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Report on the 2013 rapid assessment survey of marine species at New England bays and harbors

Authors:

Abstract and Figures

Since 2000, five rapid assessment surveys have been conducted in New England. These surveys focus on recording species at marinas, which often are in close proximity to transportation vectors (i.e., recreational boats). Species are collected from floating docks and piers because these structures are accessible regardless of the tidal cycle. Another reason for sampling floating docks and other floating structures is that marine introduced species are often found to be more prevalent on artificial surfaces than natural surfaces (Glasby and Connell 2001; Paulay et al. 2002). The primary objectives of these surveys are to: (1) identify native, introduced, and cryptogenic marine species, (2) expand on data collected in past surveys, (3) assess the introduction status and range extensions of documented introduced species, and (4) detect new introductions. This report presents the introduced, cryptogenic, and native species recorded during the 2013 survey.
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Report on the 2013
Rapid Assessment Survey of
Marine Species at New England
Bays and Harbors
June 2014
AUTHORED BY: Christopher D. Wells, Adrienne L. Pappal, Yuangyu Cao, James T. Carlton, Zara Currimjee, Jennifer A.
Dijkstra, Sara K. Edquist, Adriaan Gittenberger, Seth Goodnight, Sara P. Grady, Lindsay A. Green, Larry G. Harris,
Leslie H. Harris, Niels-Viggo Hobbs, Gretchen Lambert, Antonio Marques, Arthur C. Mathieson, Megan I. McCuller,
Kristin Osborne, Judith A. Pederson, Macarena Ros, Jan P. Smith, Lauren M. Stefaniak, and Alexandra Stevens
This report is a publication of the Massachusetts Office of Coastal Management (CZM) pursuant to the National Oceanic
and Atmospheric Administration (NOAA). This publication is funded (in part) by a grant/cooperative agreement to CZM
through NOAA NA13NOS4190040 and a grant to MIT Sea Grant through NOAA NA10OAR4170086. The views expressed
herein are those of the author(s) and do not necessarily reflect the views of NOAA or any of its sub-agencies. This project
has been financed, in part, by CZM; Massachusetts Bays Program; Casco Bay Estuary Partnership; Piscataqua Region
Estuaries Partnership; the Rhode Island Bays, Rivers, and Watersheds Coordination Team; and the Massachusetts
Institute of Technology Sea Grant College Program.
Commonwealth of Massachusetts
Deval L. Patrick, Governor
Executive Office of Energy and Environmental Affairs
Maeve Vallely Bartlett, Secretary
Massachusetts Office of Coastal Zone Management
Bruce K. Carlisle, Director
Massachusetts Office of Coastal Zone Management
251 Causeway Street, Suite 800
Boston, MA 02114-2136
(617) 626-1200
CZM Information Line: (617) 626-1212
CZM Website: www.mass.gov/czm
C R E D I T S
PHOTOS: Adriaan Gittenberger, Gretchen Lambert, Linsey Haram, and Hans Hillewaert
The New England Rapid Assessment Survey was a collaborative effort of many individuals.
In addition to the authors of this paper, who were directly involved in the identification
of the organisms, we thank those individuals that provided logistical support: Beverly
Bayley-Smith, Casco Bay Estuary Partnership, Portland, Maine; Kevin Cute, Rhode Island
Coastal Resources Management Council, Wakefield, Rhode Island; and Tracy Warncke,
Buzzards Bay National Estuary Program, East Wareham, Massachusetts. We also would
like to thank Larry Harris (University of New Hampshire) and David Murray (Brown
University) for generous use of their laboratory facilities.
We are thankful for the willingness of the owners and operators of the various marinas
who allowed access to their sites during the Rapid Assessment Survey: John Brewer,
Brewer South Freeport Marine, South Freeport, Maine; Mike Soucy, Spring Point Marina,
Portland, Maine; Noel Carlson, University of New Hampshire Coastal Marine Laboratory,
New Castle, New Hampshire; Kevin Bailey, Hampton River Marina, Hampton, New
Hampshire; Noah Flaherty, Brewer Hawthorne Cove Marina, Salem, Massachusetts;
Ryan O’Connor, Rowes Wharf Marina, Boston, Massachusetts; Michael DiMeo,
Green Harbor Marina, Marshfield, Massachusetts; Troy Lima, Sandwich Marina,
Sandwich, Massachusetts; David Remsen, Marine Biological Laboratory, Woods Hole,
Massachusetts; William Klimm, Massachusetts Maritime Academy, Buzzards Bay,
Massachusetts; Debra Yuille, Popes Island Marina, New Bedford, Massachusetts; Rich
Picard, F.L. Tripp & Sons Marina, Westport, Massachusetts; Ray Mooney, Port Edgewood
Marina, Cranston, Rhode Island; Ed Hughes, Allen Harbor Marina, North Kingstown,
Rhode Island; Richard Masse, Fort Adams State Park, Newport, Rhode Island; and
Anne Killeen, Point Judith Marina, Wakefield, Rhode Island.
This study was supported by funding from the Casco Bay Estuary Partnership; the
Massachusetts Bays National Estuary Program; the Massachusetts Institute of Technology
Sea Grant College Program; the Massachusetts Office of Coastal Zone Management;
the Piscataqua Region Estuaries Partnership; and the Rhode Island Bays, Rivers,
and Watersheds Coordination Team.
A C K N O W L E D G M E N T S
TABLE OF CONTENTS
Introduction...........................................................................1
Me thods...........................................................................3
Results...........................................................................4
Summary...........................................................................8
Appendix I: Site Descriptions..................................................10
Appendix II: Rapid Assessment Survey Participants.............14
Appendix III: Hydrographic Data...........................................15
References...........................................................................16
Lists of Introduced, Cryptogenic,
and Native Species................................................................19
1
Introduced species (i.e., non-native species that have become established in
a new location) have increasingly been recognized as a concern as they have
become more prevalent in marine and terrestrial environments (Mooney and
Cleland 2001; Simberloff et al. 2005). The ability of introduced species to alter
population, community, and ecosystem structure and function, as well as
cause significant economic damage is well documented (Carlton 1989, 1996b,
2000; Cohen and Carlton 1995; Cohen et al. 1995; Elton 1958; Meinesz et al.
1993; Occhipinti-Ambrogi and Sheppard 2007; Pimentel et al. 2005; Thresher
2000). The annual economic costs incurred from managing the approximately
50,000 introduced species in the United States alone are estimated to be over
$120 billion (Pimentel et al. 2005).
Having a monitoring network in place to track new introductions and
distributional changes of introduced species is critical for effective
management, as these efforts may be more successful when species are
detected before they have the chance to become established. A rapid
assessment survey is one such method for early detection of introduced
species. With rapid assessment surveys, a team of taxonomic experts
record and monitor marine species–providing a baseline inventory of
native, introduced, and cryptogenic (i.e., unknown origin) species (as
defined by Carlton 1996a)–and document range expansions of previously
identified species.
Since 2000, five rapid assessment surveys have been conducted in New
England. These surveys focus on recording species at marinas, which often
are in close proximity to transportation vectors (i.e., recreational boats).
Species are collected from floating docks and piers because these structures
are accessible regardless of the tidal cycle. Another reason for sampling floating
docks and other floating structures is that marine introduced species are often
found to be more prevalent on artificial surfaces than natural surfaces (Glasby
and Connell 2001; Paulay et al. 2002). The primary objectives of these surveys
are to: (1) identify native, introduced, and cryptogenic marine species,
(2) expand on data collected in past surveys, (3) assess the introduction status
and range extensions of documented introduced species, and (4) detect new
introductions. This report presents the introduced, cryptogenic, and native
species recorded during the 2013 survey.
Introduction
RAPID ASSESSMENT SURVEY: 2013 SAMPLING SITES
2
Methods
Eighteen sampling locations were chosen for the
2013 New England Rapid Assessment Survey (RAS)
in coordination with National Estuary Programs and
other partners. Factors involved in choosing sites were:
adequate access for a team of 25 scientists, even
distribution along the length of the shoreline to be
examined, and consistency with sampling conducted
during the previous five surveys (see map on page 2 for
locations). The sites were distributed as follows: three
sites in Maine, two sites in New Hampshire, four sites
in northern Massachusetts (i.e., north of Cape Cod),
five sites in southern Massachusetts (i.e., south of
Cape Cod), and four sites in Rhode Island. Locations
were marine with little freshwater influence, except
at Port Edgewood Marina in Cranston, Rhode Island.
For more detailed information on the sampling
locations, including the sampling date and time,
a brief description of the site, and the dominant
biological community, see Appendix I.
Fouling communities were sampled on permanently
installed floating docks and on associated subtidal
structures, such as the floats supporting the docks,
ropes, wires, buoys, tires, and boat fenders. Motile
animals associated with the fouling community, such
as crabs, shrimps, and fishes, were documented and
included in this report, but may be transient residents
of the fouling community. Species observed on
stationary piers, sea walls, rocky shores, pilings,
and other nearby habitats were recorded, but are not
included in the report. Other species that are difficult
to identify without reproductive structures present
(e.g., hydroids and certain algae species) were
identified to genus.
The participants in the RAS included taxonomic
experts familiar with native and introduced marine or-
ganisms, graduate and undergraduate students, and a
support team to manage logistics. See Appendix II for
details on the RAS survey team, including taxonomic
expertise and affiliations. Sampling was carried out in
early August to facilitate comparison between this RAS
and prior surveys’ efforts. Although the assemblages
found within fouling communities vary from year to
year because of environmental conditions, long-term
monitoring helps to smooth out shorter-term variations
to allow for analysis of trends through time.
