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A Rapid Assessment Survey of Non-indigenou s Species in the Shallow Waters of Puget Sound

  • Center for Research on Aquatic Bioinvasions


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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
Summary .................................................................................................................1
Non-indigenous Species Collected .........................................................................5
Future Research and Reports, Research Needs.....................................................17
Literature Cited .....................................................................................................19
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
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
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
Page 1
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 "non-
indigenous 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.
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
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 2
Overall, fewer non-indigenous species were collected in Puget Sound than were collected by
similar expeditions in San Francisco Bay. We developed an updated and corrected list of 52 non-
indigenous 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.
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 non-
indigenous 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).
5 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.
Page 3
Page 4
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 5
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 6
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 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)
B NON-INDIGENOUS SPECIES - Introduction Known but not Published for Puget Sound
C NON-INDIGENOUS SPECIES - Introduction Published for Puget Sound
E NON-INDIGENOUS SPECIES - Total (total of A through C)
Chlorophyta 0
Enteromorpha sp. x
Ulva sp. x
Rhodophyta 0
Chondria dasyphylla x
Polysiphonia paniculata x
Phaeophyceae 1
Sargassum muticum x
Anthophyta 2
Spartina anglica x
Zostera japonica x
Foraminifera 1
Trochammina hadai x
Ciliata 0
Zoothamnium sp. x
green folliculinids x
Porifera 0
Halichondria sp. x
Cnidaria: Hydrozoa 1
Cordylophora caspia (= C. lacustris) x
Cnidaria: Anthozoa 1
Diadumene lineata (= Haliplanella luciae) x
Alcyonium sp. x
Annelida: Polychaeta 2
Autolytinae sp. 1 (clear) x
Autolytinae sp. 2 (transverse bars) x
Autolytinae sp. 3 (transverse orange bars) x
Autolytinae sp. 4 (mid-dorsal orange stripe) x
Autolytinae sp. 5 (4 longitudinal lines) x
Autolytinae sp. 6 (lemon-yellow) x
Capitella sp. x
Exogone lourei x
Harmothoe imbricata x
Hobsonia florida x
Platynereis sp. x
Page 7
Table 1 (cont’d)
Annelida: Polychaeta (cont’d)
Polydora sp. x
Proceraea sp. A (of Piltz, unpubl.) x
Pseudopolydora sp. x
Typosyllis sp. x
Mollusca: Gastropoda 3
Batillaria attramentaria (= B. zonalis, = B. cumingi) x
Crepidula fornicata x
Myosotella myosotis (= Ovatella myosotis) x
Mollusca: Bivalvia 4
Crassostrea gigas x
Mya arenaria x
Mytilus sp. x
Nuttallia obscurata x
Venerupis philippinarum (= Tapes japonica) x
Arthropoda: Crustacea: Copepoda 1
Choniostomatid copepod x
Arthropoda: Crustacea: Cumacea 1
Nippoleucon hinumensis x
Arthropoda: Crustacea: Isopoda 1
Limnoria tripunctata x
Arthropoda: Crustacea: Amphipoda 9
Ampithoe valida x
Caprella mutica (= C. acanthogaster) x
Corophium acherusicum x
Corophium insidiosum x
Eochelidium sp. x
Grandidierella japonica x
Jassa marmorata x
Melita nitida x
Parapleustes derzhavini x
Bryozoa 6
Alcyonidium polyoum x
Bowerbankia “gracilis” x
Bugula stolonifera x
Bugula sp. 1 (resembling B. turbinata) x
Bugula sp. 2 (resembling B. fulva) x
Cryptosula pallasiana x
Schizoporella unicornis x
Entoprocta 1
Barentsia benedeni x
Urochordata 5
Botryllus schlosseri x
Botrylloides violaceus (earlier reported as Botrylloides sp.) x
Ciona savignyi x
Molgula manhattensis x
Styela clava x
TOTAL 11 5 23 23 39
Page 8
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 First Pacific First Puget Mechanism of
Taxon Species Native Range Coast Record Sound Record Introduction
Seaweeds Sargassum muticum Japan 1944 ? OJ
Anthophyta Spartina anglica England 1961-62 1961-62 MR
Zostera japonica W Pacific 1957 ? OJ
Foraminifera Trochammina hadai Japan 1983 1997 BW,SF,OJ
Cnidaria Cordylophora caspia Black/Caspian Seas ca. 1920 ca. 1920 BW,SF
Diadumene lineata Asia 1906 <1939 OA,SF
Annelida Hobsonia florida NW Atlantic 1940 1940 ?
Pseudopolydora sp. ? ? ? ?
Mollusca Batillaria attramentaria Japan 1924 1924 OJ
Crepidula fornicata NW Atlantic 1905 1905 OA
Myosotella myosotis Europe? 1871 1927 OA(SB,SF)
Crassostrea gigas Japan 1875 1875 OJ
Mya arenaria NW Atlantic 1874 1888-89 OA
Nuttallia obscurata Japan, Korea (China?) 1989 1991-96 BW
Venerupis philippinarum NW Pacific 1924 1924 OJ
Copepoda Choniostomatid copepod ? ? 1998 ?
