Report of the
Puget Sound Expedition
September 8-16, 1998
A Rapid Assessment Survey of Non-indigenous Species
in the Shallow Waters of Puget Sound
Prepared by
Andrew Cohen, Claudia Mills, Helen Berry, Marjorie Wonham,
Brian Bingham, Betty Bookheim, James Carlton, John Chapman,
Jeff Cordell, Leslie Harris, Terrie Klinger, Alan Kohn, Charles
Lambert, Gretchen Lambert, Kevin Li, David Secord and
Jason Toft
For the
Washington State Department of Natural Resources, Olympia, WA
United States Fish and Wildlife Service, Lacey, WA
November 1998
For additional copies of this report, contact:
Nearshore Habitat Program
Aquatic Resources Division
Washington State Department of Natural Resources
1111 Washington Street SE
PO Box 47027
Olympia, WA 98504-7027
(360) 902-1100
Report of the Puget Sound Expedition
Sept. 8-16, 1998
Contents
Summary .............................................................................................................1
Methods...............................................................................................................2
Non-indigenous Species Collected .......................................................................5
Future Research and Reports, Research Needs ................................................... 17
Acknowledgments ............................................................................................. 18
Literature Cited.................................................................................................. 19
Tables
1 Non-indigenous and Cryptogenic Species Collected .....................................6
2 Origins, First Records and Mechanisms of Introduction................................8
3 Biotic Communities at Sampling Sites ........................................................ 11
4 Puget Sound vs. San Francisco Bay Expeditions......................................... 14
5 Non-indigenous Species in Puget Sound ..................................................... 16
Figures
1 Map of Study Sites .......................................................................................3
2 Non-indigenous Species vs. Surface Salinity.................................................9
3 Non-indigenous Species vs. Surface Temperature....................................... 10
4 Non-indigenous species by Region ............................................................. 10
Appendices
1 List of Participants...................................................................................... 21
2 Expedition Schedule ................................................................................... 23
3 Description of Sampling Sites..................................................................... 25
4 Equipment List ........................................................................................... 29
5 Collections by Station of Non-indigenous Species ...................................... 31
6 Ascidian Distribution.................................................................................. 33
7 Commentary on Species of Hydrozoa, Scyphozoa and Anthozoa................ 35
Summary
A Rapid Assessment survey for non-indigenous marine organisms, based on techniques
pioneered in San Francisco Bay, was conducted in Puget Sound1 on September 8-16, 1998.
Twenty-three primary stations and nine secondary stations were surveyed, mainly consisting of
dock-fouling stations and adjacent shallow water benthic habitats. Material was sampled by a
variety of techniques and examined in the field and laboratory by a multi-institutional team
covering a broad range of taxonomic expertise. The Expedition team included core researchers
from the four San Francisco Bay Expeditions conducted in 1993-1997 and marine scientists from
the Puget Sound region. The Expedition was jointly organized by team members from the
Washington State Department of Natural Resources, the University of Washington (UW) Friday
Harbor Laboratories and the San Francisco Estuary Institute. Laboratory work was conducted at
the King County Environmental Laboratory and the UW Friday Harbor Laboratories. Direct and
in-kind contributions in support of the Expedition are listed in the Acknowledgments.
The Expedition collected and identified 39 non-indigenous species in six days of sampling. The
number of non-indigenous species collected per site showed no obvious pattern with regard to
salinity, temperature or oceanographic basin. Eleven of the non-indigenous species collected by
the Expedition are new records for Puget Sound, and at least another five are previously
unpublished. Several prior lists of non-indigenous marine species in Puget Sound and adjoining
waters have been produced:
• Carlton (1979) listed 18 species of non-indigenous invertebrates in Puget Sound. The Puget
Sound Expedition collected a total of 36 non-indigenous invertebrates, 24 of which were not
listed by Carlton.
• Elston (1997) listed 31 non-indigenous marine species in the shared inland waters of British
Columbia and Washington, 14 of which apparently represent valid established species in
Puget Sound. The Expedition collected 30 non-indigenous species that are not on Elston's list.
• Ruiz and Hines (1997) list 67 non-indigenous species in the marine and estuarine waters of
Washington and British Columbia. Based in large part on the Ruiz and Hines list, the
Washington Department of Fish and Wildlife posted on the Internet2 a list of 78 "nonindigenous marine species of Washington State and adjacent waters" (adjacent waters
apparently referring to the coastal waters of British Columbia). Fifty-one of the species on
these lists appear to reflect potentially valid records of established non-indigenous species in
this region3, 27 of which appear to reflect valid records from Puget Sound. The Expedition
collected 14 non-indigenous species that are not on either of these lists.
Overall, fewer non-indigenous species were collected in Puget Sound than were collected by
1
For the purpose of this study, the Puget Sound Ambient Monitoring Program’s definition of Puget Sound was used, it includes
Washington State’s inland marine waters east of Cape Flattery.
2
June 17, 1998 update posted at www.wa.gov/wdfw/fish/nuisance/ans4.htm
3
Species were deleted from the lists that are not established in these waters, are purely freshwater species, are
included elsewhere on the list under another name, that we judge to be better categorized as cryptogenic, or that were
apparently listed as present in these waters based solely on their inclusion in Kozloff (1987).
Page 1
similar expeditions in San Francisco Bay. We developed an updated and corrected list of 52 nonindigenous salt and brackish water species that appear to be present and established in Puget
Sound. This and other species lists in this report should be considered provisional, pending
additional taxonomic work and review.
Methods
The Rapid Assessment survey focused primarily on non-quantitative or semi-quantitative
sampling of dock fouling (organisms growing on the sides and undersides of floating docks and
associated floats, bumpers, tires, ropes, etc.). Field identification of specimens was followed by
examination of sampled material in the laboratory by a team of taxonomic experts. Sampling
dock fouling has the following advantages:
• The habitat is easily sampled at low cost and with simple equipment.
• It can be sampled without regard to the tide level.
• There is easy and quick access to a large selection of suitable sites throughout Puget Sound.
• Most sites provide an adequate working area for a sizeable team of experts to sample
simultaneously, while remaining in verbal contact.
• In many coastal regions, the dock fouling fauna has been found to include a significant nonindigenous or cryptogenic4 component.
Dock sampling sites were selected to obtain broad coverage in terms of spatial distribution, land
use, salinity and temperature (Figure 1. Map of Study Sites). Site selection was made by Claudia
Mills, assisted by Helen Berry and Betty Bookheim, and involved preliminary examination of
about 60 potential sites from the South Sound to the Canadian border.
Dock-fouling organisms were sampled by a variety of simple manual techniques. Tools included
hand scrapers, sieves, a long-handled scraper with a fine steel mesh net, and a long-handled (2.4
meter pole) net with 1 mm mesh. A sample of live bay mussels (Mytilus sp.) was obtained and
frozen from each site where they were present. At most dock sites, benthic (bottom) and
plankton samples were also taken.5 An Ekman grab was used to obtain non-quantitative bottom
samples that were sieve-washed and sorted on site; unsorted bottom samples were retained for
later examination for foraminifera and other microfauna. A custom-made cylindrical benthic
sampler fitted with 1 mm stainless steel mesh walls was thrown out on a line and retrieved by
dragging along the bottom, working like a small benthic sled to collect larger infauna. Vertical
plankton hauls were taken with a 0.5 m, 102 µm mesh net with a 211 µm mesh cod end.
Horizontal plankton tows were taken by pulling a plankton net fitted with 125 µm mesh
alongside each dock, close to the dock fouling, in an effort to obtain demersal organisms such as
4
"Cryptogenic" refers to species whose status as non-indigenous or native organisms is unknown (Carlton 1996).
Jason Toft and Marjorie Wonham were unable to participate during the first two days of the sampling. They
returned to the missed sites and collected plankton and mussel samples within two weeks.
5
Page 2
Page 3
harpacticoid copepods.
In addition, pilings were separately sampled at several sites, and nearby intertidal sites were
opportunistically sampled. In order to compare similar environments, docks were classified as
primary sites, other sites such as beaches were classified as secondary sites.
Temperature and salinity were measured at each dock site. We attempted to use two electronic
(YSI) meters to obtain depth profiles of temperature and salinity, but these devices produced
varying and unreliable readings (as was the experience of the San Francisco Bay Expeditions).
The measurements we report are near-surface measurements obtained with 2 thermometers and 2
refractometers that agreed within 0.5 parts per thousand (ppt).
From each dock site we obtained a one-liter representative voucher collection, plus additional
samples of material of interest. The samples were kept on ice on days with laboratory time
scheduled soon after the field work, and preserved on-site in formalin or alcohol on other days.
Laboratory work was conducted at the King County Environmental Laboratory (on the evening
of Sept. 8) and the University of Washington's Friday Harbor Laboratories (Sept. 11-16). The
voucher collections were all examined in the laboratory and organisms identified by team
members, with some material retained by individual team members for further study. The
voucher collections are currently held by the Washington Department of Natural Resources, and
are ultimately to be deposited in an appropriate curated facility.
Only a small portion of the data analysis was completed during the Expedition. Work completed
after the Expedition and anticipated in the future is discussed on page 17. Additional information
on participants, schedule, collecting sites and equipment is provided in Appendices 1-4.
Page 4
Non-indigenous Species Collected in Puget Sound
The information provided in this section is our best assessment of the data available at the
conclusion of the Expedition. However, a great deal of work remains in order to complete
analysis of the samples. Information given here should be considered provisional and subject to
refinement as analysis progresses.
The Expedition collected at least 39 non-indigenous salt and brackish water species in Puget
Sound (Table 1). At least 16 appear to represent new or unpublished records for Puget Sound,
however none of the species collected appears to be a new record for the Pacific Coast of North
America. Twenty-two additional species we consider to be cryptogenic. Two polychaete worms
may represent new species, previously unreported introductions, or substantial range extensions.
The status of many of these species may be clarified by further taxonomic analysis.
All of the non-indigenous species collected by the Expedition whose native range is known are
native to either the North Atlantic or the Western Pacific (Table 2), with about half from each
region. However, the importance of the two source regions has shifted over time. Sixty-one
percent of the species first recorded on the Pacific Coast before 1950 originated from the North
Atlantic, while 78% of the species first recorded after 1950 originated from the Western Pacific.