Colpomenia peregrina
The macroscopic stage of the sea potato alga Colpomenia peregrina
forms hollow brown sacks frequently found attached to other algae within
the intertidal area and just below the tide line. Colpomenia peregrinawas
not found during the 2010 New England Rapid Assessment Survey, but
was found at Sandwich Marina in Sandwich, Massachusetts, during the
2013 survey. It is becoming more and more common north of Cape Cod.
Colpomenia peregrina was initially recorded in Nova Scotia in the 1960s
(Bird and Edelstein 1978; Blackler 1964; Villard-Bohnsack 2002), followed
by a collection in the early 1980s from Grand Barachois Lagoon in
Newfoundland (Green et al. 2012). It has since spread southward from
mid-coastal Maine to Massachusetts, initially found in 2011 (Green et al.
2012). Colpomenia peregrina is originally from the northwestern Pacific,
but has been introduced to Europe and the west coast of North America
(Boo et al. 2011). Care should be taken in identifying this alga as it is
frequently confused with a similar native brown alga, Leathesia marina.
3
Colpomenia peregrina
At each location, sampling time was limited to one
hour. Scientists atop the docks collected specimens
from the vertical sides of the floats by hand or with
paint scrapers and dip nets. The undersides of the
floats were sampled by two snorkelers, except at the
Woods Hole U.S. Coast Guard Floating Dock due to se-
curity precautions. Species were brought back to labo-
ratories at the University of New Hampshire or Brown
University for identification and verification. Collected
specimens were either retained by investigators or
archived within a community voucher deposited at the
Museum of Comparative Zoology at Harvard University.
The status of invertebrates as native, introduced,
cryptogenic, or summer transient was assessed by
using the bioinvasion literature on individual species.
For other species, the criteria summarized in Chapman
and Carlton (1991, 1994) were applied, often combined
with judgments assessing the extent of recent taxo-
nomic and biogeographic work on a given species or a
given group in general. The result is that the number of
introduced species in New England remains underesti-
mated (Carlton 2003, 2009); that is, an unknown
number of species, here treated as either cryptogenic
or native, are in fact introduced. No algal species
are treated here as cryptogenic, but the same
considerations apply.
Water temperature, salinity, and dissolved oxygen
were collected through the use of two underwater
sondes (YSI30 and YSI85, YSI Inc., South Burlington,
VT). For the results of the hydrographical data,
see Appendix III.
Results
During the RAS, a total of 289 taxa were identified,
of which 218 were native, 27 were cryptogenic,
39 were introduced, and five were either summer
transient species or their establishment was unknown
(Figure 1). Hawthorne Cove Marina in Salem,
Massachusetts, had the highest percentage of
introduced species (32%). Port Edgewood Marina in
Cranston, Rhode Island, had the highest percentage
of native species (82%). Complete lists of introduced,
cryptogenic, and native taxa are provided at the end
of the document. Table 1 provides the average number
of native, cryptogenic, and introduced species found
within each state (i.e., the total number of native,
cryptogenic, and introduced species found at each
site within a state, divided by the number of survey
sites within that state). Native taxa compose similar
percentages of total taxa in all states (69-70%),
except for northern Massachusetts where they
represent a relatively low percentage (62%). The
percentage of cryptogenic taxa was highest in
northern Massachusetts (12%) and lowest in southern
Massachusetts (7%). Introduced taxa compose
similar percentages of total taxa in all states
(19-21%), except for northern Massachusetts (25%).
For a complete list of the introduced species found
within each state see Table 2.
Figure 1: Total number of native, cryptogenic, introduced,
and summer transient species found during the 2013
Rapid Assessment Survey.
218
5
39
27
Summer
Transient
Introduced
Cryptogenic
Native
The hair alga
Cladophora
sericea (green)
is a native
species.
4
Native 53 ±1 53 ±7 49 ±5 58 ±2 52 ±7
Cryptogenic 8 ±2 8 ±1 10 ±1 6 ±1 6 ±1
Introduced 14 ±1 15 ±2 20 ±1 18 ±1 16 ±4
Total 76 ±3 76 ±7 80 ±6 83 ±4 74 ±10
Table 1: Average number of native, cryptogenic, and introduced species found within each
state with standard error rounded to the nearest whole number. Massachusetts is split
between sites north and south of Cape Cod.
5
Melita palmata
SPECIES AVERAGE BY STATE
SPECIES TYPE ME NH MA (N) MA (S) RI
Melita palmata
The males of the European palmed gammarid Melita palmata have a
distinct paddle-like leg, which makes the animal very easy to distinguish
from other Melita species. During the 2013 Rapid Assessment Survey,
only one individual was found at a single site: Rowes Wharf in Boston,
Massachusetts. Although it has been introduced to the southeast coast
of South America, Melita palmata has never been recorded in New
England, and it is unknown whether there is an established population.
No additional specimens were found on a return trip to Rowes Wharf
looking specifically for this species.
Chlorophyta (green algae)
Codium fragile subsp. fragile (dead man's fingers) xxx
Heterokontophyta (brown algae)
Colpomenia peregrina (sea potato) x
Melanosiphon intestinalis (dark sea tubes) x
Rhodophyta (red algae)
Bonnemaisonia hamifera (hooked red weed) xx
Gracilaria vermiculophylla (wormleaf) xx
Grateloupia turuturu (Devil's tongue weed) xxx
Heterosiphonia japonica (siphoned feather weed) xxx
Lomentaria clavellosa (club bead-weed) xx xx
Lomentaria orcadensis (Orkney weed) x
Neosiphonia harveyi (doughball weed) xx x x x
Pyropia yezoensis (open sea nori) xx
Annelida (ringed worms)
Neodexiospira brasiliensis (Brazilian spiral tubeworm) xx
Arthropoda (crabs, shrimp, and relatives)
*Amphibalanus amphitrite (striped barnacle) x
Caprella mutica (Japanese skeleton shrimp) xx x x x
Carcinus maenas (European green crab) xx x x x
Hemigrapsus sanguineus (Asian shore crab) xx x x x
Ianiropsis serricaudis (isopod) xx xx
*Melita palmata (palmed gammarid) x
Microdeutopus gryllotalpa (tube builder) xx x
Palaemon elegans (European rock pool shrimp) xx x
*Palaemon floridanus (Florida grass shrimp) x
Palaemon macrodactylus (oriental grass prawn) x
Praunus flexuosus (bent opossum shrimp) xx x
*Stenothoe valida (gammarid amphipod) xx x
Bryozoa (bryozoans)
Bugula neritina (purple bugula) xxx
Bugula simplex (fan bugula) xx x x x
Bugula stolonifera (stoloned bugula) xx
Conopeum seurati (encrusting bryozoan) x
Membranipora membranacea (lacy crust bryozoan) xx x x
Tricellaria inopinata (unexpected bryozoan) xx xx
Chordata (tunicates and vertebrates)
Ascidiella aspersa (European tunicate) xxxx
Botrylloides violaceus (sheath tunicate) xx x x x
Botryllus schlosseri (star tunicate) xx x x x
Botryllus sp. (colonial tunicate) xxx
Didemnum vexillum (pancake batter tunicate) xx x x x
SPECIES (COMMON NAME) ME NH MA (N) MA (S) RI
Table 2: List by state of introduced species identified during the 2013 Rapid Assessment Survey.
Massachusetts is split between sites north and south of Cape Cod. Those species marked with
an asterisk (*) are either summer transient species or their establishment is unknown.
6
7
Diplosoma listerianum (gelatinous tunicate) xx x x x
Styela canopus (rough tunicate) xx
Styela clava (club tunicate) xxxx
Cnidaria (anemones, hydroids, and relatives)
Aiptasiogeton eruptaurantia (pink-spotted sea anemone) x
*Clytia linearis (hydroid) xxx
Diadumene lineata (orange striped anemone) xxxx
Hydrodendron sp. (hydroid) x
Mollusca (snails and bivalves)
Littorina littorea (common periwinkle) xx x
Ostrea edulis (European oyster) xx x x x
Total Species 22 20 30 32 29
Six introduced species were found for the first time in 2013: Aiptasiogeton eruptaurantia,
Colpomenia peregrina, Hydrodendron sp., Neodexiospira brasiliensis, Palaemon
macrodactylus, and Tricellaria inopinata. Several species found on prior surveys expanded
in range, some explosively (e.g., Heterosiphonia japonica and Palaemon elegans).
Tricellaria inopinata
Tricellaria inopinata, commonly known
as the unexpected bryozoan, has been
described as having a “crunchy” feeling
when compared to other branching bry-
ozoans, but it cannot be identified with
complete certainty without a dissecting
microscope on a high power. Tricellaria
inopinata was originally found in Eel Pond
in 2010 (Johnson et al. 2012) and has
quickly spread north as far as Gloucester
(Wells and Pappal, personal observation)
and south to Rhode Island. During the
2013 Rapid Assessment Survey, it was
found at nine of 18 sites from Salem,
Massachusetts, to Newport, Rhode Island
(see Table 5 for more details). Tricellaria
inopinata is thought to be native to the
western Pacific and has been introduced
to Europe, Australia, New Zealand, and
the West Coast of the United States. Tricellaria inopinata
Table 2 continued
SPECIES (COMMON NAME) ME NH MA (N) MA (S) RI
8
Summary
Introduced species represented a large component
of the total fouling community from Maine to Rhode
Island (up to 31%). The number of introduced species
has steadily been increasing within New England
waters; during the 2010 New England RAS, 29 intro-
duced species were found (McIntyre et al. 2013) in
contrast to the 39 detected during the 2013 survey
(Table 3) (care should be taken when comparing the
results of this survey and previous surveys as effort
has varied greatly). The total number of introduced
species recorded is far lower than similar surveys
carried out in California (Cohen et al. 2005; Boyd et
al. 2002; Lambert and Lambert 1998, 2003), Australia
(Hewitt et al. 2004), and Hawaii (Coles et al. 1999), but
is on par with results of similar surveys in the Pacific
Northwest, Eastern Atlantic, and New England (Arenas
et al. 2006; Cohen et al. 1998, 2001; Mathieson et al.