Cumacea Nippoleucon hinumensis Japan 1979 1998 BW
Isopoda Limnoria tripunctata not known 1871 or 1875 ? SF
Amphipoda Ampithoe valida NW Atlantic 1941 ? BW,OA,SF
Caprella mutica Japan to Vladivostok 1973-77 1998 BW,OJ
Corophium acherusicum not known 1905 1974-75 OA,SF
Corophium insidiosum N Atlantic 1915 1930 OA,SF
Eochelidium sp. Japan or Korea early 1990s? 1997 BW
Grandidierella japonica Japan 1966 ? BW,OJ,SF
Jassa marmorata NW Atlantic 1941 ? BW, SF
Melita nitida NW Atlantic 1938 1966 BW,OA,SB,SF
Parapleustes derzhavini W Pacific? 1904 1998 SF
Entoprocta Barentsia benedeni Europe 1929 1998 OJ,SF
Bryozoa Bowerbankia gracilis NW Atlantic? <1923 <1953 OA,SF
Bugula sp. 1 ? ? 1993 ?
Bugula sp. 2 ? ? 1998 ?
Bugula stolonifera NW Atlantic <1978 1998 SF
Cryptosula pallasiana N Atlantic 1943-44 1998 OA,SF
Schizoporella unicornis NW Pacific 1927 1927 OJ,SF
Urochordata Botrylloides violaceus Japan 1973 1977 OJ,SF
Botryllus schlosseri NE Atlantic 1944-47 ? OA,SF
Ciona savignyi Japan? 1985 1998 BW,SF
Molgula manhattensis NW Atlantic 1949 1998 BW,OA,SF
Styela clava China to Okhotsk Sea 1932-33 1998 BW,OJ,SF
Page 9
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.
Figure 2. Non-indigenous Species Collected vs. Surface Salinity
Port Townsend
Port Hadlock
Port Ludlow
Cornet Bay
Des Moines
Elliott Bay
Friday Harbor
Port Orchard
Boston Harbor
Harbor Island
Samish River
Fishermen's Terminal
0 3 21 22 25 25 29 29 30 30 31 31 31 32 32 32 32 32 32 33 33 34 34
Salinity (ppt)
Number of species
Page 10
Figure 3. Non-indigenous Species Collected vs. Surface Temperature
Figure 4. Non-indigenous Species By Region
Friday Harbor
Cornet Bay
Port Hadlock
Port Townsend
Elliott Bay
Port Ludlow
Harbor Island
Des Moines
Boston Harbor
Fishermen's Terminal
Port Orchard
Samish River
Temperature (oC)
Number of species
Boston Harbor
Harbor Island
Port Orchard
Cornet Bay
Elliott Bay
Port Hadlock
Port Ludlow
Port Townsend
Samish River
Fishermen's Terminal
Des Moines
Friday Harbor
Number of species
Sou nd Hood
Canal North
Sou nd San
Jua n
Sou th
Sou nd
Page 11
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
Site Temp.
Salinity Description of Biotic Community
(Cap Sante
15° C
31 ppt
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).
(San Juan
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).
19° C
21 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 non-
indigenous 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).
17° C
30 ppt
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).
Page 12
Table 3 (Cont’d). Characterization of the Biotic Communities at Sampling Sites
Site Temp.
Salinity Description of Biotic Community
17° C
30 ppt
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).
e Marina 17° C
32 ppt 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.)
Cornet Bay
13° C
33 ppt
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).
Des Moines
16.5° C
32 ppt
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).
(Port of
15° C
32 ppt
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).
Elliott Bay
15° C
32 ppt
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).
(Port of
16.5° C
22 ppt
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).
~18° C
0 ppt 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).
(UW Lab
12° C
32 ppt
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).
19° C
31 ppt
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
Page 13
Table 3 (Cont’d). Characterization of the Biotic Communities at Sampling Sites
Description of Biotic Community
Harbor Island
16.5° C
25 ppt
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
Port Hadlock
Bay Marina
14° C
34 ppt
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).
Port Ludlow
15.5° C
33 ppt
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.
Port Orchard
18° C
31 ppt
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).
(Boat Haven
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,
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).
Marina (Hood
19° C
29 ppt
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).
Yacht Club)
20° C
29 ppt
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).
15° C
32 ppt
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).
(South Padilla
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).
Tacoma (Ole
& Charlie's
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
Page 14
collected by either the Puget Sound Expedition of 1998 or the San Francisco Bay Expeditions of
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
6 For this analysis, the species listed as Pseudopolydora sp. in Tables 1 and 2 is assumed to be either P. kempi or P.
Codium fragile tomentosoides 0 x
Sargassum muticum x x
Cotula coronopifolia 0 x
Lepidium latifolium 0 x
Salsola soda 0 x
Spartina alterniflora 0 x
Spartina anglica x 0
Zostera japonica x 0
Trochammina hadai x x
Halichondria bowerbanki ? ?
Haliclona loosanoffi ? ?
Microciona prolifera 0 x
Cnidaria: Hydrozoa
Blackfordia virginica 0 x
Cordylophora caspia x x
Ectopleura crocea 0 x
Garveia franciscana 0 x
Gonothyraea clarki 0 ?
