov 2000 ST/Epi II III IV x Kudenov 2000<br />
Pholoe<br />
sp. "minuta"<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV x x C Austin 1985<br />
Sphaerodoropsis minuta EVOS ST/Inf II III IV x Austin 1985<br />
Sphaerodoropsis sphaerulifer EVOS ST/Inf II III IV x Austin 1985<br />
Sphaerodorum papillifer EVOS ST/Inf II x x x Austin 1985<br />
Dipolydora cf. D. bidentata Kudenov 2000 Kudenov 2000<br />
Dipolydora cf. D. giardi Kudenov 2000 Kudenov 2000<br />
Dipolydora cf. D. socialis Kudenov 2000 Kudenov 2000<br />
Laonice cirrata EVOS ST/Inf x x x x Austin 1985<br />
Berkeley and<br />
Rhynchospio glutaea Malacoceros EVOS ST/Inf II x x nr<br />
Berkeley 1952<br />
Polydora limicola Kudenov 2000 ST/Inf II III IV V Austin 1985<br />
Polydora cf. P.brachycephalata UAM ST/Inf II III IV Austin 1985<br />
Polydora ciliata P. pygidialis<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf x x x x Austin 1985<br />
Dipolydora quadrilobata EVOS I/ST/Inf II III IV C Austin 1985<br />
II III IV V VI<br />
Polydora socialis Dipolydora UAM ST/Inf VII VIII IX Austin 1985<br />
Prionospio cirrifera EVOS ST/Inf II III IV x Austin 1985<br />
Prionospio steenstrupi EVOS I/ST/Inf x x x x Austin 1985<br />
Pygospio elegans EVOS I/ST/Inf II x x Austin 1985<br />
Scolelepis squamata Nerine cirratulus EVOS ST/Inf II III IV Austin 1985<br />
Spio filicornis EVOS ST/Inf x x x x Austin 1985<br />
Spiophanes berkeleyorum EVOS ST/Inf II III IV x Austin 1985<br />
Spiophanes bombyx EVOS ST/Inf x x x x Austin 1985<br />
Feder et al.<br />
Spiophanes cirrata S. kroyeri<br />
1979 ST/Inf II x x Austin 1985<br />
Circeis<br />
spirillum<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/Epi II III IV Austin 1985<br />
Dexiospira<br />
unidentified<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/Epi II III IV Austin 1985<br />
Spirorbis<br />
unidentified<br />
Feder and<br />
Matheke 1980 ST/Epi II III IV x x Austin 1985<br />
Sternaspis<br />
scutata<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Autolytus alexandri A. verrilli UAM ST/Inf II III IV Austin 1985<br />
Autolytus cornatus A. prismaticus UAM ST/Inf II x x x Austin 1985<br />
Eusyllis<br />
bloomstrandi<br />
Feder and<br />
Matheke 1980 ST/Inf II x x Austin 1985<br />
Exogone sp. undescribed Kudenov 2000<br />
Exogone cf. E. dwisula Kudenov 2000 Kudenov 2000<br />
Exogone<br />
lourei<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 ST/Inf II III IV Austin 1985<br />
Exogone naidina E. gemmifera<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV x Austin 1985<br />
Feder and<br />
Exogone<br />
verugera<br />
Matheke 1980 ST/Inf x x x x Austin 1985<br />
Sphaerosyllis cf. C. californiensis Kudenov 2000 Kudenov 2000
Chapt 10. Biodiversity, page 10- 21<br />
Table 10.3 continued.<br />
BIOREGION<br />
GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
REFERENCE to<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS DISTRIBUTION<br />
Feder and<br />
Bryson-<br />
Schwafel 1988 ST/Inf II III IV x<br />
Banse and Hobson<br />
1968<br />
Sphaerosyllis<br />
erinaceous<br />
Feder and<br />
Syllis armillaris Typosyllis<br />
Matheke 1980 ST/Inf II III IV V VI x x x Austin 1985<br />
Syllis fasciata Typosyllis EVOS ST/Inf II III IV Austin 1985<br />
Syllis harti Typosyllis EVOS ST/Inf II III IV<br />
Feder and<br />
Syllis harti armillaris Typosyllis<br />
Matheke 1980 ST/Inf II III IV<br />
British<br />
Columbia NR Austin 1985<br />
Syllis harti unguicula Typosyllis UAM ST/Inf II III IV<br />
British<br />
Columbia NR<br />
Feder and<br />
Syllis heterochaeta Langerhansia cornuta Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Syllis hyalina Typosyllis Kudenov 2000 I/ST/Inf x x x x Austin 1985<br />
Trypanosyllis gemmipara Kudenov 2000 II III IV V Austin 1985<br />
Typosyllis stuarti Kudenov 2000 II III Austin 1985<br />
Typosyllis alternata Kudenov 2000 I/ST/Inf II III IV x Austin 1985<br />
Amphitrite cirrata Kudenov 2000 II III IV V x C Austin 1985<br />
Artacama conifera UAM ST/Inf II III IV Austin 1985<br />
Artacama<br />
proboscidea<br />
Feder et al.<br />
1979 ST/Inf II III IV x Austin 1985<br />
Lanassa venusta EVOS ST/Inf II III IV x x Austin 1985<br />
Laphania boecki EVOS ST/Inf II x x Austin 1985<br />
Leana abranchiata UAM ST/Inf II III IV x x Austin 1985<br />
Feder and<br />
Lysilla<br />
loveni<br />
Matheke 1980 ST/Inf II x x Austin 1985<br />
Pista<br />
cristata<br />
Feder and<br />
Matheke 1980 ST/Inf II x x Austin 1985<br />
Feder and<br />
Pista vinogradovi P. pacifica<br />
Jewett 1988 ST/Inf II III IV x Austin 1985<br />
Polycirrus medusa UAM ST/Inf II x x Austin 1985<br />
Proclea graffi UAM ST/Inf II III IV Austin 1985<br />
Terbellides<br />
stroemi<br />
Feder and<br />
Jewett 1988 ST/Inf x x x x Austin 1985<br />
Tomopteris septentrionalis Cooney 1987 P x x x Austin 1985<br />
Tomopteris pacificus Cooney 1987 P II III IV V x Austin 1985<br />
Tomopteris<br />
unidentified<br />
Cooney and<br />
Coyle 1988 P<br />
Trichobranchus<br />
glacialis<br />
Feder and<br />
Matheke 1980 ST/Inf II x x x Austin 1985<br />
Disoma carica Disomidae:Trichochaeta UAM ST/Inf II III IV x Austin 1985<br />
Disoma multisetosum Disomidae:Trichochaeta UAM ST/Inf II III IV x Austin 1985<br />
Typhloscolex mulleri Cooney: SEA ST II III IV x x NE Pacific NR Austin 1985<br />
Mollusca<br />
Three hundred fifteen mollusk species, representing all classes except<br />
monoplacophorans, are included in the data set (Table 10.4). There is one aplacophoran, and 108<br />
bivalves, 179 gastropods, 17 polyplacophorans, three scaphopods, and eight cephalopods.<br />
Mollusks in the UAM comprise one <strong>of</strong> the chief sources for the data set. Information on<br />
nomenclature and distribution derives form Austin (1985), Baxter (1987), Behrens (1991), Coan<br />
et al. (2000), and Turgeon et al. (1998).<br />
The shelled fauna is quite well known, but opisthobranchs, because they require special<br />
techniques to collect and preserve, have not been adequately documented in Alaska. It is not<br />
surprising to find potentially undescribed species and geographical range extensions within the<br />
Prince William Sound fauna. A paper describing range extensions for 11 species <strong>of</strong><br />
opisthobranchs is in preparation by J. Goddard and N. Foster.
Chapt 10. Biodiversity, page 10- 22<br />
Table 10.4. Mollusca.<br />
BIOREGION<br />
GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Chaetoderma robustum UAM ST/Inf II III IV Baxter 1987<br />
Pododesmus macroschisma UAM ST/Epi II III IV Coan et al. 2000<br />
Astarte borealis UAM ST/Inf II x x x Coan et al. 2000<br />
Astarte compacta A. polaris UAM ST/Inf II III IV Coan et al. 2000<br />
Astarte elliptica A. alaskensis UAM ST/Inf II x x Coan et al. 2000<br />
Astarte esquimaulti UAM ST/Inf II III x Coan et al. 2000<br />
Astarte ovata A. borealis UAM ST/Inf II x x x Coan et al. 2000<br />
Clinocardium blandum C. fucanum Coan et al. 2000 I/ST/Inf II III IV Coan et al. 2000<br />
Clinocardium californiense UAM I/ST/Inf II III IV x Coan et al. 2000<br />
Clinocardium ciliatum UAM I/ST/Inf II x x x Coan et al. 2000<br />
Clinocardium nuttallii UAM I/ST/Inf II III IV x Coan et al. 2000<br />
Serripes groenlandicus UAM I/ST/Inf II x x x Coan et al. 2000<br />
Serripes laperousii UAM I/ST/Inf II x x x Coan et al. 2000<br />
Serripes notabilis UAM ST/Inf II III IV x Coan et al. 2000<br />
Cyclocardia crebricosta UAM ST/Inf II III IV x Coan et al. 2000<br />
Cyclocardia ventricosa UAM ST/Inf II III IV Coan et al. 2000<br />
Miodontiscus prolongata Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000<br />
Cardiomya behringensis Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000<br />
Cardiomya pectinata UAM ST/Inf II III IV Coan et al. 2000<br />
Cardiomya planetica UAM ST/Inf II III IV Coan et al. 2000<br />
Glycymeris septentrionalis EVOS ST/Inf II III IV Coan et al. 2000<br />
Hiatella arctica UAM I/ST/Epi/Inf x x x x Coan et al. 2000<br />
Panomya ampla Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000<br />
Panomya norvegica P. arctica UAM ST/Inf II x x x Coan et al. 2000<br />
Kellia suborbicularis Kelliidae Coan et al. 2000 ST/Inf II III IV Coan et al. 2000<br />
Mysella planata Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000<br />
Neaeromya compressa Pseudopythina UAM I/ST/Inf II III IV Coan et al. 2000<br />
Rochefordia tumida Mysella UAM ST/Inf II III IV Coan et al. 2000<br />
Limatula attenuata L. subauriculata Coan et al. 2000 ST/Epi II III IV x Coan et al. 2000<br />
Lucina tenuisculpta Parvilucina UAM ST/Inf II III IV Coan et al. 2000<br />
Lucinoma annulatum UAM ST/Inf II III IV Coan et al. 2000<br />
Entodesma navicula E. saxicola UAM I/Epi/Inf II III IV x Coan et al. 2000<br />
Lyonsia bracteata UAM ST/Inf II III IV Coan et al. 2000<br />
Mactromeris polynyma UAM I/ST/Inf II x Coan et al. 2000<br />
Tresus capax Coan et al. 2000 I/ST/Inf II III IV Coan et al. 2000<br />
Tresus nuttallii UAM I/ST/Inf II III IV Coan et al. 2000<br />
Malletia pacifica M. cuneata UAM ST/Inf II III IV Coan et al. 2000<br />
Cryptomya californica Coan et al. 2000 I/ST/Inf II III IV x Coan et al. 2000<br />
Mya arenaria UAM I/Inf II III IV x N Atlantic definate Coan et al. 2000<br />
Mya truncata UAM I/ST/Inf II x x x Coan et al. 2000<br />
Crenella decussata UAM I/ST/Epi II x x x Coan et al. 2000<br />
Dacrydium vitreum UAM ST/Inf II x x x Coan et al. 2000<br />
Modiolus modiolus UAM ST/Inf II x x Coan et al. 2000<br />
Musculus discors UAM ST/Inf II x x Coan et al. 2000<br />
Musculus glacialis M. corrugatus UAM ST/Inf II x x x Coan et al. 2000<br />
Musculus niger UAM ST/Inf II x x x Coan et al. 2000<br />
Mytilus trossulus M. edulis UAM I/ST/Epi II III IV Coan et al. 2000<br />
Solamen columbianun Megacrenella Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000<br />
Vilasina seminuda Musculus Coan et al. 2000 ST/Epi II III IV x Coan et al. 2000<br />
Vilasina vernicosa Musculus UAM ST/Epi II III IV x Coan et al. 2000<br />
Nuculana conceptionis Perrisonota UAM ST/Inf II III IV Coan et al. 2000<br />
Nuculana minuta UAM ST/Inf II x x x Coan et al. 2000<br />
Nuculana navisa Coan et al. 2000 ST/Inf II III IV Coan et al. 2000<br />
Nuculana pernula N. fossa UAM ST/Inf II x x x Coan et al. 2000<br />
Eunucula tenuis Nucula UAM ST/Inf II x x Coan et al. 2000<br />
Crassostrea gigas UAM I/ST/Epi II III IV x NW Pacific definate Coan et al. 2000<br />
Pandora wardiana P. grandis UAM ST/Inf II III IV x Coan et al. 2000<br />
Pandora bilirata Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000<br />
Pandora filosa Coan et al. 2000 ST/Inf II III IV Coan et al. 2000<br />
Pandora glacialis UAM ST/Inf II x x x Coan et al. 2000<br />
Chlamys rubida UAM ST/Epi II III IV x Coan et al. 2000<br />
Crassdoma gigantea Hinnites Coan et al. 2000 ST/Epi II III IV Coan et al. 2000<br />
Delectopecten vancouverensis D. randolphi UAM ST/Epi II III IV x Coan et al. 2000<br />
Parvamussium alaskense UAM ST/Epi II III IV Coan et al. 2000<br />
Patinopecten caurinus UAM ST/Epi II III IV Coan et al. 2000<br />
Periploma aleuticum UAM ST/Inf II x x x Coan et al. 2000<br />
Siliqua alta Solenidae: S. sloati UAM I/ST/Inf II III IV x Coan et al. 2000<br />
Siliqua patula Solenidae Coan et al. 2000 I/Inf II III IV Coan et al. 2000<br />
Philibrya setosa Coan et al. 2000 ST/Inf II III IV Coan et al. 2000<br />
Penitella penita Coan et al. 2000 I/ST/Inf II III IV Coan et al. 2000
Chapt 10. Biodiversity, page 10- 23<br />
Table 10.4 continued.<br />
BIOREGION<br />
GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Gari californica UAM I/Inf II III IV x Coan et al. 2000<br />
Macoma dexioptera UAM ST/Inf II Coan et al. 2000<br />
Macoma elimata UAM ST/Inf II III IV Coan et al. 2000<br />
Macoma moesta UAM ST/Inf II III IV x x Coan et al. 2000<br />
Macoma balthica UAM I/Inf x x x x C Coan et al. 2000<br />
Macoma brota UAM ST/Inf II III IV Coan et al. 2000<br />
Macoma calcarea UAM ST/Inf II x x x Coan et al. 2000<br />
Macoma carlottensis UAM ST/Inf II III IV Coan et al. 2000<br />
Macoma expansa UAM I/ST/Inf II III IV Coan et al. 2000<br />
Macoma golikovi M. obliqua UAM I/ST/Inf II III IV Coan et al. 2000<br />
Macoma inquinata UAM I/ST/Inf II III IV Coan et al. 2000<br />
Macoma lipara Coan et al. 2000 ST/Inf II III IV Coan et al. 2000<br />
Macoma nasuta UAM I/ST/Inf II III IV Coan et al. 2000<br />
Tellina modesta UAM ST/Inf II III IV Coan et al. 2000<br />
Bankia setacea UAM ST/Inf II III IV x Coan et al. 2000<br />
Thracia challisiana Coan et al. 2000 ST/Inf II III IV Coan et al. 2000<br />
Thracia condoni Coan et al. 2000 ST/Inf II III IV Coan et al. 2000<br />
Thracia myopsis UAM ST/Inf II x x x Coan et al. 2000<br />
Thracia trapeziodes UAM ST/Inf II III IV Coan et al. 2000<br />
Adontorhina cyclia UAM I/ST/Inf II III IV x Coan et al. 2000<br />
Axinopsida serricata UAM I/ST/Inf II x x x Coan et al. 2000<br />
Conchocoele bisecta Thyasira Coan et al. 2000 ST/Inf II III IV x Coan et al. 2000<br />
Mendicula ferruginosa Odontogena borealis UAM ST/Inf II x x x Coan et al. 2000<br />
Thyasira flexuosa UAM ST/Inf II x x x Coan et al. 2000<br />
Turtonia minuta UAM I/Inf II x Coan et al. 2000<br />
Diplodonta impolita UAM I/Inf II III IV Coan et al. 2000<br />
Compsomyax subdiaphana UAM ST/Inf II III IV Coan et al. 2000<br />
Humilaria kennerleyi UAM I/ST/Inf II III IV Coan et al. 2000<br />
Liocyma fluctuosa UAM ST/Inf II x x x Coan et al. 2000<br />
Nutricola lordi Psephidia ovalis UAM I/Inf II III IV Coan et al. 2000<br />
Nutricola tantilla Transenella UAM I/ST/Inf II III IV Coan et al. 2000<br />
Protothaca staminea UAM I/ST/Inf II III IV x Coan et al. 2000<br />
Saxidomus gigantea UAM I/Inf II III IV Coan et al. 2000<br />
Yoldia hyperborea Y. amygdalea UAM ST/Inf II x x x Coan et al. 2000<br />
Yoldia<br />
montereyensis<br />
Megayoldia martyria/<br />
M. secunda/ M.<br />
beringiana UAM ST/Inf II III IV Coan et al. 2000<br />
Yoldia myalis UAM ST/Inf II x Coan et al. 2000<br />
Yoldia seminuda Y. scissurata UAM ST/Inf II III IV x Coan et al. 2000<br />
Yoldia thraciaeformis Megayoldia UAM ST/Inf II x x x Coan et al. 2000<br />
Gonatus<br />
unidentified<br />
Feder and Jewett<br />
1988 P<br />
Benthoctopus leioderma<br />
Scheel- possible<br />
for PWS ST/Epi II III IV Austin 1985<br />
Octopus d<strong>of</strong>leini Enteroctopus<br />
Scheel et al.<br />
1997 ST/Epi II III IV Austin 1985<br />
Octopus<br />
rubescens<br />
Scheelobservation<br />
ST/Epi II III IV Austin 1985<br />
Rossia pacifica UAM ST/Epi II III IV x Austin 1985<br />
II III IV V<br />
Chiroteuthis veranayi C. calyx Cooney 1987 P<br />
VI Austin 1985<br />
Gliteuthis armata G. phyllura Cooney 1987 P<br />
II III IV V<br />
VI x Austin 1985<br />
Acanthodoris<br />
hudsoni<br />
Lee and Foster<br />
1985 I/ST/Epi II III IV<br />
Lee and Foster<br />
1985<br />
Lee and Foster<br />
1985<br />
Acanthodoris nanaimoenisis Goddard 2000 I/ST/Epi II III IV<br />
Lee and Foster<br />
Acanthodoris pilosa<br />
1985 I/ST/Epi II x x Behrens 1991<br />
Acmaea mitra UAM I/Epi II III IV Austin 1985<br />
Aeolidia<br />
papillosa<br />
Lee and Foster<br />
1985 I/ST/Epi x x x x Behrens 1991<br />
Lee and Foster<br />
Aglaja ocelligera Melanochlamys Goddard 2000 I/ST/Epi II III IV<br />
1985<br />
Melanochlamys diomedea Aglaja<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Epi II III IV SE Alaska nr<br />
Lee and Foster<br />
1985
Chapt 10. Biodiversity, page 10- 24<br />
Table 10.4 continued.<br />
BIOREGION<br />
GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Archidoris montereyensis I/ST/Epi II III IV<br />
Lee and Foster<br />
1985<br />
Archidoris<br />
Armina<br />
odhneri<br />
californica<br />
Lee and Foster<br />
1985 I/ST/Epi II III IV<br />
Lee and Foster<br />
1985 ST/Epi II III IV<br />
Haminoea vesicula Foster 2000 I/ST/Epi II III IV<br />
Lee and Foster<br />
1985<br />
Lee and Foster<br />
1985<br />
Lee and Foster<br />
1985<br />
Lee and Foster<br />
1985<br />
Haminoea virescens Baxter 1987 I/ST/Epi II III IV<br />
Barleeia subtenuis UAM I/Inf II III IV Baxter 1987<br />
Barleeia haliotiphila Baxter 1987 I/Inf II III IV Baxter 1987<br />
Pseudodiala acuta Barleeia UAM I/Inf II III IV<br />
British<br />
Columbia NR Baxter 1987<br />
Ancistrolepis eucosmuis UAM ST/Epi II III IV Baxter 1987<br />
Beringius kennicotti UAM ST/Epi II III IV Austin 1985<br />
Buccinum baerii UAM I/Epi II III IV Austin 1985<br />
Buccinum plectrum UAM ST/Epi II III IV x x Austin 1985<br />