2008; Mills et al. 2000; Pederson et al. 2005).
There were distinct southern United States biota
identified at several of the sites during the 2013
RAS, including the striped barnacle Amphibalanus
amphitrite, the pink-spotted anemone Aiptasiogeton
eruptaurantia, and the Florida grass shrimp Palaemon
floridanus. Both A. amphitrite and P. floridanus are
summer transient species that will likely not survive
the New England winter; A. amphitrite is the only
species of the three with a history of summer appear-
ances in New England (Carlton, personal observation).
Although A. eruptaurantia does not have a long history
of presence in southern New England, it has been
observed at Point Judith Marina in Rhode Island since
the summer of 2011 (Hobbs, personal observation)
and is now considered an established introduced
species. Also present during this RAS were several
hydroids more commonly found in warmer waters:
Pennaria disticha, Schizotricha tenella, Eudendrium
glomeratum, and Hydrodendron sp. While the first
three of these hydroids have been documented in
New England previously, they are being found more
frequently. The two previous winters, which were warm
for New England, could have facilitated the increase
in warm-water species (Wells, personal observation).
These animals were likely transported to New England
by coastal recreational and commercial vessels via
ballast water or hull fouling. Future warming due to
climate change may enhance the establishment of
these and other warm-water species.
The number of species documented during the
2013 RAS is only a small portion of the known flora
and fauna of New England. For example, of the approx-
imately 500 algae species found within New England
(Mathieson, personal communication), 90 algae
species were found. Rapid assessments are limited by
time, seasonality of fouling species, and number of
habitats evaluated (Mathieson et al. 2008; Pederson et
al. 2005). While not intended to be complete surveys
of a site’s biota, the RAS effectively and economically
identifies presence/absence of species, finds newly
introduced species, and detects range expansions.
Surveys like the RAS are critical for providing a base-
line inventory of native, cryptogenic, and introduced
species, and continued monitoring of at-risk habitats
is a valuable asset for monitoring new introductions
and range expansions. The data collected during the
New England surveys can help managers prevent and
control future invasions as well as aid in the creation of
legislation for controlling and eradicating introduced
species. Future surveys will continue to build on this
wealth of knowledge and help promote understanding
of invasion processes and the ecology of New
England’s coastal habitats.
The docks at Wells Harbor Marina.
2000 34 Gloucester (MA) to Narragansett Bay (RI) 32
2003 20 Portland (ME) to Staten Island (NY) 29
2007 17 Rockland (ME) to Buzzards Bay (MA) 26
2010 20 Cape Elizabeth (ME) to Narragansett Bay (RI) 29
Table 3: Number of introduced species found on the past five New England Rapid Assessment Surveys.
Palaemon
elegans
Palaemon
macrodactylus
YEAR SITES SURVEY RANGE SPECIES
Introduced Shrimps: Palaemon elegans and P. macrodactylus
The European rock shrimp Palaemon elegans can be easily identified by its distinctive blue
and yellow claws as well as prominent blue and yellow striping and spotting on its carapace.
Palaemon elegans was first documented in New England during the 2010 New England Rapid
Assessment Survey at a single site: Hawthorne Cove Marina in Salem, Massachusetts
(McIntyre et al. 2013). It has since rapidly spread as far north as Portland, Maine, and as far
south as Provincetown, Massachusetts. It is expected to continue to spread both
north and south.
The oriental grass prawn P. macrodactylus is native to the western Pacific, but has been intro-
duced across Europe and the southeastern coast of South America. Palaemon macrodactylus
is easily confused with other shrimp such as the native marsh grass shrimp Palaemonetes
vulgaris. A faint white stripe down the center of the top of the carapace has been suggested
as an identifying characteristic of P. macrodactylus, but this character can also be present
in Palaemonetes vulgaris. The only definite way to identify this species is by looking at the
rostrum (i.e., the forward extension of the carapace in front of the eyes) with a magnifying
glass (for large individuals) or under a dissecting microscope.
Palaemon macrodactylus was originally collected in 2001 within
the estuary system of New York City (Warkentine and Rachlin
2010), and was found at one site during this survey: Fort Adams
in Rhode Island.
9
Appendix I: Site Descriptions
Brewer South Freeport Marine (BFM), South Freeport, Maine | August 4, 2013, 1:30 PM
Brewer South Freeport Marine is a relatively large marina located on the banks of the Harraseeket River on the edge of Casco Bay. It
historically served as a shipbuilding port and now provides service to both recreational craft and commercial lobstermen. The marina
provides over 100 slips and 15 moorings with dockside depths of approximately four meters. During the RAS, a total of 80 species were
identified at BFM, of which 55 were native, nine cryptogenic, 15 introduced, and one a summer transient species. The dominant species
at BFM were a mixture of the blue mussel Mytilus edulis or M. trossulus (unable to identify to species without genetic analysis), the
frilled sea anemone Metridium senile, and the introduced sheath tunicate Botrylloides violaceus.
Spring Point Marina (PHM), Portland, Maine | August 4, 2013, 10:30 AM
Spring Point Marina, formerly known as Port Harbor Marine, is the largest full-service marina in Maine. Spring Point Marina is located
at the entrance of Portland Harbor and the mouth of the Fore River. The marina was originally the site of the South Portland Shipyard,
which built Liberty Ships during World War II. Floats were low in the water and heavily fouled. During the RAS, a total of 76 species were
identified at PHM, of which 53 were native, 10 were cryptogenic, and 13 were introduced. The dominant species at PHM were a mixture
of the blue mussels M. edulis/trossulus and large, heavily fouled blades of the sugar kelp Saccharina latissima. The frilled sea anemone
M. senile and the bent opossum shrimp Praunus flexuous were abundant on the underside of the docks.
Wells Harbor Marina (WHM), Wells, Maine | August 4, 2013, 8:15 AM
Wells Harbor Marina is located at the mouth of the Webhannet River. Nearby salt marsh and beach areas provide nursery habitats for
estuary-dependent species. During the RAS, a total of 71 species were identified at WHM, of which 52 were native, four cryptogenic, 14
introduced, and one a summer transient species. The dominant species at WHM were a mixture of the blue mussels M. edulis/trossulus
and the hydroid Ectopleura larynx.
University of New Hampshire Coastal Marine Lab Floating Dock (UNH), New Castle, New Hampshire | August 5, 2013, 7:30 AM
The UNH floating dock is located at the mouth of Portsmouth Harbor adjacent to the historic Fort Constitution. The marine laboratory
provides faculty and students with access to the Gulf of Maine and includes facilities with running seawater and a 100-meter research
pier that provides berthing space for the UNH fleet of research vessels. Underneath the pier and adjacent to the floating dock there are
experimental enclosures used for marine research. Floats holding the dock up are attached farther in and are deep and were therefore
difficult to sample without the aid of divers. During the RAS, a total of 82 species were identified at the UNH floating dock, of which
59 were native, nine cryptogenic, 13 introduced, and one a summer transient species. The floating dock was dominated by the blue
mussels M. edulis/trossulus, dead remains of the hydroidE. larynx, and large individuals of the frilled sea anemone M. senile. There
was an abundant population of the medusa (i.e., jellyfish) life stage of the hydroid Clytia hemisphaerica found at this site.
Hampton River Marina (HSP), Hampton, New Hampshire | August 5, 2013, 9:30 AM
Hampton River Marina is located at the mouth of the
Piscataqua River Estuary. The marina features 144 boat slips.
The bottom is littered with bivalve shells; most appear to
be sourced from the fouling community. The area is highly
developed and supports a large amount of tourism. During
the RAS, a total of 69 species were identified at HSP, including
46 native species, seven cryptogenic species, and 16 intro-
duced species. The floating docks were dominated by the
blue mussel M. edulis/trossulus and a mixture of introduced
colonial and solitary tunicates. There was an orange-ringed
color morph of the cryptogenic sea vase tunicate Ciona
intestinalis found at this site.
The native orange sea grape
tunicate Molgula citrina was
only found north of Cape Cod.
10
Brewer Hawthorne Cove Marina (HCM),
Salem, Massachusetts | August 5, 2013, 12:15 PM
Brewer Hawthorne Cove Marina, part of the historic
Salem shipping port, is a private marina with 110 slips
located on the northern shore of Salem Harbor. Depth
ranges from 1-4 meters. During the RAS, a total of 68
species were identified at HCM, including 38 native
species, eight cryptogenic species, 21 introduced
species, and one summer transient species. This site
had the highest percentage of introduced species
(32%). Float fouling communities were dominated by
tunicates and branching bryozoans on the vertical
surfaces and mussels on the horizontal surfaces.
Hawthorne Cove Marina is the only location where
the anemone Sagartia elegans has been reported in
the Western Atlantic, but this species disappeared
after the winter of 2010-2011 (Wells 2013).
The Marina at Rowes Wharf (ROW), Boston,
Massachusetts | August 6, 2013, 9:30 AM
The Marina at Rowes Wharf is located along the highly developed waterfront of Boston’s inner harbor and is part of the Rowes Wharf
luxury hotel and condominium complex. It has 38 slips that can accommodate mega yachts. During the RAS, a total of 91 species were
identified at ROW including 57 native species, 12 cryptogenic species, 21 introduced species, and one summer transient species. The
floating docks had a light covering of the blue mussels M. edulis/trossulus and a mixture of introduced colonial and solitary tunicates
as well as encrusting sponges. One individual of the introduced amphipod Melita palmata was found at ROW (see text box on page 5).