Maeotias inexspectata 0 x
Turritopsis nutricula 0 x
Cnidaria: Anthozoa
Diadumene "cincta" 0 x
Diadumene franciscana 0 x
Diadumene leucolena 0 x
Diadumene lineata x x
Page 15
Annelida: Polychaeta
Ficopomatus enigmaticus 0 x
Heteromastus filiformis 0 x
Hobsonia florida x 0
Marenzelleria viridis 0 x
Neanthes succinea * x
Polydora ligni 0 ?
Pseudopolydora kempi ? ?
Pseudopolydora paucibranchiata ? ?
Streblospio benedicti * x
Typosyllus sp. 1 0 x
Mollusca: Gastropoda
Batillaria attramentaria x 0
Crepidula fornicata x 0
Ilyanassa obsoleta 0 x
Littorina saxatilis 0 x
Myosotella myosotis x x
Okenia plana 0 x
Philine auriformis 0 x
Tenellia adspersa 0 x
Mollusca: Bivalvia
Crassostrea gigas x 0
Gemma gemma 0 x
Geukensia demissa 0 x
Macoma petalum 0 x
Musculista senhousia 0 x
Mya arenaria x x
Nuttallia obscurata x 0
Potamocorbula amurensis 0 x
Teredo navalis 0 x
Theora fragilis 0 x
Venerupis philippinarum x x
Arthropoda: Crustacea: Copepoda
Choniostomatid copepod x 0
Arthropoda: Crustacea: Cirripedia
Balanus amphitrite 0 x
Balanus improvisus 0 x
Arthropoda: Crustacea: Nebaliacea
Epinebalia sp. 1 0 x
Arthropoda: Crustacea: Cumacea
Nippoleucon hinumensis x x
Arthropoda: Crustacea: Isopoda
Dynoides dentisinus 0 x
Iais californica 0 x
Ianiropsis serricaudis 0 x
Limnoria tripunctata x ?
Munna sp. 1 0 x
Paranthura sp. 1 0 x
Sphaeroma quoyanum 0 x
Sphaeroma walkeri 0 x
Synidotea laevidorsalis 0 x
Arthropoda: Crustacea: Tanaidacea
tanaid sp. 1 ? x
Arthropoda: Crustacea: Amphipoda
Ampelisca abdita 0 x
Ampithoe valida x x
Caprella mutica x x
Corophium acherusicum x x
Corophium alienense 0 x
Corophium heteroceratum 0 x
Corophium insidiosum x x
Eochelidium sp. 1 x x
Gammarus daiberi 0 x
Grandidierella japonica x x
Jassa marmorata x 0
Leucothoe sp. 1 0 x
Melita nitida x x
Melita sp. 1 0 x
Paradexamine sp. 0 x
Parapleustes derzhavini x x
Stenothoe valida 0 x
Transorchestia enigmatica 0 x
Arthropoda: Crustacea: Decapoda
Carcinus maenas 0 x
Eriocheir sinensis 0 x
Palaemon macrodactylus 0 x
Rhithropanopeus harrisii 0 x
Anguinella palmata 0 x
Bowerbankia “gracilis” x x
Bugula neritina 0 x
Bugula stolonifera x x
Bugula sp. 1 x 0
Bugula sp. 2 x 0
Conopeum tenuissimum ? x
Cryptosula pallasiana x x
Schizoporella unicornis x x
Watersipora "subtorquata" 0 x
Zoobotryon verticillatum 0 x
Barentsia benedeni x x
Ascidia zara 0 x
Botrylloides cf. diegensis 0 x
Botryllus schlosseri x x
Botrylloides violaceus x x
Ciona intestinalis 0 x
Ciona savignyi x x
Molgula manhattensis x x
Styela clava x x
Chordata: Pisces
Acanthogobius flavimanus 0 x
Lucania parva 0 x
Morone saxatilis 0 x
Tridentiger trigonocephalus 0 x
Page 16
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
non-indigenous 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.
Sargassum muticum
Spartina alterniflora
Spartina anglica
Spartina patens
Zostera japonica
Trochammina hadai
Cnidaria: Hydrozoa
Cladonema radiatum
Cordylophora caspia (= C. lacustris)
Cnidaria: Anthozoa
Diadumene lineata (= Haliplanella luciae)
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
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
Bowerbankia “gracilis”
Bugula stolonifera
Bugula sp. 1
Bugula sp. 2
Cryptosula pallasiana
Schizoporella unicornis
Barentsia benedeni
Botryllus schlosseri
Botrylloides violaceus
Ciona savignyi
Molgula manhattensis
Styela clava
Page 17
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 18
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
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 19
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
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 20
Page 21
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)
work phone: 360-902-1052
Brian Bingham, PhD
Huxley College of Environmental Studies
Western Washington University
Bellingham, WA 98225
(expertise in general invertebrates)
work phone: 360-650-2845
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)
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)
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)
work phone: 541-867-0235
Andrew Cohen, PhD
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)
work phone: 510-231-9423
Jeff Cordell, MS
University of Washington
Fisheries Research Institute
Box 357980
Seattle, WA 98195-7980
(expertise in copepods and other small crustaceans)
work phone: 206-543-7532
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)
work phone: 213-763-3234
Terrie Klinger, PhD
University of Washington
Friday Harbor Laboratories
620 University Road
Friday Harbor, WA 98250
(expertise in algae)
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)
work phone: 206-543-1629
Eugene Kozloff, PhD
University of Washington
Page 22
Friday Harbor Laboratories
620 University Road
Friday Harbor, WA 98250
(expertise in general invertebrates)
Charles Lambert, PhD
12001 11th Ave. NW
Seattle, WA 98177
(CSU Fullerton, Prof. Emeritus)
(expertise in tunicates)
phone: 206-365-3734
Gretchen Lambert, MS
12001 11th Ave. NW
Seattle, WA 98177
(formerly at CSU Fullerton)
(expertise in tunicates;
San Francisco Bay team 1997)