Colus aphelus C. hypolispus UAM ST/Epi II III IV x x<br />
Turgeon et al.<br />
1998<br />
Colus halli UAM ST/Epi II III IV Austin 1985<br />
Lirabuccinum dira Searlesia UAM I/ST/Epi II III IV Baxter 1987<br />
Lussivolutopsius filosa Volutopsius UAM I/ST/Epi II III IV Baxter 1987<br />
Neptunea lyrata UAM ST/Epi II III IV x x Austin 1985<br />
Neptunea pribil<strong>of</strong>fensis UAM ST/Epi II III IV x Austin 1985<br />
Pyrul<strong>of</strong>usus harpa Volutopsius UAM I/ST/Epi II III IV Austin 1985<br />
Volutopsius undescribed sp. Baxter 1987 I/ST/Epi II III IV<br />
Turgeon et al.<br />
1998<br />
Caecum crebricinctum Micranellum UAM I/ST/Inf II III IV Baxter 1987<br />
Caecum occidentale Fartulum UAM I/Inf II III IV Baxter 1987<br />
Calliostoma ligatum Trochidae UAM I/ST/Epi II III IV Baxter 1987<br />
Calyptraea fastigiata UAM I/ST/Epi II III IV Baxter 1987<br />
Crepidula perforans Baxter 1987 I/ST/Epi II III IV Austin 1985<br />
Crepipatella lingulata<br />
Calyptraea fastigiata/<br />
Crepiptella dorsata UAM I/ST/Epi II III IV Baxter 1987<br />
Admete viridula A. couthouyi UAM ST/Inf II x x x Baxter 1987<br />
Neadmete modesta UAM ST/Inf II III IV Baxter 1987<br />
Trichotropis borealis Trichotropidae UAM I/ST/Epi II x Baxter 1987<br />
Trichotropis cancellata Trichotropidae UAM I/ST/Epi II III IV Baxter 1987<br />
Trichotropis insignis Trichotropidae UAM I/ST/Epi II III IV x Baxter 1987<br />
Stylidium eschrichti Bittium Baxter 1987 I/Inf II III IV Austin 1985<br />
II III IV V<br />
Cadlina luteomarginata Rosenthal 1977 ST/Epi VI SE Alaska nr Behrens 1991<br />
Cooney and<br />
Wrobel and Mills<br />
Clione<br />
limacina<br />
Coyle 1988 P x x x x<br />
1998<br />
Cocculina baxteri UAM ST/Epi II III Baxter 1987<br />
Alia carinata Mitrella I/ST/Epi II III IV Baxter 1987<br />
Amphissa columbiana UAM I/ST/Inf II III IV Baxter 1987<br />
Amphissa reticulata UAM ST/Epi II III IV Baxter 1987<br />
Astrys gauspata Alia UAM I/ST/Inf II III IV Baxter 1987<br />
Grantotoma incisula Turridae: Oenopota UAM ST/Inf II x Baxter 1987<br />
Kurtziella plumbea Turridae EVOS I/ST/Epi II III IV Austin 1985<br />
Oenopota bicarinata Turridae UAM ST/Inf II x Austin 1985<br />
Oenopota excurvata Turridae UAM ST/Inf II III IV Austin 1985<br />
Oenopota krausei Turridae UAM ST/Inf II III IV Austin 1985<br />
Oenopota rugulata Turridae UAM ST/Inf II x Austin 1985<br />
Oenopota turricula Turridae UAM ST/Inf II x Austin 1985<br />
Propebela arctica<br />
Turridae: Oenopota<br />
viridula UAM ST/Inf II x Baxter 1987<br />
Pseudotaranis strongi Turridae: Tarais EVOS ST/Epi II III IV Austin 1985<br />
Doridella steinbergi Suhinia EVOS ST/Epi II III IV Behrens 1991<br />
Flabellina trophina F. fusca<br />
Lee and Foster<br />
1985 ST/Epi II III IV<br />
Lee and Foster<br />
1985 ST/Epi II x x x<br />
Lee and Foster<br />
1985<br />
Lee and Foster<br />
1985<br />
Flabellina salmonacea<br />
Euclio pyamidata Clio Cooney 1987 P x x x x Austin 1985<br />
Granulina margaritula Marginellidae UAM I/ST/Epi II III IV Baxter 1987
Chapt 10. Biodiversity, page 10- 25<br />
Table 10.4 continued.<br />
GENUS SPECIES OTHER NAMES<br />
Dendronotus<br />
Dendronotus<br />
Dendronotus<br />
Diaphana<br />
albus<br />
frondosus<br />
iris<br />
minuta<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Lee and Foster<br />
1985 ST/Epi II III IV Behrens 1991<br />
Lee and Foster<br />
1985 ST/Epi II x Behrens 1991<br />
Lee and Foster<br />
1985 ST/Epi II III IV Behrens 1991<br />
Lee and Foster<br />
1985 ST/Inf II x<br />
Diaphana brunnea UAM II III IV<br />
Turgeon et al.<br />
1998<br />
Lee and Foster<br />
1985<br />
Dirona<br />
albolineata<br />
Lee and Foster<br />
1985 ST/Epi II III IV Behrens 1991<br />
Dirona<br />
aurantia<br />
Lee and Foster<br />
1985 ST/Epi II III IV Behrens 1991<br />
Anisodoris lentigenosa<br />
Lee and Foster<br />
1985 ST/Epi II III IV Behrens 1991<br />
Anisodoris nobilis<br />
Lee and Foster<br />
1985 ST/Epi II III IV Behrens 1991<br />
Dialula<br />
sandiegensis<br />
Lee and Foster<br />
1985 ST/Epi II III IV Behrens 1991<br />
Geitodoris heathi Goddard 2000 ST/Epi II III IV<br />
British<br />
Columbia NR Behrens 1991<br />
Eubranchus olivaceous Goddard 2000 ST/Epi II III IV SE Alaska nr Behrens 1991<br />
Melanella columbiana Vitreolina UAM II III IV Baxter 1987<br />
Melanella micrans Balcis UAM ST/Epi II III IV Baxter 1987<br />
Hermissenda crassicornis Goddard 2000 I/ST/Epi II III IV Behrens 1991<br />
Diadora aspera Baxter 1987 I/ST/Epi II III IV Austin 1985<br />
Puncturella cooperi UAM ST/Epi II III IV Austin 1985<br />
Puncturella galeata UAM ST/Epi II III IV Austin 1985<br />
Puncturella noachina UAM ST/Epi II III IV x x Austin 1985<br />
Feder and Bryson-<br />
Gastropteron pacificum<br />
Schwafel 1988 I/Epi II III IV Behrens 1991<br />
Ancula pacifica Goddard 2000 ST/Epi II III IV SE Alaska nr Behrens 1991<br />
Alderia modesta Goddard 2000 ST/Epi II x<br />
British<br />
Columbia NR Behrens 1991<br />
Cryptobranchia alba UAM I/ST/Epi II III IV x Baxter 1987<br />
Cryptobranchia concentrica UAM I/ST/Epi II III IV x Baxter 1987<br />
Lepeta caeca UAM ST/Epi II III IV x Baxter 1987<br />
Limacina<br />
helicina<br />
Cooney and<br />
Coyle 1988 P x x x x<br />
Wrobel and Mills<br />
1998<br />
Lacuna carinata Lacunidae UAM I/ST/Epi II III IV Austin 1985<br />
Lacuna marmorata Lacunidae Baxter 1987 I/ST/Epi II III IV Baxter 1987<br />
Lacuna variegata Lacunidae Baxter 1987 I/ST/Epi II III IV Baxter 1987<br />
Lacuna vincta Lacunidae UAM I/ST/Epi II x Austin 1985<br />
Littorina scutulata UAM I/Epi II III IV Austin 1985<br />
Littorina sitkana UAM I/Epi II III IV Austin 1985<br />
Lottia borealis UAM I/Epi II III IV Austin 1985<br />
Lottia instabilis Baxter 1987 I/ST/Epi II III IV Austin 1985<br />
Lottia ochracea UAM I/ST/Epi II III IV Baxter 1987<br />
Lottia pelta UAM I/ST/Epi II III IV Austin 1985<br />
Lottia triangularis Baxter 1987 I/ST/Epi II III IV Austin 1985<br />
Lottia digitalis UAM I/Epi II III IV Baxter 1987<br />
Tectura fenestrata UAM I/Epi II III IV Austin 1985<br />
Tectura persona UAM I/Epi II III IV Austin 1985<br />
Tectura scutum UAM I/ST/Epi II III IV Austin 1985<br />
Tectura testudinalis Baxter 1987 I/ST/Epi II x Austin 1985<br />
Boreotrophon clathratus UAM I/ST/Epi II x x x Baxter 1987<br />
Boreotrophon multicostata UAM I/ST/Epi II III IV<br />
Turgeon et al.<br />
1998<br />
Boreotrophon truncatus B. pacificus/ B. beringi UAM I/ST/Epi II x x x<br />
Turgeon et al.<br />
1998<br />
Nucella canaliculata UAM I/Epi II III IV Baxter 1987<br />
Nucella lamellosa UAM I/Epi II III IV Austin 1985<br />
Nucella lima UAM I/Epi II III IV x Baxter 1987<br />
Ocenebrina interfossa Ocenebra UAM I/Epi II III IV Baxter 1987<br />
Ocenebrina lurida Ocenebra UAM I/Epi II III IV Austin 1985<br />
Scabrotrophon maltzan Trophonopsis lasius UAM I/Epi II III IV Baxter 1987
Chapt 10. Biodiversity, page 10- 26<br />
Table 10.4 continued.<br />
BIOREGION<br />
GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Nassarius mendicus UAM I/ST/Inf II III IV Austin 1985<br />
Cryptonatica affinis<br />
Natica russa/ Natica<br />
aleutica/ Natica clausa UAM I/ST/Inf II III IV x x x Austin 1985<br />
Euspira pallida Polinices pallidus UAM I/ST/Inf II x x x Austin 1985<br />
Olea hansinensis Goddard 2000 ST/Epi II III IV<br />
British<br />
Columbia NR Behrens 1991<br />
Olivella baetica UAM I/ST/Inf II III IV Baxter 1987<br />
Onchidella<br />
borealis<br />
Lee and Foster<br />
1985 I/Epi II III IV<br />
Lee and Foster<br />
1985<br />
British<br />
Columbia NR Behrens 1991<br />
Lee and Foster<br />
1985<br />
Adalaria jannae Goddard 2000 ST/Epi II III IV<br />
Lee and Foster<br />
Adalaria<br />
proxima<br />
1985 ST/Epi II x x<br />
Adalaria sp. 1 <strong>of</strong> Behrens Goddard 2000 ST/Epi II III IV SE Alaska nr Behrens 1991<br />
Onchidoris<br />
bilamellata<br />
Lee and Foster<br />
1985 ST/Epi II x x<br />
Lee and Foster<br />
1985<br />
Lee and Foster<br />
1985<br />
Onchidoris muricata Goddard 2000 ST/Epi II x x<br />
Policera zosterae Goddard 2000 ST/Epi II III IV SE Alaska nr Behrens 1991<br />
Lee and Foster<br />
Lee and Foster<br />
Triopha catalinae Triophidae<br />
1985 ST/Epi II III IV x<br />
1985<br />
Cyclostremella concordia Cyclostremellidae UAM II III IV Baxter 1987<br />
Odostomia angularis UAM I/ST/Inf II III IV Austin 1985<br />
Odostomia arctica UAM I/ST/Inf II x x Arctic nr<br />
Turgeon et al.<br />
1998<br />
Odostomia iliuliukensis UAM I/ST/Inf II III IV Austin 1985<br />
Odostomia cassandra<br />
Feder and<br />
Matheke 1980 I/ST/Inf II III IV Austin 1985<br />
Odostomia pesa Baxter 1987 I/ST/Inf II III IV Baxter 1987<br />
Turbonilla canadensis Baxter 1987 I/ST/Inf II III IV Austin 1985<br />
Turbonilla eyerdami Baxter 1987 I/ST/Inf II III IV Austin 1985<br />
Turbonilla lordii Baxter 1987 I/ST/Inf II III IV Austin 1985<br />
Turbonilla lyalli Baxter 1987 I/ST/Inf II III IV Austin 1985<br />
Turbonilla middendorffi Baxter 1987 I/ST/Inf II III IV Austin 1985<br />
Turbonilla rinella Baxter 1987 I/ST/Inf II III IV Austin 1985<br />
Fusitriton oregonensis Cyamatiidae UAM I/ST/Epi II III IV Baxter 1987<br />
Retusa obtusa R. perteuis/ R. semen UAM I/ST/Inf II x Bering Sea nr<br />
Lee and Foster<br />
1985<br />
"Rissoella" alaskensis<br />
Feder and<br />
Bryson-Schwafel<br />
1988 I/ST/Inf II III IV Austin 1985<br />
Alvania compacta UAM I/ST/Inf II III IV Baxter 1987<br />
Alvania moerchi Baxter 1987 I/ST/Inf II x x x Baxter 1987<br />
Alvania rosana UAM I/ST/Inf II III IV Baxter 1987<br />
Feder and Bryson-<br />
Boreocingula martyni Cingula katherenae Schwafel 1988 I/ST/Inf II III IV Austin 1985<br />
Falsicingula aleutica Falsicingulidae UAM I/ST/Inf II III IV Baxter 1987<br />
Onoba dalli Alvania Baxter 1987 I/ST/Inf II III IV Baxter 1987<br />
Onoba cerinella Baxter 1987 I/ST/Inf II III IV Austin 1985<br />
Onoba kyskensis Baxter 1987 I/ST/Inf II III IV Austin 1985<br />
Onoba montereyensis Alvania carpenteri Baxter 1987 I/ST/Inf II III IV Austin 1985<br />
Acteocina culcitella Cylichnella EVOS I/ST/Inf II III IV Austin 1985<br />
Acteocina harpa A. oldroydi<br />
Lee and Foster<br />
1985 I/ST/Inf II III IV Behrens 1991<br />
Cylichna attonsa UAM I/ST/Inf II III IV Austin 1985<br />
Lee and Foster<br />
Cylichna occulta UAM I/ST/Inf II x<br />
1985<br />
Cylichna alba UAM I/ST/Inf II x x x Austin 1985<br />
Siphonaria thirsites UAM I/Epi II III IV Baxter 1987<br />
Hermaea vancouverensis Hermaeidae Foster 1987 ST/Epi II III IV Behrens 1991<br />
Cuthona albocrusta Goddard 2000 ST/Epi II III IV Washington NR Goddard 2000<br />
Cuthona pustulata Goddard 2000 ST/Epi II III IV<br />
British<br />
Columbia NR Goddard 2000<br />
Melibe<br />
leonina<br />
Lee and Foster<br />
1985 P II III IV<br />
Lee and Foster<br />
1985
Chapt 10. Biodiversity, page 10- 27<br />
Table 10.4 continued.<br />
GENUS SPECIES OTHER NAMES<br />
Tritonia<br />
Tritonia<br />
diomedea<br />
festiva<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Lee and Foster<br />
1985 ST/Epi II III IV x<br />
Lee and Foster<br />
1985 ST/Epi II III IV<br />
Lee and Foster<br />
1985 ST/Epi II III IV x<br />
Lee and Foster<br />
1985<br />
Lee and Foster<br />
1985<br />
Lee and Foster<br />
1985<br />
Trochuina tetraquetra<br />
Cidarina cidaris UAM ST/Epi II III IV Baxter 1987<br />
Lirularia succincta UAM I/Inf II III IV Austin 1985<br />
Margarites beringensis UAM I/Inf II III IV Baxter 1987<br />
Margarites helicinus UAM ST/Inf II x x x Austin 1985<br />
Margarites pupillus UAM ST/Inf II III IV Austin 1985<br />
Solariella varicosa UAM ST/Inf II x Baxter 1987<br />
Solariella obscura UAM ST/Inf II x Austin 1985<br />
Spiromoellaria kachemakens UAM I/ST/Epi II III IV Baxter 1987<br />
Aforia circinata UAM ST/Inf II III IV Baxter 1987<br />
Tachyrhynchus erosus Baxter 1987 ST/Inf II x Austin 1985<br />
Tachyrhynchus reticulatus Baxter 1987 ST/Inf II III IV Austin 1985<br />
Velutina plicatilis Baxter 1987 ST/Epi II x Austin 1985<br />
Velutina prolongata Baxter 1987 ST/Epi II III IV Austin 1985<br />
Velutina rubra Foster 2000 I/Epi II III IV Austin 1985<br />
Velutina undata Baxter 1987 ST/Epi II x x x Austin 1985<br />
Volumitra alaskana UAM ST/Epi II III IV Baxter 1987<br />
Arctomelon stearnsii UAM ST/Epi II III IV Austin 1985<br />
Janolus fuscus Goddard 2000 ST/Epi II III IV SE Alaska nr Goddard 2000<br />
Turgeon et al.<br />
Clio pyramidata Cooney 1987 P x x x<br />
1998<br />
Cryptochiton stelleri UAM I/ST/Epi II III IV Baxter 1987<br />
Lepidozona interstincta Ischnochiton UAM ST/Epi II III IV Baxter 1987<br />
Stenonemus albus Ischnochiton UAM ST/Epi II x Baxter 1987<br />
Leptochiton rugatus UAM I/ST/Epi II III IV x Austin 1985<br />
Katherina tunicata UAM I/Epi II III IV Baxter 1987<br />
Mopalia laevior UAM I/ST/Epi II III IV Austin 1985<br />
Mopalia lignosa UAM I/ST/Epi II III IV Baxter 1987<br />
Mopalia ciliata UAM I/ST/Epi II III IV Baxter 1987<br />
Mopalia cirrata UAM I/ST/Epi II III IV Baxter 1987<br />
Mopalia mucosa UAM I/ST/Epi II III IV Baxter 1987<br />
Mopalia spectabilis Baxter 1987 I/ST/Epi II III IV Baxter 1987<br />
Mopalia swanii UAM I/ST/Epi II III IV Austin 1985<br />
Placiphorella rufa UAM I/ST/Epi II III IV Austin 1985<br />
Schizoplax brandtii UAM I/Epi II III IV x Austin 1985<br />
Tonicella insignis Ischnochitonidae Foster 2000 I/ST/Epi II III IV Baxter 1987<br />
Tonicella lineata Ischnochitonidae UAM I/ST/Epi II III IV Baxter 1987<br />
Tonicella rubra Ischnochitonidae UAM I/ST/Epi II x Baxter 1987<br />
Rhabdus rectius Dentalium dalli UAM ST/Inf II III IV Baxter 1987<br />
Pulsellum salishorum UAM ST/Inf II III IV Baxter 1987<br />
Polyschides tolmei Cadulus stearnsi UAM ST/Inf II III IV Baxter 1987
Chapt 10. Biodiversity, page 10- 28<br />
Arthropoda<br />
Chelicerata and Pycnogonida are relatively scarce in Prince William Sound compared to<br />
Crustacea. Seven <strong>of</strong> the ten pycnogonid species are included in the data set based on UAM<br />
specimens, One Acarina and one pseudoscorpion are frequently observed in the high intertidal<br />
zone <strong>of</strong> southeastern and south-central Alaska.<br />
Crustacea make up about 28% <strong>of</strong> the animal species listed in the data set (Table 10.5).<br />
We list 172 copepods, (including planktonic species from the Gulf <strong>of</strong> Alaska which may be<br />
present occasionally in Prince William Sound), 105 decapods, 105 amphipods, 21 cumacean, 19<br />
isopods, and smaller numbers <strong>of</strong> others.<br />
Several taxa (e.g. Ostracoda) are probably very much under-counted. Six copepods, one<br />
amphipod, and two ostracods are identified to the family level, and no further. A large number<br />
<strong>of</strong> small crustacea, mostly harpacticoid copepods and amphipods have not been identified to<br />
species. Two species, the amphipod Jassa sp./marmorata and the copepod Leimia vaga are<br />
possible NIS, but this number probably represents an underestimate. Cordell (2000) pointed out<br />
that “Harpacticoid copepods may be particularly likely to be transported and introduced because<br />
as a group they have successfully occupied almost all benthic and epibenthic habitats. … The<br />
paucity <strong>of</strong> studies <strong>of</strong> harpacticoid taxonomy in the northeastern Pacific makes it nearly<br />
impossible to determine whether or not a given species has been introduced without extensive<br />
distribution or genetic studies.” We have not included an inventory <strong>of</strong> parasitic cirripedia or<br />
isopods, but several species <strong>of</strong> these were recorded by Hines et al. (2000).<br />
Sources for information <strong>of</strong> Crustacea include Barnard 1969 (amphipods), Butler 1980<br />
(shrimp), Hart 1982 (crabs), and Schultz 1969 and Squires and Figueria 1974 (isopods). J.<br />
Chapman (2000) contributed to the information on Pericarida, and J. Cordell (2000) to our<br />
knowledge <strong>of</strong> the harpacticoid Copepoda. Records for the planktonic crustacea derive from<br />
unpublished data compiled from T. Cooney’s Sound Ecosystem Analysis project <strong>of</strong> 1994-1998.<br />
We found eleven new records for the occurrence <strong>of</strong> Crustacea.<br />
The third major group <strong>of</strong> Arthropoda, marine and brackish water insects, have not been<br />
surveyed in any detail in the area. For insects, Austin (1985) lists eight Coleoptera, and 12<br />
Diptera with ranges that include Alaska.