Green Harbor Marina (GHM), Marshfield, Massachusetts | August 6, 2013, 12:45 PM
Green Harbor Marina is located on the eastern side of Green Harbor located on the South Shore of Massachusetts. Green Harbor
Marina has approximately 135 slips. Pontoons were composed of polystyrene foam. During the RAS, a total of 71 species were identified
at GHM, including 45 native species, seven cryptogenic species, 18 introduced species, and one summer transient species. The fouling
community was dominated by solitary and colonial tunicates and branching bryozoans. The blue mussels Mytilus edulis/trossulus and
barnacles were rare.
Sandwich Marina (SWM), Sandwich, Massachusetts | August 8, 2013, 8:30 AM
Sandwich Marina is an important site for monitoring the northerly progression of introduced species via the Cape Cod Canal. The
marina features 164 slips for recreation ships and 42 commercial slips. During the RAS, a total of 90 species were found at SWM,
including 57 native species, 11 cryptogenic species, 21 introduced species, and one summer transient species. The fouling community
was dominated by solitary and colonial tunicates.
The introduced bryozoan Bugula neritina hosts a bacterium that
produces a chemical that may have anti-cancer properties.
The native hair alga
Cladophora sericea
was very common
at Hawthorne Cove
Marina where it forms
dense clumps on the
floats just beneath
the surface.
11
US Coast Guard Station Floating Dock (WHC), Woods Hole, Massachusetts | August 8, 2013, 1:45 PM
The Woods Hole Coast Guard Station has been in existence since 1857. The station is located on a highly developed shoreline on the
Vineyard Sound side of Cape Cod. This site was surveyed for 30 minutes per the request of the Coast Guard; divers were not allowed
in the water for security reasons and therefore organisms found deep on the vertical surfaces and on the horizontal surfaces are likely
underrepresented. During the RAS, a total of 79 species were found at WHC, including 55 native species, four cryptogenic species,
18 introduced species, and two summer transient species. The fouling community was dominated by solitary and colonial tunicates
growing over the abundant common jingle shell Anomia simplex. One specimen of the Florida grass shrimp Palaemon floridanus was
collected at the docks amongst the marsh grass shrimp Palaemonetes vulgaris. Palaemon floridanus has only been documented as
far north as Fort Pierce, Florida, prior to this record and is unlikely to survive the winter.
Marine Biological Laboratory Floating Dock (MBL), Woods Hole, Massachusetts | August 8, 2013, 2:30 PM
The Marine Biological Laboratory floating dock is located on the southwestern shore of Eel Pond, an embayment in Woods Hole,
Massachusetts. During the RAS, a total of 70 species were identified at MBL, including 51 native species, three cryptogenic species,
15 introduced species, and one summer transient species. The fouling community was dominated by solitary and colonial tunicates
as well as the blue mussels M. edulis/trossulus and branching bryozoans.
Massachusetts Maritime Academy Floating Dock (MMA), Buzzards Bay, Massachusetts | August 8, 2013, 10:15 AM
The Massachusetts Maritime Academy is located at the southern end of the Cape Cod Canal. The MMA features one permanent floating
dock of approximately 70 meters. Several large vessels are docked nearby, including the 160-meter-long USTS Kennedy. The floating
docks are located on the Buzzards Bay end of the Cape Cod Canal. During the RAS, a total of 90 species were identified at MMA, includ-
ing 61 native species, nine cryptogenic species, 19 introduced species, and one summer transient species. There was a patchy base of
the blue mussels M. edulis/trossulus and colonial and solitary tunicates. The hydroid E. larynx was very common at this site.
Pope’s Island Marina (POP), New Bedford, Massachusetts | August 9, 2013, 8:15 AM
Pope’s Island Marina is a public boat facility with 198 boat slips located on the south side of Pope’s Island in the upper region of New
Bedford Harbor. The marina was opened in 1993 with assistance from the Massachusetts Department of Conservation and Recreation
and is currently maintained and operated by the New Bedford Harbor Development Commission. During the RAS, a total of 88 species
were identified at POP, including 59 native species, seven cryptogenic species, 20 introduced species, and two summer transient
species. The fouling community was dominated by the tube-building worm Hydroides dianthus and the encrusting bryozoan
Schizoporella variabilis. On the last set of slips at POP, a grouping of the striped barnacle Amphibalanus amphitrite was found.
Amphibalanus amphitriteis an introduced barnacle normally found much farther south and will likely not survive the winter.
F.L. Tripp & Sons Marina (TRM), Westport, Massachusetts | August 9, 2013, 10:30 AM
Tripp’s Marina is located on the Westport River between Rhode Island Sound and Buzzards Bay and features 178 slips. The floating
pontoons at TRM are made of polystyrene foam. During the RAS, a total of 90 species were identified at TRM, including 62 native
species, eight cryptogenic species, 18 introduced species, and two summer transient species. The fouling community was dominated
by introduced colonial tunicates, such as the sheath tunicate Botrylloides violaceus, the star tunicate Botryllus schlosseri, and the
pancake batter tunicate Didemnum vexillum.
Close-up photograph
of the sheath tunicate
Botrylloides violaceus.
The lighter orange cir-
cles are the individual
animals living within a
shared tunic. The darker
orange circles are
developing embryos,
nearly the same size
as their parents.
12
13
Port Edgewood Marina (PEW), Cranston, Rhode Island | August 7, 2013, 1:45 PM
Port Edgewood Marina is located in the northern region of Providence Harbor adjacent to a large industrial park. The marina includes
three large, wooden floats and 160 seasonal and transient boat slips. This site had the lowest average salinity of any site sampled
during the 2013 RAS. During the RAS, a total of 50 species were identified at PEW, including 41 native species, four cryptogenic species,
four introduced species, and one summer transient species. Port Edgewood Marina had a heavy freshwater influence and hosted a
significantly different community of organisms, as indicated by the presence of the American oyster Crassostrea virginica. This site had
the highest percentage of native species (82%). The fouling community was dominated by the orange striped anemone Diadumene
lineata and the ivory barnacle Amphibalanus eburneus. Two individuals of a rare green color morph of the ghost anemone D. leucolena
were found at PEW.
Allen Harbor Marina (ALM), North Kingstown, Rhode Island | August 7, 2013, 10:45 AM
Allen Harbor Marina is located on the western side of Narragansett Bay on the corner of the former Quonset-Davisville Naval Base. The
marina features a wooden dock system with 80 moorings and 66 slips and is surrounded by heavily industrialized lands. Allen Harbor
features polystyrene foam pontoons. During the RAS, a total of 64 species were identified at ALM, including 40 native species, nine
cryptogenic species, 14 introduced species, and one summer transient species. The fouling community was dominated by solitary and
colonial tunicates. Sponges, specifically of the genera Halichondria and Haliclona, were relatively common at ALM.
Fort Adams State Park Floating Dock (FAD), Newport, Rhode Island | August 9, 2013, 10:30 AM
Fort Adams State Park is located at the mouth of Newport Harbor facing the East Passage of Narragansett Bay. Construction of the fort
began in 1824 and was completed 30 years later. The fort was deeded to the state of Rhode Island in 1966 and has since become a
major public access point into Narragansett Bay and a recreational attraction. The 105-acre park includes a small, semi-enclosed marina
at the southern-most point. During the RAS, a total of 89 species were found at FAD, including 60 native species, five cryptogenic
species, 23 introduced species, and one summer transient species. The base of the fouling community was composed of the blue
mussels M. edulis/trossulus, commonly with branching bryozoans Bugula spp. and the sea lettuce Ulva lactuca. The bryozoan
Tricellaria inopinata was collected at FAD; this is the southernmost record for this species and the first for Rhode Island.
Point Judith Marina (PTJ), Wakefield, Rhode Island | August 7, 2013, 8:30 AM
The Point Judith Marina is located on the southern end of Narragansett Bay within the sheltered enclosure of Point Judith Pond.
The marina features a large system of floating docks and wooden pilings. During the RAS, a total of 94 species were identified at PTJ,
including 66 native species, seven cryptogenic species, and 21 introduced species; PTJ had the most documented species during
the 2013 RAS. The fouling community was dominated by solitary and colonial tunicates and the bryozoans Bugula spp.,
with patchy clumps of the blue mussels M. edulis/trossulus. The pink-spotted anemone Aiptasiogeton eruptaurantia was found at
PTJ; this species is normally found much farther to the south, but has established a population over the last several years
(Hobbs, personal observation).
The hydromedusan
jellyfish Clytia
hemisphaerica
was only found at
the University of
New Hampshire
Coastal Marine
Laboratory floating
docks where it was
highly abundant.
The field team for the 2013 RAS included marine scientists with varying specialties.