phone: 206-365-3734
Kevin Li
King County Environmental Lab
322 W. Ewing St.
Seattle, WA 98119-1507
(expertise in amphipods and other crustaceans)
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)
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)
work phone: 360-378-9105; 206-543-1484
David Secord, PhD
University of Washington, Tacoma
1900 Commerce Street, Mailstop #358436
Tacoma, WA 98402-3100
(expertise in sea anemones and general invertebrates)
work phone: 253-692-5659
Jason Toft
University of Washington
Fisheries Research Institute
Box 357980
Seattle, WA 98195-7980
(graduate student in the School of Fisheries)
work phone: 206-543-7567; 685-3395
Marjorie Wonham
University of Washington
Department of Zoology
Box 351800
Seattle, WA 98195-1800
(graduate student in the Department of Zoology)
work phone: 206-685-8615
Page 23
Puget Sound Expedition Schedule
Low tide +0.9 ' at 1151 (Seattle prediction).
1. Port of Everett Marina 9:30-10:20 am
2. Port of Edmonds Marina 11:15-12:05 pm
3. City of Des Moines Marina 2:30-3:25 pm
4. Harbor Island Marina, near mouth of Duwamish River 4:25-5:10 pm
5. Elliott Bay Marina, Seattle 5:40-6:20 pm
6. Fishermen's Terminal, Seattle 6:45-7:10 pm
Evening work in laboratory at the King County Environmental Lab.
Day 2. Sept. 9 (Wednesday) SOUTH PUGET SOUND
Low tide +1.9 ' at 1237 (Seattle prediction).
7. Ole & Charlie's Marina, Tacoma 9:50-10:40 am
8. Steilacoom Marina 12:10-12:55 pm
9. Boston Harbour Marina, near Olympia 3:05-4:10 pm
10. Port of Shelton Marina 5:20-6:05 pm
11. Fairharbor Marina, Grapeview 6:45-7:20 pm
Low tide +3.1 ' at 1326 (Seattle prediction).
12. Kitsap Marina, Port Orchard 8:00-8:45 am
13. Brownsville Marina 9:30-10:30 am
14. Seabeck Marina, Hood Canal 11:50-12:50 pm
15. Port Ludlow Marina 3:15-4:00 pm
16. Port Hadlock Bay Marina 5:00-5:30 pm
17. Boat Haven Marina, Port Townsend 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 8:50-9:40 am
19. Blaine Marina 11:15-12:10 pm
20. Squalicum Harbor, Port of Bellingham 2:25-2:45 pm
21. Samish Bay - small float in Samish River near Edison 3:35-4:05 pm
22. Padilla Bay - Vaucheria flats east of the Swinomish Channel 4:30-4:45 pm
23. Cap Sante Boat Haven, Anacortes 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 3:30-4:45 pm
C. and G. Lambert sampled 5 additional marinas on San Juan Island for tunicates.
Day 7. Sept. 14 (Monday) Friday Harbor Laboratories, SAN JUAN ISLAND.
Page 24
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 7:15-8:15 am
C. and G. Lambert sampled 6 marinas on Orcas Island for tunicates.
Sept. 15, 1998 (Tuesday) At the Friday Harbor Laboratories, SAN JUAN ISLAND.
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 ISLAND.
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 25
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 non-
indigenous 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,
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 26
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
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
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.