The first record of collection on the Pacific Coast for any of the non-indigenous species collected
by the Expedition is in 1871. For the species for which we have data, half were first recorded on
the Pacific Coast in the latter half of the 1871-1998 period, while 70% have first records for
Puget Sound in the latter half of the period. These data suggest no obvious trend over time with
regard to initial appearance on the Pacific Coast, but suggest an increase in the rate at which they
are appearing or being discovered in Puget Sound.
Possible mechanisms of introduction were assigned in Table 2 based on historic records, known
associations of organisms with transport mechanisms, the organisms’ biological characteristics,
etc. The assigned mechanisms show trends over time. Ship fouling appears to have declined in
importance between the earlier and later halves of the period from 1871 to 1998. It is listed as a
possible mechanism for 71% of species with first records in the earlier period, and 53% of
species in the later period (note that in many cases more than one mechanism is listed for each
species). Introductions with Atlantic or Japanese oysters declined slightly between the two
periods, while ballast water increased in importance (ballast water listed for 18% of species in
the earlier period, 53% of species in the later period).
Page 5
Table 1. Non-indigenous and Cryptogenic Species and other New Records Collected by the
Puget Sound Expedition
This list of species is provisional pending further taxonomic work and review by expedition members and
associates.
A
B
C
D
E
NON-INDIGENOUS SPECIES - New Record of an Introduction for Puget Sound (not previously published or
known; Puget Sound defined as the inland marine waters of Washington State east of Cape Flattery)
NON-INDIGENOUS SPECIES - Introduction Known but not Published for Puget Sound
NON-INDIGENOUS SPECIES - Introduction Published for Puget Sound
CRYPTOGENIC SPECIES
NON-INDIGENOUS SPECIES - Total (total of A through C)
A
Chlorophyta
Enteromorpha sp.
Ulva sp.
Rhodophyta
Chondria dasyphylla
Polysiphonia paniculata
Phaeophyceae
Sargassum muticum
Anthophyta
Spartina anglica
Zostera japonica
Foraminifera
Trochammina hadai
Ciliata
Zoothamnium sp.
green folliculinids
Porifera
Halichondria sp.
Cnidaria: Hydrozoa
Cordylophora caspia (= C. lacustris)
Cnidaria: Anthozoa
Diadumene lineata (= Haliplanella luciae)
Alcyonium sp.
Annelida: Polychaeta
Autolytinae sp. 1 (clear)
Autolytinae sp. 2 (transverse bars)
Autolytinae sp. 3 (transverse orange bars)
Autolytinae sp. 4 (mid-dorsal orange stripe)
Autolytinae sp. 5 (4 longitudinal lines)
Autolytinae sp. 6 (lemon-yellow)
Capitella sp.
Exogone lourei
Harmothoe imbricata
Hobsonia florida
Platynereis sp.
Page 6
B
C
D
E
0
x
x
0
x
x
1
x
2
x
x
1
x
0
x
x
0
x
1
x
1
x
x
2
x
x
x
x
x
x
x
x
x
x
x
Table 1 (cont’d)
A
Annelida: Polychaeta (cont’d)
Polydora sp.
Proceraea sp. A (of Piltz, unpubl.)
Pseudopolydora sp.
Typosyllis sp.
Mollusca: Gastropoda
Batillaria attramentaria (= B. zonalis, = B. cumingi)
Crepidula fornicata
Myosotella myosotis (= Ovatella myosotis)
Mollusca: Bivalvia
Crassostrea gigas
Mya arenaria
Mytilus sp.
Nuttallia obscurata
Venerupis philippinarum (= Tapes japonica)
Arthropoda: Crustacea: Copepoda
Choniostomatid copepod
x
Arthropoda: Crustacea: Cumacea
Nippoleucon hinumensis
x
Arthropoda: Crustacea: Isopoda
Limnoria tripunctata
Arthropoda: Crustacea: Amphipoda
Ampithoe valida
Caprella mutica (= C. acanthogaster)
x
Corophium acherusicum
Corophium insidiosum
Eochelidium sp.
Grandidierella japonica
Jassa marmorata
Melita nitida
Parapleustes derzhavini
x
Bryozoa
Alcyonidium polyoum
Bowerbankia “gracilis”
Bugula stolonifera
x
Bugula sp. 1 (resembling B. turbinata)
Bugula sp. 2 (resembling B. fulva)
x
Cryptosula pallasiana
x
Schizoporella unicornis
Entoprocta
Barentsia benedeni
x
Urochordata
Botryllus schlosseri
Botrylloides violaceus (earlier reported as Botrylloides sp.)
Ciona savignyi
x
Molgula manhattensis
x
Styela clava
x
TOTAL
11
B
C
D
E
x
x
x
x
3
x
x
x
4
x
x
x
x
x
1
1
1
x
9
x
x
x
x
x
x
x
6
x
x
x
x
1
5
x
x
5
23
23
39
Page 7
Table 2. Origins, First Records and Mechanisms of Introduction of Non-indigenous
Species Collected by the Puget Sound Expedition
This list of species is provisional pending further taxonomic work and review by expedition members and
associates.
Native ranges, dates of first record (planting, collection, observation or report) in Puget Sound and on the Pacific
Coast of North America, and possible initial mechanisms of introduction to the Pacific Coast are given. First records
consisting of written accounts that do not state the date of planting, collection or observation are preceded by the
symbol "<". Mechanisms given in parentheses indicate less likely mechanisms. Mechanisms are listed as:
OA - with shipments of Atlantic oysters
OJ - with shipments of Japanese oysters
SF - in ship fouling or boring
SB - in solid ballast
BW - in ship ballast water or seawater system
MR - planted for marsh restoration or erosion control
General
Taxon
Seaweeds
Anthophyta
Foraminifera
Cnidaria
Annelida
Mollusca
Copepoda
Cumacea
Isopoda
Amphipoda
Entoprocta
Bryozoa
Urochordata
Page 8
Species
Native Range
First Pacific
Coast Record
Sargassum muticum
Spartina anglica
Zostera japonica
Trochammina hadai
Cordylophora caspia
Diadumene lineata
Hobsonia florida
Pseudopolydora sp.
Batillaria attramentaria
Crepidula fornicata
Myosotella myosotis
Crassostrea gigas
Mya arenaria
Nuttallia obscurata
Venerupis philippinarum
Choniostomatid copepod
Nippoleucon hinumensis
Limnoria tripunctata
Ampithoe valida
Caprella mutica
Corophium acherusicum
Corophium insidiosum
Eochelidium sp.
Grandidierella japonica
Jassa marmorata
Melita nitida
Parapleustes derzhavini
Barentsia benedeni
Bowerbankia gracilis
Bugula sp. 1
Bugula sp. 2
Bugula stolonifera
Cryptosula pallasiana
Schizoporella unicornis
Botrylloides violaceus
Botryllus schlosseri
Ciona savignyi
Molgula manhattensis
Styela clava
Japan
England
W Pacific
Japan
Black/Caspian Seas
Asia
NW Atlantic
?
Japan
NW Atlantic
Europe?
Japan
NW Atlantic
Japan, Korea (China?)
NW Pacific
?
Japan
not known
NW Atlantic
Japan to Vladivostok
not known
N Atlantic
Japan or Korea
Japan
NW Atlantic
NW Atlantic
W Pacific?
Europe
NW Atlantic?
?
?
NW Atlantic
N Atlantic
NW Pacific
Japan
NE Atlantic
Japan?
NW Atlantic
China to Okhotsk Sea
1944
1961-62
1957
1983
ca. 1920
1906
1940
?
1924
1905
1871
1875
1874
1989
1924
?
1979
1871 or 1875
1941
1973-77
1905
1915
early 1990s?
1966
1941
1938
1904
1929
<1923
?
?
<1978
1943-44
1927
1973
1944-47
1985
1949
1932-33
Possible
First Puget
Mechanism of
Sound Record Introduction
?
1961-62
?
1997
ca. 1920
<1939
1940
?
1924
1905
1927
1875
1888-89
1991-96
1924
1998
1998
?
?
1998
1974-75
1930
1997
?
?
1966
1998
1998
<1953
1993
1998
1998
1998
1927
1977
?
1998
1998
1998
OJ
MR
OJ
BW,SF,OJ
BW,SF
OA,SF
?
?
OJ
OA
OA(SB,SF)
OJ
OA
BW
OJ
?
BW
SF
BW,OA,SF
BW,OJ
OA,SF
OA,SF
BW
BW,OJ,SF
BW, SF
BW,OA,SB,SF
SF
OJ,SF
OA,SF
?
?
SF
OA,SF
OJ,SF
OJ,SF
OA,SF
BW,SF
BW,OA,SF
BW,OJ,SF
The number of non-indigenous species collected and identified per site, not counting the annelids
or peracarids, ranged from zero to eight (Appendix 5). Initial analysis of the distribution of the
non-indigenous species collected by the Expedition reveals no obvious trends in the number of
non-indigenous species collected at each dock-fouling or benthic site with regard to salinity,
temperature or region (Figures 1-3). The annelid and peracarid material for dock and benthic
sites has not been worked-up in the laboratory for all sampled sites; partial data are included in
Appendix 5.