Participant Name Specialty/Role Affiliation
Yuangyu Cao Algal taxonomy University of New Hampshire
James T. Carlton General taxonomy Williams College-Mystic Seaport
Zara Currimjee Lab and field assistance Williams College-Mystic Seaport
Kevin Cute Logistics Rhode Island Coastal Resource Management Council
Jennifer Dijkstra Ascidian taxonomy University of New Hampshire
Sara K. Edquist Worm taxonomy University of New Hampshire
Megan Flenniken Lab and field assistance Stony Brook University
Adriaan Gittenberger General taxonomy, photography GiMaRIS (Netherlands)
Seth Goodnight Algal taxonomy University of New Hampshire
Sara P. Grady General taxonomy, dockmaster Massachusetts Bays Program/
North and South Rivers Watershed Association
Lindsay A. Green Algal taxonomy University of New Hampshire
Larry G. Harris General taxonomy University of New Hampshire
Leslie H. Harris Worm taxonomy Natural History Museum of Los Angeles
Niels-Viggo Hobbs Peracarid taxonomy University of Rhode Island
Gretchen Lambert Ascidian taxonomy, photography University of Washington
Antonio C. Marques Medusazoan taxonomy Universidade de São Paulo (Brazil)
Arthur C. Mathieson Algal taxonomy University of New Hampshire
Megan I. McCuller Lab and field assistance University of New Hampshire
Kristin Osborne Lab and field assistance University of Massachusetts, Boston
Adrienne L. Pappal Co-Organizer and logistics Massachusetts Office of Coastal Zone Management
Judith A. Pederson General taxonomy Massachusetts Institute of Technology Sea Grant
Macarena Ros Caprellid taxonomy Universidad de Sevilla (Spain)
Jan P. Smith Co-Organizer and logistics Massachusetts Office of Coastal Zone Management
Lauren M. Stefaniak Ascidian taxonomy University of Connecticut
Alexandra Stevens Lab and field assistance Williams College-Mystic Seaport
Christopher D. Wells Anthozoan taxonomy and logistics University of New Hampshire/Massachusetts
Office of Coastal Zone Management
Appendix II: Rapid Assessment Survey Participants
An exceptional color morph of the
star tunicate Botryllus schlosseri
was found at Massachusetts
Maritime Academy.
14
15
Appendix III: Hydrographic Data
Surface-water temperatures showed a north-south pattern, with coldest temperatures in Maine (18.2 ± 3.6 °C) and New
Hampshire (17.1 ± 0.8 °C) and highest temperatures in southern Massachusetts (i.e., south of Cape Cod, 23.2 ± 1.2 °C).
Bottom-water temperature followed a similar pattern with coldest temperatures in Maine (16.9 ± 2.7 °C) and New Hampshire
(16.5 ± 1.2 °C) and warmest temperatures in southern Massachusetts (22.8 ± 0.9 °C). Surface and bottom-water salinities
were similar across states with no inherent pattern (27.8-30.3 ppt and 28.9-30.3 ppt respectively). Surface-water dissolved
oxygen was highest in Maine (7.71 ± 0.67 mg/L) and decreased southerly, with dissolved oxygen lowest in southern
Massachusetts (5.84 ± 1.14 mg/L) and Rhode Island (6.09 ± 0.72 mg/L). Bottom-water dissolved oxygen followed a
similar pattern to surface-water dissolved oxygen, although it was much more variable. Bottom-water dissolved oxygen
was highest in Maine (7.22 ± 0.93 mg/L) and lowest in Rhode Island (5.30 ± 2.08 mg/L). For depth, temperature, salinity,
and oxygen concentration for each site, see table below.
S. Freeport, ME (BFM) 5.3 21.1 18.6 28.8 28.9 7.56 7.26
Portland, ME (PHM) 4.5 19.5 16.7 24.5 28.1 7.13 7.16
Wells, ME (WHM) 5 14.2 14 30.2 30.1 8.81 8.29
New Castle, NH (UNH) 5.5 17.3 17.2 27.9 27.9 7.25 7.1
Hampton, NH (HSP) 3.8 16.8 16.8 29.3 29.8 6.57 7.87
Salem, MA (HCM) 4.5 15.8 14 30.1 30.3 7.1 6.89
Boston, MA (ROW) 6 19.6 17.9 28.5 29.5 5.81 6.43
Marshfield, MA (GHM) 4.5 21.3 20.2 29.1 29.5 7.02 6.32
Sandwich, MA (SWM) 3.5 19.4 18.6 29 30 6.06 5.97
Woods Hole, MA (WHC) 3.8 24.4 23.2 30.4 30.6 5.56 5.11
Woods Hole, MA (MBL) 2.5 24.8 23.8 30.4 30.4 6.46 6.68
Buzzards Bay, MA 5.3 22.1 22 29.9 29.8 6.34 6.22
New Bedford, MA (POP) 4 23.9 23.7 30.1 30.2 4.4 4.96
Westport, MA (TRM) 2.5 22.2 22.1 30.8 30.8 6.6 6.45
Cranston, RI (PEW) 2 23.7 22.7 25.2 26 nd nd
N. Kingstown, RI (ALM) 4.5 23.5 21.9 28.6 29.3 5.43 3.64
Newport, RI (FAD) 3 20 19.2 30.3 30.6 5.98 5.75
Wakefield, RI (PTJ) 4.3 20.6 20.5 30.4 30.5 6.86 7.2
LOCATION MAXIMUM SURFACE BOTTOM SURFACE BOTTOM SURFACE BOTTOM
DEPTH (m) TEMP (°C) TEMP (°C) SALINITY SALINITY OXYGEN OXYGEN
(ppt) (ppt) (mg/L) (mg/L)
The cryptogenic sea vase tunicate Ciona intestinalis formed dense aggregations at Hampton River Marina.
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19 19
Hemigrapsus sanguineus
(Asian shore crab)
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Lists of Introduced, Cryptogenic, and Native Species
The following tables contain the species found during the 2013 Rapid Assessment Survey, excluding species that could not be identified to genus or lower. Within the tables, the sampling sites have been abbreviated as
follows: Brewer South Freeport Marine (BFM), Sprin
g
Point Marina (PHM), Wells Harbor Marina (WHM), UNH Coastal Marine Laboratory Floatin
g
Dock (UNH), Hampton River Marina (HSP), Brewer Hawthorne Cove
Marina (HCM), Rowes Wharf Marina (ROW), Green Harbor Marina (GHM), Sandwich Marina (SWM), Woods Hole US Coast Guard Floating Dock (WHC), Marine Biological Laboratory Floating Dock (MBL),
Massachusetts Maritime Academy Floating Dock (MMA), Pope's Island Marina (POP), F.L. Tripp & Sons Marina (TRM), Port Edgewood Marina (PEW), Allen Harbor Marina (ALM), Fort Adams State Park Floating
Dock (FAD), and Point Judith Marina (PTJ). Sites are listed from northeast to southwest following the coast.
Introduced Species Identified During the 2013 Rapid Assessment Survey by Sampling Site
Species marked with an asterisk (*) are likely summer transient species or their establishment status is unknown.
SPECIES (COMMON NAME) BFM PHM WHM UNH HSP HCM ROW GHM SWM WHC MBL MMA POP TRM PEW ALM FAD PTJ
Chlorophyta (green algae)
Codium fragile subsp. fragile (dead man's fingers) x x xxx xxx
Heterokontophyta (brown algae)
Colpomenia peregrina (sea potato) x
Melanosiphon intestinali
s
(dark sea tubes) x
Rhodophyta (red algae)
Bonnemaisonia hamifera (hooked red weed) x x
Gracilaria vermiculophylla (wormleaf) xxx
Grateloupia turuturu (Devil's tongue weed) x xx xxx xxx
Heterosiphonia japonica (siphoned feather weed) xxx x
Lomentaria clavellosa (club bead-weed) xx x x x
Lomentaria orcadensis (Orkney weed) x
Neosiphonia harveyi (doughball weed) xxxxxxxxxxxxxxxxxx
Pyropia yezoensi
s
(open sea nori) x x
Annelida (ringed worms)
Neodexiospira brasiliensi
s
(Brazilian spiral tubeworm) xxx
Arthropoda (crabs, shrimp, and relatives)
*Amphibalanus amphitrite (striped barnacle) x
Caprella mutica (Japanese skeleton shrimp) xxxxxxxxxxxxxx xxx
Carcinus maenas (European green crab) xxxxxxxxxx xxx xxx
Hemigrapsus sanguineus
(Asian shore crab)
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Ianiropsis serricaudis (isopod) xxxxxxxxxx xx
*Melita palmata (palmed gammarid) x
Microdeutopus gryllotalpa (tube builder) xx xx
Palaemon elegans (European rock pool shrimp) xx xx xx
*Palaemon floridanus (Florida grass shrimp) x
Palaemon macrodactylus (oriental grass prawn) x
Praunus flexuosus (bent opossum shrimp) xxx xx xx
*Stenothoe valida (gammarid amphipod) xxx x
Bryozoa (bryozoans)
Bugula neritina (purple bugula) xx xx x xx
Bugula simplex (fan bugula) x xxxxxxxxxx xxx
Bugula stolonifera (stoloned bugula) x x
Conopeum seurati (encrusting bryozoan) x
Membranipora membranacea (lacy crust bryozoan) xxxx x x x
Tricellaria inopinat
a
(unexpected bryozoan) xxxxxxxx x
Chordata (tunicates and vertebrates)
Ascidiella aspersa (European tunicate) xx xxxxxxxx xx
Botrylloides violaceus (sheath tunicate) xxxxxxxxxxxxxx xxx
Botryllus schlosseri (star tunicate) xxxxxxxxxxxxxx xxx
Botryllus sp. (colonial tunicate) xx xxxx x
Didemnum vexillum (pancake batter tunicate) x xx xxxxxxx x xx
Diplosoma listerianum (gelatinous tunicate) xxxxxxxxxxxxx xxx
Styela canopus (rough tunicate) xxx xx
Styela clava (club tunicate) x xxxxxxxxx xxx
Introduced Species continue
d
SPECIES (COMMON NAME) BFM PHM WHM UNH HSP HCM ROW GHM SWM WHC MBL MMA POP TRM PEW ALM FAD PTJ
Cnidaria (anemones, hydroids, and relatives)
Aiptasiogeton eruptaurantia (pink-spotted sea anemone) x
*Clytia linearis (hydroid) x xxxxxxxxxx
Diadumene lineata (orange striped anemone) x xxxx xxxxxxx
Hydrodendron sp. (hydroid) x
Mollusca (snails and bivalves)
Littorina littorea (common periwinkle) xxxx
Ostrea edulis (European oyster) xxxxxxxx x x
20 20
21 21
Moll (snails bi es and relati
Cryptogenic Species Identified During the 2013 Rapid Assessment Survey by Sampling Site
SPECIES (COMMON NAME) BFM PHM WHM UNH HSP HCM ROW GHM SWM WHC MBL MMA POP TRM PEW ALM FAD PTJ
Annelida (ringed worms)
Capitella teleta (telete worm) x
Lepidonotus sp. (scale worm) xx x
Perkinsiana s
p
1.