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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
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
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Equipment list for the Puget Sound Expedition, Sept. 8-16, 1998
2 vans DNR
2 radios and cell phone DNR
vouchers for WA State ferries DNR
48 liter-jars for voucher samples DNR
300 scintillation vials DNR
24, 125ml. jars DNR
18, 250 ml jars DNR
24, 500 ml jars DNR
Ziploc bags in various sizes DNR
duct tape for labels DNR
pencils, Sharpies for labels DNR
YSI salinity/temperature meter DNR
cameras and film DNR
Rubbermaid tubs - 2 about 15 gal DNR
2 buckets DNR
2 clipboards and large rubber bands DNR
3-ring notebook for field notes DNR
200 sheets punched lined paper for 3-ring DNR
books: Kozloff’s Keys to NW Invertebrates and Light’s Manual DNR
3 coolers DNR
boat ramp/ marina guides DNR
salinity/temperature meter Lamberts
refractometer Lamberts
lab thermometer Lamberts
vehicle Lamberts
evening lab facility with microscopes King County Environmental Lab
dip net sampler on 8 foot (2.4 meter) pole King County Environmental Lab
benthic sampler with 1 mm steel screen King County Environmental Lab
additional bottles King County Environmental Lab
vehicle King County Environmental Lab
Ricoh digital camer King County Environmental Lab
coolers King County Environmental Lab
cellular phone King County Environmental Lab
ID info for barnacles, littorines Jim Carlton
original field notes from 1970s Jim Carlton
label paper Leslie Harris
3 buckets Claudia Mills
3 dishpans Claudia Mills
3 dipnets Claudia Mills
10% formalin, about 5 gallons Claudia Mills
6 gallons 70% ETOH Claudia Mills
squeeze bottle for formalin Claudia Mills
squeeze bottle for ethanol Claudia Mills
squeeze bottle for water Claudia Mills
1 lab thermometer Claudia Mills
camera and film Claudia Mills
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40 pre-printed data cover sheets Claudia Mills
15 copies of the agenda Claudia Mills
15 maps for participants notes Claudia Mills
maps for drivers and participants Claudia Mills
bowls and petri dishes for IDs Claudia Mills
vials for Aurelia samples Claudia Mills
jellyfish scoop Claudia Mills
1 gallon container for seawater for evening labwork Claudia Mills
custom-built scraper for pilings John Chapman
steel sieves John Chapman
5 gallon plastic rectangular tub John Chapman
field scope and light John Chapman
field thermometer John Chapman
Ekman benthic grab sampler John Chapman
small sieve for formalin samples Andy Cohen
refractometer Andy Cohen
102-125 µm mesh plankton nets and associated gear Jeff Cordell/UW
YSI salinity/temperature meter Jeff Cordell/UW
gloves, scrapers everyone
personal dissecting tools everyone
additional vials or bags for individual use everyone
sleeping bags everyone
specialized literature for identifications everyone
access to FHL library provided on last 3 days UW Friday Harbor Labs
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Collections by Station of Non-indigenous Species
Insert spreadsheet here
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Insert spreadsheet here
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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.
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Page 35
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.
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 West-
Atlantic 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.
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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.
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.
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 37
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:
Bougainvillia muscus (= B. ramosa) C
Cladonema radiatum NIS
Cordylophora caspia (= Cordylophora lacustris) NIS
Obelia dichotoma C
Obelia longissima C
Diadumene lineata (=Haliplanella luciae) NIS or C
Nematostella vectensis 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
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 (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.
... In this article we explore patterns, mechanisms and hypotheses associated with marine invasions. Although new data on marine invasions have increased rapidly, they have never been summarized, beyond analysis for single bays or estuaries (11,29,30,69,117) (JT Carlton 2000, unpublished checklist; Carlton & Wonham 2000, unpublished manuscript). Moreover, the complexities and potential biases of these data, and inferences that can be drawn from the data, have not been evaluated critically. ...
... Our review and synthesis relied on four main sources of information. The primary source was published information, including especially some existing analyses of NIS for particular bays (11,29,30,69,117; JT Carlton 2000, unpublished checklist; Carlton & Wonham 2000, unpublished manuscript) as well as a diffuse collection of literature. We also reviewed unpublished reports, theses, and records from long-term monitoring efforts along each coast. ...
... Considerable variation exists in the number and overlap of known NIS among estuaries (11,29,30,67,69,117 ...
Full-text available
The known extent of biological invasions exhibits strong variation with respect to space, time, taxonomic group, and vector. Using a synthesis of nonindigenous species (NIS) occurrences in North America, we characterized the invasion history for coastal marine ecosystems by invertebrates and algae through 2010, to evaluate variation and existing patterns at a continental scale. This study updates a previous analysis of invasions for the same taxonomic groups in North America, providing a first assessment of changes in the last 11 year period (2000–2010). Overall, we documented 450 marine and estuarine NIS that are considered to have established populations in tidal waters of North America, representing a 51% increase in NIS richness compared to the earlier analysis. Of the 152 species added, 71 species (47%) have first documented records since the year 1999, and 81 species (53%) were additions attributed to earlier time periods due to recent reports and further analysis. Across all time periods, taxonomic groups with the largest contribution were Crustaceans (112 species) and Molluscs (80 species), together providing 43% of the total species richness for North America. Species richness was unevenly distributed among coasts, with most documented on the Pacific Coast (310 species) and fewer on the Atlantic Coast (189 species) and Gulf Coast (88 species). Commercial ships have contributed between (a) 44–78% of the initial (primary) invasions of all nonindigenous species to North America and (b) 52–82% of NIS in the last 30-year time interval, being driven by transfers associated with ballast water and hull biofouling. Importantly, invasion dynamics are a shifting landscape, where the past may not predict the future, especially with emerging trade patterns and global to local environmental changes. Thus, effective management to reduce future invasions requires a dynamic and multi-vector approach, instead of single vector strategies based on past history alone.
... The Pacific transparent ascidian Ciona savignyi Herdman (Cionidae) is a Japanese native with an expanding invasive range that currently includes Argentina, British Columbia, California, New Zealand, Puget Sound, and Spain (Fofonoff et al., 2003; Lambert & Lambert, 1998; Smith, Cahill & Fidler, 2010). Ciona savignyi is recognised as a problematic biofouling organism; it reproduces rapidly in invaded environments and can dominate man-made and natural substrates (e.g., Cohen et al., 1998; Zvyagintsev, Sanamyan & Kashenko, 2007). ...