Chapt 10. Biodiversity, page 10- 29<br />
Table 10.5. Arthropoda.<br />
BIOREGION<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Arachnida: Acarina Nemolgus littoralis<br />
Foster/Hobergobservation<br />
I/Epi<br />
Arachnida:<br />
Pseudoscorpionida unidentified unidentified UAM I/epi<br />
Crustacea: Amphipoda Acanthonotozomatidae Odius carinatus EVOS ST/Epi II x Austin 1985<br />
Crustacea: Amphipoda Ampeliscidae Ampelisca birulai<br />
Feder and<br />
Matheke 1980 I/ST/Inf II III IV x x<br />
Coyle and Highsmith<br />
1994<br />
Crustacea: Amphipoda Ampeliscidae Ampelisca brevisimulata EVOS ST II III IV Austin 1985<br />
Crustacea: Amphipoda Ampeliscidae Ampelisca caryei EVOS ST II III IV Austin 1985<br />
Crustacea: Amphipoda Ampeliscidae Ampelisca pugetica EVOS ST II III IV Austin 1985<br />
Crustacea: Amphipoda Ampeliscidae Ampelisca eschrichti<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV x Austin 1985<br />
Feder and<br />
Crustacea: Amphipoda Ampeliscidae Ampelisca macrocephala A. careyi<br />
Matheke 1980 ST/Inf II III IV Austin 1985<br />
Feder and<br />
Crustacea: Amphipoda Ampeliscidae Haploops tubicola<br />
Matheke 1980 ST/Inf II x x Austin 1985<br />
Crustacea: Amphipoda Amphitoidae Ampithoe kussakini Chapman 2000 I/Inf II III IV x Chapman 2000<br />
Crustacea: Amphipoda Amphitoidae Ampithoe sectimanus Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Amphitoidae Ampithoe simulans<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST x x x x Austin 1985<br />
Crustacea: Amphipoda Amphitoidae Peramphithoe eoa Ampithoe mea Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Amphitoidae Peramphithoe humeralis Ampithoe Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Anisogammaridae Anisogammarus pugettensis EVOS ST/Epi II III IV x Chapman 2000<br />
Crustacea: Amphipoda Anisogammaridae Eogammarus confervicolus Anisogammarus Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Anisogammaridae Eogammarus oclairi Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Anisogammaridae Eogammarus unidentified Chapman 2000 I/ST<br />
Crustacea: Amphipoda Anisogammaridae Locustogammarus locustoides Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Anisogammaridae Spinulogammarus subcarinatus Chapman 2000 I II III IV Chapman 2000<br />
Crustacea: Amphipoda Aoridae Aoroides columbiae EVOS ST II III IV x Austin 1985<br />
Crustacea: Amphipoda Atylidae Atylus collingi EVOS ST II III IV x Austin 1985<br />
Crustacea: Amphipoda Calliopiidae Calliopiella unidentified Paracalliopiella Chapman 2000 I/Inf
Chapt 10. Biodiversity, page 10- 30<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Amphipoda Calliopiidae Calliopius behringi Cooney 1987 I/ST II III x x x Gur’yanova 1951, 1962<br />
Crustacea: Amphipoda Calliopiidae Calliopius carinatus Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Calliopiidae Calliopius laeviuscula Cooney 1987 P III IV V VI x x Austin 1985<br />
Crustacea: Amphipoda Caprellidae Caprella drepanochir Chapman 2000 I/Inf II III IV x C Chapman 2000<br />
Crustacea: Amphipoda Caprellidae Caprella laeviuscula Chapman 2000 I/Inf II III IV x Chapman 2000<br />
Crustacea: Amphipoda Caprellidae Caprella unidentified Chapman 2000 I/Inf<br />
Crustacea: Amphipoda Caprellidae Metacaprella kennerlyi Chapman 2000 I/Inf II III IV Chapman 2000<br />
Feder and<br />
Crustacea: Amphipoda Corophidae Neohela unidentified<br />
Matheke 1980 ST/Inf<br />
Crustacea: Amphipoda Corophidae Americorophium brevis Corophium Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Corophidae Americorophium salmonis Corophium Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Corophidae Americorophium spinicore Corophium Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Corophidae Corophium unidentified EVOS ST<br />
Crustacea: Amphipoda Corophidae Monocorophium carlottensis Chapman 2000 I/Inf II III IV C Chapman 2000<br />
Crustacea: Amphipoda Dexamidae Guernea unidentified EVOS ST<br />
Crustacea: Amphipoda Eusiridae Eusiriella multicalceola Gracilipes Cooney 1987 P II III Austin 1985<br />
Crustacea: Amphipoda Eusiridae Pontogeneia unidentified EVOS ST<br />
Crustacea: Amphipoda Eusiridae Rhachotropis natator Cooney 1987 P II III IV V x Austin 1985<br />
Crustacea: Amphipoda Gammaridae Lagunogammarus setosus Chapman 2000 I/Inf II III IV x Chapman 2000<br />
Feder and<br />
Crustacea: Amphipoda Haustoriidae Ericthonius hunteri E. rubricornis<br />
Matheke 1980 I/ST/Inf II x x Austin 1985<br />
Crustacea: Amphipoda Hyalellidae Najna unidentified Najnidae EVOS I/St<br />
Crustacea: Amphipoda Hyalidae Allorchestes angusta Chapman 2000 I/Inf II III IV x Chapman 2000<br />
Crustacea: Amphipoda Hyalidae Hyale frequena Chapman 2000 I/Inf II III IV Chapman 2000
Chapt 10. Biodiversity, page 10- 31<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Amphipoda Hyalidae Hyale plumulosa Chapman 2000 I/Inf II x x C Chapman 2000<br />
Crustacea: Amphipoda Hyperiidae Euprimno macropa Primno abyssalis Cooney: SEA P<br />
II III IV V<br />
VI VII VIII<br />
IX x x Austin 1985<br />
Crustacea: Amphipoda Hyperiidae Euprimno unidentified Primno Cooney 1987 P<br />
Crustacea: Amphipoda Hyperiidae Hyperia medusarum hystryx Cooney 1987 P<br />
Crustacea: Amphipoda Hyperiidae Hyperia unidentified Cooney 1987 P<br />
Crustacea: Amphipoda Hyperiidae Hyperoche medusarum Cooney 1987 P<br />
II III IV V<br />
VI x Austin 1985<br />
II III IV V<br />
VI x Austin 1985<br />
Crustacea: Amphipoda Hyperiidae Hyperoche unidentified Cooney 1987 P<br />
Crustacea: Amphipoda Hyperiidae Parathemisto gracilipes Thermisto Cooney 1987 P II III x x Austin 1985<br />
Crustacea: Amphipoda Hyperiidae Parathemisto libellula Thermisto Cooney 1987 P II III IV V x Austin 1985<br />
Crustacea: Amphipoda Hyperiidae Parathemisto pacifica Thermisto Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Amphipoda Isaeidae Photis unidentified EVOS ST/Inf<br />
Crustacea: Amphipoda Ischyroceridae Ischyrocerus unidentified EVOS I/ST/Inf<br />
Crustacea: Amphipoda Ischyroceridae Jassa sp./J. marmorata Chapman 2000 I/Inf II possible Austin 1985<br />
Crustacea: Amphipoda Ischyroceridae Jassa staudel Chapman 2000 I/Inf II III IV C Chapman 2000<br />
Feder and<br />
Crustacea: Amphipoda Ischyroceridae Protomedeia unidentified<br />
Matheke 1980 I/ST/Inf<br />
Crustacea: Amphipoda Lanceolidae Lanceola pacifica Cooney 1987 P II III x x Gur’yanova 1951, 1962<br />
Crustacea: Amphipoda Lycaeidae Tryphana malmii Cooney 1987 P II III IV V Austin 1985<br />
Crustacea: Amphipoda Lysianassidae Andaniexis subabyssis Cooney 1987 P II III IV x Gur’yanova 1951, 1962<br />
Crustacea: Amphipoda Lysianassidae Anonyx unidentified<br />
Feder and<br />
Matheke 1980 I/ST/Inf<br />
Feder and<br />
Crustacea: Amphipoda Lysianassidae Crybelocephalus unidentified<br />
Matheke 1980 P x x x x<br />
Crustacea: Amphipoda Lysianassidae Cyphocaris anonyx C. micronyx Cooney 1987 P<br />
II III IV V<br />
VI VII VIII x Austin 1985
Chapt 10. Biodiversity, page 10- 32<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
Crustacea: Amphipoda Lysianassidae Cyphocaris challengeri<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Cooney and Coyle<br />
1988 S/P x x x x Austin 1985<br />
Crustacea: Amphipoda Lysianassidae Hippomedon kurilicus H. abyssi UAM ST II III IV x Gur’yanova 1951, 1962<br />
Crustacea: Amphipoda Lysianassidae Koroga megalops Cooney 1987 P II III x x x Austin 1985<br />
Crustacea: Amphipoda Lysianassidae Lepidepecerium unidentified EVOS ST<br />
Crustacea: Amphipoda Lysianassidae Onismius unidentified EVOS ST<br />
Crustacea: Amphipoda Lysianassidae Orchomene obtusa Cooney: SEA ST<br />
II III IV V<br />
VI VII Gur’yanova 1951, 1962<br />
Crustacea: Amphipoda Lysianassidae Orchomene pacifica EVOS ST II III IV x Austin 1985<br />
Crustacea: Amphipoda Lysianassidae Orchomene unidentified EVOS ST<br />
Crustacea: Amphipoda Lysianassidae Paracallisoma alberti Cooney 1987 P II III x Gur’yanova 1951, 1962<br />
Feder and<br />
Crustacea: Amphipoda Melitidae Maera loveni M.danae<br />
Matheke 1980 ST/Inf II x x Austin 1985<br />
Crustacea: Amphipoda Melitidae Melita dentata EVOS ST II III IV x Austin 1985<br />
Crustacea: Amphipoda Melphidippidae unidentified unidentified Cooney: SEA ST II III x NE Pac NR Austin 1985<br />
Feder and<br />
Crustacea: Amphipoda Oedicerotidae Aceroides latipes<br />
Matheke 1980 I/ST/Inf II III IV x Austin 1985<br />
Crustacea: Amphipoda Oedicerotidae Bathymedon unidentified EVOS ST/Inf<br />
Feder and<br />
Crustacea: Amphipoda Oedicerotidae Monoculodes diamesus<br />
Matheke 1980 I/ST/Inf II III IV Gur’yanova 1951, 1962<br />
Crustacea: Amphipoda Oedicerotidae Synchelidium rectipalmatum EVOS ST/Inf II III IV Austin 1985<br />
Crustacea: Amphipoda Oedicerotidae Westwoodillia rectangulata W. caecula Cooney 1987 P II x Gur’yanova 1951, 1962<br />
Crustacea: Amphipoda Oxycephalidae Streesia unidentified Cooney 1987 P<br />
Crustacea: Amphipoda Paraphronimidae Paraphronima crassipes Cooney 1987 P<br />
II III IV V<br />
VI VII x Austin 1985<br />
Feder and<br />
Crustacea: Amphipoda Pardaliscidae Nicippe tumida<br />
Matheke 1980 I/ST/Inf II III IV x Austin 1985<br />
Feder and<br />
Crustacea: Amphipoda Phoxocephalidae Harpiniopsis sanpedroensis Harpinia<br />
Matheke 1980 I/ST/Inf II III IV Austin 1985<br />
Crustacea: Amphipoda Phoxocephalidae Paraphoxus homilis EVOS ST/Inf II III IV Austin 1985<br />
Crustacea: Amphipoda Phoxocephalidae Paraphoxus similis EVOS ST/Inf II III IV Austin 1985<br />
II III IV V<br />
Crustacea: Amphipoda Phronimidae Phronima sedentaria Cooney 1987 P<br />
VI x Austin 1985
Chapt 10. Biodiversity, page 10- 33<br />
Table 10.5 continued.<br />
BIOREGION<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
II III IV V<br />
Crustacea: Amphipoda Phrosinidae Primno macropa P. abyssalis Cooney 1987 P<br />
VI x x Austin 1985<br />
Crustacea: Amphipoda Pleustidae Pleustes cataphractus EVOS I/Inf x x x Austin 1985<br />
Crustacea: Amphipoda Pontogeneiidae Megamorea subiener Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Pontogeneiidae Paramoera bousfieldi Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Pontogeneiidae Paramoera columbiana<br />
Feder and Bryson-<br />
Schwafel 1988 I/Inf II III IV Austin 1985<br />
Crustacea: Amphipoda Pontogeneiidae Paramoera mohri Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Pontogeneiidae Pontogeneia ivanovae P. ivanovi Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Pontogeneiidae Pontogeneia rostrata Chapman 2000 I/Inf II III IV C Chapman 2000<br />
Crustacea: Amphipoda Pontoporeiidae Pontoporeia femorata Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Amphipoda Proscinidae Proscina birsteini Cooney 1987 P II x Gur’yanova 1951, 1962<br />
II III IV V<br />
Crustacea: Amphipoda Scinidae Scina borealis UAM I/ST/Inf VI x x Austin 1985<br />
Crustacea: Amphipoda Scinidae Scina rattayi Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Amphipoda Scinidae Scina unidentified Cooney: SEA P<br />
Crustacea: Amphipoda Scinidae Scina stebbingi Cooney 1987 P II x Gur’yanova 1951, 1962<br />
Feder and<br />
Crustacea: Amphipoda Stenothoidae Metopa unidentified<br />
Matheke 1980 I/ST/Inf<br />
Crustacea: Amphipoda Stenothoidae Metopelloides erythrophthalmus EVOS II III IV Austin 1985<br />
Feder and<br />
Crustacea: Amphipoda Synopiidae Syrrhoe crenulata<br />
Matheke 1980 I/ST/Inf II x x Austin 1985<br />
Crustacea: Amphipoda Synopiidae Tiron biocellata EVOS II III IV Austin 1985<br />
Crustacea: Amphipoda Talitridae Talitrus unidentified<br />
Crustacea: Amphipoda Urothoidae Urothoe denticulata<br />
Feder and Bryson-<br />
Schwafel 1988 I/Inf<br />
Feder and<br />
Matheke 1980 I/ST/Inf II III IV Gur’yanova 1951, 1962<br />
Crustacea: Amphipoda Vibiliidae Viblia australis Cooney 1987 P II III IV x Austin 1985<br />
II III IV V<br />
Crustacea: Cladocera Podonidae Evadne nordmanni Cooney 1987 P<br />
VI x x x Austin 1985<br />
Crustacea: Cladocera Podonidae Evadne unidentified Cooney 1987 P
Chapt 10. Biodiversity, page 10- 34<br />
Table 10.5 continued.<br />
BIOREGION<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Cladocera Podonidae Evadne tergistina Pseudevadne Cooney 1987 P<br />
II III IV V<br />
VI Austin 1985<br />
Crustacea: Cladocera Podonidae Podon leuckarti Cooney 1987 P II III IV x x Austin 1985<br />
II III IV V<br />
Crustacea: Cladocera Podonidae Podon polyphmoies Cooney 1987 P<br />
VI x x x Austin 1985<br />
Crustacea: Cladocera Podonidae Podon unidentified Cooney 1987 P<br />
Crustacea: Copepoda:<br />
Calanoida Acartiidae Acartia cf. A. clausi Cordell 2000 P II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Calanoida Acartiidae Acartia longiremis<br />
Crustacea: Copepoda:<br />
Calanoida Acartiidae Acartia unidentified Cooney: SEA P<br />
Crustacea: Copepoda:<br />
Calanoida Acartiidae Acartia tumida<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Aetideus divergens A. armatus Cooney 1987 P<br />
Cooney and Coyle<br />
1988 P II x x Austin 1985<br />
Cooney and Coyle<br />
1988 P II III IV x Austin 1985<br />
II III IV V<br />
VI Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Aetideus pacificus Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Aetideus unidentified Cooney 1987 P<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Bradyidius saanichi Cooney 1987 P II III IV Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Chiridiella unidentified<br />
Cooney and Coyle<br />
1988 P<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Chiridius gracilis Cooney 1987 P II III x x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Chiridius poppei Cooney 1987 P II x Brodsky 1950<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Chiridius unidentified<br />
Cooney and Coyle<br />
1988 P<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Gaetanus intermedius Gaidius Cooney 1987 P II III IV Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Gaidius tenuispinus G. minutus Cooney 1987 P II III IV Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Gaidius variabilis Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Pseudochirella sarsi Euchaetidae: Euchaeta Cooney 1987 P II III IV Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae Pseudochirella unidentified Cooney 1987 P<br />
Crustacea: Copepoda:<br />
Calanoida Aetideidae unidentified unidentified Cooney: SEA P<br />
Crustacea: Copepoda:<br />
Calanoida Augaptilidae Haloptilus pseudooxycephalus Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Augaptilidae Pachyptilis pacificus Cooney 1987 P II III x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Calanidae Calanus glacialis E. marshallae Cooney 1987 P II III IV x Austin 1985
Chapt 10. Biodiversity, page 10- 35<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
Crustacea: Copepoda:<br />
Calanoida Calanidae Calanus marshallae C. glacialis<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Cooney and Coyle<br />
1988 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Calanidae Calanus pacificus Cooney 1987 P II III IV Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Calanidae Neocalanus cristatus Cooney 1987 P II III IV V x x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Calanidae Neocalanus flemingeri Cooney: SEA P II III IV Miller 1988<br />
Crustacea: Copepoda:<br />
Calanoida Calanidae Neocalanus plumchrus<br />
Cooney and Coyle<br />
1988 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Calanidae Neocalanus unidentified Cooney: SEA P<br />
Crustacea: Copepoda:<br />
Calanoida Candaciidae Candacia bipinnata Cooney: SEA P II III IV V x x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Candaciidae Candacia columbiae<br />
Crustacea: Copepoda:<br />
Calanoida Centropagidae Centropages abdominalis<br />
Cooney and Coyle<br />
1988 P II III IV x Cooney 1987<br />
Cooney and Coyle<br />
1988 P II III IV x Cordell 2000<br />
Crustacea: Copepoda:<br />
Calanoida Clausocalanidae Clausocalanus arcuicornis Cooney 1987 P II III IV Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Clausocalanidae Clausocalanus unidentified Cooney: SEA P<br />
Crustacea: Copepoda:<br />
Calanoida Clausocalanidae Mesocalanus tenuicornis Cooney 1987 P II x x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Clausocalanidae Microcalanus unidentified Cooney 1987 P<br />
Crustacea: Copepoda:<br />
Calanoida Clausocalanidae Pseudocalanus unidentified Cooney: SEA P<br />
Crustacea: Copepoda:<br />
Calanoida Corycaeidae Corycaeus anglicus Cooney 1987 P II III IV V x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Corycaeidae Corycaeus unidentified Cooney: SEA P<br />
Crustacea: Copepoda:<br />
Calanoida Ectinosomatidae Microsetella rosea Cooney 1987 P<br />
Crustacea: Copepoda:<br />
Calanoida Ectinosomatidae Microsetella unidentified Cooney: SEA P<br />
Crustacea: Copepoda:<br />
Calanoida Eucalanidae Eucalanus bungii<br />
Crustacea: Copepoda:<br />
Calanoida Euchaetidae Euchaete elongata<br />
II III VI V<br />
VI x x Austin 1985<br />
Cooney and Coyle<br />
1988 P II III IV Austin 1985<br />
Cooney and Coyle<br />
1988 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Heterhabdidae Heterorhabdus compactus Cooney 1987 P II x x Brodsky 1950<br />
Crustacea: Copepoda:<br />
Calanoida Heterhabdidae Heterorhabdus robustoides Cooney 1987 P II III x<br />
Gardner and Szabo<br />
1982<br />
Crustacea: Copepoda:<br />
Calanoida Heterhabdidae Heterorhabdus tanneri Cooney 1987 P II III IV x x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Heterhabdidae Heterostylites longicornis Cooney: SEA P III VI V VI x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Heterhabdidae Heterostylites major Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Lucicutiidae Lucicutia flavicornis Cooney 1987 P<br />
II III IV V<br />
VI x Austin 1985
Chapt 10. Biodiversity, page 10- 36<br />
Table 10.5 continued.<br />
BIOREGION<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Copepoda:<br />
Calanoida Lucicutiidae Lucicutia ovalis Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Lucicutiidae Lucicutia unidentified Cooney: SEA P<br />
Crustacea: Copepoda:<br />
Calanoida Metridiidae Metridia curticauda<br />
Crustacea: Copepoda:<br />
Calanoida Metridiidae Metridia okhotensis M. longa<br />
Crustacea: Copepoda:<br />
Calanoida Metridiidae Metridia pacifica<br />
Crustacea: Copepoda:<br />
Calanoida Metridiidae Metridia princeps<br />
Crustacea: Copepoda:<br />
Calanoida Metridiidae Metridia unidentified<br />
Cooney and Coyle<br />
1988 P II III IV x Austin 1985<br />
Cooney and Coyle<br />
1988 P II III IV x Austin 1985<br />
Cooney and Coyle<br />
1988 P II III IV x Austin 1985<br />
Cooney and Coyle<br />
1988 P<br />
Cooney and Coyle<br />
1988 P<br />
II III IV V<br />
VI x x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Metridiidae Pleuromamma robusta Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Metridiidae Pleuromamma scutullata Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Monstrillidae Cymbasoma rigidum Cooney 1987 P II III x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Monstrillidae Monstrilla canadensis Cooney 1987 P II III IV x<br />
Crustacea: Copepoda:<br />
Calanoida Monstrillidae Monstrilla helgolandica Cooney 1987 P II III IV x<br />
Crustacea: Copepoda:<br />
Calanoida Monstrillidae Monstrilla longiremus Cooney 1987 P II III IV<br />
Crustacea: Copepoda:<br />
Calanoida Monstrillidae Monstrilla unidentified Cooney: SEA P<br />
Crustacea: Copepoda:<br />
Calanoida Monstrillidae Monstrilla wandlii Cooney 1987 P II III IV x<br />
Crustacea: Copepoda:<br />
Calanoida Paracalanidae Paracalanus parvus Cooney 1987 P<br />
Crustacea: Copepoda:<br />
Calanoida Paracalanidae Paracalanus unidentified Cordell 2000 P<br />
Crustacea: Copepoda:<br />
Calanoida Phaennidae Gaetanus unidentified<br />
Cooney and Coyle<br />
1988 P<br />
Gardner and Szabo<br />
1982<br />
Gardner and Szabo<br />
1982<br />
Gardner and Szabo<br />
1982<br />