(
sabellid worm
)
x
Polycirrus sp. (terebellid worm) x
Arthropoda (shrimp, crabs, and relatives)
Caprella penantis (caprellid amphipod) x
Microdeutopus anomalus (amphipod) x
Tanais dulongii (tanaid) xx
Bryozoa (bryozoans)
Bowerbankia imbricata (tiled moss hydroid) xxxx
Cryptosula pallasiana (orange crust bryozoan) x x x x x x x x x x x x
Cercozoa (some amoebas and relatives)
Gromia oviformis (rhizopod) x
Chordata (tunicates and vertebrates)
Ciona intestinalis (sea vase tunicate) x x xxxx xxxx xx
Cnidaria (anemones, hydroids, and relatives)
Bougainvillia muscus (sticky moss hydroid) x x x x x
Clava multicornis (club hydroid) x x x
Clytia gracilis (hydroid) xxx xx x x
Clytia hemisphaerica (hydroid) x
Dynamena pumila (sea oak hydroid) x x xx
Ectopleura crocea (pink-hearted hydroid) x x x
Ectopleura larynx (ringed tubularian) xxxxxxxxx x x x
Gonothyraea loveni (Hydroid) xx x
Hydractinia sp. (snail fur hydroid) x x x x
Laomedea calceolifera (hydroid) xx x
Obelia dichotoma (sea thread hydroid) x x xxxxxx xxxxxx
Obelia longissima (bushy wine-glass hydroid) x x x xxx
Pennaria disticha (Christmas tree hydroid) x
Mollusca (snails bivalves and relatives)
u
sca
, v
al
v , v
es)
Tenellia adspersa (miniature aeolis) x
Porifera (sponges)
Halichondria sp. (sponge) xxxxxxxxxx xxxxxxx
Haliclona sp. (sponge) xxxx xxx x x x
1This species has been tentatively identified as Perkinsiana sp., but the specimens collected may be an undescribed (i.e., new) species.
22 22
Ascophyllum nodosum
x
x
x
x
x
x
x
x
x
Native Species Identified During the 2013 Rapid Assessment Survey by Sampling Site
SPECIES BFM PHM WHM UNH HSP HCM ROW GHM SWM WHC MBL MMA POP TRM PEW ALM FAD PTJ
Chlorophyta (green algae)
Acrosiphonia arcta x
Acrosiphonia spinescens xx x
Blidingia minima x xx xxxx x
Bryopsis hypnoides xx xx
Bryopsis plumosa x xxxx x x x
Capsosiphon fulvescens xx x
Chaetomorpha ligustica xxxxxx x
SPECIES BFM PHM WHM UNH HSP HCM ROW GHM SWM WHC MBL MMA POP TRM PEW ALM FAD PTJ
Chlorophyta (green algae)
Chaetomorpha linum xx
Chaetomorpha picquotiana xxxx
Cladophora albida x xxx
Cladophora sericea x xxxxxx x x
Percursaria percursa xx
Pseudendoclonium submarinum xx
Rhizoclonium riparium x x
Ulva compressa xx x x x
Ulva flexuosa subsp. flexuosa xxx x
Ulva flexuosa subsp. paradoxa x
Ulva intestinalis xx x x x xx x
Ulva lactuca xxxxxxx xxx xx xx
Ulva linza x xxxx x
Ulva prolifera xx x x x
Ulvaria obscura xx x
Urospora penicilliformis xx
Cyanobacteria (blue-green algae)
Lyngbya confervoides xx x xxxx x
Spirulina major x
Heterokontophyta (brown algae)
Ascophyllum nodosum
x
x
x
x
x
x
x
x
x
Berkeleya rutilans xxxx
Chordaria flagelliformis x
Desmarestia aculeata x
Ectocarpus fasciculatus xx
Ectocarpus siliculosus xx xx
Elachista fucicola x
Fragilaria sp. xx x
Fucus spiralis x xxx x
Fucus vesiculosus xxxxxxxxx x x x
Fucus vesiculosus ecad volubilis x
Hincksia granulosa x x
Isthmia nervosa xxxx
Melosira nummuloides xx
Microspongium gelatinosum
1
x
Petalonia fascia x x
Protectocarpus speciosus x
Ralfsia verrucosa x
Saccharina latissima xx x x x
Saccharina longicruris x
Sargassum filipendula xxxx x
Sphacelaria cirrosa x
Rhodophyta (red algae)
Acrochaetium secundatum xx
Agardhiella subulata xxxxxxxxxx
Aglaothamnion halliae
1
xx
Antithamnion hubbsii x x
Antithamnion plumula x
23 23
Annelida (ringed worms)
Native Species continue
d
SPECIES BFM PHM WHM UNH HSP HCM ROW GHM SWM WHC MBL MMA POP TRM PEW ALM FAD PTJ
Rhodophyta (red algae)
Antithamnionella floccosa xx
Callithamnion corymbosum x
Ceramium deslongchampsii xx x xx x xxxx
Ceramium virgatum xxx xxx xx x x xx
Champia parvula xxx xx
Chondria capillaris x xxx
Chondrus crispus xxx xxx x x x x
Cystoclonium purpureum x
Dasya baillouviana xx x x
Erythrotrichia carnea xx xxx
Gracilaria tikvahiae xxx
Grinnellia americana xx x x
Hypnea musciformis x
Lomentaria divaricata x xxx xx
Mastocarpus stellatus xxx x
Palmaria palmata xx
Polysiphonia elongata xx
Polysiphonia flexicaulis xx
Polysiphonia fucoides xxxx
Polysiphonia nigra x
Polysiphonia schneideri x xxxxxx xxx
Polysiphonia stricta xxxxx x x
Porphyra purpurea x xxxx x x
Porphyra umbilicalis xx
Spermothamnion repens xxxxxx
Spyridia filamentosa xxxx x
Stylonema alsidii x x
Titanoderma pustulatum x x xxxx x
Vertebrata lanosa xx
Annelida (ringed worms)
Alitta virens xxxxxxxxx
Amphitrite ornata xxxxx x
Cirriformia grandis xx
Dodecaceria concharum x
Eulalia viridis xx
Eumida sanguinea x
Harmothoe extenuata x x xxxx x xx
Harmothoe imbricata xxxxx xxx x xx
Harmothoe spinulosa x
Hydroides dianthus xxxxxxxxxx
Lepidonotus squamatus xxxxx xxxxxx xx
Neanthes succinea xxx xxxxxxx
Neoamphitrite figulus xx x xx x
Nereis pelagica x xxx x x x
Oxydromus obscurus x xxxx xxxx
Parasabella microphthalma xx xx xx
Pista palmata xx
Platynereis dumerilii xx
Polydora cornuta xx x x xxx
Polydora websteri x
Syllis gracilis xx
Terebella verrilli xx x
Arthropoda (shrimp, crabs, and relatives)
Amphibalanus eburneus xx xx x xx
Amphibalanus improvisus xx x xxxx
Ampithoe longimana xx xx
Ampithoe rubricata xx
Ampithoe valida x x xx xxx
24 24
Stenothoe minuta
x
Native Species continue
d
SPECIES BFM PHM WHM UNH HSP HCM ROW GHM SWM WHC MBL MMA POP TRM PEW ALM FAD PTJ
Arthropoda (shrimp, crabs, and relatives)
Anurida maritima xxx
Apocorophium acutum xx x
Balanus crenatus xxxxxxxxx x x
Callinectes sapidus x xxx
Cancer borealis x x
Cymadusa compta xx
Dyspanopeus sayi x xxx x
Elasmopus levis xx xx xxx
Eurypanopeus depressus xx
Gammarus mucronatus xxx x xxx
Globosolembos smithi xxxx
Idotea balthica xxxxxxx
Idotea phosphorea xxxx x
Jaera marina x
Jassa marmorata xxxxxxxxxx xxx xx
Libinia dubia xxx
Libinia emarginata xxxx
Limnoria lignorum x x
Melita nitida xx
Monocorophium acherusicum xx x
Monocorophium insidiosum x xxxxxxxx xx x
Palaemonetes pugio xxx
Palaemonetes vulgaris xxx xxxxxx
Panopeus herbstii xxxxxxx xx
Paracaprella tenuis x x
Phoxichilidium femoratum x
Ptilohyale plumulosus x xxx
Semibalanus balanoides xxxxxxxxxxxxxxxxxx
Stenothoe minuta
x
Bryozoa (bryozoans)
Amathia dichotoma x
Bowerbankia gracilis x xxxxxxxxx
Bowerbankia tertia xx
Bugula turrita xx xxx xxxxx x
Celleporella hyalina xxxxx
Electra pilosa xxxx xx x
Microporella sp. xx
Schizoporella variabilis xx xxxxxx
Scrupocellaria scabra xxxxxx
Chordata (tunicates and vertebrates)
Aplidium constellatum xx x
Aplidium glabrum xxxx x
Aplidium pallidum x
Fundulus heteroclitus xx
Molgula citrina xxxxx x
Molgula manhattensis xx x xxxxxx
Molgula provisionalis xx
Myoxocephalus sp. x
Osmerus mordax x
Perophora viridis x
Pholis gunnellus xx
Pungitius pungitius x x
Tautoga onitis xxx
Tautogolabrus adspersus xxxxxxx xx xxx
25 25
Crepidula fornicata
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Native Species continue
d
SPECIES BFM PHM WHM UNH HSP HCM ROW GHM SWM WHC MBL MMA POP TRM PEW ALM FAD PTJ
Cnidaria (anemones, hydroids, and relatives)
Astrangia poculata xxxxx xx
Aurelia sp. xxxx
Cyanea capillata x
Diadumene leucolena x xx xxxx xx
Edwardsiella lineata x xx xx xxxxx
Eudendrium album x
Eudendrium glomeratum x
Halopteris tenella xx
Metridium senile xxxxxxxxxx xxx xxx
Obelia geniculata xx
Tubularia indivisa xx
Ctenophora (comb jellies)
Mnemiopsis leidyi x x xx xxx x
Echinodermata (sea stars, urchins, and relatives)
Arbacia punctulata xx
Asterias forbesi xxx
Asterias rubens xxx xx
Strongylocentrotus droebachiensis x
Entoprocta (nodding heads)
Barentsia laxa x
Mollusca (snails, bivalves, and relatives)
Aeolidia papillosa xx
Anadara transversa x
Anomia simplex xxxxx xxxx x xx
Astyris lunata xxxx
Corambe obscura x x
Costoanachis sp. x
Crassostrea virginica x
Cratena pilata x
Crepidula fornicata
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Crepidula plana xxx xx x xxx xxx
Cuthona gymnota xx
Dendronotus frondosus x
Eubranchus exiguus xx
Facelina bostoniensis xxx
Geukensia demissa xx
Heteranomia squamula xxx xxx x
Hiatella arctica xxxx x
Lacuna vincta xx
Lunarca ovalis xx
Modiolus modiolus xx
Mya arenaria xxx
Mytilus edulis /trossulus
2
xxxxxxxxxxxxxx xxx
Onchidoris bilamellata x
Onchidoris muricata x
Placida dendritica x
Polycerella emertoni xx
Urosalpinx cinerea xx x
Nemertea (ribbon worms)
Cyanophthalma cordiceps x
Lineus arenicola x
Lineus ruber xxx
Ramphogordius sanguineus x
Tenuilineus bicolor x
Zygonemertes virescens x
Native Species continue
d
SPECIES BFM PHM WHM UNH HSP HCM ROW GHM SWM WHC MBL MMA POP TRM PEW ALM FAD PTJ
Platyhelminthes (flatworms)
Euplana gracilis xx
Notoplana atomata xx
Pleioplana atomata xx
Stylochus ellipticus xxx
Porifera (sponges)
Clathria prolifera xxxx
Cliona sp. x xxx
Leucosolenia sp. xx xx xxxx x xx
Sycon ciliatum xx xx x
1The identifications of Microspongium gelatinosum and Aglaothamnion halliae are tentative.