... The Pacific transparent ascidian Ciona savignyi Herdman (Cionidae) is a Japanese native with an expanding invasive range that currently includes Argentina, British Columbia, California, New Zealand, Puget Sound, and Spain (Fofonoff et al., 2003; Lambert & Lambert, 1998; Smith, Cahill & Fidler, 2010). Ciona savignyi is recognised as a problematic biofouling organism; it reproduces rapidly in invaded environments and can dominate man-made and natural substrates (e.g., Cohen et al., 1998; Zvyagintsev, Sanamyan & Kashenko, 2007). This hermaphroditic species can spawn year-round in temperate regions (Nomaguchi et al., 1997; P Cahill, pers. ...
Full-text available
In this study, the effects of environmental variables on larval metamorphosis of the solitary ascidian Ciona savignyi were investigated in a laboratory setting. The progression of metamorphic changes were tracked under various temperature, photoperiod, substrate, larval density, and vessel size regimes. Metamorphosis was maximised at 18 °C, 12:12 h subdued light:dark, smooth polystyrene substrate, and 10 larvae mL(-1) in a twelve-well tissue culture plate. Eliminating the air-water interface by filling culture vessels to capacity further increased the proportion of metamorphosed larvae; 87 ± 5% of larvae completed metamorphosis within 5 days compared to 45 ± 5% in control wells. The effects of the reference antifouling compounds polygodial, portimine, oroidin, chlorothalonil, and tolylfluanid on C. savignyi were subsequently determined, highlighting (1) the sensitivity of C. savignyi metamorphosis to chemical exposure and (2) the potential to use C. savignyi larvae to screen for bioactivity in an optimised laboratory setting. The compounds were bioactive in the low ng mL(-1) to high µg mL(-1) range. Polygodial was chosen for additional investigations, where it was shown that mean reductions in the proportions of larvae reaching stage E were highly repeatable both within (repeatability = 14 ± 9%) and between (intermediate precision = 17 ± 3%) independent experiments. An environmental extract had no effect on the larvae but exposing larvae to both the extract and polygodial reduced potency relative to polygodial alone. This change in potency stresses the need for caution when working with complex samples, as is routinely implemented when isolating natural compounds from their biological source. Overall, the outcomes of this study highlight the sensitivity of C. savignyi metamorphosis to environmental variations and chemical exposure.
... During a TUBITAK project (number 111Y268), specimens of Styela clava Herdman, 1881 were encountered and photographed in one locality, Karamürsel, located in Izmit Bay. This sessile and solitary ascidian species is native to the north-western Pacific but now occurs worldwide, due to anthropogenic transport (Carlisle 1954;Millar 1960;Holmes 1976;Christiansen and Thomsen 1981;Berman et al. 1992;Cohen et al. 1998;Lambert and Lambert 1998;Minchin et al. 2006;Ross et al. 2007;Hayward and Morley 2009). It is mainly characterized by its tunic shape and long stalk. ...
Full-text available
During the implementation of a large project aimed to investigate the benthic community structures of the Sea of Marmara, specimens of the invasive ascidian species Styela clava were collected on natural substrata (rocks) at 10 m depth at one locality (Karamürsel) in İzmit Bay. The specimens were mature, containing gametes, indicating that the species had become established in the area. The Sea of Marmara seems to provide suitable conditions for this species to survive and form proliferating populations.
... We can now say with certainty that no region of the world's oceans has remained free of alien (non-indigenous) marine species (e.g., Carlton 1979, Cranfield et al. 1998, Cohen et al. 1998, Coles et al. 1999, Coles and Eldredge 2002, Hewitt 2002, Oresanz et al. 2002, Leppä koski et al. 2002, Lewis et al. 2003, Castilla et al. 2005, Wyatt et al. 2005. Similarly, alien marine species from all major animal, plant and algal phyla have been detected. ...
Macroalgae are a significant component of historic and modern invasions, with association to a wide variety of transport mechanisms. These transport mechanisms pose specific constraints on the ways by which species can be taken up, transported and released into a new environment. Currently operating transport mechanisms for marine macroalgae are either associations with intentional introductions (translocations for aquaculture, aquarium or live seafood trade) or accidental introductions (mainly as hull-fouling). A number of potential management options exist, including the development of international instruments and regional agreements. The development of treatment options for hull fouling, the most significant and poorly managed transport mechanism for macroalgae, is of urgent need. Our current ability to identify which species are likely to invade next is limited. However, an examination of the synergies between species' functional traits, transport constraints, and recipient community attributes will likely provide possible options in the future. © 2007 by Walter de Gruyter GmbH & Co. KG, 10785 Berlin All rights reserved.
... Además se ha señalado que los impactos causados por las EANI en general son irreversibles y que en el ambiente marino este problema se agudiza por el hecho de que virtualmente todas las especies marinas tienen en su ciclo de vida uno o varios estadios planctónicos , que son los que se transportan con más facilidad en el agua de lastre de los buques (Carlton, 1985). Se ha calculado que alrededor de 100 especies marinas trasladadas desde otras regiones han resultado invasoras desde 1990 (34ª Sesión del Comité de Protección del Ambiente Marino, de la Organización Marítima Internacional, Londres, julio de 1993) y se menciona que la velocidad de nuevas invasiones por agua de lastre se ha incrementado en las últimas décadas (Carlton, 1987; Cohen et al., 1998). De hecho se señala que más de 50% de las especies consideradas amenazadas o en peligro de extinción han sido afectadas por especies no nativas invasoras. ...