9
0
0
Port Townsend
Port Hadlock
Port Ludlow
Brownsville
Cornet Bay
Steilacoom
Des Moines
Elliott Bay
Edmonds
Grapeview
Port Orchard
Blaine
Anacortes
Friday Harbor
1
Harbor Island
2
Tacoma
3
Everett
4
Samish River
5
Bellingham
6
Fishermen's Terminal
Number of species
7
Shelton
Boston Harbor
Seabeck
8
3 21 22 25 25 29 29 30 30 31 31 31 32 32 32 32 32 32 33 33 34 34
Salinity (ppt)
Figure 2. Non-indigenous Species Collected vs. Surface Salinity
Page 9
Page 10
Number of species
Number of species
Edmonds
Elliott Bay
Brownsville
Hood Admiralty
Canal
Inlet
Des Moines
Seabeck
Port Hadlock
Port Ludlow
Port Townsend
Bellingham
North
Sound
Anacortes
Samish River
Blaine
San
Juan
Friday Harbor
Port Hadlock
Port Townsend
14
Anacortes
15
Edmonds
15
Elliott Bay
15
Steilacoom
Tacoma
15.5
Port Ludlow
15.5
16.5
16.5
16.5
Everett
Harbor Island
Des Moines
Boston Harbor
17
Blaine
17
Brownsville
17
18
Fishermen's Terminal
18
Port Orchard
18.5
Samish River
19
19
Bellingham
Seabeck
19
Grapeview
20
Shelton
9
Cornet Bay
Cornet Bay
14
15
8
Port Orchard
7
Tacoma
6
Harbor Island
Friday Harbor
13
Temperature (oC)
Central Sound
Region
Figure 4. Non-indigenous Species By Region
Everett
5
Fishermen's Terminal
4
Shelton
3
Grapeview
Figure 3. Non-indigenous Species Collected vs. Surface Temperature
Boston Harbor
2
12
1
0
9
8
7
6
5
4
3
2
1
0
Steilacoom
South
Sound
Steilacoom
The general biotic community at each sampling site is described in Table 3. Exotic species were
common or dominant organisms at several sites (although generally less common than has been
observed by similar expeditions in San Francisco Bay). At many sites bay mussels (Mytilus sp.)
were the dominant organisms in dock fouling, sometimes approaching 100% cover. These
mussels may consist in part of one or two exotic species from the Mediterranean and the Atlantic
or their hybrids (the genetic composition of mussels collected from these sites will be analyzed
by M. Wonham). At the low salinity Samish River site (3 ppt), the hydroid Cordylophora caspia,
native to the Black and Caspian seas, covered the shaded sides and undersides of the dock and
pilings, and could be collected by the tubful. The Japanese colonial tunicate Botrylloides
violaceus was abundant in dock fouling at many sites, while the solitary tunicates Ciona savignyi
(from Japan) and Molgula manhattensis (from the western Atlantic) were abundant at,
respectively, the Des Moines and Shelton marinas. The small, orange-striped Asian anemone
Diadumene lineata was also common at Shelton. Schizoporella unicornis (from Asia) and
Cryptosula pallasiana (from the North Atlantic) were among the most frequently collected
encrusting bryozoans. The varnish clam Nuttallia obscurata, a recent arrival from Asia, was
common intertidally at Argyle Lagoon, Blaine and Port Ludlow. The Japanese amphipod
Grandidierella japonica comprised nearly 100% of the organisms present at the two high
intertidal mudflat areas sampled. At three other sites G. japonica was found in subtidal grabs or
fouling samples, where it would be expected only if dense intertidal mudflat populations were
nearby. The native amphipods Corophium salmonis and Corophium brevis are normally present
on intertidal mudflats in the Pacific Northwest, but were not found in the intertidal mudflats
where G. japonica was abundant.
Table 3. Characterization of the Biotic Communities at Sampling Sites
Sampling
Site
Temp.
Salinity
Anacortes
(Cap Sante
Boat
Haven)
15° C
31 ppt
Argyle
Lagoon
(San Juan
Island)
Description of Biotic Community
The fouling community on wood/styrofoam and concrete floats was diverse, dominated by
the kelp Laminaria saccharina with thecate hydroids growing on its blades, Metridium sp.
sea anemones, Mytilus sp. mussels, halichondria-like sponge, colonial tunicates especially
Botrylloides violaceus and sabellid worms. Extensive sheets of Perophora annectens
tunicates were found on submerged floats (primary site).
The beach and lagoon were surveyed for Spartina sp. and Nuttallia obscurata. The marsh
gastropod Myosotella myosotis was common under pieces of wood in the Salicornia
virginica marsh (secondary site).
Bellingham
(Squalicum
Harbor)
19° C
21 ppt
Blaine
(Blaine
Marina)
17° C
30 ppt
The fauna was depauperate, new and old floats supported sparse Enteromorpha sp. and
filamentous diatoms, few mussels and barnacles. This site was sampled for nonindigenous species by Jim Carlton in 1976 and also found to be of very low diversity at
that time. We sampled old and new floats in covered and uncovered areas (primary site).
The fouling community was dominated by Mytilus sp, with abundant Botrylloides
violaceus, halichondria-like yellow sponge and Metridium sp.; Styela clava common in
patches. Water silty due to harbor dredging; some resultant dieoff evident on floats.
Sediments collected beneath the floats were black and sulphurous. We sampled the older
wood and styrofoam floats (primary site) and the marine park north of the marina
(secondary site).
Table 3 (Cont’d). Characterization of the Biotic Communities at Sampling Sites
Page 11
Sampling
Site
Temp.
Salinity
Boston
Harbor
(Budd
Inlet)
17° C
30 ppt
Brownsvill
e Marina
17° C
32 ppt
Cornet Bay
(Deception
Pass
Marina,
Whidbey
Island)
Des Moines
(City
Marina)
Edmonds
(Port of
Edmonds
Marina)
Elliott Bay
Marina
(Seattle)
13° C
33 ppt
16.5° C
32 ppt
15° C
32 ppt
15° C
32 ppt
Everett
(Port of
Everett
Marina)
16.5° C
22 ppt
Fishermen's
Terminal
(Seattle)
~18° C
0 ppt
Friday
Harbor
(UW Lab
dock)
Grapeview
(Fairharbor
Marina)
12° C
32 ppt
19° C
31 ppt
Description of Biotic Community
The stryofoam and tire-covered wooden docks were dominated by small (typically less
than 2 cm) Mytilus sp., which formed a general crust about 6 cm thick. Distaplia tunicates,
Metridium sp. were common. Enteromorpha sp. was present at the waterline. We sampled
submerged and surface floats (primary site), and the adjacent sand beach (secondary site).
Floats were dominated by Mytilus sp., white or brown Metridium sp., Botrylloides
violaceus, the thecate hydroid Obelia dichotoma, sabellid worms, barnacles and patchy
strands of filamentous diatoms. Small, reproductive hydromedusae of Phialidium sp.
were abundant near the concrete and wooden floats (primary site.)
The fouling community was dominated by scyphozoan scyphistomae (most likely
Aurelia sp.) in the shaded dock areas, along with Laminaria saccharina and other kelps,
thecate hydroids, and short diatom strands. This marina was unique in not having any
colonial tunicates, including the nearly ubiquitous non-indigenous species Botrylloides
violaceus. We sampled covered and uncovered wooden docks with styrofoam floats
(primary site).
In covered areas, floats were dominated by the solitary tunicate Ciona savignyi, with smaller
numbers of other tunicates, especially the colonial Botrylloides violaceus. Metridium sp. were
common at the waterline and few Mytilus sp. Open docks had a similar assemblage, but
Ciona savigni was confined to shaded float bottoms. The wood pilings were covered mostly
by barnacles, with a few mussels. We sampled covered and uncovered floats on ‘L’ dock,
made of wood with exposed stryrofoam flotation (primary site).
A relatively diverse float community, dominated by barnacles (primarily Semibalanus
cariosus), colonial tunicates (primarily Botrylloides violaceus and Distaplia occidentalis)
and mixed algae including abundant filamentous diatoms. Pilings were dominated by
Semibalanus cariosus, with some Mytilus sp. We sampled the concrete floats at the Guest
Dock, the adjacent covered dock (dominated by the native solitary tunicate Corella
inflata) and a submerged, heavily-fouled power cable sheathed in black plastic (primary
site).
The concrete floats and pilings supported a diverse fauna and flora, but were dominated by
barnacles, mussels, and large sabellid worms near the bases of the floats. Other common
residents included limpets, Pododesmus sp. rock scallops, Myxicola infundibulum worms
and several species of tunicates (primary site).
Floats were dominated by filamentous diatoms, patchy barnacles, and mussels. Pilings were
primarily covered by barnacles. We sampled covered and open areas of the concrete floats,
and lines and bumpers on the ‘I’, ‘J’ and ‘B’ docks (primary site).
The floats, pilings and lines were characterized by small sponges, small feathery green
algae, and two freshwater species of bryozoans. We sampled a work float on the outer
edge of one of the central piers (primary site).
The fauna on the concrete pontoons and tires was diverse and patchy. Large Metridium
sp. were common. Mixed algae was common including large kelps (especially Alaria sp.,
Laminaria sp. and Costaria sp.), as well as many species of red bladed algae. Other
common species included Balanus nubilis (some covered with Hydractinia hydroids), the
bryozoan Dendrobeania sp., thecate and athecate hydroids, halichondria-like sponges and
several species of tunicates (primary site).
Dense (up to 100% cover) Mytilus sp. was growing on the floats, the mussels were
covered by Botrylloides violaceus, bryozoans and Metridium sp. The covered and
uncovered portions of the dock (primary site) and the adjacent beach (secondary site) were
sampled.
Table 3 (Cont’d). Characterization of the Biotic Communities at Sampling Sites
Page 12
Sampling
Site
Harbor Island
Marina
Port Hadlock
Bay Marina
Port Ludlow
Marina
Port Orchard
(Kitsap
Marina)
Port
Townsend
(Boat Haven
Marina)
Temp.
Salinity
16.5° C
25 ppt
14° C
34 ppt
15.5° C
33 ppt
18° C
31 ppt
Description of Biotic Community
The concrete floats were dominated by Mytilus sp., barnacles, and extensive mats of
filamentous diatoms. We sampled in the center of the Harbor Island Marina (primary
site).
The concrete floats and a dinghy were fouled by Mytilus sp., Metridium sp., the branched
thecate hydroid Obelia longissima, sabellid worms, nudibranchs, tunicates and abundant
caprellids. At the waterline, Ulva sp. and Enteromorpha sp. algae were common.
Nereocystis leutkeana and other kelps were attached to the docks (primary site).
The concrete float surfaces were dominated by mussels, ascidians, Metridium sp., and
sponges. We sampled the floats (primary site), the adjacent man-made lagoon
(secondary site) behind the marina parking lot, and the nearby debris-covered beach
(secondary site). The lagoon supported masses of filamentous green algae and abundant
Haminoea sp. . On the beach, Nuttallia obscurata shells were common.
This marina was composed of concrete floats and wooden floats fitted with tires. The
dominant organisms were Mytilus sp., small Metridium sp., barnacles, and clumps of the
thecate hydroid Obelia longissima. Large and small Aurelia labiata jellyfish were
common (approximately 3” and 12” diameter) (primary site).
14° C
34 ppt
The concrete floats were dominated by barnacles (mixed Balanus crenatus and
Semibalanus cariosus,) Mytilus sp., sabellid worms, Metridium sp., Botrylloides
violaceus, and Ulva sp. (primary site).