Gardner and Szabo<br />
1982<br />
II III IV V<br />
VI x x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Pontellidae Epilabidocera longipedata E. amphitrites Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Pseudocalanidae Pseudocalanus unidentified Cordell 2000 P<br />
Crustacea: Copepoda:<br />
Calanoida Scolecitrichidae Lophothrix frontalis Cooney 1987 P<br />
II III IV V<br />
VI x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Scolecitrichidae Racovitzanus antarcticus Cooney 1987 P III IV x x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Scolecitrichidae Scaphocalanus brevicornis Cooney 1987 P II III IV x x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Scolecitrichidae Scaphocalanus magnus Cooney 1987 P II III IV x x Austin 1985
Chapt 10. Biodiversity, page 10- 37<br />
Table 10.5 continued.<br />
BIOREGION<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Copepoda:<br />
Calanoida Scolecitrichidae Scolecithricella minor Cooney 1987 P II x x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Scolecitrichidae Scolecithricella ovata S. subdentata Cooney: SEA P<br />
II III IV V<br />
VI Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Spinocalanidae Spinocalanus brevicaudatus Cooney 1987 P II III x x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Temoridae Eurytemora americana Cooney 1987 P II III x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Temoridae Eurytemora herdmani E. pacifica Cordell 2000 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Calanoida Temoridae Eurytemora unidentified Cooney 1987 P<br />
Crustacea: Copepoda:<br />
Calanoida Tharydidae Undinella unidentified Cooney 1987 P<br />
Crustacea: Copepoda:<br />
Calanoida Tortanidae Tortanus discaudata<br />
Crustacea: Copepoda:<br />
Calanoida Amallothryx inornata P<br />
Crustacea: Copepoda:<br />
Cyclopoida Cyclopidae Euryte unidentified Cordell 2000 P<br />
Cooney and Coyle<br />
1988 P II x x Austin 1985<br />
Crustacea: Copepoda:<br />
Cyclopoida Cyclopidae Halicyclops unidentified Cordell 2000 P<br />
Crustacea: Copepoda:<br />
Cyclopoida Cyclopinidae Cyclopina unidentified Cooney 1987 P<br />
Crustacea: Copepoda:<br />
Cyclopoida Cyclopinidae unidentified unidentified Cordell 2000 P<br />
Crustacea: Copepoda:<br />
Cyclopoida Oithonidae Oithona similis Cordell 2000 P x x x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Cyclopoida Oithonidae Oithona unidentified Cooney: SEA P<br />
Crustacea: Copepoda:<br />
Cyclopoida Oithonidae Oithona spinirostris Cordell 2000 P x x x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Ameiridae Ameira longipes Cordell 2000 I/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Ameiridae Ameira sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Ameiridae Ameira sp. 2 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Ameiridae unidentified sp. 1 Cordell 2000 I/Inf
Chapt 10. Biodiversity, page 10- 38<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Copepoda:<br />
Harpacticoida Ancorabilidae Arthropsyllus serratus Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Canthocamptidae Mesochra pygmaea<br />
Feder and Bryson-<br />
Schwafel 1988 I/Inf II x x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Canthocamptidae Mesochra sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Canthocamptidae unidentified incertae sedis Cordell 2000 II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Cletodidae Acrenhydrosoma unidentified Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Cletodidae Leimia vaga Cordell 2000 I/Inf II x x possible Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Cletodidae Nannopus palustris<br />
Crustacea: Copepoda:<br />
Harpacticoida Cletodidae Rhizothrix unidentified<br />
Crustacea: Copepoda:<br />
Harpacticoida Danielsseniidae Danielsennia typica<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Inf II x x Austin 1985<br />
Feder and Bryson-<br />
Schwafel 1988<br />
I/ST/Inf<br />
Feder and Bryson-<br />
Schwafel 1988 I/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Amonardia normani Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Amonardia perturbata Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Amphiascoides cf. D. debilis Cordell 2000 I/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Amphiascoides sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Amphiascopis cinctus Cordell 2000 I/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Amphiascus minutus Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Amphiascus sp. 1 Cordell 2000 I/Inf
Chapt 10. Biodiversity, page 10- 39<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Diosaccus sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Diosaccus spinatus Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Robertsonia unidentified Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Stenhelia peniculata Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Diosaccidae Stenhelia unidentified<br />
Feder and Bryson-<br />
Schwafel 1988<br />
I/ST/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Ectinosomatidae Ectinosoma unidentified Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Ectinosomatidae Halectinosoma finmarchium<br />
Crustacea: Copepoda:<br />
Harpacticoida Ectinosomatidae Halectinosoma gothiceps<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Inf II x x Austin 1985<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Inf II x x Austin 1985<br />
Crustacea: Copepoda:<br />
Harpacticoida Ectinosomatidae Halectinosoma sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Ectinosomatidae Halectinosoma sp. 2 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Ectinosomatidae Halectinosoma sp. 3 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Ectinosomatidae Microarthridion norvegica Cordell 2000 I/Inf x x x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Harpacticidae Harpacticus compressus Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Harpacticidae Harpacticus septentrionalis Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Harpacticidae Harpacticus<br />
sp.- obscurus group<br />
1 Cordell 2000 I/Inf II III IV Cordell 2000
Chapt 10. Biodiversity, page 10- 40<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Copepoda:<br />
Harpacticoida Harpacticidae Harpacticus<br />
Crustacea: Copepoda:<br />
Harpacticoida Harpacticidae Harpacticus<br />
sp.- obscurus group<br />
2 Cordell 2000 I/Inf II III IV Cordell 2000<br />
sp.- uniremis group<br />
1 Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Harpacticidae Harpacticus superflexus<br />
Crustacea: Copepoda:<br />
Harpacticoida Harpacticidae Harpacticus uniremis<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Inf II III Cooney 1987<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Harpacticidae Zaus unidentified Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Echinolaophonte unidentified Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Heterolaophonte discophora Cordell 2000 I/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Heterolaophonte longisetigera Cordell 2000 I/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Heterolaophonte sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Heterolaophonte variabilis Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Laophonte applanata Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Laophonte elongata Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Laophonte sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Paralaophonte cf. L. congenera Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Paralaophonte hyperborea Cordell 2000 I/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Paralaophonte pacifica Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Paralaophonte perplexa<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV x Cordell 2000
Chapt 10. Biodiversity, page 10- 41<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Paralaophonte sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Laophontidae Pseudonychocamptus spinifer Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Longipediidae Longipedia unidentified Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Parastenheliidae Parastenhelia sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Parastenheliidae Parastenhelia sp. 2 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Tachidiidae Microarthridion littorale<br />
Feder and Bryson-<br />
Schwafel 1988 I/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Tegastidae Tegestes sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Tegastidae Tegestes sp. 2 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Dactylopausia cf. D. glacialis Dactylopodia Cordell 2000 I/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Dactylopausia glacialis Dactylopodia Cordell 2000 I/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Dactylopausia paratisboides Dactylopodia Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Dactylopausia sp. 1 Dactylopodia Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Dactylopausia vulgaris Dactylopodia Cordell 2000 I/Inf II x x Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Diarthrodes sp. 2 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Idomene unidentified Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Paradactylopodia latipes<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV Austin 1985
Chapt 10. Biodiversity, page 10- 42<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Paradactylopodia latipes<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV Austin 1985<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Paradactylopodia sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Parathalestris sp. 1 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Parathalestris sp. 2 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Thalestridae Parathalestris sp. 3 Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Harpacticoida Tisbidae Scutellidium arthuri Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Tisbidae Tisbe cf. T. furcata Cordell 2000 I/Inf II III IV Cordell 2000<br />
Crustacea: Copepoda:<br />
Harpacticoida Tisbidae Tisbe gracilis Cooney 1987 I/Inf Cooney 1987<br />
Crustacea: Copepoda:<br />
Harpacticoida Tisbidae Tisbe inflata <br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Inf II III IV Austin 1985<br />
Crustacea: Copepoda:<br />
Harpacticoida Tisbidae Tisbe unidentified Cordell 2000 I/Inf<br />
Crustacea: Copepoda:<br />
Poecilostomatoida Oncaeidae Lubbockia wilsonae Cooney 1987 P II III x Austin 1985<br />
Crustacea: Copepoda:<br />
Poecilostomatoida Oncaeidae Oncaea borealis Copepoda: Cyclopoida Cooney 1987 P<br />
II III IV V<br />
VI x Austin 1985<br />
Crustacea: Copepoda:<br />
Poecilostomatoida Oncaeidae Oncaea conifera Cooney 1987 P II III IV x x x Austin 1985<br />
Crustacea: Copepoda:<br />
Poecilostomatoida Oncaeidae Oncaea prolata O. notopus Cooney 1987 P II III IV Austin 1985<br />
Crustacea: Copepoda:<br />
Poecilostomatoida Oncaeidae Oncaea unidentified Cooney: SEA P<br />
Crustacea: Copepoda:<br />
Poecilostomatoida Oncaeidae Pseudolubbokia dilatata Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Copepoda:<br />
Poecilostomatoida unidentified unidentified Copepoda: Cyclopoida Cordell 2000<br />
Crustacea: Cumacea Bodoptriidae Vaunthompsonia pacifica EVOS ST<br />
II III IV V<br />
VI Austin 1985
Chapt 10. Biodiversity, page 10- 43<br />
Table 10.5 continued.<br />
BIOREGION<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Cumacea Diastylidae Brachydiastylis unidentified<br />
Feder and<br />
Matheke 1980 I/ST/Inf<br />
Crustacea: Cumacea Diastylidae Diastylis alaskensis EVOS ST II III IV x Austin 1985<br />
Crustacea: Cumacea Diastylidae Diastylis bidentata EVOS ST II III IV x Austin 1985<br />
Crustacea: Cumacea Diastylidae Diastylis koreana EVOS ST II III IV x Austin 1985<br />
Crustacea: Cumacea Diastylidae Diastylis paraspinulosa Hoberg 1986 ST<br />
II III IV V<br />
VI Austin 1985<br />
Crustacea: Cumacea Diastylidae Diastylopsis dawsoni Hoberg 1986 ST<br />
II III IV V<br />
VI Austin 1985<br />
Crustacea: Cumacea Diastylidae Leptostylis unidentified EVOS ST II III IV x x Austin 1985<br />
Crustacea: Cumacea Lampropidae Lamprops beringi Chapman 2000 I/Inf II III IV Chapman 2000<br />
Crustacea: Cumacea Lampropidae Lamprops quadriplicata EVOS ST II x x Chapman 2000<br />
Crustacea: Cumacea Lampropidae Lamprops sarsi EVOS ST II III IV x Austin 1985<br />
Crustacea: Cumacea Leuconiidae Eudorella emarginata<br />
Feder and Jewett<br />
1988 ST/Inf II x x Austin 1985<br />
Crustacea: Cumacea Leuconiidae Eudorellopsis biplicata EVOS ST/Inf II x x Austin 1985<br />
Crustacea: Cumacea Leuconiidae Eudorellopsis deformis EVOS ST/Inf II III IV Austin 1985<br />
Crustacea: Cumacea Leuconiidae Eudorellopsis dezhavini EVOS ST/Inf II III IV x Austin 1985<br />
Crustacea: Cumacea Leuconiidae Eudorellopsis integra<br />
Feder and<br />
Matheke 1980 ST/Inf II x x Austin 1985<br />
Crustacea: Cumacea Leuconiidae Leptocuma unidentified<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST<br />
Crustacea: Cumacea Leuconiidae Leucon nasica<br />
Feder and<br />
Matheke 1980 ST/Inf II x x x Austin 1985<br />
Crustacea: Cumacea Leuconiidae Leucon nasica orientalis UAM ST II x x x Austin 1985<br />
Crustacea: Cumacea Nannastacidae Campylaspis rubicunda UAM II x x x Schultz 1969<br />
Crustacea: Cumacea Nannastacidae Cumella vulgaris<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Epi II III IV C Chapman 2000<br />
Crustacea: Decapoda Atelecyclidae Telmessus cheiragonus Hart 1968 I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Callianassidae Callinassa unidentified Hart 1968 I/ST<br />
Crustacea: Decapoda Cancridae Cancer branneri Hart 1968 I/ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Cancridae Cancer gracilis Hart 1968 I/ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Cancridae Cancer magister Hart 1968 I/ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Cancridae Cancer oregonensis Hart 1968 I/ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Cancridae Cancer productus Hart 1968 I/ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Crangonidae Argis alaskensis UAM ST/Epi II III IV Butler 1980<br />
Crustacea: Decapoda Crangonidae Argis lar EVOS ST/Epi II III IV x Butler 1980
Chapt 10. Biodiversity, page 10- 44<br />
Table 10.5 continued.<br />
BIOREGION<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Decapoda Crangonidae Argis dentata<br />
Squires and<br />
Figueira 1974 I/ST II III IV x Butler 1980<br />
Crustacea: Decapoda Crangonidae Argis levior<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV Butler 1980<br />
Crustacea: Decapoda Crangonidae Crangon alaskensis Cooney 1987 ST/Epi<br />
II III IV V<br />
VI Austin 1985<br />
Crustacea: Decapoda Crangonidae Crangon communis<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV x Butler 1980<br />
Crustacea: Decapoda Crangonidae Crangon dalli<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV x Butler 1980<br />
Crustacea: Decapoda Crangonidae Crangon franciscorum<br />
Squires and<br />
Figueira 1974 I/ST II III IV Butler 1980<br />
Crustacea: Decapoda Crangonidae Crangon nigricauda<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV Butler 1980<br />
Crustacea: Decapoda Crangonidae Lissocrangon stylirostris Crangon<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV Butler 1980<br />
Crustacea: Decapoda Crangonidae Mesocrangon minuitella EVOS ST/Epi II III IV Butler 1980<br />
Crustacea: Decapoda Crangonidae Neocrangon alaskensis Crangon<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV x Butler 1980<br />
Crustacea: Decapoda Crangonidae Paracrangon echinata<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV x Butler 1980<br />
Crustacea: Decapoda Crangonidae Rhynchocramgon alata EVOS ST/Epi II III IV x Butler 1980<br />
Squires and<br />
Crustacea: Decapoda Crangonidae Sclerocrangon boreas<br />
Figueira 1974 ST/Epi II III IV x Butler 1980<br />
Crustacea: Decapoda Galatheidae Munida quadrispina UAM ST/Epi II III IV Butler 1980<br />
Crustacea: Decapoda Grapsidae Hemigrapsus nudus Hart 1968 I/ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Grapsidae Hemigrapsus oregonensis<br />
Crustacea: Decapoda Hippolytidae Eualus fabricii<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Epi II III IV Hart 1968<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV x Butler 1980<br />
Crustacea: Decapoda Hippolytidae Eualus macropthalmalmus UAM ST/Epi II III IV Butler 1980<br />
Crustacea: Decapoda Hippolytidae Eualus suckleyi UAM ST/Epi II III IV x Butler 1980<br />
Crustacea: Decapoda Hippolytidae Eualus avinus UAM ST/Epi II III IV Butler 1980<br />
Squires and<br />
Crustacea: Decapoda Hippolytidae Eualus gaimardii<br />
Figueira 1974 ST/Epi II III IV x Austin 1985
Chapt 10. Biodiversity, page 10- 45<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
Crustacea: Decapoda Hippolytidae Eualus pusiolus<br />
Crustacea: Decapoda Hippolytidae Eualus townsendi<br />
Crustacea: Decapoda Hippolytidae Heptacarpus decorus<br />
Crustacea: Decapoda Hippolytidae Heptacarpus brevirostris<br />
Crustacea: Decapoda Hippolytidae Heptacarpus camtschaticus<br />
Crustacea: Decapoda Hippolytidae Heptacarpus carinatus<br />
Crustacea: Decapoda Hippolytidae Heptacarpus paludicola<br />
Crustacea: Decapoda Hippolytidae Heptacarpus pictus<br />
Crustacea: Decapoda Hippolytidae Heptacarpus sitchensis<br />
Crustacea: Decapoda Hippolytidae Heptacarpus stimpsoni<br />
Crustacea: Decapoda Hippolytidae Heptacarpus stylus<br />
Crustacea: Decapoda Hippolytidae Heptacarpus tridens<br />
Crustacea: Decapoda Hippolytidae Hippolyte clarki<br />
Crustacea: Decapoda Hippolytidae Lebbeus groenlandicus<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Squires and<br />
Figueira 1974 ST/Epi II x x Butler 1980<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV x Butler 1980<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV Butler 1980<br />
Squires and<br />
Figueira 1974 I/ST II III IV Butler 1980<br />
Squires and<br />
Figueira 1974 I/ST II III IV x Butler 1980<br />
Squires and<br />
Figueira 1974 I/ST II III IV Butler 1980<br />
Squires and<br />
Figueira 1974 I/ST II III IV Butler 1980<br />
Squires and<br />
Figueira 1974 I/ST II III IV Austin 1985<br />
Squires and<br />
Figueira 1974 I/ST II III IV Butler 1980<br />
Squires and<br />
Figueira 1974 I/ST II III IV Butler 1980<br />
Squires and<br />
Figueira 1974 I/ST II III IV Butler 1980<br />
Squires and<br />
Figueira 1974 I/ST II III IV Butler 1980<br />
Squires and<br />
Figueira 1974 I/ST II III IV Butler 1980<br />
Squires and<br />
Figueira 1974 I/ST II III IV x Butler 1980<br />
Crustacea: Decapoda Hippolytidae Spirontocaris arcuata UAM I/ST II III IV x Butler 1980<br />
Crustacea: Decapoda Hippolytidae Spirontocaris lamellicornis UAM I/ST II III IV Butler 1980<br />
Crustacea: Decapoda Hippolytidae Spirontocaris ochotensis<br />
Squires and<br />
Figueira 1974 I/ST II III IV x Butler 1980<br />
Crustacea: Decapoda Hippolytidae Spirontocaris phippsi<br />
Squires and<br />
Figueira 1974 I/ST II x x Austin 1985<br />
Crustacea: Decapoda Lithodidae Acantholithodes hispidus Hart 1968 I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Lithodidae Cryptolithodes sitchensis Hart 1968 I/ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Lithodidae Cryptolithodes typicus Hart 1968 I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Lithodidae Hapalogaster cavicauda Austin 1985 I/ST/Epi II III IV Austin 1985<br />
Crustacea: Decapoda Lithodidae Hapalogaster grebnitskii Hart 1968 I/ST II III IV x Hart 1968<br />
Crustacea: Decapoda Lithodidae Hapalogaster mertensii Hart 1968 I/ST/Epi II III IV Hart 1968