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Pathways are the natural and artificial introduction processes associated to human activities that result in the introduction of exotic species. In aquatic environments, aquaculture is particularly important be- cause the species are selected for their resistance and adaptability, thus presenting a higher probability of becoming invasive. In Mexico, aquaculture relies mainly on exotic species that are grown extensively. Tila- pia, carp and rainbow trout support continental aquaculture production and their propagation is carried out by government agencies (SAGARPA). The aquarium trade is a very important pathway considering the high number of species that are traded internationally, the frequency of imports and their production. Several species that have entered through this pathway are invading the country’s water bodies (e.g., ar- mored catfish, jewel cichlid, convict cichlid, lionfish). Recreational fishing is another important pathway that primarily introduces predators (e.g., bass), plus several potential invaders used as live bait or feed (e.g., red crayfish). Moreover, ballast water is recognized as the major unintended pathway with conse- quences for the environment, economy and health. It is estimated that through this process around 12 billion m3 of water are transported per year. Virtually all marine species have planktonic stages that can be moved by this pathway and it is likely that more than 3 000 are transported daily in ballast water. Unfortunately, of the various proposed treatments for ballast water, none has proved to be completely effective, considering efficiency, environmental safety, convenience and cost. Assessing the risk for ballast water in Mexico has been difficult, and to date, the introduction of about 200 non-indigenous aquatic species by this pathway has been reported. Similarly, fouling and artificial channels are pathways that cause the unintentional introduction of a large number of species that are causing economic and ecologi- cal problems. Other pathways to consider are dams, drilling platforms, boat trailers, recreational activities, and natural dispersal of floating material.
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Biological invasions can pose a severe threat to coastal ecosystems, but are difficult to track due to inaccurate species identifications and cryptic diversity. Here, we clarified the cryptic diversity and introduction history of the marine amphipod Ampithoe valida by sequencing a mtDNA locus from 683 individuals and genotyping 10,295 single-nucleotide polymorphisms (SNPs) for 349 individuals from Japan, North America and Argentina. The species complex consists of three cryptic lineages: two native Pacific and one native Atlantic mitochondrial lineage. It is likely that the complex originated in the North Pacific and dispersed to the north Atlantic via a trans-arctic exchange approximately 3 MYA. Non-native A. valida in Argentina have both Atlantic mitochondrial and nuclear genotypes, strongly indicating an introduction from eastern North America. In two eastern Pacific estuaries, San Francisco Bay and Humboldt Bay, California, genetic data indicate human-mediated hybridization of Atlantic and Pacific sources, and possible adaptive introgression of mitochondrial loci, nuclear loci, or both. The San Francisco Bay hybrid population periodically undergoes population outbreaks and profoundly damages eelgrass Zostera marina thalli via direct consumption, and these ecological impacts have not been documented elsewhere. We speculate that novel combinations of Atlantic and Pacific lineages could play a role in A. valida’s unique ecology in San Francisco Bay. Our results reinforce the notion that we can over-estimate the number of non-native invasions when there is cryptic native structure. Moreover, inference of demographic and evolutionary history from mitochondrial loci may be misleading without simultaneous survey of the nuclear genome.
The solitary ascidian, Ciona savignyi (Ascidiacea, Enterogona) is a notorious marine invader still expanding its habitat range worldwide. This species is considered native to the North West Pacific, but its indigeneity in Korean coastal waters has been questioned because of outdated taxonomic records and its inhabitation of oceanographically marginal areas. To clarify their cryptic invasion state, 247 individual C. savignyi samples were collected from 12 harbors and marinas on the Korean coast, and a 744 bp region of mitochondrial DNA (mtDNA) cytochrome c oxidase subunit I gene was sequenced and analyzed. Our analyses of population genetic structure and demographic history provided considerable pieces of evidence supporting their long-term establishment on the Korean coasts: differentiated population genetic structure, sequentially arrayed star-shape haplotype network, neutrality test results of past population expansions, and post-glacial colonization pattern of demography. Consequently, we concluded that C. savignyi populations on the Korean Coast are indigenous rather than exotic. These results could be used as reference data for further phylogeographic and demographic studies of problematic Ciona species, and to clarify and resolve similar cryptic invasion states of the other Korean coastal marine organisms. This study is the first to resolve the cryptic invasion state of Korean marine organisms using genetic analysis.
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Until very recently the Madeira Archipelago registered a total of eight Bugulidae species. In the present study we include descriptions of seven Bugulidae species, now with Scanning Electron Microscopy images, with four new records for the Archipelago: Bugulina fulva and Bugulina simplex for Madeira Island and Bugula neritina and Crisularia gracilis for the neighbouring Island of Porto Santo. Furthermore, we report the correction of the previous identification of Bugulina calathus minor earlier reported from Funchal harbour in 1998, now as Bugulina flabellata. This study is part of an ongoing monitoring program for detecting non-indigenous species in all marinas from the Madeira archipelago, between 2007 and 2015. Specimens were collected in previously deployed PVC settling plates, marina pontoons, and also on recreational hull vessels while performing dry dock inspections at a local shipyard. Our study reveals that the Madeira archipelago now registers a total of ten Bugulidae species, contributing therefore to the total bryozoan fauna of the Archipelago, now with more than 100 records. These numbers could increase, as Madeira is considered to be a “hotspot” for bryozoan diversity when compared to other nearby regions. Finally, hull fouling is considered as the most likely vector of introduction for the non-indigenous species of Bugulidae detected in Madeira.