Samish River
(private dock,
Edison)
18.5° C
3 ppt
The wooden floats and pilings were dominated by strands of diatoms, with large clumps
of the hydroid Cordylophora caspia in shaded places, few barnacles, and a narrow band
of Enteromorpha sp. at the waterline (primary site).
Seabeck
Marina (Hood
Canal)
19° C
29 ppt
Shelton
(Shelton
Yacht Club)
20° C
29 ppt
Steilacoom
Marina
15° C
32 ppt
Swinomish
Channel
(South Padilla
Bay)
Tacoma (Ole
& Charlie's
Marina)
The wood and styrofoam floats (some wrapped in black plastic sheeting) were dominated
by Mytilus sp., the colonial tunicates Botryllus schlosseri and Botrylloides violaceus, the
thecate hydroid Obelia longissima, and thick Enteromorpha sp. We sampled the floats
(primary site) and the adjacent gravel upper beach (secondary site).
The floats were thickly encrusted with Mytilus sp. and dense anemones (Diadumene
lineata at the waterline and Metridium sp. below), tunicates (especially Botryllus
schlosseri and Molgula manhattensis) and a halichondria-like sponge. We sampled the
main portion of the dock, lined and shaded by individual boathouses (primary site).
Floats and tires in the water at the Steilacoom Marina were dominated by Mytilus sp.,
Metridium sp., the thecate hydroid Obelia dichotoma branching out several cm, and Ulva
sp., with large, patchy clumps of sabellid worms. The pilings were covered with
barnacles and pandalid shrimp were common several feet below the waterline (primary
site). We also sampled the adjacent gravel beach (secondary site).
This middle and high intertidal mud lagoon on the Swinomish Channel is adjacent to the
Swinomish Casino. The mudflat is dominated by Vaucheria sp. We sampled only
Spartina anglica (secondary site).
15.5° C
25 ppt
The wood and styrofoam float surfaces were dominated by Mytilus sp. and filamentous
diatoms up to one meter long. Enteromorpha sp. was common at the waterline. Bankia
sp. shipworms were found in older wood. We sampled floats and lines in open and
covered areas (primary site).
Current data indicate that of the 113 non-indigenous species listed as collected or potentially
collected by either the Puget Sound Expedition of 1998 or the San Francisco Bay Expeditions of
Page 13
1993, 1994, 1996 and 1997 (Table 4), the Puget Sound Expedition collected at least 39 species6
and may have collected another four species (pending further taxonomic analysis). John
Chapman reported collecting an additional two listed species from Puget Sound immediately
after the Expedition. The San Francisco Bay Expeditions collected at least 95 of the listed
species, and may have collected another seven (Cohen and Carlton 1995). Including the two
Chapman records, 28 of the species were collected by expeditions in both regions, 13 are listed
as collected by the Puget Sound but not by the San Francisco Bay Expeditions, and 67 are listed
as collected by the San Francisco Bay but not the Puget Sound Expedition. (Due to unresolved
taxonomic issues, for another five listed species it remains unclear whether they were collected
by any of the expeditions.) Even taking into account the fact that these data represent a greater
collecting effort over a longer period of time in San Francisco Bay than in Puget Sound, they
nevertheless indicate that a substantially greater number of non-indigenous species are present in
San Francisco Bay.
Table 4. Non-indigenous Species from Salt or Brackish Water Collected or Observed
by the Puget Sound or San Francisco Bay Expeditions
This list of species is provisional pending further taxonomic work and review by expedition members and
associates.
PSX = Puget Sound Expedition (1998); SFX = San Francisco Bay Expeditions (1993, 1994, 1996, 1997)
x = collected or observed; 0 = not collected or observed; ? = unclear whether collected pending further taxonomic
work or review; * = reported by J. W. Chapman from Mud Bay near Olympia after the Puget Sound Expedition
Chlorophyta
Codium fragile tomentosoides
Phaeophyceae
Sargassum muticum
Anthophyta
Cotula coronopifolia
Lepidium latifolium
Salsola soda
Spartina alterniflora
Spartina anglica
Zostera japonica
Foraminifera
Trochammina hadai
Porifera
Halichondria bowerbanki
Haliclona loosanoffi
Microciona prolifera
Cnidaria: Hydrozoa
Blackfordia virginica
6
PSX
SFX
0
x
x
x
0
0
0
0
x
x
x
x
x
x
0
0
x
x
?
?
0
PSX
?
?
x
SFX
0
x
Cordylophora caspia
Ectopleura crocea
Garveia franciscana
Gonothyraea clarki
Maeotias inexspectata
Turritopsis nutricula
Cnidaria: Anthozoa
Diadumene "cincta"
Diadumene franciscana
Diadumene leucolena
Diadumene lineata
x
0
0
0
0
0
x
x
x
?
x
x
0
0
0
x
x
x
x
x
For this analysis, the species listed as Pseudopolydora sp. in Tables 1 and 2 is assumed to be either P. kempi or P.
paucibranchiata.
Page 14
Annelida: Polychaeta
Ficopomatus enigmaticus
Heteromastus filiformis
Hobsonia florida
Marenzelleria viridis
Neanthes succinea
Polydora ligni
Pseudopolydora kempi
Pseudopolydora paucibranchiata
Streblospio benedicti
Typosyllus sp. 1
Mollusca: Gastropoda
Batillaria attramentaria
Crepidula fornicata
Ilyanassa obsoleta
Littorina saxatilis
Myosotella myosotis
Okenia plana
Philine auriformis
Tenellia adspersa
Mollusca: Bivalvia
Crassostrea gigas
Gemma gemma
Geukensia demissa
Macoma petalum
Musculista senhousia
Mya arenaria
Nuttallia obscurata
Potamocorbula amurensis
Teredo navalis
Theora fragilis
Venerupis philippinarum
Arthropoda: Crustacea: Copepoda
Choniostomatid copepod
Arthropoda: Crustacea: Cirripedia
Balanus amphitrite
Balanus improvisus
Arthropoda: Crustacea: Nebaliacea
Epinebalia sp. 1
Arthropoda: Crustacea: Cumacea
Nippoleucon hinumensis
Arthropoda: Crustacea: Isopoda
Dynoides dentisinus
Iais californica
Ianiropsis serricaudis
Limnoria tripunctata
Munna sp. 1
Paranthura sp. 1
Sphaeroma quoyanum
Sphaeroma walkeri
Synidotea laevidorsalis
PSX
SFX
0
0
x
0
*
0
?
?
*
0
x
x
0
x
x
?
?
?
x
x
x
x
0
0
x
0
0
0
0
0
x
x
x
x
x
x
x
0
0
0
0
x
x
0
0
0
x
0
x
x
x
x
x
0
x
x
x
x
x
0
0
0
x
x
0
x
x
x
0
0
0
x
0
0
0
0
0
x
x
x
?
x
x
x
x
x
PSX
SFX
Arthropoda: Crustacea: Tanaidacea
tanaid sp. 1
Arthropoda: Crustacea: Amphipoda
Ampelisca abdita
Ampithoe valida
Caprella mutica
Corophium acherusicum
Corophium alienense
Corophium heteroceratum
Corophium insidiosum
Eochelidium sp. 1
Gammarus daiberi
Grandidierella japonica
Jassa marmorata
Leucothoe sp. 1
Melita nitida
Melita sp. 1
Paradexamine sp.
Parapleustes derzhavini
Stenothoe valida
Transorchestia enigmatica
Arthropoda: Crustacea: Decapoda
Carcinus maenas
Eriocheir sinensis
Palaemon macrodactylus
Rhithropanopeus harrisii
Bryozoa
Anguinella palmata
Bowerbankia “gracilis”
Bugula neritina
Bugula stolonifera
Bugula sp. 1
Bugula sp. 2
Conopeum tenuissimum
Cryptosula pallasiana
Schizoporella unicornis
Watersipora "subtorquata"
Zoobotryon verticillatum
Entoprocta
Barentsia benedeni
Urochordata
Ascidia zara
Botrylloides cf. diegensis
Botryllus schlosseri
Botrylloides violaceus
Ciona intestinalis
Ciona savignyi
Molgula manhattensis
Styela clava
Chordata: Pisces
Acanthogobius flavimanus
Lucania parva
Morone saxatilis
Tridentiger trigonocephalus
?
x
0
x
x
x
0
0
x
x
0
x
x
0
x
0
0
x
0
0
x
x
x
x
x
x
x
x
x
x
0
x
x
x
x
x
x
x
0
0
0
0
x
x
x
x
0
x
0
x
x
x
?
x
x
0
0
x
x
x
x
0
0
x
x
x
x
x
x
x
0
0
x
x
0
x
x
x
x
x
x
x
x
x
x
x
0
0
0
0
x
x
x
x
Page 15
Based on the data developed by the Puget Sound Expedition, and a brief review of the extant lists
of non-indigenous species that include the Puget Sound area (including reviews of the Cnidaria
by Claudia Mills and Polychaeta by Leslie Harris), we offer a provisional list of 52 nonindigenous species that have been collected from and appear to be established in the salt or
brackish waters of Puget Sound (Table 5). In addition, a single specimen of an Asian copepod,
Pseudodiaptomus inopinus, was collected in the fall of 1991 in the Snohomish River estuary, and
a few specimens of another Asian copepod, Pseudodiaptomus marinus were collected in the
spring of 1998 in Elliott Bay, but it is unclear whether either of these specimens is established in
Puget Sound (information from J.F. Cordell).
Table 5. Non-indigenous Species in Puget Sound
This list of species is provisional pending further taxonomic work and review by expedition members and
associates.
For the purposes of this list, Puget Sound is defined as the inland marine waters of Washington State east of Cape
Flattery. * = species reported by J. W. Chapman from Mud Bay near Olympia after the Puget Sound Expedition.
Phaeophyceae
Sargassum muticum
Anthophyta
Spartina alterniflora
Spartina anglica
Spartina patens
Zostera japonica
Foraminifera
Trochammina hadai
Cnidaria: Hydrozoa
Cladonema radiatum
Cordylophora caspia (= C. lacustris)
Cnidaria: Anthozoa
Diadumene lineata (= Haliplanella luciae)
Platyhelminthes
Pseudostylochus ostreophagus
Annelida: Polychaeta
Hobsonia florida
Neanthes succinea *
Pseudopolydora sp.