Chapt 10. Biodiversity, page 10- 46<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Decapoda Lithodidae Lithodes aequispina UAM ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Lithodidae Lithodes couesi UAM ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Lithodidae Lopholithodes foraminatus UAM ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Lithodidae Oedignathus inermis Hart 1968 I/ST/Epi II III IV x Hart 1968<br />
Crustacea: Decapoda Lithodidae Paralithodes camtschatica UAM ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Lithodidae Phyllolithodes papillosus UAM I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Lithodidae Placetron wosnessenskii UAM ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Lithodidae Rhinolithodes wosnessenskii UAM ST/Epi II III IV Hart 1968<br />
Feder and Jewett<br />
Crustacea: Decapoda Majidae Chionecetes bairdi<br />
1988 ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Majidae Chorilia longipes Hart 1968 ST/Epi II III IV x Hart 1968<br />
Crustacea: Decapoda Majidae Hyas lyratus Hart 1968 I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Majidae Oregonia gracilis Hart 1968 I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Majidae Pugettia gracilis Hart 1968 I/ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Majidae Pugettia producta Hart 1968 I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Majidae Pugettia richii Hart 1968 I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Majidae Scyra acutifrons Hines et al. 2000 I/ST II III IV x Hart 1968<br />
II III IV V<br />
Crustacea: Decapoda Oplophoridae Hymenadora frontalis Cooney 1987 P<br />
VI x Austin 1985<br />
Crustacea: Decapoda Paguridae Discorsopagurus schmitti UAM I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Paguridae Elassochirus cavimanus UAM ST/Epi II III IV x Hart 1968<br />
Crustacea: Decapoda Paguridae Elassochirus gilli UAM I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Paguridae Elassochirus tenimanius UAM I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Paguridae Labidochirus splendescens UAM I/ST II III IV x x Hart 1968<br />
Crustacea: Decapoda Paguridae Pagurus aleuticus UAM I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Paguridae Pagurus beringanus UAM I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Paguridae Pagurus capillatus UAM I/ST II III IV Hart 1968
Chapt 10. Biodiversity, page 10- 47<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Decapoda Paguridae Pagurus cornutus UAM ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Paguridae Pagurus granosimanus UAM I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Paguridae Pagurus<br />
hirsutiusculus<br />
hirsutiusculus<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Paguridae Pagurus kennerleyi UAM I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Paguridae Pagurus samuelis UAM I/ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Paguridae Pagurus caurinus Hart 1968 I/ST II III IV V Hart 1968<br />
Crustacea: Decapoda Paguridae Pagurus confragosus Feder et al. 1979 ST/Epi II III IV Hart 1968<br />
Crustacea: Decapoda Paguridae Pagurus ochotensis<br />
Crustacea: Decapoda Pandalidae Pandalopsis dispar<br />
Crustacea: Decapoda Pandalidae Pandalus danae<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST II III IV Hart 1968<br />
Feder and Jewett<br />
1988 ST/Epi II III IV Butler 1980<br />
Squires and<br />
Figueira 1974 I/ST II III IV Butler 1980<br />
Crustacea: Decapoda Pandalidae Pandalus jordani UAM ST/Epi II III IV Butler 1980<br />
Crustacea: Decapoda Pandalidae Pandalus unidentified Cooney: SEA ST/Epi<br />
Crustacea: Decapoda Pandalidae Pandalus borealis<br />
Feder and Jewett<br />
1988 ST/Epi II x x Butler 1980<br />
Crustacea: Decapoda Pandalidae Pandalus goniurus<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV x Butler 1980<br />
Crustacea: Decapoda Pandalidae Pandalus hypsinotus<br />
Squires and<br />
Figueira 1974 I/ST II III IV x Butler 1980<br />
Crustacea: Decapoda Pandalidae Pandalus montagui tridens<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV Butler 1980<br />
Crustacea: Decapoda Pandalidae Pandalus platyceros<br />
Squires and<br />
Figueira 1974 I/ST II III IV x Butler 1980<br />
Crustacea: Decapoda Pandalidae Pandalus stenolepis<br />
Squires and<br />
Figueira 1974 ST/Epi II III IV Butler 1980<br />
Crustacea: Decapoda Pasiphaeidae Pasiphaea pacifica UAM P II III IV x Butler 1980<br />
Crustacea: Decapoda Pinnotheridae Pinnixa littoralis Hart 1968 I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Pinnotheridae Pinnixa occidentalis Hart 1968 ST/Epi II III IV Hart 1968<br />
Feder and<br />
Crustacea: Decapoda Pinnotheridae Pinnixa schmitti<br />
Matheke 1980 ST/Epi II III IV Hart 1968
Chapt 10. Biodiversity, page 10- 48<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Crustacea: Decapoda Porcellanidae Petrolisthes eriomerus Hart 1968 I/ST II III IV Hart 1968<br />
Crustacea: Decapoda Sergestidae Sergestes similis Eusergestes Cooney 1987 P<br />
II III IV V<br />
VI x Austin 1985<br />
Crustacea: Decapoda Upogebiidae Upogebia pugettensis<br />
Squires and<br />
Figueira 1974 I/Inf II III IV Hart 1968<br />
Crustacea: Decapoda Xanthidae Lophopanopeus bellus bellus Hart 1968 I/ST II III IV Hart 1968<br />
Crustacea:<br />
Euphausiacea Euphausiidae Euphausia pacifica Cooney 1987 P<br />
II III IV V<br />
VI x Austin 1985<br />
Crustacea:<br />
Euphausiacea Euphausiidae Stylocherion unidentified Cooney 1987 P<br />
Crustacea:<br />
Euphausiacea Euphausiidae Tessarabranchion oculata Cooney 1987 P II III IV V x Austin 1985<br />
Crustacea:<br />
Euphausiacea Euphausiidae Thysanoessa inermis Cooney 1987 P<br />
II III IV V<br />
VI x Austin 1985<br />
Crustacea:<br />
Euphausiacea Euphausiidae Thysanoessa longipes Cooney 1987 P III IV x Austin 1985<br />
Crustacea:<br />
Euphausiacea Euphausiidae Thysanoessa spinifera Cooney 1987 P<br />
II III IV V<br />
VI Austin 1985<br />
Crustacea:<br />
Euphausiacea Euphausiidae Thysanoessa inspinata Cooney 1987 P II III IV x Austin 1985<br />
Crustacea:<br />
Euphausiacea Euphausiidae Thysanoessa raschii<br />
Cooney and Coyle<br />
1988 P II x x Austin 1985<br />
Crustacea: Isopoda Aegidae Rocinela angustana UAM ST II III IV Schultz 1969<br />
Crustacea: Isopoda Dajidae Holophryxus alascensis UAM ST II III IV Austin 1985<br />
Crustacea: Isopoda Gnathiidae Gnathia tridens EVOS ST II III IV California NR Schultz 1969<br />
Crustacea: Isopoda Gnathiidae Gnathia unidentified<br />
Feder and<br />
Matheke 1980 ST/Inf Schultz 1969<br />
Crustacea: Isopoda Idoteidae Idotea fewkesi EVOS I/ST<br />
II III IV V<br />
VI Schultz 1969<br />
Crustacea: Isopoda Idoteidae Idotea obscura Chapman 2000 I/Epi II III IV Chapman 2000<br />
Crustacea: Isopoda Idoteidae Pentidotea wosnessenskii Idotea<br />
Feder and Bryson-<br />
Schwafel 1988 I/Epi II III IV Schultz 1969<br />
Crustacea: Isopoda Idoteidae Synidotea ritteri UAM ST/Epi II III IV V California NR Schultz 1969<br />
Crustacea: Isopoda Janiridae Janiropsis kincaldi J. pugettensis Chapman 2000 I II III IV Chapman 2000<br />
Crustacea: Isopoda Ligiidae Ligia pallasii Chapman 2000 I/Epi II III IV Chapman 2000<br />
Crustacea: Isopoda Limnoriidae Limnoria lignorum EVOS I/ST II III IV Schultz 1969<br />
cf. M.<br />
Crustacea: Isopoda Munnidae Munna<br />
chromocephala UAM ST II III IV Washington NR Schultz 1969<br />
Crustacea: Isopoda Munnidae Munna ubiquita EVOS ST II III IV Washington NR Schultz 1969<br />
Crustacea: Isopoda Munnidae Pleurogonium unidentified I/Epi Schultz 1969<br />
Crustacea: Isopoda Sphaeromatidae Dynamenella sheari UAM I/Epi II III IV Austin 1985<br />
Crustacea: Isopoda Sphaeromatidae Exosphaeroma amplicauda UAM I/Epi II III IV Schultz 1969<br />
Crustacea: Isopoda Sphaeromatidae Exosphaeroma unidentified UAM I/Epi Schultz 1969<br />
Crustacea: Isopoda Sphaeromatidae Gnorimosphaeroma lutea G. insulare Chapman 2000 I/Epi II III IV Chapman 2000
Chapt 10. Biodiversity, page 10- 49<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
Crustacea: Isopoda Sphaeromatidae Gnorimosphaeroma oregonensis<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST II III IV Schultz 1969<br />
Crustacea: Leptostraca Nebaliidae Nebalia unidentified EVOS P<br />
Crustacea: Mysidacea Lophogastridae Gnatahophausia gigas Cooney 1987 P<br />
Crustacea: Mysidacea Mysidae Acanthomysis nephropthalma Cooney 1987 P<br />
II III IV V<br />
VI VII VIII<br />
IX x x Austin 1985<br />
II III IV V<br />
VI Austin 1985<br />
Crustacea: Mysidacea Mysidae Acanthomysis pseudomacraspis Cooney 1987 P II III IV x Austin 1985<br />
II III IV V<br />
Crustacea: Mysidacea Mysidae Holmsiella anomala Cooney 1987 P<br />
VI x Austin 1985<br />
Crustacea: Mysidacea Mysidae Meterythrops robusta Cooney 1987 P II III IV x Austin 1985<br />
Crustacea: Mysidacea Mysidae Mysis littoralis Chapman 2000 P II x x x Chapman 2000<br />
Crustacea: Mysidacea Mysidae Mysis oculata M. littoralis Cooney 1987 P II III IV x x Austin 1985<br />
Crustacea: Mysidacea Mysidae Neomysis kadiakensis Cooney 1987 P II III IV Austin 1985<br />
Crustacea: Mysidacea Mysidae Neomysis rayi Cooney 1987 P II III IV V x Austin 1985<br />
II III IV V<br />
Crustacea: Mysidacea Mysidae Pseudomma truncatus Cooney 1987 P<br />
VI Austin 1985<br />
Crustacea: Ostracoda Halocyprididae Conchoecia alata minor Alacia<br />
Cooney and Coyle<br />
1988 ST/P II III<br />
Crustacea: Ostracoda Halocyprididae Conchoecia elegans Paraconchoecia UAM ST/P II III<br />
Cooney and Coyle<br />
Crustacea: Ostracoda Halocyprididae Conchoecia unidentified<br />
1988 ST/P<br />
British<br />
Columbia NR Austin 1985<br />
British<br />
Columbia NR Austin 1985<br />
Crustacea: Ostracoda Parodoxostomatinae unidentified unidentified UAM ST/P<br />
Crustacea: Ostracoda Philomedidae Philomedes dentata UAM ST/Inf II III<br />
Crustacea: Ostracoda Philomedidae Scleroconcha trituberculata UAM ST/Inf II III<br />
British<br />
Columbia NR Austin 1985<br />
British<br />
Columbia NR Austin 1985<br />
Crustacea: Ostracoda Podocopidae unidentified unidentified UAM<br />
Crustacea: Tanaidacea Paratanaidae Leptochelia savignyi Chapman 2000 I/Epi II x x C Chapman 2000<br />
Crustacea: Thoracica Balanamorpha Balanus crenatus<br />
Crustacea: Thoracica Balanamorpha Balanus glandula<br />
Crustacea: Thoracica Balanamorpha Chirona evermani<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST II III IV x Austin 1985<br />
Feder and Bryson-<br />
Schwafel 1988 I/Epi II III IV Austin 1985<br />
Feder and<br />
Matheke 1980 ST/Inf II III IV Austin 1985
Chapt 10. Biodiversity, page 10- 50<br />
Table 10.5 continued.<br />
HIGHER TAXON FAMILY GENUS SPECIES OTHER NAMES<br />
Crustacea: Thoracica Balanamorpha Semibalanus balanoides Balanus<br />
Crustacea: Thoracica Balanamorpha Semibalanus cariosus Balanus<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Feder and Bryson-<br />
Schwafel 1988 I/Epi II III IV Austin 1985<br />
Feder and Bryson-<br />
Schwafel 1988 I/Epi II III IV x Austin 1985<br />
Crustacea: Thoracica Chthamalidae Chthamalus dalli EVOS I/Epi II III IV x Austin 1985<br />
Crustacea: Thoracica Scalpellidae Scaphellum pacificum<br />
Feder and<br />
Matheke 1980 ST/Epi II III IV x Austin 1985<br />
Pycnogonida Ammotheidae Achelia alaskensis Child 1995 ST/Epi II III IV x Child 1995<br />
Pycnogonida Ammotheidae Achelia borealis UAM ST/Epi II III IV x x Child 1995<br />
Pycnogonida Ammotheidae Achelia chaelata UAM ST/Epi II III IV Washington NR Austin 1985<br />
Pycnogonida Ammotheidae Achelia gracilipes Ammothea UAM ST/Epi II III IV<br />
British<br />
Columbia NR Austin 1985<br />
Pycnogonida Ammotheidae Achelia latifrons Child 1995 ST/Epi II III IV x Child 1995<br />
Pycnogonida Ammotheidae Achelia megova Child 1995 ST/Epi II III IV Child 1995<br />
Pycnogonida Ammotheidae Achelia pribil<strong>of</strong>ensis Ammothea UAM ST/Epi II III IV Bering Sea nr Child 1995<br />
Pycnogonida Phoxochilidiidae Phoxichilidium femoratum UAM ST/Epi II x x Austin 1985<br />
Pycnogonida Phoxochilidiidae Phoxichilidium quadrodentatum UAM ST/Epi II III IV Austin 1985<br />
Pycnogonida Tanystylidae Tanystylum anthostomasti UAM ST/Epi II III IV x Austin 1985
Chapt 10. Biodiversity, page 10- 51<br />
Echinodermata<br />
Ninety-nine echinoderm species, representing all five classes are listed (Table 10.6).<br />
Principal sources for taxonomic and distributional information are D‘yakonov (1954)<br />
(ophiuroids), Lambert (1981, 1997) (Asteroidea and Holothuroidea). No range extensions or<br />
possibly undescribed species were noted for Prince William Sound.<br />
Table 10.6. Echinodermata.<br />
CLASS FAMILY GENUS SPECIES<br />
BIOREGION<br />
OTHER<br />
NAMES SPECIMEN or SOURCE HABITAT NEP NWP AR NWA<br />
REFERENCE to<br />
DISTRIBUTION<br />
Asteroidea Asteriidae Evasterias troschelii<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Epi II III IV Lambert 1981<br />
Asteroidea Asteriidae Leptasterias hexactis UAM I/ST/Epi II III IV Lambert 1981<br />
Asteroidea Asteriidae Orthasterias koehleri UAM ST II III IV Lambert 1981<br />
Asteroidea Asteriidae Pisaster ochraceus UAM I/ST/Epi II III IV Lambert 1981<br />
Asteroidea Asteriidae Pycnopodia helianthoides<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Epi II III IV Lambert 1981<br />
Asteroidea Asteriidae Rathbunaster californianus UAM ST II III IV Lambert 1981<br />
Asteroidea Asteropseidae Dermasterias imbricata Poraniidae UAM I/ST/Epi II III IV Lambert 1981<br />
Asteroidea Astropectinidae Dipsacaster borealis UAM ST II III IV Austin 1985<br />
Asteroidea Astropectinidae Leptychaster anomalus Feder and Matheke 1980 ST II III IV Lambert 1981<br />
Asteroidea Astropectinidae Leptychaster arcticus UAM ST II III IV x x x Lambert 1981<br />
Asteroidea Astropectinidae Leptychaster pacificus Lambert 1981 ST II III IV Lambert 1981<br />
Asteroidea Benthopectinidae Luidiaster dawsoni UAM ST II III IV Lambert 1981<br />
Asteroidea Benthopectinidae Nearchaster aciculosus UAM ST II III IV Austin 1985<br />
Asteroidea Echinasteridae Henricia aspera UAM ST II III IV Lambert 1981<br />
Asteroidea Echinasteridae Henricia asthenactis Lambert 1981 ST II III IV Lambert 1981<br />
Asteroidea Echinasteridae Henricia leviuscula UAM ST II III IV Lambert 1981<br />
Asteroidea Echinasteridae Henricia longispina Lambert 1981 ST II III IV x Lambert 1981<br />
Asteroidea Echinasteridae Henricia sanguinolenta UAM ST II III x x Lambert 1981<br />
Asteroidea Echinasteridae Poraniopsis inflata UAM ST II III IV x Lambert 1981<br />
Asteroidea Goniasteridae Ceramaster arcticus UAM ST II III IV Lambert 1981<br />
Asteroidea Goniasteridae Ceramaster patagonicus Lambert 1981 ST<br />
II III IV V VI<br />
VII VIII IX Lambert 1981<br />
Asteroidea Goniasteridae Gephyreaster swifti Radiasterridae Feder and Jewett 1987 ST II III IV Lambert 1981<br />
Asteroidea Goniasteridae Hippasteria spinosa UAM ST II III IV Lambert 1981<br />
Asteroidea Goniasteridae Mediaster aequalis UAM ST II III IV Lambert 1981<br />
Asteroidea Goniasteridae Pseudarchaster parelii UAM ST II x x Lambert 1981<br />
Asteroidea Goniopectinidae Ctenodiscus crispatus Ctenodiscidae UAM ST<br />
I II III IV V<br />
VI VII IX x x x Lambert 1981<br />
Asteroidea Luidiidae Luidia foliolata UAM ST II III IV Lambert 1981<br />
Asteroidea Pedicellasteridae Pedicellaster magister Heliasteridae UAM ST II III IV x Austin 1985<br />
Asteroidea Pterasteridae Diplopteraster multipes UAM ST II III IV V x x x Lambert 1981<br />
Asteroidea Pterasteridae Pteraster militaris UAM ST II III IV x Lambert 1981<br />
Asteroidea Pterasteridae Pteraster tesselatus UAM ST II III IV Lambert 1981<br />
Asteroidea Solasteridae Crossaster papposus UAM ST II III IV x x x Lambert 1981<br />
Asteroidea Solasteridae Lophaster furcilliger UAM ST II III IV Lambert 1981<br />
Asteroidea Solasteridae Solaster dawsoni UAM ST II III IV x Lambert 1981<br />
Asteroidea Solasteridae Solaster endeca UAM ST II III IV V x x Lambert 1981<br />
Asteroidea Solasteridae Solaster paxillatus UAM ST II III IV x Lambert 1981<br />
Asteroidea Solasteridae Solaster stimpsoni Lambert 1981 ST II III IV x Lambert 1981<br />
Criniodea Antedonidae Florometra serratissima Austin 1985 ST II III IV V Austin 1985<br />
Criniodea Antedonidae Florometra asperrima<br />
Helimetra<br />
gracilis UAM ST II III IV V x Austin 1985<br />
Criniodea Antedonidae Psathyrometra fragilis Austin 1985 ST II III IV V x Austin 1985<br />
Criniodea Antedonidae Retiometra alascana Austin 1985 ST II III IV Austin 1985<br />
Echinoidea Echinarachniidae Echinarachnius parma Foster and Hines 2000 ST II III IV x Pavloskii 1955<br />
Echinoidea Schizasteridae Brisaster townsendi Feder and Matheke 1980 ST II III IV x Pavloskii 1955<br />
Echinoidea Strongylocentrotidae Allocentrotus fragilis Feder and Jewett 1987 ST II III IV Austin 1985<br />
Echinoidea Strongylocentrotidae Strongylocentrotus droebachiensis<br />
Feder and Bryson-<br />
Schwafel 1988 I/ST/Epi II x x Austin 1985<br />
Echinoidea Strongylocentrotidae Strongylocentrotus franciscanus Rosenthal 1977 ST/Epi II III IV V Austin 1985<br />
Holothuroidea Cucumariidae Cucumaria frondosa japonica Lambert 1997 ST II III IV Lambert 1997<br />
Holothuroidea Cucumariidae Cucumaria miniata Lambert 1997 ST II III IV Lambert 1997<br />
Holothuroidea Cucumariidae Cucumaria vegae Lambert 1997 ST II III IV x Lambert 1997<br />
Holothuroidea Cucumariidae Ekmania diomedeae Lambert 1997 ST II III IV x Lambert 1997<br />
Holothuroidea Cucumariidae Pentamera calcigera UAM ST II x x Austin 1985<br />
Holothuroidea Cucumariidae Thyonidium kurilensis Lambert 1997 ST II III IV x Lambert 1997<br />
Holothuroidea Molpadiidae Molpadia intermedia UAM ST II III IV Lambert 1997<br />
Holothuroidea Phyllophoridae Pentamera populifera Lambert 1997 ST II III IV Lambert 1997<br />
Holothuroidea Phyllophoridae Thyone cf. T. benti UAM ST II III IV Lambert 1997<br />
Holothuroidea Psolidae Psolus chitonoides Lambert 1997 ST II III IV Lambert 1997<br />
Holothuroidea Psolidae Psolus squamatus Lambert 1997 ST II x x Lambert 1997<br />
Holothuroidea Sclerodactylidae Eupentacta pseudoquinquesemita Lambert 1997 ST II III IV Lambert 1997<br />
Holothuroidea Stichopodidae Parastichopus californicus Lambert 1997 ST II III IV Lambert 1997<br />
Holothuroidea Synallactidae Pseudostichopus mollis Lambert 1997 ST x x x x Lambert 1997<br />
Holothuroidea Synallactidae Synallactes challengeri Lambert 1997 ST x x x x Lambert 1997<br />
Holothuroidea Synaptidae Leptosynapta clarki UAM ST II III IV x Austin 1985<br />
Ophiuroidea Amphiuridae Amphipolis squamata Feder and Jewett 1987 ST x x x x Austin 1985<br />
Ophiuroidea Amphiuridae Diamphiodia craterodmeta Amphiodia Feder and Matheke 1980 ST II III IV x D’Yakanov 1954<br />
Ophiuroidea Amphiuridae Unioplis macraspis Amphioplus Feder and Matheke 1980 ST II III IV x Austin 1985<br />
Ophiuroidea Ophiactidae Ophiopholis aculeatata O. caryi Rosenthal 1977 ST II II IV x Austin 1985<br />
Ophiuroidea Ophiuridae Ophiura sarsi Feder and Matheke 1980 ST II x x x Austin 1985<br />
Ophiuroidea Ophiuridae Ophiura quadrispina Feder and Matheke 1980 ST II III IV D’Yakanov 1954
Chapt 10. Biodiversity, page 10- 52<br />
Bryozoa<br />
Bryozoa are abundant in Alaskan waters and prevalent in the fouling community. The<br />
data set consists <strong>of</strong> 74 species (Table 10.7), which is probably an underestimate. Nineteen<br />
identifications are confident only to the generic level. Sources for distribution data include Dick<br />
and Ross (1988), Hines and Ruiz (2000), Kluge (1962), as well as cataloged specimens in the<br />
UAM collection. J. Winston (2000) contributed the identifications and distributional information<br />
for 22 species. Schizoporella unicornis is regarded as a nonindigenous species, native to the<br />
northwestern Pacific. Ten species are possible new records for Alaska, range extensions form<br />
either the Arctic or Pacific region.<br />
Table 10.7. Bryozoa.<br />
ORDER FAMILY GENUS SPECIES<br />
OTHER<br />
NAMES<br />
Cheilostomata:<br />
Anasca Beaniidae Beania mirabilis UAM ST<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