2It is unclear at this time which species was found during the Rapid Assessment Survey as they are morphologically similar and may require genetic analysis for identification.
26 26
... It was particularly abundant in 2000 and, therefore, was suspected to have been introduced some years prior. Subsequent rapid assessment surveys in 2003,2007, and 2010 also found S. elegans at HCM (Pederson et al., 2005;Wells et al., 2014). The introduced population of S. elegans in Salem Harbor was seasonally abundant; it appeared as early as June but died back completely by January. ...
... Depth at HCM ranges from 1 to 4 m. HCM is known for its exceptionally high load of introduced species compared to other sites in the Gulf of Maine (Pederson et al., 2005;Wells et al., 2014). ...
... comm.). Additional surveys in 2013 and 2018 did not locate any S. elegans individuals at HCM (Wells et al., 2014;A. Pappal, Massachusetts Coastal Zone Management, pers. ...
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Failed invasions can be a key component for understanding and controlling introduced populations because understanding mechanisms behind failures can improve effective controls. In 2000, the non-native sea anemone Sagartia elegans was first found in Salem, Massachusetts, and it recol-onized each summer. No individuals of S. elegans have been found after 2010, despite intensive search efforts. A mismatch between the species' thermal tolerance and winter water temperature is the most likely mechanism for this failed invasion. In both laboratory-and field-based temperature growth studies , S. elegans began regressing at 11 7C, stopped asexually reproducing at 9 7C, and died by 4 7C. These temperatures are above the average winter sea surface temperature in the Gulf of Maine, therefore suggesting that S. elegans requires a warm-water refuge. Another potential contributor to the disappearance of S. elegans is low genetic diversity as a result of establishment of only females (likely clones) and no males.
... In the New England-New York region, these surveys began in 2000 focusing on sampling biofouling communities on floating docks in marinas (Pederson et al., 2021). Seven surveys have been held, generally every two to three years, between 2000 and 2019, largely including sites from Maine to Connecticut (with New York being sampled as well in 2003 and 2019) (Pederson et al., 2005(Pederson et al., , 2021Mathieson et al., 2008;McIntyre et al., 2013;Wells et al., 2014;Kennedy et al., 2020). The success of these surveys has resulted in similar surveys being deployed in European waters (Arenas et al., 2006;Ashton et al., 2006;Nall et al., 2015;O'Shaughnessy et al., 2023) and elsewhere. ...
... We first reviewed the original species inventory spreadsheets that were compiled for each survey since 2000, in addition to reviewing summaries of surveys that have been published in the grey literature by the Massachusetts Office of Coastal Zone Management (McIntyre et al., 2013;Wells et al., 2014;Kennedy et al., 2020). Historical records and species statuses were updated based on recent taxonomic revisions, DNA barcoding studies, updated biogeographical analyses and name/rank changes obtained from the World Register of Marine Species (WoRMS Editorial Board, 2024) database. ...
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This study presents the first comprehensive annotated checklist of polychaetes collected from floating dock communities across New England and adjacent New York areas, emphasizing the significance of rapid biodiversity assessment surveys in understanding marine biodiversity. With 61% of the identified species classified as cryptogenic, the research highlights the complexity of marine biogeography and the challenges of distinguishing between native and non-native species. The incorporation of DNA barcoding has significantly improved species identification and revealed the presence of cryptic species complexes. The study's findings illustrate the prevalence of tubicolous families (Spionidae, Serpulidae, Sabellidae, and Terebelllidae) that are susceptible to anthropogenic dispersal mechanisms. It also addresses the challenge posed by the high proportion of cryptogenic species, calling for enhanced taxonomic and genetic analyses to resolve their origins and ecological roles. Despite the temporal variation in polychaete composition across years, the absence of distinct community assemblages suggests a level of resilience within floating dock communities. Our study advocates for the continuation and expansion of rapid assessment surveys, coupled with the integration of genetic methodologies, to provide a clearer picture of marine biodiversity.
... We then examined the effect of estimated climate-induced shifts in reproduction on early succession in communities that span a temperature gradient. In the GOM, the biological resistance of communities to invasions will likely depend upon latitude as the assemblages of species and the amount of free space, strong determinants for the successful establishment of invasions, that make up fouling communities are not equal across latitudes (Pederson et al., 2005;Wells et al., 2014). Northern GOM benthic communities have lower numbers of introduced species compared to more southern GOM communities (Dijkstra, Sherman, & Harris, 2007b;Pederson et al., 2005;Wells et al., 2014). ...
... In the GOM, the biological resistance of communities to invasions will likely depend upon latitude as the assemblages of species and the amount of free space, strong determinants for the successful establishment of invasions, that make up fouling communities are not equal across latitudes (Pederson et al., 2005;Wells et al., 2014). Northern GOM benthic communities have lower numbers of introduced species compared to more southern GOM communities (Dijkstra, Sherman, & Harris, 2007b;Pederson et al., 2005;Wells et al., 2014). This may stem from environmental resistance to invasion due to cold-water seasonal temperatures or the ability of the communities to resist invasion. ...
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Full-text available
Aim Couple field and modelling studies to evaluate the effects of rising water temperature on reproduction of a dominant marine species and how temperature induced changes in a species' phenology may affect fouling communities that occur along a temperature gradient. Location North western Atlantic. Methods We examined the interaction between maximum and seasonal elevations in seawater temperature and reproductive phenology of a global invader to forecast the effect of climate warming on traits associated with the spread and establishment (i.e., growth and reproduction) of an invasive species. We then coupled these data with a study of early succession along a temperature gradient to forecast invasion success in species assemblages across a range of latitudes. Results Elevated seasonal temperature will lead to greater asexual reproduction and to multi‐annual sexual reproduction in areas that currently have bi‐annual or annual reproduction. The output from our model combined with our field studies of early succession in communities suggest that in colder environments, communities may be more affected by climate‐induced shifts in reproductive phenology as they have more free space at the beginning of the growing season and fewer competitively superior species. Main conclusions Predicted elevated water temperature will eliminate the thermal barriers that limit reproductive success of a cosmopolitan invasive species. Increased reproduction combined with limited biological resistance in regions with cooler water temperatures may lead to a community state change.
... Cordell et al. (2013) collected janirids while surveying the fouling communities of a number of sites in Puget Sound in Washington State. A team of researchers taking part in a series of Rapid Assessment Surveys of non-indigenous species of New England in 2000and 2013(Pederson et al. 2005McIntyre et al. 2013;Wells et al. 2014) collected samples at numerous coastal sites in New England and New York, focused mostly on floating docks, but including some surveys of rocky intertidal sites during 2010 (McIntyre et al. 2013). A similar rapid assessment survey of dock fouling communities conducted in England in 2004 (Arenas et al. 2006) produced janirid isopods of interest. ...