Pygospio elegans
Streblospio benedicti *
Mollusca: Gastropoda
Batillaria attramentaria (= B. zonalis,
= B. cumingi)
Ceratostoma inornatum (= Ocenebra japonica)
Crepidula fornicata
Crepidula plana
Myosotella myosotis (= Ovatella myosotis)
Urosalpinx cinerea
Mollusca: Bivalvia
Crassostrea gigas
Musculista senhousia (= Musculus senhousia)
Mya arenaria
Mytilus galloprovincialis
Page 16
Mollusca: Bivalvia (con’t)
Nuttallia obscurata
Venerupis philippinarum (= Tapes japonica)
Arthropoda: Crustacea: Copepoda
Choniostomatid copepod
Mytilicola orientalis
Arthropoda: Crustacea: Cumacea
Nippoleucon hinumensis
Arthropoda: Crustacea: Isopoda
Limnoria tripunctata
Arthropoda: Crustacea: Amphipoda
Ampithoe valida
Caprella mutica (= C. acanthogaster)
Corophium acherusicum
Corophium insidiosum
Eochelidium sp.
Grandidierella japonica
Jassa marmorata
Melita nitida
Parapleustes derzhavini
Bryozoa
Bowerbankia “gracilis”
Bugula stolonifera
Bugula sp. 1
Bugula sp. 2
Cryptosula pallasiana
Schizoporella unicornis
Entoprocta
Barentsia benedeni
Urochordata
Botryllus schlosseri
Botrylloides violaceus
Ciona savignyi
Molgula manhattensis
Styela clava
Future Research and Reports, and Research Needs
Follow-up work
Jeff Cordell and Jason Toft of the University of Washington's Fisheries Research Institute
identified the zooplankton. Mary McGann of the U. S. Geological Survey in Menlo Park, CA has
completed a preliminary examination and identification of the foraminifera. John Chapman of
Oregon State University will complete identification of peracaridan crustacean species. Marjorie
Wonham will examine mussels with molecular genetic techniques as part of her University of
Washington dissertation research to determine if two nonindigenous mussels (Mytilus
galloprovincialis and possibly M. edulis) are present in Puget Sound in addition to the native bay
mussel (M. trossulus). Additional taxonomic work remains, which will be completed by
expedition participants and associates as time allows.
Anticipated additional reports/presentations
Claudia Mills plans to present a summary of the Expedition at the National Conference on
Marine Bioinvasions at the Massachusetts Institute of Technology in January 1999. John
Chapman has submitted a paper to the same conference titled “Climate and Non-indigenous
Species Introductions in Northern Hemisphere Estuaries.” Charles and Gretchen Lambert will
include tunicate data from the Expedition in their final report to California Sea Grant on the
tunicates of San Diego Bay, as all of the non-indigenous tunicates collected by the Puget Sound
Expedition are also known from southern California.
Research needs
As noted in this report, the results reported for the Expedition are provisional and incomplete.
This is primarily due to the lack of adequate funding for the Expedition as a whole. Most of the
Expedition members participated in the project without benefit of funding for their time, out of
an interest in the marine ecology of Puget Sound and the phenomenon of biological invasions.
However, judging from past experience with the San Francisco Bay Expeditions, much of the
desired follow-up work -- including the time-consuming task of identifying the organisms that
could not be identified to species in the immediate round of laboratory work, and compiling,
analyzing and reporting on the data -- will likely not take place without additional funding. Much
of the potential value of the Expedition could therefore be lost. Funding would make needed
research possible, including: taxonomic analysis, publication of results, surveys to monitor
invasions over time, further sampling in Puget Sound, and expansion of sampling to British
Columbia and other areas.
The substantial ecological and economic impacts of biological invasions in aquatic ecosystems
have been well documented, and are increasingly referenced by Washington State agencies and
discussed by the public and the press. However, the overall level of funding provided for
research into both the nature of these invasions and potential solutions remains a small fraction
of the cost of the impacts. Unless this situation changes, efforts to reduce the rate of biological
invasions or mitigate their impacts are likely to be hampered.
Page 17
Acknowledgments
We wish to thank:
•
the Washington State Department of Natural Resources, the U. S. Fish and Wildlife Service,
and the Puget Sound Water Quality Action Team for financial support to cover the direct
expenses of the Expedition;
•
the Washington State Department of Natural Resources for providing equipment and
logistical support;
•
the University of Washington/Friday Harbor Laboratories and the King County
Environmental Lab for providing laboratory space and equipment;
•
the Western Washington University/Shannon Point Marine Laboratory, the University of
Washington/Friday Harbor Laboratories, and Gretchen and Charles Lambert for providing
housing;
•
Eugene Kozloff (University of Washington), Bruno Pernet (University of Washington),
Scottie Henderson (University of Washington), Wim Vervoort (National Museum of Natural
History, Leiden, Netherlands), Mary McGann (USGS) and Doris Sloan (University of
California, Berkeley) for assistance with species identifications;
•
the California Sea Grant and Oregon Sea Grant College Programs, the Los Angeles County
Museum of Natural History, the Padilla Bay National Estuarine Research Reserve Fellowship
Program, the San Francisco Estuary Institute, the Switzer Environmental Leadership Grant
Program/San Francisco Foundation, the U. S. Fish and Wildlife Service San Francisco Bay
Program, the Wetland Ecosystem Team at the University of Washington, Williams College Mystic Seaport, Tom Schroeder and Amy Chapman for providing financial or other support
to enable Expedition team members and associated taxonomists to participate in the work of
the Expedition;
•
Tom Mumford (Washington State Department of Natural Resources), John Armstrong (U.S.
Environmental Protection Agency) and Mary Mahaffy (U.S. Fish and Wildlife Service) for
their support and assistance with the development of the Expedition; and
•
All of the marinas who allowed us to sample at their facilities.
Page 18
Literature Cited
Carlton, J. T. 1979. History, Biogeography and Ecology of the Introduced Marine and Estuarine
Invertebrates of the Pacific Coast of North America. Ph. D. thesis, University of California, Davis
CA.
Carlton, J. T. 1996. Biological invasions and cryptogenic species. Ecology 77(6): 1653-1655.
Cohen, A. N. and J. T. Carlton 1995. Non-indigenous Aquatic Species in a United States Estuary:
A Case Study of the Biological Invasions of the San Francisco Bay and Delta. Prepared for the U.S.
Fish and Wildlife Service, Washington DC.
Elston, R. 1997. Pathways and Management of Marine Non-indigenous Species in the Shared
Waters of British Columbia and Washington. Puget Sound/Georgia Basin Environmental Report
Series No. 5. Puget Sound Water Quality Action Team, Olympia WA.
Kozloff, E. N. 1987. Key to the Marine Invertebrates of Puget Sound, the San Juan Archipelago, and
Adjacent Regions. University of Washington Press, Seattle and London.
Lambert, C.C. and G. Lambert 1998. Non-indigenous ascidians in southern California harbors and
marinas. Marine Biology 130: 675-688.
Ruiz, G. M. and A. H. Hines 1997. The Risk of Non-indigenous Species Invasion in Prince William
Sound Associated with Oil Tanker Traffic and Ballast Water Management: Pilot Study. Prepared for
the Regional Citizens Advisory Committee of Prince William Sound, Valdez AK.
Page 19
Page 20
APPENDIX 1
List of Participants
Helen Berry, MS
Washington DNR, Aquatic Resources Divison
Nearshore Habitat Group
1111 Washington St. SE, PO Box 47027
Olympia, WA 98504-7027
(expertise in Puget Sound nearshore habitats)
e-mail:
[email protected]
work phone: 360-902-1052
San Francisco Estuary Institute
180 Richmond Field Station
1325 South 46th Street
Richmond, CA 94804
(expertise in general invertebrates and exotics;
San Francisco Bay team 1993-1997)
e-mail:
[email protected]
work phone: 510-231-9423
Brian Bingham, PhD
Huxley College of Environmental Studies
Western Washington University
Bellingham, WA 98225
(expertise in general invertebrates)
e-mail:
[email protected]
work phone: 360-650-2845
Jeff Cordell, MS
University of Washington
Fisheries Research Institute
Box 357980
Seattle, WA 98195-7980
(expertise in copepods and other small crustaceans)
e-mail:
[email protected]
work phone: 206-543-7532
Betty Bookheim
Washington DNR, Aquatic Resources Divison
Nearshore Habitat Group
1111 Washington St. SE, PO Box 47027
Olympia, WA 98504-7027
(expertise in Puget Sound nearshore habitats)
e-mail:
[email protected]
work phone: 360-902-1076
James Carlton, PhD
Maritime Studies Program
Williams College - Mystic Seaport
Post Office Box 6000
Mystic, CT 06355
(expertise in general invertebrates and exotics;
San Francisco Bay team 1993-1997)
e-mail:
[email protected]
work phone: 860-572-5359
John Chapman, PhD
Department of Fisheries and Wildlife
Oregon State University
Hatfield Marine Science Center
2030 S. Marine Science Dr.
Newport, Oregon 97365-5296
(expertise in small crustaceans and exotics;
San Francisco Bay team 1993-1997)
e-mail:
[email protected]
work phone: 541-867-0235
Andrew Cohen, PhD
Leslie Harris
LACM-Allan Hancock Foundation
Polychaete Collection
Los Angeles County Museum of Natural History
900 Exposition Boulevard
Los Angeles, California 90007
(expertise in polychaetes;
San Francisco Bay team 1996, 1997)
e-mail:
[email protected]
work phone: 213-763-3234
Terrie Klinger, PhD
University of Washington
Friday Harbor Laboratories
620 University Road
Friday Harbor, WA 98250
(expertise in algae)
e-mail:
[email protected]
work phone: 206-543-1484
Alan Kohn, PhD
University of Washington
Department of Zoology
Box 351800
Seattle, WA 98195-1800
(expertise in gastropods and general invertebrates)
e-mail:
[email protected]
work phone: 206-543-1629
Eugene Kozloff, PhD
University of Washington
Page 21
Friday Harbor Laboratories
620 University Road
Friday Harbor, WA 98250
(expertise in general invertebrates)
Tacoma, WA 98402-3100
(expertise in sea anemones and general invertebrates)
e-mail:
[email protected]
work phone: 253-692-5659
Charles Lambert, PhD
12001 11th Ave. NW
Seattle, WA 98177
(CSU Fullerton, Prof. Emeritus)
(expertise in tunicates)
e-mail:
[email protected]
phone: 206-365-3734
Jason Toft
University of Washington
Fisheries Research Institute
Box 357980
Seattle, WA 98195-7980
(graduate student in the School of Fisheries)
e-mail:
[email protected]
work phone: 206-543-7567; 685-3395
Gretchen Lambert, MS
12001 11th Ave. NW
Seattle, WA 98177
(formerly at CSU Fullerton)
(expertise in tunicates;
San Francisco Bay team 1997)
e-mail:
[email protected]