II III IV V VI<br />
VII VIII IX x x<br />
British<br />
Columbia nr Austin 1985<br />
Cheilostomata:<br />
Anasca Bugulidae Bugula californica Hines and Ruiz 2000 I II III IV Austin 1985<br />
Cheilostomata:<br />
Anasca Bugulidae Bugula pacifica Hines and Ruiz 2000 I II III IV Austin 1985<br />
Cheilostomata:<br />
Anasca Bugulidae Bugula unidentified Hines and Ruiz 2000 I<br />
Cheilostomata:<br />
Anasca Bugulidae Dendrobeania curvirostrata Hines and Ruiz 2000 I II III IV V VI<br />
British<br />
Columbia NR Austin 1985<br />
Cheilostomata:<br />
Anasca Bugulidae Dendrobeania lichenoides UAM I II III IV Dick and Ross 1988<br />
Cheilostomata:<br />
Anasca Bugulidae Dendrobeania murrayana UAM ST II x x x Kluge 1962<br />
Cheilostomata:<br />
Anasca Calloporidae Calloporella craticula Callopora Hines and Ruiz 2000 I/ST II x x x Kluge 1962<br />
Cheilostomata:<br />
Anasca Calloporidae Calloporella lineata Callopora UAM I/ST II x x x C Kluge 1962<br />
Cheilostomata:<br />
Anasca Calloporidae Calloporella "lineata" Hines and Ruiz 2000 I/ST II x x Austin 1985<br />
Cheilostomata:<br />
Anasca Calloporidae Tegella aquilirostris Hines and Ruiz 2000 I/ST II III IV x Austin 1985<br />
Cheilostomata:<br />
Anasca Calloporidae Tegella armifera Hines and Ruiz 2000 I/ST II III IV x Austin 1985<br />
Cheilostomata:<br />
Anasca Calloporidae Tegella robertsonae UAM ST II III IV V VI x Osburn 1950<br />
Cheilostomata:<br />
Anasca Cryptosulidae Cryptosula okadai Hines and Ruiz 2000 I II III IV x Dick and Ross 1988<br />
Cheilostomata:<br />
Anasca Cryptosulidae Harmeria scutulata Hippothoidae UAM I/ST II x x x Arctic nr Dick and Ross 1988<br />
Cheilostomata:<br />
Anasca Electridae Cauloramphus pseudospinifer Hines and Ruiz 2000 I II III IV x Dick and Ross 1988<br />
Cheilostomata:<br />
Anasca Electridae Electra unidentified UAM I<br />
Cheilostomata:<br />
Anasca Flustridae Termin<strong>of</strong>lustra membranaceotruncata UAM I/ST II x x x Dick and Ross 1988<br />
Cheilostomata:<br />
Anasca Hincksinidae Cauloramphus "variegata" C. spiniferum UAM I/ST II III IV V x x Austin 1985<br />
Cheilostomata:<br />
Anasca Membraniporidae Conopeum sp. (chesapeakensis) Hines and Ruiz 2000 I/ST II III IV V x x x Austin 1985<br />
Cheilostomata:<br />
Anasca Membraniporidae Membranipora membranipora Hines and Ruiz 2000 I/ST II III x x x Kluge 1962<br />
Cheilostomata:<br />
Anasca Membraniporidae Membranipora serrilamella UAM I/ST x x x x N Pacific nr Austin 1985<br />
Cheilostomata:<br />
Anasca Microporidae Micropora unidentified UAM I/ST<br />
Cheilostomata:<br />
Anasca Microporinidae Microporina articulata UAM I II x x x Dick and Ross 1988<br />
Cheilostomata:<br />
Anasca Scrupariidae Brettia unidentified<br />
Bicellariidae/C<br />
orynoprella UAM I/ST<br />
Cheilostomata:<br />
Anasca Scrupocellariidae Scrupocellaria unidentified UAM I/ST<br />
Cheilostomata:<br />
Anasca Scrupocellariidae Tricellaria gracilis UAM I/ST II x x x Kluge 1962<br />
Cheilostomata:<br />
Anasca Scrupocellariidae Tricellaria occidentalis UAM ST II III IV V VI<br />
British<br />
Columbia nr Austin 1985<br />
Cheilostomata:<br />
Anasca Scrupocellariidae Tricellaria ternata UAM I II x x x Dick and Ross 1988<br />
Cheilostomata:<br />
Ascophora Cribrilinidae Cribrilina annulata UAM I/ST II x x x Kluge 1962<br />
Cheilostomata:<br />
Ascophora Cribrilinidae Cribrilina corbicula Hines and Ruiz 2000 I II III IV C Austin 1985<br />
Cheilostomata:<br />
Ascophora Cribrilinidae Cribrilina unidentified Hines and Ruiz 2000 I/ST<br />
Cheilostomata:<br />
Ascophora Hippoporinidae Lepralia unidentified Hippothoa UAM I/ST<br />
Cheilostomata:<br />
Ascophora Hippothoidae Cellepora craticula Costazia Hines and Ruiz 2000 I/ST x C Kluge 1962
Chapt 10. Biodiversity, page 10- 53<br />
Table 10.7 continued.<br />
ORDER FAMILY GENUS SPECIES<br />
OTHER<br />
NAMES<br />
BIOREGION<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN<br />
REFERENCE to<br />
NIS STATUS DISTRIBUTION<br />
Cheilostomata:<br />
Ascophora Hippothoidae Celleporella hyalina Hippothoa UAM I/ST x x x x C Dick and Ross 1988<br />
Cheilostomata:<br />
Ascophora Hippothoidae Hippothoa unidentified UAM I<br />
Cheilostomata:<br />
Ascophora Microporellidae Fenestruloides eopacifica Hines and Ruiz 2000 I II III IV V VI California NR Soule et al. 1996<br />
Cheilostomata:<br />
Ascophora Microporellidae Microporella germana Hines and Ruiz 2000 I II III IV Dick and Ross 1988<br />
Cheilostomata:<br />
Ascophora Mucronellidae Cystisella bicornia Osburn 1952 ST II III IV x Osburn 1952<br />
Cheilostomata:<br />
Ascophora Mucronellidae Cystisella saccata Osburn 1952 ST II III x Arctic nr Osburn 1952<br />
Cheilostomata:<br />
Ascophora Mucronellidae Parasmittina trispinosa Smittinidae Hines and Ruiz 2000 I x x x x C Dick and Ross 1988<br />
Cheilostomata:<br />
Ascophora Mucronellidae Porella acutirostris Smittinidae UAM I/ST II III IV V VI x x x Dick and Ross 1988<br />
Cheilostomata:<br />
Ascophora Mucronellidae Porella acutirostris<br />
Cheilostomata:<br />
Ascophora Mucronellidae Rhamphostomella bilaminata<br />
Cheilostomata:<br />
Ascophora Mucronellidae Rhamphostomella gigantea<br />
Cheilostomata:<br />
Ascophora Mucronellidae Rhamphostomella hincksi<br />
P. major<br />
Smittinidae Hines and Ruiz 2000 ST II III IV V VI x x x Osburn 1952<br />
Rhamphostom<br />
ellidae UAM I/ST x x Kluge 1962<br />
Rhamphostom<br />
ellidae Hines and Ruiz 2000 ST x x Arctic nr Osburn 1952<br />
Rhamphostom<br />
ellidae Hines and Ruiz 2000 ST II x x Kluge 1962<br />
Cheilostomata:<br />
Ascophora Mucronellidae Smittina unidentified Smittinidae UAM I/ST<br />
Cheilostomata:<br />
Ascophora Reteporidae Phidolopora pacifica P. labiata Hines and Ruiz 2000 I II III IV Austin 1985<br />
Cheilostomata:<br />
Ascophora Schizoporellidae Hippodiplosia unidentified UAM I/ST II x x x Kluge 1962<br />
Cheilostomata:<br />
Ascophora Schizoporellidae Hippoporina vulgaris Hines and Ruiz 2000 I/ST II<br />
Cheilostomata:<br />
Ascophora Schizoporellidae Schizomavella porifera Hines and Ruiz 2000 I/ST II x x x Arctic nr Osburn 1952<br />
Cheilostomata:<br />
Ascophora Schizoporellidae Schizoporella "unicornis" Hines and Ruiz 2000 I x x x x NW Pacific Definate Hines and Ruiz 2000<br />
Cheilostomata:<br />
Ascophora Umbonulidae Umbonula unidentified UAM I/ST<br />
Ctenostomata Alcyonidiidae Alcyonidium hirsutum Hines and Ruiz 2000 I/ST II x x x NE Atlantic NR Kluge 1962<br />
Ctenostomata Alcyonidiidae Alcyonidium polynoum A. mytili Hines and Ruiz 2000 I/ST II III IV V x x x C Austin 1985<br />
Ctenostomata Alcyonidiidae Alcyonidium unidentified Hines and Ruiz 2000 I/ST<br />
III IV V VI<br />
Cyclostomata Crisiidae Crisia occidentalis UAM I/ST VII VIII IX x x Washington NR Austin 1985<br />
Cyclostomata Crisiidae Crisia serrulata Hines and Ruiz 2000 I II III IV Austin 1985<br />
Cyclostomata Crisiidae Filicrisia fanciscana Hines and Ruiz 2000 I II III IV x Austin 1985<br />
Cyclostomata Crisiidae Filicrisia geniculata UAM I/ST II x x Kluge 1962<br />
Cyclostomata Crisiidae Filicrisia smitti Hines and Ruiz 2000 ST II III x x x Kluge 1962<br />
Cyclostomata Diaperoeciidae Diaperoecia unidentified UAM ST<br />
Cyclostomata Diastoporidae Disporella unidentified UAM I/ST<br />
Cyclostomata Heteroporidae Heteropora magna UAM ST II III IV<br />
British<br />
Columbia nr Austin 1985<br />
Cyclostomata Heteroporidae Heteropora pacifica UAM ST II III IV Austin 1985<br />
Cyclostomata Heteroporidae Heteropora unidentified UAM I/ST<br />
Cyclostomata Lichenoporidae Lichenopora unidentified UAM I/ST<br />
Cyclostomata Oncousoeciidae Oncousiecia unidentified UAM I/ST<br />
Cyclostomata Oncousoeciidae Stomatopora granulata UAM I/ST II III IV x Kluge 1962<br />
Cyclostomata Tubuliporidae Idmonea unidentified Idmoneidae UAM I/ST<br />
Cyclostomata Tubuliporidae Platonea unidentified UAM I/ST<br />
Cyclostomata Tubuliporidae Tubulipora flabellaris UAM I/ST II III IV x Kluge 1962<br />
Cyclostomata Tubuliporidae Tubulipora sp. (tuba) Hines and Ruiz 2000 I/ST II III IV<br />
Cyclostomata Tubuliporidae Tubulipora tuba<br />
T.<br />
occidentalis,<br />
T. fasciculifera Hines and Ruiz 2000 I/ST II III IV V<br />
British<br />
Columbia NR Austin 1985<br />
Miscellaneous Invertebrates<br />
Ninety-five taxa, representing twelve different phyla make up this data set (Table 10.8).<br />
Nemerteans, chaetognaths, brachiopods, and urochordates , and are fairly well-known or have<br />
received the attention <strong>of</strong> the project’s taxonomists. (Lambert 2000; Mills 2000). However, it is<br />
clear that sponges, flatworms, and nematodes are undercounted. The nonindigenous sponge,<br />
Cliona thosina was recorded by the fouling community survey (Hines and Ruiz 200).<br />
Phylum<br />
taxa reported in this study<br />
Porifera 4<br />
Platyhelminthes none identified to species<br />
Nemertea 54<br />
Sipunculida 3<br />
Priapulida 1<br />
Echiuridae 1
Chapt 10. Biodiversity, page 10- 54<br />
Phoronidae none identified to species<br />
Brachiopoda 3<br />
Chaetognatha 5<br />
Tardigrada 1<br />
Urochordata 28<br />
Gnathostomulida, Gastrotrichida, Kinorhyncha have not been documented.<br />
Table 10.8. Miscellaneous invertebrates.<br />
Bioregion<br />
PHYLUM/ CLASS FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Brachiopoda:<br />
Articulata Cancellothyrididae Terebratulina unguicula UAM I/ST/Epi II III IV Austin 1985<br />
Brachiopoda:<br />
Articulata Hemithyridae Hemithiris psittacea UAM ST II III IV x x Austin 1985<br />
Brachiopoda:<br />
Articulata Laqueidae Terebratalia transversa UAM ST II III IV SE Alaska nr Austin 1985<br />
Brachiopoda:<br />
Articulata Laqueidae Laqueus californianus UAM ST II III IV x Austin 1985<br />
Chaetagnatha Eukrohniidae Eukrohnia fowleri Cooney 1987 P II III IV x x x Austin 1985<br />
Chaetagnatha Eukrohniidae Eukrohnia bathypelagica Cooney 1987 P II III IV x Austin 1985<br />
Chaetagnatha Eukrohniidae Eukrohnia hamata<br />
Cooney and Coyle<br />
1988 P x x x x<br />
Smith and Johnson<br />
1996<br />
Chaetagnatha Sagittidae Sagitta elegans<br />
Cooney and Coyle<br />
1988 P II III IV<br />
Smith and Johnson<br />
1996<br />
Chaetagnatha Sagittidae Sagitta scrippsae Cooney 1987 P<br />
II III IV V VI<br />
VII VIII x Austin 1985<br />
Echiurida Echiuridae Echiuris<br />
echiuris<br />
alaskensis<br />
Feder and Bryson-<br />
Schwafel 1988 I II III IV x Austin 1985<br />
Nematoda unidentified unidentified<br />
Feder and<br />
Matheke 1980 Inf<br />
Nemertea Amphiporidae Amphiporus angulatus Coe 1910 I II III IV V VI x Coe 1904<br />
Nemertea Amphiporidae Amphiporus bimaculatus Coe 1910 I<br />
II III IV V VI<br />
VII VIII Coe 1904<br />
Nemertea Amphiporidae Amphiporus exilis A. formidabilis Coe 1910 I II III IV V VI Coe 1904<br />
Nemertea Amphiporidae Amphiporus gelatinosus Austin 1985 I II III x Austin 1985<br />
Nemertea Amphiporidae Amphiporus imparispinosus Austin 1985 I<br />
II III IV V VI<br />
VII VIII Austin 1985<br />
Nemertea Amphiporidae Amphiporus lactifloreus I I<br />
Nemertea Amphiporidae Amphiporus leuciodes Coe 1910 I III Coe 1904<br />
Nemertea Amphiporidae Amphiporus nebulosus Coe 1910 I II III Coe 1904<br />
Nemertea Amphiporidae Amphiporus tigrinus Coe 1910 I III IV Coe 1904<br />
Nemertea Amphiporidae Zygonemertes albida Coe 1910 I III IV V VI Coe 1904<br />
Nemertea Amphiporidae Zygonemertes thalassina Coe 1910 I II III Coe 1904<br />
Nemertea Amphiporidae Zygonemertes virescens Austin 1985 I<br />
I II III IV V VI<br />
VII VIII IX x Austin 1985<br />
III IV V VI<br />
Nemertea Carinomidae Carinoma griffini C. mutabilis Coe 1910 I<br />
VII VIII Coe 1904<br />
Nemertea Carinomidae Carinomella lactea Tubulanidae Austin 1985 I I II III IV V VI Austin 1985<br />
Nemertea Cephalothricidae Cephalothrix linearis Procephalothrix spiralis Coe 1910 I II III IV V VI Coe 1904<br />
Nemertea Cerebratulidae Cerebratulus albifrons Coe 1910 I III IV V VI Coe 1904<br />
Nemertea Cerebratulidae Cerebratulus herculeus Coe 1910 I III IV V VI Coe 1904<br />
Nemertea Cerebratulidae Cerebratulus longiceps Coe 1910 I II III IV V Coe 1904<br />
Nemertea Cerebratulidae Cerebratulus marginatus Coe 1910 I III Coe 1904<br />
Nemertea Cerebratulidae Cerebratulus montgomeryi Coe 1910 I II III IV V Coe 1904<br />
Nemertea Cerebratulidae Cerebratulus occidentalis Coe 1910 I II III Coe 1904<br />
Nemertea Cerebratulidae Cerebratulus unidentified<br />
Feder and Matheke<br />
1980 ST<br />
Nemertea Emplectonematidae Emplectonema buergeri Coe 1910 I III Coe 1904<br />
Nemertea Emplectonematidae Emplectonema gracile Coe 1910 I II III IV V Coe 1904<br />
Nemertea Emplectonematidae Paranemertes carnea Coe 1910 I II III IV Coe 1904<br />
Nemertea Emplectonematidae Paranemertes pallida Coe 1910 I II III Coe 1904<br />
Feder and Bryson-<br />
Nemertea Emplectonematidae Paranemertes peregrina<br />
Schwafel 1988 I II III IV x Austin 1985<br />
Nemertea Lineidae Lineus torquatus Coe 1910 I II III IV V VI Coe 1904<br />
Nemertea Lineidae Micrura alaskensis Coe 1910 I<br />
III IV V VI<br />
VII VIII Coe 1904<br />
Nemertea Lineidae Micrura verrilli Coe 1910 I II III IV V VI Coe 1904<br />
Nemertea Lineidae Myoisophagus sanguineus Lineus socialis/vegetus Austin 1985 I II III IV V VI x x Austin 1985<br />
Nemertea Tetrastemmatidae Tetrastemma aberrans Coe 1910 I II III Coe 1904<br />
Nemertea Tetrastemmatidae Tetrastemma bicolor Coe 1910 I II III Coe 1904<br />
Nemertea Tetrastemmatidae Tetrastemma caecum Amphinemertes Coe 1910 I/ST II III Coe 1904<br />
Nemertea Tetrastemmatidae Tetrastemma candidum Austin 1985 I<br />
II III IV V VI<br />
VII VIII x x Austin 1985<br />
Nemertea Tetrastemmatidae Tetrastemma nigrifrons Austin 1985 I<br />
I II III IV V VI<br />
VII VIII IX Austin 1985<br />
Nemertea Tubulanidae Carinella albocinctus Tubulanus Austin 1985 ST II III IV V VI Austin 1985<br />
Nemertea Tubulanidae Carinella annulatus Tubulanus Austin 1985 I II III x Austin 1985<br />
Nemertea Tubulanidae Carinella capistrata Tubulanus capistratus Coe 1910 I II Coe 1904<br />
Nemertea Tubulanidae Carinella cingulatus Tubulanus Austin 1985 I II III IV V VI Austin 1985<br />
Nemertea Tubulanidae Carinella dinema Tubulanus sexlineatus Coe 1910 I II III Coe 1904<br />
Nemertea Tubulanidae Carinella frenatus Tubulanus Austin 1985 I II III IV V VI Austin 1985<br />
Nemertea Tubulanidae Carinella nothus Tubulanus Austin 1985 I II III x Austin 1985<br />
Nemertea Tubulanidae Carinella speciosa Tubulanus polymorphus Coe 1910 I II III Coe 1904<br />
Nemertea Valenciniidae Taeniosoma princeps Baseodiscus Coe 1910 I II III IV Coe 1904<br />
Phoronida unidentified unidentified<br />
Feder and Matheke<br />
1980 ST
Chapt 10. Biodiversity, page 10- 55<br />
Table 10.8 continued.<br />
Bioregion<br />
PHYLUM/ CLASS FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Platyhelminthes unidentified unidentified<br />
Feder and Bryson-<br />
Schwafel 1988 Inf<br />
Porifera Myxillidae Myxilla incrustans UAM ST II III IV x Austin 1985<br />
Porifera Subertidae Suberites ficus<br />
Scheel, Foster, and<br />
Hough, 1997 I/ST/Epi II III IV Austin 1985<br />
Porifera Spirastrellidae Cliona thosina Hines et al. 2000 boring II unkown definate<br />
Porifera Spirastrellidae Cliona celata Rosenthal 1977 Epi II III IV V Austin 1985<br />
Priapulida Priapulidae Priapulus caudatus<br />
Feder and Matheke<br />
1980 ST II x x x Austin 1985<br />
Sipunculida Golfingiidae Golfingia margaritacea<br />
Feder and Matheke<br />
1980 I/ST/Inf II x x Austin 1985<br />
Sipunculida Golfingiidae Golfingia vulgaris<br />
Feder and Matheke<br />
1980 ST II x x Austin 1985<br />
Sipunculida Golfingiidae Phascolion strombi<br />
Feder and Matheke<br />
1980 ST x x x x Austin 1985<br />
Sipunculida Phascolosomatidae Phascolosoma agassizii Fisher 1952 I/ST/Inf<br />
II III IV V VI<br />
VIII IX Austin 1985<br />
Tardigrada Hypsibiidae Hypsibius appelloefi<br />
Feder and Bryson-<br />
Schwafel 1988 ST x x x x<br />
Schuster and<br />
Grigarick 1965<br />
Urochordata:<br />
Ascidiacea Ascidiidae Ascidia adhaerens Lambert 2000 I II x x Lambert 2000<br />
Urochordata:<br />
Ascidiacea Ascidiidae Ascidia columbiana A. callosa Lambert 2000 I II III IV x Lambert 2000<br />
Urochordata:<br />
Ascidiacea Ascidiidae Ascidia unidentified Lambert 2000 I/ST/Epi<br />
Urochordata:<br />
Ascidiacea Clavelinidae Distaplia alaskensis Lambert 2000 ST/Epi<br />
Urochordata:<br />
Ascidiacea Clavelinidae Distaplia sp. undescribed Lambert 2000 ST/Epi possible Lambert 2000<br />
Urochordata:<br />
Ascidiacea Corellidae Chelysoma columbiana Austin 1985 ST/Epi II III IV Austin 1985<br />
Urochordata:<br />
Ascidiacea Corellidae Chelysoma productum Lambert 2000 ST/Epi II III IV V VI Austin 1985<br />
Urochordata:<br />
Ascidiacea Corellidae Corella inflata Lambert 2000 I II III IV Austin 1985<br />
Urochordata:<br />
Ascidiacea Corellidae Corella unidentified Lambert 2000 I<br />
Urochordata:<br />
Ascidiacea Corellidae Corella willmeriana Lambert 2000 I II III IV Austin 1985<br />
Urochordata:<br />
Ascidiacea Molgulidae Molgula retortiformis Lambert 2000 I II x x Austin 1985<br />
Urochordata:<br />
Ascidiacea Polyclinidae Ritterella unidentified Lambert 2000 I<br />
Urochordata:<br />
Ascidiacea Pyruridae Boltenia echinata EVOS ST/Epi II III IV x Austin 1985<br />
Urochordata:<br />
Ascidiacea Pyruridae Boltenia ovifera UAM ST/Epi II III IV x Pavloskii 1955<br />
Urochordata:<br />
Ascidiacea Pyruridae Halocynthia igoboja UAM ST/Epi II III IV x Austin 1985<br />
Urochordata:<br />
Ascidiacea Pyruridae Halocynthia auranticum Tethyum<br />
Scheel, Foster, and<br />
Hough, 1997 ST/Epi II III IV x Austin 1985<br />
Urochordata:<br />
Ascidiacea Pyruridae Pyura haustor Lambert 2000 ST/Epi II III IV V VI Austin 1985<br />
Urochordata:<br />
Ascidiacea Styelidae Botrylloides violaceus Lambert 2000 ST/Epi x Japan definate<br />
Urochordata:<br />
Ascidiacea Styelidae Cnemidocarpa finmarkiensis Stylea Rosenthal 1977 ST/Epi II III IV V x x Austin 1985<br />
Urochordata:<br />
Ascidiacea Styelidae Metandrocarpa taylori Rosenthal 1977 ST/Epi II III IV V Austin 1985<br />
Urochordata:<br />
Ascidiacea Styelidae Styela truncata Lambert 2000 I II III IV Austin 1985<br />
Urochordata:<br />
Ascidiacea Clavelinidae Distaplia occidentalis Lambert 2000 ST/Epi III IV V VI<br />
British<br />
Columbia NR Austin 1985<br />
Urochordata:<br />
Larvacea Fritillariidae Fritillaria borealis Cooney 1987 P II III IV V VI x x x Austin 1985<br />
Urochordata:<br />
Larvacea Fritillariidae Fritillaria unidentified Cooney: SEA P<br />
Urochordata:<br />
Larvacea Oikopleuridae Oikopleura labradorensis Cooney 1987 P II III IV V x x x<br />
Urochordata:<br />
Larvacea Oikopleuridae Oikopleura unidentified<br />
Cooney and Coyle<br />
1988 P<br />
Urochordata:<br />
Larvacea Oikopleuridae Oikopleura vanhoeffeni Cooney 1987 P II III x x x<br />
Urochordata Oikopleuridae Oikopleura dioica Cooney 1987 P II III IV V<br />
II III IV V VI<br />
Urochordata Salpida Salpa fusiformis Cooney 1987 P<br />
VII VIII IX x x<br />
II III IV V VI<br />
Urochordata Salpida Salpa maxima Cooney 1987 P<br />
VII VIII IX x x<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Wrobel and Mills<br />
1998<br />
Fishes<br />
The 175 species we list represent 33 families (Table 10.9). The fish data set uses records<br />
for fish specimens catalogued in the University <strong>of</strong> Alaska Museum, as well as published and<br />
manuscript sources. (Baxter unpublished manuscript, Hart 1973). Name changes derive from and<br />
Humann (1996). Two species, the American shad, Alosa sapidissima, and the Atlantic Salmon,<br />
Samo salar are nonindigenous.<br />
Mecklenberg, (letter <strong>of</strong> 10/30/00) recognizes an undescribed cottid, possibly a<br />
Malacocottus. It has been identified incorrectly as Thecopterus aleuticus.