Recent surveys of invertebrates in many coastal areas throughout the world demonstrate the ever-increasing rate of introduction of nonindigenous ascidians. Harbors, with their protected waters and large marinas, are usually the entry points for organisms carried in ballast water or on boat hulls. Marina floats, harbor buoys and boat hulls provide a huge surface area for settlement. Introductions may be further aided by transport of contaminated shells or live bivalves between mariculture facilities. Many ascidians reach sexual maturity and produce a second generation in just a few weeks. This review summarizes the results of recent studies of coastal areas, ballast water and harbor structures in many parts of the world, and sampling of U.S. Navy drydocks recently moved between major Pacific ports. Some of these studies include companion surveys of neighboring natural shallow environments. Most natural areas appear to have resisted invasions of nonindigenous ascidians except where perturbation has resulted in a decrease in species diversity.
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Samples of invertebrate communities on buoys and floating docks at Vancouver, Esquimalt, Nanaimo, Port Alberni and Prince Rupert, British Columbia, were obtained in August and September 2005. The purpose of the preliminary survey was to provide baseline data on the taxonomic composition of the invertebrates, with specific reference to possible aquatic invasive species. Thirty-five invasive or cryptogenic species were found. Conopeum seurati (Canu, 1928), a bryozoan native to the Atlantic Ocean, was confirmed from Port Alberni and may be the first record of the species from the northeast Pacific.
The small, burrowing, edwardsiid sea anemone Nematostella vectensis is widely distributed in estuaries and bays. Most typically it occurs in pools in marshes though it may occur subtidally as well. We have compiled records of its occurrence in North America from Nova Scotia to Georgia along the shores of the Atlantic Ocean, from Florida to Louisiana in the Gulf of Mexico and from California to Washington on the Pacific coast. To date we have found no records of its presence in Alabama or Texas, though it is present in all other of the contiguous coastal states of the United States. The species also occurs in England. We have obtained living specimens from many locations and have crossed females from England, Maryland, Georgia, California, Oregon, and Washington with males from Nova Scotia, Maryland, Georgia, and Oregon. These 24 crosses all yielded viable first-generation anemones that in turn produced second-generation animals. We accept this as proof that this widely distributed anemone is a single species. We have obtained living N. vectensis from 11 areas. Of these, only samples from Maine, Maryland, Georgia, and Oregon contained both sexes. The sample from Nova Scotia was all male and our samples from England, New Hampshire, California, and Washington were all female. We hypothesize that the unisexual samples were from clones resulting from asexual reproduction in this species. *** DIRECT SUPPORT *** A01BY066 00015
Southern California's many large harbors form an important coastal ecosystem, yet they are also a␣major destination for thousands of pleasure craft and cargo vessels that have often traveled great distances. Many groups of marine organisms, including ascidians, have now been documented as undergoing range extensions as a consequence of rapid ship-transport between distant harbors phenomenon. This has resulted in a rapid increase in the rate of introductions of non-indigenous species worldwide, yet these effects of boat traffic remain largely unstudied in southern California. Ascidians are sessile marine filter-feeders, hermaphroditic, and often self-fertilizing; many species are tolerant of a wide range of environmental conditions, can reach sexual maturity in just a few weeks, and have a long breeding season. This paper documents the arrival of 14␣non-indigenous species in southern California harbors␣during this century, 13 of which have persisted:␣four prior to the 1960s (Cionaintestinalis, Styelaclava, S.␣plicata, Botryllusschlosseri), another by 1972 (S.␣canopus, formerly S. partita), and 8 since 1983 [C.␣savignyi, Ascidia zara, Ascidia sp., Polyandrocarpa zorritensis, Symplegma brakenhielmi (formerly S. oceania, and S. reptans, Microcosmus squamiger, and Molgula␣manhattensis)]. We estimate the relative abundance and seasonal fluctuations of both non-indigenous and native ascidians in all harbors in southern California from San Diego to Santa Barbara based upon the historical record, our 35 yr of field notes, and our recent surveys carried out during fall 1994, spring and fall 1995, fall 1996 and spring 1997. Possible points of origin of the exotics and predictions on further U.S. Pacific coast range-extensions are included. The concomitant decline in numbers and species of native ascidians in the harbors of southern California during this century is also reviewed.
Thesis (Ph. D.)--University of California, Davis, 1979. Includes bibliographical references (leaves 749-857). Microfiche. s
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. 22. Padilla Bay -Vaucheria flats east of the
  • G M Ruiz
  • A H Hines
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. 22. Padilla Bay -Vaucheria flats east of the Swinomish Channel 4:30-4:45 pm 23. Cap Sante Boat Haven, Anacortes 4:50-5:40 pm
Key to the Marine Invertebrates of Puget Sound, the San Juan Archipelago, and Adjacent Regions
  • E N Kozloff
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.