phone: 206-365-3734
Kevin Li
King County Environmental Lab
322 W. Ewing St.
Seattle, WA 98119-1507
(expertise in amphipods and other crustaceans)
e-mail:
[email protected]
work phone: 206-684-2344
Claudia Mills, PhD
University of Washington
Friday Harbor Laboratories
620 University Road
Friday Harbor, WA 98250
(expertise in hydromedusae and ctenophores;
wrote the Pacific NW keys to hydroids;
San Francisco Bay team 1993-1997)
e-mail:
[email protected]
work phone: 360-378-9105; 206-543-1484
Bruno Pernet, PhD
University of Washington
Friday Harbor Laboratories
620 University Road
Friday Harbor, WA 98250
(expertise in polychaetes and general invertebrates)
e-mail:
[email protected]
work phone: 360-378-9105; 206-543-1484
David Secord, PhD
University of Washington, Tacoma
1900 Commerce Street, Mailstop #358436
Page 22
Marjorie Wonham
University of Washington
Department of Zoology
Box 351800
Seattle, WA 98195-1800
(graduate student in the Department of Zoology)
e-mail:
[email protected]
work phone: 206-685-8615
APPENDIX 2
Puget Sound Expedition Schedule
Day 1 . Sept. 8 (Tuesday) EAST-CENTRAL PUGET SOUND
Low tide +0.9 ' at 1151 (Seattle prediction).
1. Port of Everett Marina
2. Port of Edmonds Marina
3. City of Des Moines Marina
4. Harbor Island Marina, near mouth of Duwamish River
5. Elliott Bay Marina, Seattle
6. Fishermen's Terminal, Seattle
Evening work in laboratory at the King County Environmental Lab.
TIME SAMPLED
9:30-10:20 am
11:15-12:05 pm
2:30-3:25 pm
4:25-5:10 pm
5:40-6:20 pm
6:45-7:10 pm
Day 2. Sept. 9 (Wednesday) SOUTH PUGET SOUND
Low tide +1.9 ' at 1237 (Seattle prediction).
7. Ole & Charlie's Marina, Tacoma
8. Steilacoom Marina
9. Boston Harbour Marina, near Olympia
10. Port of Shelton Marina
11. Fairharbor Marina, Grapeview
9:50-10:40 am
12:10-12:55 pm
3:05-4:10 pm
5:20-6:05 pm
6:45-7:20 pm
Day 3. Sept. 10 (Thursday) KITSAP PENINSULA AND OLYMPIC PENINSULA
Low tide +3.1 ' at 1326 (Seattle prediction).
12. Kitsap Marina, Port Orchard
13. Brownsville Marina
14. Seabeck Marina, Hood Canal
15. Port Ludlow Marina
16. Port Hadlock Bay Marina
17. Boat Haven Marina, Port Townsend
8:00-8:45 am
9:30-10:30 am
11:50-12:50 pm
3:15-4:00 pm
5:00-5:30 pm
6:10-6:45 pm
Day 4. Sept. 11 (Friday) NORTH PUGET SOUND
Low tide +4.0 ' at 1328 (Pt. Townsend prediction).
18. Deception Pass Marina, Cornet Bay
19. Blaine Marina
20. Squalicum Harbor, Port of Bellingham
21. Samish Bay - small float in Samish River near Edison
22. Padilla Bay - Vaucheria flats east of the Swinomish Channel
23. Cap Sante Boat Haven, Anacortes
8:50-9:40 am
11:15-12:10 pm
2:25-2:45 pm
3:35-4:05 pm
4:30-4:45 pm
4:50-5:40 pm
Day 5. Sept. 12 (Saturday) Friday Harbor Laboratories, San Juan Island.
Work in laboratory.
C. & G. Lambert sampled 2 marinas on San Juan Island for tunicates
Day 6. Sept. 13 (Sunday) Friday Harbor Laboratories, SAN JUAN ISLAND.
Low tides +0.2 ' at 0322 and +5.5 ' at 1610 (Pt. Townsend prediction).
Work in laboratory.
24. UW Friday Harbor Laboratories dock, San Juan Island
C. and G. Lambert sampled 5 additional marinas on San Juan Island for tunicates.
3:30-4:45 pm
Day 7. Sept. 14 (Monday) Friday Harbor Laboratories, SAN JUAN ISLAND.
Page 23
Low tides +0.2 ' at 0430 and +5.6 ' at 1740 (Pt. Townsend prediction).
Work in laboratory.
25. Argyle Lagoon and adjacent beach on North, San Juan Island
C. and G. Lambert sampled 6 marinas on Orcas Island for tunicates.
7:15-8:15 am
Sept. 15, 1998 (Tuesday) At the Friday Harbor Laboratories, SAN JUAN I SLAND.
Work in laboratory and begin report.
26. Mud Bay, at base of Eld Inlet, near Olympia sampled by J. Chapman noon for 45 minutes at around noon.
C. and G. Lambert sampled 2 additional marinas on San Juan Island for tunicates.
Sept. 16, 1998 (Wednesday) At the Friday Harbor Laboratories, SAN JUAN I SLAND.
Complete work in laboratory and continue work on report.
C. & G. Lambert sampled 1 marina on Lopez Island and 3 additional marinas in Anacortes for tunicates.
Sept. 17-20 1998 (Thursday-Sunday)
Complete work on report.
Page 24
APPENDIX 3
Description of Sampling Sites
(listed in the order they were sampled)
Site 1. Port of Everett Marina
Everett is in Possession Sound, in the central Puget Sound basin. The marina is located near the mouth of the
Snohomish River in an industrial area north of the Kimberley-Clark facility and the U.S. Naval Base. Salinity in the
area is lowered by fresh water input from the Snohomish River, making this site potentially susceptible to nonindigenous brackish water species.
Site 2. Port of Edmonds Marina
Edmonds is located north of Seattle in the central basin. The marina is south of the Edmonds-Kingston ferry
terminal. Many of the concrete floats at the marina were replaced after suffering heavy snow damage in December,
1996.
Site 3. City of Des Moines Marina
Des Moines was settled in 1870. The large marina is located in the central basin, approximately mid-way between
Elliott Bay, Seattle, and Commencement Bay, Tacoma. The area is heavily used by recreational boats.
Site 4. Harbor Island Marina, Seattle
This small facility with commercial and recreational boats is located in central Seattle, near the mouth of the
Duwamish River. The primary land use in southern Elliott Bay is water-based commerce and industry. Reduced
salinity makes it a potential introduction site for brackish water species.
Site 5. Elliott Bay Marina, Seattle
Built 6 years ago, this large 1,200 slip marina is situated at the north end of Elliott Bay, near Seattle’s commercial /
industrial area. Elliott Bay has been a major shipping center in Puget Sound since the mid-1800s.
Site 6. Fishermen's Terminal, Seattle
This large marina for commercial fishing boats lies just inside the Hiram M. Chittenden Locks that connect the
freshwater lakes (Lake Washington and Lake Union) with Puget Sound. The site was examined primarily for
potential freshwater species such as the hydroid Cordylophora.
Site 7. Ole & Charlie's Marina, Tacoma
The marina is located near the mouth of the Hylebos Waterway in Commencement Bay, Tacoma. Tacoma was first
settled by Euro-Americans in 1852. Commencement Bay is one of the earliest and most heavily used areas for
water-based commerce and industry. Natural mudflat and marsh habitat were dredged and filled in order to create
the current waterway system. Significant contamination exists as a result of historical uses. Near the marina, the area
is currently dominated by commercial shipping, including log yards and scrap steel.
Site 8. Steilacoom Marina
This small private marina is the northernmost site in the south sound basin. It lies south of the McNeil and Anderson
Island ferry terminal in Steilacoom. Steilacoom was established in 1851, and was an early center for West Coast
shipping in Puget Sound. The marina is closed to business, yet some boats remain in the area.
Site 9. Boston Harbor Marina, near Olympia
This small, neighborhood marina is located at the northeast end of Budd Inlet, near Olympia. Olympia was an early
population center, with a population larger than Seattle during early settlement years. Boston Harbor was
recommended to us by Erik Thuesen who teaches marine science at the Evergreen State College, as more diverse
than marinas deeper inside Budd Inlet. The site was selected to represent a south sound site that maintained higher
salinity and lower temperature than the heads of the bays in Eld Inlet, Budd Inlet and Totten Inlet.
Page 25
Site 10. Port of Shelton Marina
The Shelton Marina in Oakland Bay at the end of Hammersley Inlet was selected to represent a geographic endpoint
of habitats farthest from the entrance to Puget Sound. Activities in the area that might lead to species introduction
include aquaculture, log shipping, and recreational boating. Currently, logs are stored in much of the nearshore area
in the Shelton embayment. Oyster growing areas were severely impacted by sulfite from pulp mills in the 1920s and
1930s.
Site 11. Fairharbor Marina, Grapeview
Grapeview is located in the northern portion of Case Inlet, in the south sound. Fairharbor Marina is a relatively
small neighborhood marina for local and visiting recreational boats. Aquaculture in the area is common. Grapeview
was originally platted in 1891.
Site 12. Kitsap Marina, Port Orchard
The Kitsap Marina is a small facility near the Bremerton Naval Shipyard in Sinclair Inlet. Sites in Port Orchard
were deemed more accessible than sites in Bremerton. A lumber mill in Port Orchard was functioning for shipping
as early as 1855. Heavy historic use of the water for commerce and defense makes the area a potential site of species
introductions.
Site 13. Brownsville Marina
Brownsville is located on the Kitsap Peninsula facing western Bainbridge Island. The marina is a small, residential
Yacht Club. It was decided to sample at Brownsville, rather than Poulsbo, which is a little further north and at the
end of Liberty Bay, based on preliminary investigations by Claudia Mills, who found that the Poulsbo fauna was
more depauperate than that on the floats at Brownsville.
Site 14. Seabeck Marina, Hood Canal
Seabeck is on the Kitsap Peninsula on Hood Canal. The Seabeck Marina is the only marina in this primarily
residental area. The site represents an approximate mid-point in energy, temperature and salinity gradients found in
Hood Canal. Euro-American use of Seabeck began in 1857 with the establishment of a company-owned mill town.