Chapt 10. Biodiversity, page 10- 56<br />
The following habitat descriptors used in the data sets are based on usage in Eschmeyer<br />
et al (1983)and Eschmeyer (1990) and Hart (1973).<br />
Table 10.9. Fishes.<br />
ANA anadromous BW brackish D dermesal<br />
E eelgrass Epip epipelagic Est estuaries<br />
FW freshwater I intertidal KB kelp bed<br />
NS nearshore Off <strong>of</strong>fshore Pel pelagic<br />
PB pebble bottom RI rocky intertidal<br />
RST rocky subtidal SAB sand bottom<br />
SB s<strong>of</strong>tbottom ST subtidal<br />
SW saltwater<br />
BIOREGION<br />
FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Acipenseridae Acipenser medirostris Hart 1973 BW, SW, NS IV x x Hart 1973<br />
Acipenseridae Acipenser transmontanus Hart 1973 ANA IV Hart 1973<br />
Agonidae Anoplagonus inermis UAM D IV x Hart 1973<br />
Agonidae Aspidophoroides bartoni UAM SB, SA IV x Baxter 1987ms<br />
Agonidae Aspidophoroides olrikii UAM SB, SA IV x Baxter 1987ms<br />
Agonidae Bathyagonus infraspinata Asterotheca UAM D, SB IV Hart 1973<br />
Agonidae Bathyagonus nigripinnis UAM D, SB IV x x Hart 1973<br />
Agonidae Bathyagonus pentacanthus Asterotheca Hart 1973 D, SB IV Hart 1973<br />
Agonidae Bathyagonus alascanus<br />
Asterotheca<br />
alascana Hart 1973 RST, D<br />
II III<br />
IV Hart 1973<br />
Agonidae Bothragonus swani Hart 1973 I, ST IV Hart 1973<br />
Agonidae Hypsagonus quadricornis Hart 1973 D, SB IV x x Hart 1973<br />
Agonidae Occella verrucosa Hart 1973 D, SB IV Hart 1973<br />
Agonidae Odontopyxis trispinosa UAM D, SB IV SE Alaska nr Hart 1973<br />
Agonidae Podothecus acipenserinus Agonus Hart 1973 D IV x Hart 1973<br />
Ammodytidae Ammodytes hexapterus UAM I, ST, NS, P IV x x Hart 1973<br />
Anarhichadidae Anarrhichthys ocellatus Hart 1973 RST, D IV x Hart 1973<br />
Anoplopomatidae Anoplopoma fimbria Feder et al. 1979 ST, D IV x x Hart 1973<br />
1, ST, SAB, KB, II III<br />
Aulorhynchidae Aulorhynchus flavidus UAM<br />
E<br />
IV Hart 1973<br />
Bathylagidae Leuroglossus callorhinus L. schmidti UAM P IV x Hart 1973<br />
Bathymasteridae Bathylagus milleri Hart 1973 P IV x Hart 1973<br />
Bathymasteridae Bathylagus ochotensis Hart 1973 P IV x x Hart 1973<br />
Bathymasteridae Bathymaster caerule<strong>of</strong>asciatus Rogers et al. 1986 RST<br />
II III<br />
IV<br />
II III<br />
IV<br />
Eschmeyer et al.<br />
1983<br />
Eschmeyer et al.<br />
1983<br />
Bathymasteridae Bathymaster leurolepis Rogers et al. 1986 I/ST<br />
x<br />
Bathymasteridae Bathymaster signatus UAM D, SB IV x x Hart 1973<br />
Bathymasteridae Ronquilus jordani Hart 1973 RST, D IV Hart 1973<br />
Bothidae Citharichthys sordidus Hart 1973 D, SAB IV Hart 1973<br />
Carcharhinidae Prionace glauca UAM Epip x x x x Hart 1973<br />
Clupeidae Alosa sapidissima UAM ANA III IV x Atlantic definate Hart 1973<br />
Clupeidae Clupea pallasii UAM P, Est II x x Hart 1973<br />
Cottidae Artedius fenestralis Hart 1973 I, ST IV Hart 1973<br />
Cottidae Artedius harringtoni Hart 1973 I, RST IV Hart 1973<br />
Cottidae Artedius lateralis UAM I, ST IV Hart 1973<br />
Cottidae Blepsias bilobus UAM I, ST IV x x Hart 1973<br />
Cottidae Blepsias cirrhosus Hart 1973 I, ST IV x Hart 1973<br />
RI, ST, SAB, E, II III<br />
Cottidae Clinocottus acuticeps UAM<br />
KB<br />
IV Hart 1973<br />
Cottidae Clinocottus embryum UAM RI IV Hart 1973<br />
Cottidae Clinocottus globiceps Hart 1973 RI IV Hart 1973<br />
Cottidae Dasycottus setiger UAM SB IV x Hart 1973<br />
Cottidae Enophrys bison Hart 1973 RST, SB IV Hart 1973<br />
Cottidae Gymnocanthus galeatus UAM SB, NS IV Hart 1973<br />
Cottidae Hemilepidotus hemilepidotus UAM I, RST, NS IV x Hart 1973<br />
Cottidae Hemilepidotus jordani UAM ST IV x Hart 1973<br />
Cottidae Hemilepidotus spinosus Hart 1973 I, ST IV SE Alaska nr Hart 1973<br />
Cottidae Icelinus borealis UAM SB IV Hart 1973<br />
Cottidae Icelus spiniger Hart 1973 SB IV x x Hart 1973<br />
Cottidae Leptocottus armatus UAM I, ST IV Hart 1973<br />
Cottidae Malacocottus kincaidi UAM SB IV x Hart 1973<br />
Cottidae Malacocottus zonurus UAM SB IV x Hart 1973<br />
Cottidae Myoxocephalus polyacanthocephalus Hart 1973 I, NS, SB, SAB IV x Hart 1973<br />
Cottidae Myoxocephalus scorpius<br />
M.<br />
groenlandicus Rogers et al. 1986 ST<br />
II III<br />
IV x x Baxter 1987ms
Chapt 10. Biodiversity, page 10- 57<br />
Table 10.9 continued.<br />
BIOREGION<br />
FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Cottidae Myoxocephalus verrucosus UAM ST IV x Baxter 1987ms<br />
Cottidae Nautichthys ocul<strong>of</strong>asciatus Hart 1973 RST IV x x Hart 1973<br />
Cottidae Oligocottus maculosus Hart 1973 I, RST IV x x Hart 1973<br />
Cottidae Oligocottus rimensis UAM I, RST IV Hart 1973<br />
Cottidae Psychrolutes paradoxus Hart 1973 NS, RST, SB IV x x Hart 1973<br />
Cottidae Psychrolutes sigalutes Gilbertidia Hart 1973 RST, NS, SB IV Hart 1973<br />
Cottidae Radulinus asprellus UAM NS IV Hart 1973<br />
Cottidae Rhamphocottus richardsoni UAM I, RST, SAB IV Hart 1973<br />
Cottidae Malacocottus sp.<br />
Thecopterus<br />
aleuticus UAM RST II Bering Sea nr Baxter 1987ms<br />
Cottidae Triglops macellus Hart 1973 D IV Hart 1973<br />
Cottidae Triglops pingelii UAM D II x x x Hart 1973<br />
Cryptacanthodidae Delolepis giganteus Hart 1973 SB IV Hart 1973<br />
Cryptacanthodidae Lyconectes aleutensis UAM ST, NS, SAB IV Hart 1973<br />
Cyclopteridae Aptocyclus ventricosus Hart 1973 I, ST IV x x Hart 1973<br />
Cyclopteridae Careproctus colletti UAM SB, SA II x Hart 1973<br />
Cyclopteridae Careproctus gilberti Hart 1973 SB, SA IV Hart 1973<br />
Cyclopteridae Careproctus melanurus UAM ST, SB, SAB IV Hart 1973<br />
Cyclopteridae Eumicrotremus orbis UAM I, RST IV x Hart 1973<br />
Cyclopteridae Liparis callyodon Hart 1973 I IV Hart 1973<br />
Cyclopteridae Liparis cyclopus Hart 1973 NS IV Hart 1973<br />
Cyclopteridae Liparis dennyi UAM SB, SA IV Hart 1973<br />
Cyclopteridae Liparis florae UAM I IV Hart 1973<br />
Cyclopteridae Liparis pulchellus Hart 1973 SB IV x Hart 1973<br />
Cyclopteridae Liparis rutteri Hart 1973 I, ST IV Hart 1973<br />
Cyclopteridae Nectoliparis pelagicus UAM P IV x Hart 1973<br />
Cyclopteridae Paraliparis deani UAM D IV x x SE Alaska nr Hart 1973<br />
Gadidae Eleginus gracilis UAM P, D IV x Hart 1973<br />
Gadidae Gadus macrocephalus UAM P, D IV x x Hart 1973<br />
Gadidae Merluccius productus Hart 1973 P, D IV x Hart 1973<br />
Gadidae Microgadus proximus UAM D IV Hart 1973<br />
Gadidae Theragra chalcogramma UAM Off, D IV x x Hart 1973<br />
Gasterosteidae Gasterosteus aculeatus Hart 1973 FW/SW, NS II x Hart 1973<br />
Hexagrammidae Hexagrammos decagrammus Hart 1973 RST, KB, SAB IV Hart 1973<br />
Hexagrammidae Hexagrammos lagocephalus Hart 1973 RST IV x Hart 1973<br />
Hexagrammidae Hexagrammos octogrammus UAM RST IV x x Hart 1973<br />
Hexagrammidae Hexagrammos stelleri UAM RST, NS, E IV x x Hart 1973<br />
Hexagrammidae Ophiodon elongatus Hart 1973<br />
NS, RST, SB,<br />
SAB<br />
II III<br />
IV Hart 1973<br />
Lamnidae Cetorhinus maximus Hart 1973 Epip, P, NS x x x x Hart 1973<br />
Lamnidae Lamna ditropus Hart 1973 Off, Epip, NS IV x x Hart 1973<br />
Osmeridae Hypomesus pretiosus pretiosus Hart 1973 NS IV x Hart 1973<br />
Osmeridae Mallotus villosus Hart 1973 Oceanic II x x x Hart 1973<br />
Osmeridae Spirinchus thaleichthys Hart 1973 ANA IV Hart 1973<br />
Osmeridae Thaleichthys pacificus UAM ANA IV x Hart 1973<br />
Petromyzontidae Lampetra japonica Rogers et al. 1986 ANA IV Baxter 1987ms<br />
Petromyzontidae Lampetra tridentata Hart 1973 ANA IV x x Hart 1973<br />
Pholidae Apodichthys flavidus Hart 1973 I/RT IV Hart 1973<br />
Pholidae Pholis gilli UAM I/RT IV Hart 1973<br />
Pholidae Pholis laeta UAM I, ST IV Hart 1973<br />
Pleuronectidae Atheresthes stomias UAM SB, Off IV Hart 1973<br />
Pleuronectidae Eopsetta jordani Hart 1973 SAB IV Hart 1973<br />
Pleuronectidae Glyptocephalus zachirus UAM SB, SAB IV Hart 1973<br />
Pleuronectidae Hippoglossoides elassodon UAM SB IV x x Hart 1973<br />
Pleuronectidae Hippoglossus stenolepis Rogers et al. 1986 ST, NS<br />
II III<br />
IV x<br />
Eschmeyer et al.<br />
1983<br />
Pleuronectidae Iopsetta ischyra Hart 1973 NS, Off IV Hart 1973<br />
Pleuronectidae Limanda aspera UAM NS IV x x Hart 1973<br />
Pleuronectidae Limanda proboscidea UAM SB IV Hart 1973<br />
Pleuronectidae Microstomus pacificus UAM SB IV Hart 1973<br />
Pleuronectidae Platichthys stellatus UAM NS, Est IV x x Hart 1973<br />
Pleuronectidae Pleuronectes bilineata Lepidosetta UAM NS, Off, PB IV x x Hart 1973<br />
Pleuronectidae Pleuronectes vetulus Parophrys UAM I, ST IV Hart 1973<br />
Pleuronectidae Pleuronichthys decurrens Feder et al. 1979 SB IV Hart 1973<br />
Pleuronectidae Psettichthys melanostictus Hart 1973 NS IV Hart 1973<br />
Pleuronectidae Reinhardtius hippoglossoides Hart 1973 SAB, PB II x x x Hart 1973<br />
Rajidae Bathyraja kincaidi Raja UAM D IV Hart 1973<br />
Rajidae Bathyraja aleutica Wilimovsky 1958 D II III x Wilimovsky 1958<br />
Rajidae Bathyraja interrupta Escheyer et al. 1983<br />
Rajidae Raja binoculata Escheyer et al. 1983 D<br />
II III<br />
IV<br />
II III<br />
IV<br />
Escheyer et al.<br />
1983<br />
Escheyer et al.<br />
1983<br />
Escheyer et al.<br />
1983<br />
Rajidae Raja parmifera Escheyer et al. 1983 D II III x<br />
Rajidae Raja rhina Bathyraja Feder et al. 1979 D IV Hart 1973<br />
Rajidae Bathyraja stellulata Raja Hart 1973 D IV Hart 1973<br />
Salmonidae Oncorhynchus clarki Salmo clarki Hart 1973 ANA, FW IV Hart 1973<br />
Salmonidae Oncorhynchus gorbuscha UAM ANA IV x Hart 1973<br />
Salmonidae Oncorhynchus keta Hart 1973 ANA IV x x Hart 1973
Chapt 10. Biodiversity, page 10- 58<br />
Table 10.9 continued.<br />
BIOREGION<br />
FAMILY GENUS SPECIES OTHER NAMES<br />
SPECIMEN or<br />
SOURCE HABITAT NEP NWP AR NWA ORIGIN NIS STATUS<br />
REFERENCE to<br />
DISTRIBUTION<br />
Salmonidae Oncorhynchus kisutch Hart 1973 ANA IV x Hart 1973<br />
Salmonidae Oncorhynchus mykiss Salmo gardnerii Hart 1973 ANA IV x Hart 1973<br />
Salmonidae Oncorhynchus nerka Hart 1973 ANA IV x x Hart 1973<br />
Salmonidae Oncorhynchus tshawytscha Hart 1973 ANA IV x x Hart 1973<br />
Salmonidae Salmo salar Hart 1973 ANA III IV x<br />
British<br />
Columbia definate Hines et al. 2000<br />
Salmonidae Salvelinus malma Hart 1973 ANA IV x x Hart 1973<br />
Scombridae Sarda chiliensis Hart 1973 NS IV x Hart 1973<br />
Scorpaenidae Sebastes aleutianus UAM D IV Hart 1973<br />
Scorpaenidae Sebastes alutus UAM Off IV x Hart 1973<br />
II III<br />
Eschmeyer et al.<br />
Scorpaenidae Sebastes auriculatus Rogers et al. 1986 ST, NS IV SE Alaska nr<br />
1983<br />
Scorpaenidae Sebastes borealis UAM SB IV x Hart 1973<br />
Scorpaenidae Sebastes brevispinis Hart 1973 D IV Hart 1973<br />
Scorpaenidae Sebastes caurinus Hart 1973 RST, SAB, NS IV Hart 1973<br />
Scorpaenidae Sebastes ciliatus Hart 1973 D, NS IV x Hart 1973<br />
Scorpaenidae Sebastes crameri Hart 1973 SB IV Hart 1973<br />
Scorpaenidae Sebastes diploproa Hart 1973 Off, SB IV Hart 1973<br />
Scorpaenidae Sebastes elongatus Hart 1973 RST, SB IV Hart 1973<br />
Scorpaenidae Sebastes emphaeus Rogers et al. 1986 ST, NS IV Baxter 1987ms<br />
Scorpaenidae Sebastes flavidus Hart 1973 P IV Hart 1973<br />
Scorpaenidae Sebastes maliger Hart 1973 RST, NS IV Hart 1973<br />
Scorpaenidae Sebastes melanops Hart 1973 D, NS, RR IV Hart 1973<br />
Scorpaenidae Sebastes mystinus Hart 1973 D, RR, KB IV Hart 1973<br />
Scorpaenidae Sebastes nebulosus Rogers et al. 1986 NS, RST, RR IV SE Alaska nr Baxter 1987ms<br />
Scorpaenidae Sebastes nigrocinctus Rogers et al. 1986 RST IV SE Alaska nr Baxter 1987ms<br />
Scorpaenidae Sebastes paucispinis Hart 1973 D, RR IV Hart 1973<br />
Scorpaenidae Sebastes proriger Hart 1973 D IV Hart 1973<br />
Scorpaenidae Sebastes ruberrimus Feder et al. 1979 RR IV Hart 1973<br />
Scorpaenidae Sebastes variegatus UAM D IV Hart 1973<br />
Scorpaenidae Sebsatsolobus alascanus Hart 1973 SB IV x Hart 1973<br />
Scorpaenidae Sebsatsolobus altivelis Hart 1973 Off, SB IV Hart 1973<br />
Scytalinidae Scytalina cerdale Hart 1973 I, NS SAB, PB IV Hart 1973<br />
Sphyraenidae Sphyraena argentea Hart 1973 NS IV Hart 1973<br />
Squalidae Somniosus pacificus Hart 1973 I, P, D IV x x Hart 1973<br />
Squalidae Squalus acanthias Hart 1973 P, D II x x Hart 1973<br />
Stichaeidae Anoplarchus insignis Hart 1973 RST IV Hart 1973<br />
Stichaeidae Anoplarchus purpurescens UAM I, RST IV Hart 1973<br />
Stichaeidae Chirolophis decoratus Hart 1973 RST, KB IV Hart 1973<br />
Stichaeidae Chirolophis nugator Rogers et al. 1986 RST<br />
II III<br />
IV<br />
Eschmeyer et al.<br />
1983<br />
Stichaeidae Lumpenella longirostris UAM Off IV Hart 1973<br />
Stichaeidae Lumpenus maculatus UAM SAB IV x Hart 1973<br />
Stichaeidae Lumpenus sagitta UAM Off, NS IV x Hart 1973<br />
Stichaeidae Phytichthys chirus Hart 1973 I, RST IV Hart 1973<br />
Stichaeidae Poroclinus rothrocki Hart 1973 SB, SA IV Hart 1973<br />
Stichaeidae Stichaeus punctatus UAM RST, SAB II x x x Hart 1973<br />
Stichaeidae Xiphister mucosus UAM I, RST IV Hart 1973<br />
Trichodontidae Trichodon trichodon UAM NS, SB SAB IV x x Hart 1973<br />
Zaproridae Zaprora silenus Hart 1973 D IV x x Hart 1973<br />
Zoarcidae Bothrocara molle UAM SB, SA IV x Hart 1973<br />
Zoarcidae Lycodapus mandibularis UAM P IV Hart 1973<br />
Zoarcidae Lycodes brevipes UAM SB, SAB IV x x Hart 1973<br />
Zoarcidae Lycodes diapterus UAM SB IV x x Hart 1973<br />
Zoarcidae Lycodes palearis UAM SB- SAB IV x Hart 1973<br />
Zoarcidae Lycodopsis pacifica Feder et al. 1979 P IV Hart 1973<br />
Stichaeidae Xiphister atropurpureus Hart 1973 I, RST IV Hart 1973<br />
Birds<br />
The rich bird fauna <strong>of</strong> Prince William Sound, the northern Gulf <strong>of</strong> Alaska coast and<br />
Copper River Delta has been described in detail by Isleb and Kessel (1973, 1989). From their<br />
checklist, we have selected 114 species with habitats designated “ beaches and tidal flats, rocky<br />
shores, and reefs, inshore waters, and <strong>of</strong>fshore waters” (Table 10.10).<br />
The American Ornithologists’ Union’s (1983) Check-list <strong>of</strong> North American Birds, was<br />
used to update common and scientific names. Usage <strong>of</strong> terms that define status: migrant, visitor,<br />
breeder and resident is derived from Isleb and Kessel (1973).<br />
Nsh nearshore m migrant<br />
Pel pelagic v visitor<br />
Int intertidal r rare<br />
b breeder
Chapt 10. Biodiversity, page 10- 59<br />
Table 10.10. Birds.<br />
FAMILY GENUS SPECIES COMMON NAME<br />
OTHER<br />
NAMES SOURCE HABITAT STATUS<br />
Gaviidae Gavia immer Common Loon Isleib and Kessel 1973 nsh r<br />
Gaviidae Gavia adamsii Yellow-billed Loon Isleib and Kessel 1973 nsh r<br />
Gaviidae Gavia arctica Arctic Loon Isleib and Kessel 1973 nsh r<br />
Gaviidae Gavia stellata Red-throated Loon Isleib and Kessel 1973 nsh r<br />
Podicipedidae Podiceps grisegena Red-necked Grebe Isleib and Kessel 1973 nsh r<br />
Podicipedidae Podiceps auritis Horned Grebe Isleib and Kessel 1973 nsh r<br />
Porcellariidae Fulmarus glacialis Fulmar Isleib and Kessel 1973 pel v<br />
Porcellariidae Puffinus creatopus Pink-footed Shearwater Isleib and Kessel 1973 pel v<br />
Porcellariidae Puffinus carneipes Pale-footed Shearwater Isleib and Kessel 1973 pel v<br />
Porcellariidae Puffinus griseus Sooty Shearwater Isleib and Kessel 1973 pel v<br />