Seabeck was the site of a major port facility, shipyards, two sawmills producing 80,000 board feet per day, and a
logging camp with 600-1,000 people. The mill was destroyed by fire in 1886.
Site 15. Port Ludlow Marina
Port Ludlow is the name of both a small inlet and the community occupying its shores on the east side of the
Olympic Peninsula. Between the 1850s and 1938, Port Ludlow had a mill and shipyard and was one of the most
active shipping ports. Closure of the mill changed the nature of this town, which is now predominantly a residential
and resort development with little commercial traffic. In addition to visiting the marina area, we walked an adjacent
beach strewn with what is said to be old solid ballast.
Site 16. Port Hadlock Bay Marina
Port Hadlock is approximately mid-way between Port Ludlow and Port Townsend. It is a small historic working
port on the eastern shore of the Olympic Peninsula, which was the site of an alcohol plant. The Old Alcohol Plant is
currently a lodge with a fairly small, private marina.
Site 17. Boat Haven Marina, Port Townsend
Port Townsend, located on the northeast tip of the Olympic Peninsula, was a thriving port in the early decades of
settlement of Puget Sound. The port’s regional importance decreased when the cross-continental railroad chose
Tacoma as its terminus. The city is a popular area for recreational boating and tourism.
Site 18. Deception Pass Marina, Cornet Bay
This small, private marina is situated on Cornet Bay, just east of Deception Pass. The small marina serves private
pleasure boats in a small residential community that is situated near the smaller of the two passes through which
marine water enters Puget Sound. Spartina infestations are known west of the marina.
Page 26
Site 19. Blaine Marina
The Blaine Marina in Drayton Harbor is run by the Port of Bellingham and is adjacent to the Canada-U.S. border. It
was the northernmost point in our expedition and might more accurately be described as Strait of Georgia than Puget
Sound. The town of Blaine was platted in 1884. The Blaine marina serves both a small fishing fleet and
recreational boaters; there are several fish processing plants and a shipyard at the port facility. The marina is
currently being expanded; the water was silty due to dredging and some dieoff on the floats was evident.
Site 20. Squalicum Harbor, Port of Bellingham
This is a very large marina facility run by the Port of Bellingham primarily for pleasure boats. The downtown area
is a center for water-dependent commerce. Industrial shipping as well as a large ferry terminal for the Alaska State
ferry are located further south in Fairhaven. A number of rivers drain into Bellingham Bay. Bellingham Bay was an
early settlement site in Puget Sound. Contamination is known in the bay, stemming from historic activities. Shellfish
growing areas were severely impacted by sulfite from pulpmills in the 1930’s and 1940’s.
Site 21. Samish Bay - small float in Samish River near Edison
This site consisted of a small series of wooden floats in line along the bank of the Samish River near its mouth into
Samish Bay. It is used by recreational boaters and fishermen.
Site 22. Swinomish Channel, South Padilla Bay
Spartina populations are known to inhabit the mudflat adjacent to the Swinomish Casino on the Swinomish
Reservation. We stopped to survey the status of control efforts there. We collected Spartina shoots, no other
sampling took place.
Site 23. Cap Sante Boat Haven, Anacortes
The city of Anacortes contains a number of marinas serving recreational boaters, many of which are located on the
west side of Fidalgo Bay. Commercial shipping facilities are primarily located either in Guemes Channel to the
north, or at the oil refineries on March Point across Fidalgo Bay, with some commercial activity also utilizing the
west side of Fidalgo Bay. In addition to oil, Anacortes is the site of log and petroleum coke shipping. The Cap
Sante Boat Haven on Fidalgo Bay was selected for its diverse fouling community and its proximity to commercial
activities.
Site 24. UW Friday Harbor Laboratories dock, San Juan Island
The University of Washington dock facility is composed of 3 large concrete pontoons, whose sides are ringed with
old tires. For nearly one hundred years, many animals and plants that have been studied at the Friday Harbor
Laboratories have been returned to the sea near the lab dock. Nearby in Friday Harbor, the Washington State ferry
terminal transports cars and passengers north to Sidney, British Columbia, and south to Anacortes. The harbor also
receives substantial private passenger ferry traffic from Seattle, recreational boaters from both Puget Sound to the
south and the Strait of Georgia to the north, and tour boats. The San Juan Archipelago lies along the route of
commercial vessels from the Seattle/Tacoma and Bellingham/Vancouver areas. Ballast water exchange has been
observed by residents living on the west side of San Juan Island during 1998.
Site 25. Argyle Lagoon and adjacent beach on San Juan Island
This site was chosen based on known non-indigenous populations. It was invaded by the cordgrass Spartina
alterniflora in the early 1990s. The cordgrass has been controlled by hand-pulling by classes from the UW Friday
Harbor Laboratories, and application of Roundup herbicide -- no shoots were spotted during our rapid survey.
Earlier this summer, Claudia Mills discovered that Argyle Lagoon and the adjacent beach on North Bay were host to
a several-year old population of the purple varnish clam, Nuttallia obscurata, including the largest known specimen,
measuring 62 mm in length. The site was surveyed only for Spartina alterniflora, Myosotella myosotis and Nuttallia
obscurata.
Page 27
Page 28
APPENDIX 4
Equipment list for the Puget Sound Expedition, Sept. 8-16, 1998
2 vans
2 radios and cell phone
vouchers for WA State ferries
48 liter-jars for voucher samples
300 scintillation vials
24, 125ml. jars
18, 250 ml jars
24, 500 ml jars
Ziploc bags in various sizes
duct tape for labels
pencils, Sharpies for labels
YSI salinity/temperature meter
cameras and film
Rubbermaid tubs - 2 about 15 gal
2 buckets
2 clipboards and large rubber bands
3-ring notebook for field notes
200 sheets punched lined paper for 3-ring
books: Kozloff’s Keys to NW Invertebrates and Light’s Manual
3 coolers
boat ramp/ marina guides
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
DNR
salinity/temperature meter
refractometer
lab thermometer
vehicle
Lamberts
Lamberts
Lamberts
Lamberts
evening lab facility with microscopes
dip net sampler on 8 foot (2.4 meter) pole
benthic sampler with 1 mm steel screen
additional bottles
vehicle
Ricoh digital camer
coolers
cellular phone
King County Environmental Lab
King County Environmental Lab
King County Environmental Lab
King County Environmental Lab
King County Environmental Lab
King County Environmental Lab
King County Environmental Lab
King County Environmental Lab
ID info for barnacles, littorines
original field notes from 1970s
Jim Carlton
Jim Carlton
label paper
Leslie Harris
3 buckets
3 dishpans
3 dipnets
10% formalin, about 5 gallons
6 gallons 70% ETOH
squeeze bottle for formalin
squeeze bottle for ethanol
squeeze bottle for water
1 lab thermometer
camera and film
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
Page 29
40 pre-printed data cover sheets
15 copies of the agenda
15 maps for participants notes
maps for drivers and participants
bowls and petri dishes for IDs
vials for Aurelia samples
jellyfish scoop
1 gallon container for seawater for evening labwork
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
Claudia Mills
custom-built scraper for pilings
steel sieves
5 gallon plastic rectangular tub
field scope and light
field thermometer
Ekman benthic grab sampler
John Chapman
John Chapman
John Chapman
John Chapman
John Chapman
John Chapman
small sieve for formalin samples
refractometer
Andy Cohen
Andy Cohen
102-125 µm mesh plankton nets and associated gear
YSI salinity/temperature meter
Jeff Cordell/UW
Jeff Cordell/UW
gloves, scrapers
personal dissecting tools
additional vials or bags for individual use
sleeping bags
specialized literature for identifications
access to FHL library provided on last 3 days
everyone
everyone
everyone
everyone
everyone
UW Friday Harbor Labs
Page 30
APPENDIX 6
Ascidians in Puget Sound and the San Juan Islands
by Gretchen Lambert, Charles Lambert, and Claudia Mills
September 1998
Ascidians are prominent members of the fouling community and are often present in very high densities
on marina floats and boat hulls. We found that, after mussels and barnacles, they comprised perhaps the
most important group of dock inhabitants in Puget Sound. Boat harbors were surveyed from a northern
extreme of Blaine to a southernmost point near Olympia on the Washington mainland, as well as sites on
Fidalgo and Whidbey Islands within the context of the Puget Sound Expedition. Sampling was
accomplished by reaching under floating docks to remove adherent ascidians and by raising ropes, hoses
and wires to examine attached organisms. Salinity and temperature were measured at each collection site.
Summarizing our findings from north to south, Drayton Harbor at Blaine contained the native species
Distaplia occidentalis and Corella inflata along with the introduced species Botrylloides violaceus and
Styela clava. This is the first report of S. clava in American waters north of San Francisco Bay in the NE
Pacific. Squalicum Harbor in Bellingham, with reduced salinity, had only a few colonies of Botrylloides
violaceus. The Edmonds Marina had the introduced Botrylloides violaceus as well as the native Distaplia
occidentalis, Corella inflata, Styela gibbsii and Diplosoma listerianum. Similar ascidians were
encountered in the Elliott Bay Marina. At the Des Moines Marina several native species were common,
but in addition there were huge numbers of Ciona savignyi, a species first recorded in California in 1985
but never before noted north of San Francisco Bay. The Des Moines Marina also contained Botryllus
schlosseri and Botrylloides violaceus. In Tacoma at Ole and Charlie’s Marina, we found B. schlosseri to
be numerous and only found a few native S. gibbsii and Distaplia occidentalis. At the Shelton Marina
Yacht Club floats, both B. violaceus and B. schlosseri were very numerous as was a third introduced
ascidian Molgula manhattensis which was very abundant, covering all submerged objects. At the
Brownsville Yacht Club we found a number of native species along with B. violaceus and a few Ciona
savignyi. The Boat Haven Marina at Port Townsend had all the previously mentioned native species and
also the native Perophora annectens and Ascidia callosa. In addition, the non-native Botrylloides
violaceus was present in large numbers.
Ascidians are thus an important component of nearly every float-fouling community in Puget Sound.
Non-indigenous species comprise a substantial part of this ascidian load, in places forming the dominant
biota. At the Des Moines Marina, Ciona savignyi occupied a roughly-estimated 90% of the concrete float
surface area under covered (shaded) docks, although it was much less abundant in uncovered areas and
limited to float bottoms. At many sites, the non-indigenous B. violaceus probably accounted for 10-20%
of the float cover, often overgrowing other organisms.
The Lamberts examined an additional seven boat harbors on San Juan Island, six more on Orcas Island,
and one on Lopez Island in the San Juans on the days immediately following the group collections.
Primarily native species were found in most harbors, but Botrylloides violaceus was in Roche Harbor,
Mitchell Bay and Westcott Bay on San Juan Island, in most harbors on Orcas Island, and in Fishermen’s
Bay on Lopez. Ciona savignyi, Molgula manhattensis and Styela clava seem to be absent from the San
Juan Archipelago at this time.
Introductions of ascidian species seem to fall on a north-south gradient. In Puget Sound we found five
introduced species; in San Francisco Bay we found eight and in southern California we found 14
(Lambert & Lambert 1998). It would be very instructive to sample southern California for non-ascidian
species to make this north-south comparison comprehensive.
Page 33
Page 34
APPENDIX 7
Commentary on species of Hydrozoa, Scyphozoa and Anthozoa (Cnidaria)
Sometimes Listed as Non-Indigenous in Puget Sound
by Claudia Mills
October 1998
A number of lists of non-indigenous marine and aquatic species for Washington State have recently been
developed (see page 1 of this report) without the specific guidance of taxonomic experts for various groups. I
offer here my assessment of the cnidarian species that have been included on some of those lists, ending with a
list of those species that should henceforth be included as either non-indigenous or cryptogenic in Puget Sound.
HYDROZOA
Bougainvillia muscus (Allman, 1863). Sometimes listed under its junior synonym B. ramosa. I collected
hydroids of this species on floats of the town docks of Friday Harbor in 1978. Medusae from the hydroids
were raised in the laboratory in order to positively identify this species. The status of this species, which has
been recorded from all over the world, is uncertain, as it may in fact represent more than one cryptic species
(Calder, 1988). It should be be listed as cryptogenic in Washington.
Cladonema radiatum Dujardin, 1843. This very small hydromedusa and its polyp have been very abundant
amongst Zostera marina on the east shore of Padilla Bay during the past decade. It is considered to be a WestAtlantic boreal species, although it has also been collected at least in Bermuda, the Bahamas, Florida and
perhaps Japan (Kramp, 1959; Calder, 1988). The species was first collected in Washington in 1988 by Eugene
Kozloff, Judy Friesen and Francis Ambrose and identified by Claudia Mills. The population was probably
established for several to many years before it was noticed. I have continued to collect it regularly in Padilla
Bay since 1988 and have made some unpublished observations on its natural history.
Cordylophora caspia (Pallas, 1771). Sometimes referred to by its junior synonym Cordylophora lacustris,
this species is reported to have been collected in Lake Union by Trevor Kincaid about 1920 (Carlton, 1979).
We found luxurious growths of this species at Edison, near the mouth of the Samish River. The Puget Sound
Expedition site at Fishermen’s Terminal was included in order to look for this species and I further searched
during October 1998 at several sites on Lake Union and in the Montlake cut, but none was found. This
species is generally considered to be native to the Caspian and Black Seas, but has been found worldwide
including a variety of low salinity sites from British Columbia to California (Carlton, 1979).
Ectopleura crocea (L. Agassiz, 1862). There are records of this Atlantic species in the San Juan Islands from
the 1930s (see Carlton, 1979), but except for an unsuccessful intentional attempt to introduce this species
using East Coast material in the early 1980s, I know of no recent collections. Confusion with several of what
are thought to be native species of Ectopleura, Tubularia or Hybocodon is likely, and species names should
only be assigned to this group with great care. This species should be removed from the list of non-indigenous
species in Washington until such time as new collections show that it is really present and established.
Gonothyraea clarki (Marktanner-Turneretscher, 1895). This is one of several Gonothyraea species
reported from the Pacific Coast by Fraser (1937), who records it from numerous locations between the
Aleutian Islands and southern California including the San Juan Islands. If all are a single species, it could
be either native eastern-Pacific or circumpolar. This genus can be separated from other members of the
family Campanulariidae based on its reproductive stuctures, which are also needed to make positive
identification to species. This species should be removed from lists of non-indigenous west coast species
pending further study.
Page 35
Obelia spp. I remain skeptical that Obelia is an exotic genus in the eastern Pacific; in a poll of my colleagues
at the September 1998 Workshop of the Hydrozoan Society in California, most hydrozoan taxonomists
present felt confidant that this genus is cosmopolitan. Species identification within this genus is highly
problematic and the number of valid species is hotly debated by specialists. At the Hydrozoan Workshop,
Professor Wim Vervoort of the National Museum of Natural History in Leiden, Netherlands, identified both
Obelia dichotoma (Linneaus, 1758) and Obelia longissima (Pallas, 1766) from the Puget Sound Expedition
samples. Until someone does an in-depth molecular and genetic study of this genus, both O. dichotoma and
O. longissima might be listed as cryptogenic in Washington.
Sarsia tubulosa (M. Sars, 1835). This species should not be on any list of non-indigenous west coast species,
as it almost certainly occurs natively on the west coast. All of the problems related to Sarsia identification,
including early references to Syncoryne mirabilis, Coryne rosaria and others (see Carlton, 1979), actually
reflect purely taxonomic (rather than biogeographic) problems. There are at least 4 species of Sarsia that seem
to be native to the Puget Sound/Strait of Georgia region, includingS. tubulosa, which appears to be a
circumboreal species (see Arai and Brinckmann-Voss, 1980). Although the genus is well known and easily
recognized, most species of Sarsia are very difficult to identify correctly, and detailed morphology of both the
medusa and its polyp must be known in order to apply a species name in most cases.
SCYPHOZOA
Aurelia aurita (Linnaeus, 1758) and Aurelia labiata Chamisso and Eysenhardt, 1821. Recent unpublished work
by Lisa Gershwin of the University of California at Berkeley has convinced me that both species are good and
that both have been found in recent times on the west coast. Certainly both species can presently be found on
display in public aquariums on the west coast, sometimes in the same tank. Although the name A. labiata has
been used in various publications including Light’s Manual Third Edition, it is Gershwin’s new morphological
observations that indicate this species to be distinctly different from A. aurita. New unpublished molecular data
by a number of scientists appears to support that conclusion. I summarize what are believed to be valid
morphological species descriptions in Wrobel and Mills (1998). Most Aurelia medusae on the west coast,
including all specimens known from Puget Sound, seem to be A. labiata, which was originally described from
near San Francisco. Specimens of A. labiata collected in Port Orchard during the Puget Sound Expedition were
sent to Werner Schroth of Johann Wolfgang Goethe-University, Frankfurt am Main, Germany, for molecular
analysis. Aurelia aurita should be removed from the list of non-indigenous species in Washington until such
time as new collections show that a second species is also present here.
ANTHOZOA
Diadumene lineata Merrill, 1870. Often listed as its junior synonym Haliplanella luciae.
This anemone with worldwide distribution was collected by the Puget Sound Expedition at the Shelton
Marina where it occurred in great numbers at the waterline of the floats. It has also been collected by Jim
Carlton, Art Siebert, and Claudia Mills on separate occasions in the 1970s on rocks in the high intertidal at the
Friday Harbor Laboratories, below the library. Bruno Pernet collected it in Padilla Bay in May 1998 on
cobbles just beyond the boulders supporting the beginning of the dyke trail, along with lots of non-indigenous
Batillaria snails. Carlton (1979) gives a fairly cohesive argument that it is likely to be non-indigenous to
Washington State, favoring the theory of an Asian Pacific origin, but since its real origin may never be
understood, it might fairly be listed as either non-indigenous or cryptogenic in Washington.
Nematostella vectensis Stephenson, 1935. Known in Puget Sound from Jaekle’s Lagoon on San Juan Island
Page 36
and a lagoon near Camp Casey on Whidbey Island. This estuarine species has an extremely broad worldwide
distribution, not so different from Diadumene lineata. Hand and Uhlinger (1994) have located it in all but two
of the United States with coastlines. There is some question of whether or not this species with widespread
global distribution is native or non-indigenous, but Hand and Uhlinger weigh in slightly favoring the spread
of this low salinity, quiet water species with mariculture and shipping. Listing this species as cryptogenic in
Washington seems most appropriate.
In summary, the following non-indigenous (NIS) and cryptogenic (C) species should be listed as present
and established in Puget Sound:
HYDROZOA
Bougainvillia muscus (= B. ramosa)
Cladonema radiatum
Cordylophora caspia (= Cordylophora lacustris)
Obelia dichotoma
Obelia longissima
C
NIS
NIS
C
C
SCYPHOZOA (NONE)
ANTHOZOA
Diadumene lineata (=Haliplanella luciae)
Nematostella vectensis
NIS or C
C
Key References
Arai, M. N. and A. Brinckmann-Voss, 1980. Hydromedusae of British Columbia and Puget Sound. Canadian
Bulletin of Fisheries and Aquatic Sciences, Bulletin 204: 1-192.
Calder, D. R. 1988. Shallow-water hydroids of Bermuda: the Athecatae. Royal Ontario Museum Life
Sciences Contributions 148: 1-107.
Carlton, J. T. 1979. History, Biogeography, and Ecology of the Introduced Marine and Estuarine Invertebrates
of the Pacific Coast of North America. Pd. D. dissertation, University of California, Davis, 903 pages.
Fraser, C. M. 1937. Hydroids of the Pacific Coast of Canada and the United States. University of Toronto
Press, Toronto, 207 pages and 44 plates.
Hand, C. and K. R. Uhlinger, 1994. The unique, widely distributed, estuarine sea anemone, Nematostella
vectensis Stephenson: a review, new facts, and questions. Estuaries 17: 501-508.
Kramp, P. L. 1959. The hydromedusae of the Atlantic Ocean and adjacent waters. Dana Report 46: 1-283.
Website for Non-indigenous Aquatic Nuisance Species (ANS), Washington State Department of Fish and
Wildlife, located at http://www.wa.gov/wdfw/fish/nuisance/ans4.htm (as of October 28, 1998).
Wrobel, D. and C. Mills, 1998. Pacific Coast Pelagic Invertebrates: a Guide to the Common Gelatinous
Animals. Sea Challengers and the Monterey Bay Aquarium, Monterey, California, 108 pages.
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