Porcellariidae Puffinus tenuirostris Slender-billed Shearwater Isleib and Kessel 1973 pel v<br />
Porcellariidae Oceandroma furcata Fork-tailed Storm Petrel Isleib and Kessel 1973 pel r<br />
Phalacrocoracidae Phalacrocorax auritius Double-crested Cormorant Isleib and Kessel 1973 nsh/int r<br />
Phalacrocoracidae Phalacrocorax penicillatus Brandt’s Cormorant Isleib and Kessel 1973 nsh/int b<br />
Phalacrocoracidae Phalacrocorax peligicus Pelagic Cormorant Isleib and Kessel 1973 nsh/int r<br />
Phalacrocoracidae Phalacrocorax urile Red-faced Cormorant Isleib and Kessel 1973 nsh/int r<br />
Ardeidae Ardea herodias Great blue Heron Isleib and Kessel 1973 int r<br />
Anatidae Cygnus columbianus Whistling Swan Olor Isleib and Kessel 1973 int m<br />
Anatidae Cygnus buccinator Trumpeter Swam Olor Isleib and Kessel 1973 int r<br />
Anatidae Branta canadensis Canada Goose Isleib and Kessel 1973 int r<br />
Anatidae Branta bernicula Brant B. nigricans Isleib and Kessel 1973 int m<br />
Anatidae Chen canagica Emperor Goose Philacate Isleib and Kessel 1973 int m<br />
Anatidae Anser albifrons Greater White-fronted Goose Isleib and Kessel 1973 int m<br />
Anatidae Chen caerulescens Snow Goose<br />
C.<br />
hyperborea Isleib and Kessel 1973 int m<br />
Anatidae Anas platyrhynchos Mallard Isleib and Kessel 1973 int r<br />
Anatidae Anas strepera Gadwall Isleib and Kessel 1973 int r<br />
Anatidae Anas acuta Northern Pintail Isleib and Kessel 1973 int r<br />
Anatidae Anas crecca Green-winged Teal<br />
A.<br />
carolinensis Isleib and Kessel 1973 int r<br />
Anatidae Anas discors Blue-winged Teal Isleib and Kessel 1973 int m<br />
Anatidae Anas penelope Europeam Widgeon Mareca Isleib and Kessel 1973 int m<br />
Anatidae Anas americana American Widgeon Mareca Isleib and Kessel 1973 int r<br />
Anatidae Anas clypeata Northern Shoveler Spatula Isleib and Kessel 1973 int m/b<br />
Anatidae Aythya valisineria Canvasback Isleib and Kessel 1973 int m/b<br />
Anatidae Aythya marila Greater Scaup Isleib and Kessel 1973 int r<br />
Anatidae Bucephala clangula Common Goldeneye Isleib and Kessel 1973 int/nsh r<br />
Anatidae Bucephala islandica Barrow’s Goldeneye Isleib and Kessel 1973 int/nsh r<br />
Anatidae Bucephala albeola Bufflehead Isleib and Kessel 1973 int/nsh r<br />
Anatidae Clangula hymenalis Oldsquaw Isleib and Kessel 1973 int/nsh r<br />
Anatidae Histrionicus histrionicus Harlequin Duck Isleib and Kessel 1973 int/nsh r<br />
Anatidae Polysticta stelleri Steller’s Eider Isleib and Kessel 1973 int/nsh v<br />
Anatidae Somateria mollissima Common Eider Isleib and Kessel 1973 int/nsh v<br />
Anatidae Somateria spectabilis King Eider Isleib and Kessel 1973 int/nsh v<br />
Anatidae Melanitta fusca White-winged Scoter M. deglandi Isleib and Kessel 1973 int/nsh r<br />
Anatidae Melanitta perspicillata Surf Scoter Isleib and Kessel 1973 int/nsh r<br />
Anatidae Melanitta nigra Black Scoter Oidema Isleib and Kessel 1973 int/nsh r<br />
Anatidae Lophodytes cucullatus Hooded Merganser Isleib and Kessel 1973 int/nsh r<br />
Anatidae Mergus merganser Common Merganser Isleib and Kessel 1973 int/nsh<br />
Anatidae Mergus serrator Red-breated Merganser Isleib and Kessel 1973 int/nsh r<br />
Accipitridae Haliaeetus leucocephalus Bald Eagle Isleib and Kessel 1973 int r<br />
Accipitridae Circus cyaneus Marsh Hawk Isleib and Kessel 1973 int r<br />
Accipitridae Pandion haliaetus Osprey P. rusticolus Isleib and Kessel 1973 int r<br />
Accipitridae Falco peregrinus Peregrine Falcon Isleib and Kessel 1973 int r<br />
Accipitridae Falco columbarius Merlin Isleib and Kessel 1973 int r<br />
Gruidae Grus canadensis Sandhill Crane Isleib and Kessel 1973 int r<br />
Hematopodidae Haematopus bachmani Black Oystercatcher Isleib and Kessel 1973 int r<br />
Caradriidae Charadrius semipalmatus Semipalmated Plover Isleib and Kessel 1973 int m/b<br />
Caradriidae Charadrius vociferus Killldeer Isleib and Kessel 1973 int v<br />
Caradriidae Pluvialis dominica American Golder Plover Isleib and Kessel 1973 int m/b<br />
Caradriidae Pluvialis squatarola Black-bellied Plover Squatarola Isleib and Kessel 1973 int m<br />
Caradriidae Aphriza virgata Surfbird Isleib and Kessel 1973 int r<br />
Caradriidae Arinaria interpres Ruddy Turnstone Isleib and Kessel 1973 int m
Chapt 10. Biodiversity, page 10- 60<br />
Table 10.10 continued.<br />
FAMILY GENUS SPECIES COMMON NAME<br />
OTHER<br />
NAMES SOURCE HABITAT STATUS<br />
Caradriidae Arinaria melanocephala Black Turnstone Isleib and Kessel 1973 int r<br />
Caradriidae Galligano galligano Common Snipe Capella Isleib and Kessel 1973 int r<br />
Caradriidae Numenius phaeopus Whimbrel Isleib and Kessel 1973 int m<br />
Caradriidae Actitus macularia Spotted Sandpiper Isleib and Kessel 1973 int m<br />
Caradriidae Trigna solitaria Solitary Sandpiper Isleib and Kessel 1973 int m<br />
Caradriidae Heteroscelus incanus Wandering Tattler Isleib and Kessel 1973 int m<br />
Caradriidae Tringa melanoleuca Greater Yellowlegs Tatonus Isleib and Kessel 1973 int m<br />
Caradriidae Tringa flavipes Lesser Yellowlegs Tatonus Isleib and Kessel 1973 int m<br />
Caradriidae Calidris canutus Red Knot Isleib and Kessel 1973 int m<br />
Caradriidae Calidris ptilocnemis Rock Sandpiper Erolia Isleib and Kessel 1973 int m<br />
Caradriidae Calidris acuminata Sharp-tailed Sandpiper Erolia Isleib and Kessel 1973 int m<br />
Caradriidae Calidris melanotos Pectoral Sandpiper Erolia Isleib and Kessel 1973 int m<br />
Caradriidae Calidris baridii Baird’s Sandpiper Erolia Isleib and Kessel 1973 int m<br />
Caradriidae Calidris minutilla Least Sandpiper Erolia Isleib and Kessel 1973 int m/b<br />
Caradriidae Calidris alpina Dunlin Erolia Isleib and Kessel 1973 int r<br />
Caradriidae Limnodromus griseus Short-billed Dowitcher Isleib and Kessel 1973 int m/b<br />
Caradriidae Limnodromus scolopaceus Long-billed Dowitcher Isleib and Kessel 1973 int m<br />
Caradriidae Calidris pusilla Semipaliated Sandpiper Ereunetes Isleib and Kessel 1973 int m<br />
Caradriidae Calidris mauri Western Sandpiper Ereunetes Isleib and Kessel 1973 int m<br />
Caradriidae Limosa lapponica Bar-tailed Godwit Isleib and Kessel 1973 int m<br />
Caradriidae Limosa haemastica Hudsonian Godwit Isleib and Kessel 1973 int<br />
Caradriidae Calidris alba Sanderling Crocethia Isleib and Kessel 1973 int m<br />
Caradriidae Phalaropus fulicarius Red Phalarope Isleib and Kessel 1973 int/nsh/pel m<br />
Caradriidae Lobipes lobatus Northern Phalarope Isleib and Kessel 1973 int/nsh/pel m/b<br />
Stercorariidae Stercorarius pomarinus Pomarine Jaeger Isleib and Kessel 1973 pel m<br />
Stercorariidae Stercorarius parasiticus Parasitic Jaeger Isleib and Kessel 1973 pel m/b<br />
Stercorariidae Stercorarius longicaudus Long-tailed Jaeger Isleib and Kessel 1973 pel m<br />
Stercorariidae Catharacta skua Greater Skua Isleib and Kessel 1973 pel v<br />
Laridae Larus hyperboreus Glaucus Gull Isleib and Kessel 1973 int/nsh/pel r<br />
Laridae Larus glaucesens Glaucous-winged Gull Isleib and Kessel 1973 int/nsh/pel r<br />
Laridae Larus argentatus Herring Gull Isleib and Kessel 1973 int/nsh/pel r<br />
Laridae Larus canus Mew Gull Isleib and Kessel 1973 int/nsh/pel r<br />
Laridae Larus philadelphia Bonaparte’s Gulls Isleib and Kessel 1973 int/nsh/pel m/b<br />
Laridae Rissa tridactyla Black-legged Kittywake Isleib and Kessel 1973 int/nsh/pel r<br />
Laridae Xema sabini Sabine’s Gull Isleib and Kessel 1973 int/nsh/pel m<br />
Laridae Sterna paradisaea Arctic Tern Isleib and Kessel 1973 int/nsh/pel m/b<br />
Laridae Sterna aleutica Aleutian Tern Isleib and Kessel 1973 int/nsh/pel b<br />
Alcidae Uria aalge Common Murre Isleib and Kessel 1973 nsh/pel r<br />
Alcidae Uria lomvia Thick-billed Murre Isleib and Kessel 1973 nsh/pel r<br />
Alcidae Cepphus columba Pigeon Guillemont Isleib and Kessel 1973 nsh/pel r<br />
Alcidae Brachyramphus marmoratus Marbled Murrelet Isleib and Kessel 1973 nsh/pel r<br />
Alcidae Brachyramphus brevirostis Kitlitz’s Murrelet Isleib and Kessel 1973 nsh/pel r<br />
Alcidae Synthliboramphus antiquum Ancient Murrelet Isleib and Kessel 1973 nsh/pel r<br />
Alcidae Aethia psittacula Parakeet Auklet<br />
Synthlibora<br />
mphus Isleib and Kessel 1973 nsh/pel b<br />
Alcidae Aethia cristatella Crested Auklet Isleib and Kessel 1973 nsh/pel v<br />
Alcidae Cerorhica monocrata Rhinoceros Auklet Isleib and Kessel 1973 nsh v/b<br />
Alcidae Fratercula corniculata Horned Puffin Isleib and Kessel 1973 nsh/pel r<br />
Alcidae Lunda cirrhata Tufted Puffin Isleib and Kessel 1973 nsh/pel r<br />
Alcedinidae Megaceryle alcyon Belted Kingfisher Isleib and Kessel 1973 int r<br />
Corvidae Cyanocitta stelleri Steller’s Jay Isleib and Kessel 1973 int r<br />
Corvidae Corvus corax Common Raven Isleib and Kessel 1973 int r<br />
Corvidae Corvus caurinus Northwestern Crow Isleib and Kessel 1973 int r<br />
Mammals<br />
Eleven marine mammal species are well known from Prince William Sound. We also<br />
include the river otter, and black and brown bear because <strong>of</strong> their use <strong>of</strong> marine resources<br />
(salmon, intertidal animals) (Table 10.11). Wynne (1992) and UAM mammal collection records<br />
were used as a source for species names. Distribution, community, and status are inferred from<br />
range maps in Wynne (1992).
Chapt 10. Biodiversity, page 10- 61<br />
Table 10.11. Mammals.<br />
FAMILY GENUS SPECIES COMMON NAME<br />
BIOREGION<br />
SPECIMEN or<br />
SOURE HABITAT STATUS NEP NWP AR NWA Source<br />
Balaenopteridae Balaenoptera acutorostrata Minke Whale Wynne 1992 nsh/pel m x x x x Wynne 1992<br />
Balaenopteridae Balaenoptera physalus Fin Whale Wynne 1992 nsh/pel m x x x x Wynne 1992<br />
Balaenopteridae Megaptera novaeangliae Humpback Whale Wynne 1992 nsh m x x x x Wynne 1992<br />
Delphinidae Orcinus orca Killer Whale Wynne 1992 nsh/pel r/m x x x x Wynne 1992<br />
Eschrichtiidae Eschrichtius robustus Grey Whale Wynne 1992 pel m II III IV Wynne 1992<br />
Mustelidae Enhydra lutris Sea Otter Wynne 1992 int/nsh r II III IV x Hall 1981<br />
Mustelidae Lontra canadensis River Otter UAM Mammals int r x x x x Hall 1981<br />
Otariidae Callorhinus ursinus Northern Fur Seal UAM Mammals pel m II III IV x Hall 1981<br />
Otariidae Eumetopias jubatus Steller’s Sea Lion UAM Mammals nsh r II III IV x Hall 1981<br />
Phocidae Phoca vitulina Harbor Seal UAM Mammals nsh r II III IV Hall 1981<br />
Phoecoenidae Phocoena phocoena Harbor Porpoise Wynne 1992 nsh m/r II x x Wynne 1992<br />
Phoecoenidae Phocoenoides dalli Dall’s Porpoise Wynne 1992 nsh/pel m/r II III IV x Wynne 1992<br />
Ursidae Ursus americanus Black Bear UAM Mammals int r II III Hall 1981<br />
Ursidae Ursus arctos Brown Bear UAM Mammals int r II III Hall 1981<br />
10E. Discussion<br />
The data sets contain entries for 1878 taxa, <strong>of</strong> which all but 180 are species-level<br />
identifications. Thirty-nine species are recognized as possibly undescribed. Eighty-nine species<br />
had not previously been reported in Prince William Sound, and represent distributional range<br />
extension. (Table 10.12.) These species were probably overlooked in previous environmental<br />
surveys, because <strong>of</strong> their small size, cryptic appearance, similarity to more well-known species,<br />
or lack <strong>of</strong> taxonomic experts to work with them. Eighteen species are likely to represent<br />
nonindigenous species. At least 97 species are uncertain in origin (cryptogenic). Over 70% <strong>of</strong> the<br />
animals (1343) are invertebrates. Arthropods 24.3%, mollusks 16.8%, and polychaete annelids<br />
12.4% make up the largest number <strong>of</strong> total species. (Table 10.13. Figure 10.4).<br />
Plants<br />
Cnidaria/<br />
Ctenophora<br />
Nemertea<br />
Polychaeta<br />
Mollusca<br />
Arthropoda<br />
probable NIS 5 1 2 1 9<br />
definate NIS 1 1 2 1 1 1 2 9<br />
range extension<br />
within Alaska 4 9 1 8 1 8 31<br />
range extension to<br />
Alaska 17 2 19 7 10 2 1 58<br />
cryptogenic 68 7 1 6 82<br />
potential new or<br />
undescribed species 1 6 2 1 1 11<br />
not identified to<br />
species 10 27 1 7 1 109 19 5 3 182<br />
total spcies-level<br />
taxa 231 102 45 233 315 455 68 74 30 23 178 113 14 1881<br />
Echinodermata<br />
Bryozoa<br />
Urochordata<br />
Other<br />
invertebrates<br />
Fishes<br />
Birds<br />
Mammals<br />
TOTALS<br />
Table 10.12. Summary <strong>of</strong> Prince William Sound marine biota by NIS status and identification confidence.
Chapt 10. Biodiversity, page 10- 62<br />
Plants<br />
Cnidaria/ Ctenophor<br />
Mollusca<br />
numer <strong>of</strong> species 231 102 315 233 455 66 74 98 178 113 14<br />
percent <strong>of</strong> total fauna 6.2% 19.1% 14.2% 27.7% 4.0% 4.5% 6.0% 10.6% 6.9% 0.9%<br />
percent <strong>of</strong> total biota 12.3% 5.4% 16.8% 12.4% 24.3% 3.5% 3.9% 5.2% 9.3% 6.0% 0.7%<br />
total invertebrates 1343<br />
total vertebrates 302<br />
total plants 233<br />
total biota considered 1876<br />
Polychaeta<br />
Arthropoda<br />
Echinodermata<br />
Bryozoa<br />
Other invertebrates<br />
Fishes<br />
Birds<br />
Mammals<br />
Table 10.13. Summary <strong>of</strong> the plants and animals considered.<br />
Marine Plants<br />
Ascomycota<br />
Cyanophyta<br />
Xanthophyta<br />
Chlorophyta<br />
Rhodophyta<br />
Phaeophyta<br />
vascular plants<br />
Marine Biota<br />
total invertebrates<br />
total vertebrates<br />
total plants<br />
Marine Invertebrates<br />
Cnidaria/Ctenophora<br />
Polychaeta<br />
Mollusca<br />
Arthropoda<br />
Echinodermata<br />
Bryozoa<br />
other invertebrate phyla<br />
Vertebrates<br />
fishes<br />
birds<br />
mammals<br />
Figure 10.4. Species composition <strong>of</strong> Prince William Sound marine biota by major taxon.
Chapt 10. Biodiversity, page 10- 63<br />
The marine biota <strong>of</strong> Prince William Sound is a mix <strong>of</strong> species with ranges that overlap<br />
several biogeographic provinces. Forty-nine percent <strong>of</strong> the biota for which reliable data are<br />
available are northeast Pacific species, not found north or west <strong>of</strong> Bering Strait. Over one third<br />
<strong>of</strong> the species (43.4 %) have geographic ranges that extend into the northwestern Pacific. Fewer<br />
species have distributions that overlap the north Atlantic 19.6.5% and/or Arctic 16.9%. The<br />
biogeographic affinities vary greatly among the phyla.<br />
We caution users <strong>of</strong> these data sets that in spite <strong>of</strong> our best efforts the information is only<br />
as good as our sources. We were unable to obtain data, or found few reliable records, for five<br />
major groups <strong>of</strong> animals: sponges, Anthozoa, Nematoda, Oligochaeta , Hirudinea, Ostracoda and<br />
insects. Two factors account for this, lack <strong>of</strong> taxonomic experts, and the need for specialized<br />
collection and preservation techniques. Further, even among several well-known taxonomic<br />
groups, we found few sources <strong>of</strong> authentic information and so we have had to rely on<br />
unpublished reports and manuscripts. We hope that using these data sets will not perpetuate<br />
erroneous records. In critical situations, users are advised to rely on museum records and<br />
consultation with other experts.<br />
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