... Ianiropsis serricaudis now ranges widely across New England, with records in hand from Maine to Connecticut (Pederson et al. 2005;McIntyre et al. 2013;Wells et al. 2014). The first indication of the fuller range of this species came from a Rapid Assessment Survey (RAS) for nonindigenous species on floating dock sites in southern New England in the summer of 2000, when it was found in dock fouling at a few sites in estuarine Rhode Island waters. ...
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The Asian isopod Ianiropsis serricaudis is now well established in fouling communities, often associated with introduced ascidians, throughout the Northern Hemisphere but has gone largely unnoticed because of its diminutive size (typically less than 3 mm in length) and the difficulties of identifying small peracarid crustaceans. Known locations include the northeastern Pacific (Puget Sound, San Francisco Bay, and Monterey Bay), the northwestern Atlantic (from the Gulf of Maine to Barnegat Bay, NJ), and the northeastern Atlantic (England and the Netherlands). We predict that this species is widespread along North America and European coasts, and may already be introduced to cold temperate waters of the Southern Hemisphere as well.
... Fouling communities are also populated with many introduced species (Cohen and Carlton, 1996;Ruiz et al., 2000), which has led to a homogenization of communities (Simberloff, 2013). Specifically, many non-native species have flourished in the Gulf of Maine in the last fifty years, particularly within fouling communities (Berman et al., 1992;Kennedy et al., 2020;Wells et al., 2013). Additionally, climate change has had a significant impact on communities in the Gulf of Maine over the last thirty years. ...
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The Boston Harbor Islands is the only coastal drumlin archipelago in the USA, featuring a distinctive and uncommon geological intertidal habitat known as mixed coarse substrate, which supports a range of coastal species and ecological processes. Recently designated as one of America’s 11 most endangered historic places due to climate change impacts, coastal adaptation and restoration efforts are crucial to their preservation. Such efforts can benefit from historic and current knowledge of endemic and emergent biodiversity. To investigate broad trends in coastal biodiversity, we compiled an inventory of marine coastal macroalgae, macroinvertebrates, fish, mammals, and shorebirds observed in the harbor since 1861. Records span 159 years, consisting of 451 unique taxa from 19 phyla. Analysis of average taxonomic distinctness (AvTD) revealed increases in diversity towards the end of the twentieth and early twenty-first century, likely associated with improved water quality (dissolved oxygen; AvTD > 85, p = 0.01) due to harbor restoration in the 1980s. Macroinvertebrates comprised 50% of the records, making this the most diverse taxonomic group in the time series. A significant increase of non-indigenous species, primarily macroinvertebrates and macroalgae, was observed over the last 20 years near human infrastructure and across multiple islands, a consequence of global change and characteristic of most urban harbors. The mixed coarse intertidal habitat, which makes up > 70% of Boston Harbor’s inner islands and supports high macroinvertebrate and macroalgal diversity (47% of species records), is not routinely monitored; our findings serve as a foundational resource for climate adaptation projects and decision-making.
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Widespread non-native species tend to demonstrate an apparent lack of selectivity in habitat requirements, feeding regimes, and reproductive needs, while displaying a tendency to thrive in human-modified habitats. The high phenotypic plasticity typical of sessile, substrate-attached marine species may enhance their chances of survival and spread in a new region. Anthropogenic activities have changed marine habitats over a wide range of phenomena, including water temperature, community species composition, and the types of available substrates, creating new physical and biotic regimes that may contribute to the potential for successful species introduction. Here we examine ten species of sea anemones that have been introduced outside of their native range, and elucidate specific characteristics that are common among globally introduced sea anemones. Various life history strategies enable these species to survive and flourish through transport, introduction, establishment and spread, leading to the successful colonization of a new geographic area. Considering life history strategies and weighing of vector potential, we suggest conditions that facilitate introduction of these species, and identify species of sea anemones that may be introduced in the future in the face of changing climate and increased anthropogenic activities.
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Criteria for distinguishing introduced from endemic peracaridan crustaceans were used to deduce that a human-borne global invasion by the Oriental isopod Synidotea laevidorsalis (Miers, 1881) has occurred in the past 100 years. These criteria concern the ecological, evoluntionary, and geographical attributes of introduced species. The criteria were used first to hypothesize that Synidotea laticauda is an introduced species in the eastern Pacific that arrived on the hulls of nineteenth-century sailing ships. This hypothesis was tested by searching for previously described conspecifics throughout the world. The search culminated in discoveries that Synidotea laticauda Benedict, 1897 of the eastern Pacific and Synidotea marplatensis (Giambiagi, 1922) of the Atlantic coast of South America are misidentified populations and thus synonymies of S. laevidorsalis. Synidotea brunnea Pires and Moreira, 1975, of central Brazil is also a probable junior synonym of S. laevidorsalis. The discovery of these synonymies was thus based upon predictive criteria rather than inductive classical, taxonomic revisions. These errors in species identifications indicate that the prevalence of marine and estuarine introductions has been underestimated and that the extent of many introductions remains poorly resolved.
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Most species of bryozoans have short-lived larvae with limited dispersal potential, yet many of these species possess global distributions. In this study, we report the first occurrence from the western Atlantic Ocean of the widely distributed arborescent bryozoan Tricellaria inopinata d'Hondt and Occhipinti-Ambrogi, 1985. This species was collected in Eel Pond, Woods Hole, Massachusetts, in September 2010. At that time, T. inopinata colonies had already formed dense conspecific aggregations at some collection sites, despite the presence of several other arborescent bryozoans. Sites were monitored throughout 2011 to track the success of this introduction, and to assess the reproductive timing of T. inopinata in Eel Pond. To determine the likelihood of T. inopinata persisting in Eel Pond and competing with previously established bryozoans, rates of metamorphic initiation, metamorphic completion, and overall offspring survivability were compared to one of the other dominant arborescent species. Finally, we provide taxonomic details to aid in identifying these animals, consider the potential mode of transport, and discuss the potential ecological implications resulting from this introduction.
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Colpomenia, a small genus with 11 species of globular to bullate form, occurs in temperate to tropical waters world-wide. Because morphology is highly diverse, the species-level taxonomy requires re-evaluation. We analyzed the mitochondrial cox3 gene from 50 samples of the genus. A new species, Colpomenia claytonii sp. nov., is described based on samples collected from Korea, Japan, Hong Kong, Australia, New Zealand, South Africa, and the USA and compared with similar congeners. Compared to others, the new species is larger, and has a more irregular thallus often with a deeply infolded surface. It is usually epilithic in tide pools and in the lower intertidal and subtidal zones. In all phylogenetic analyses of cox3 sequences, C. claytonii was consistently distinct from congeners. Colpomenia expansa is closely related to C. claytonii, and the clade containing these two species is closely related to C. peregrina. A total of 14 cox3 haplotypes was found in C. claytonii, indicating high haplotype diversity and a recent dispersal pattern. The pres-ent study shows that C. claytonii is a widely distributed spe-cies across the Pacific Ocean to South Africa; it was previously misidentified as a variant of C. peregrina.
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Rapid assessment surveys of fouling seaweed populations were conducted at 67 sites between Downeast Maine and Staten Island, New York during August of 2000, 2003, and 2005, plus July-August of 2007. A total of 126 taxa were recorded, including 29 Chlorophyceae, 31 Phaeophyceae, 55 Rhodophyceae, four Cyanophyceae, one Xanthophyceae, and six macroscopic colonial diatoms (Bacillariophyceae). Several species were fast-growing nuisance organisms (e.g., Ulva spp.), while seven were introduced taxa. Four introduced species originated from Asia either directly or secondarily (Codium fragile subsp. fragile, Grateloupia turuturu, Neosiphonia harveyi, and Bonnemaisonia hamifera), two were from Europe (Lomentaria clavellosa and L. orcadensis), and one from the North Pacific (Melanosiphon intestinalis). Neosiphonia harveyi was the most widely distributed introduced taxon, occurring at 48 sites (71.6%), while L. clavellosa and L. orcadensis were only found at 3 sites (4.5%). Repeated observations (i.e., 2000, 2003, and 2007) at three sites in Massachusetts documented a recent rapid expansion of the invasive red alga G. turuturu into the Gulf of Maine through the Cape Cod Canal. The numbers of taxa per site were highest between Maine and Massachusetts (28-42 taxa) and lowest (1-13 taxa) in southern New England and New York, presumably because of increased loading of various pollutants within Long Island Sound and near New York City. The highest mean (± SD) number of taxa per state was recorded in New Hampshire (28.8 ± 8.0) and the lowest in Rhode Island (7.1 ± 3.6). The green and red algae exhibited peak numbers in New Hampshire (i.e. 9.0 ± 3.1 and 11.6 ± 5.1), while brown algae were maximal in New Hampshire (8.2 ± 1.1) and Maine (7.8 ± 3.4) and much lower in Connecticut (1.0 ± 1), Rhode Island (1.1 ± 2.9), and New York (1.2 ± 1.0). Fifty-four species were limited to 1-3 sites (1.5-4.7%), while only 7 occurred at > 50% of the sites. The most ubiquitous seaweeds included three opportunistic green algae (U. intestinalis, U. lactuca, and Blidingia minima), two perennial browns (Fucus vesiculosus and Ascophyllum nodosum), and the introduced Asiatic red alga, N. harveyi. Thirty-six taxa were restricted to individual states (31.3%), with Massachusetts exhibiting the highest number of unique taxa (17 or 14.8%) and Connecticut and New York the lowest (one taxon each or 0.9%). Massachusetts also had the highest number of total and mean shared species (86 and 43.3 ± 10.7, respectively), while Connecticut had the opposite pattern (18 total and 15.0 ± 7.5 shared taxa). Cheney's floristic ratio indicated that seaweeds restricted to the north of the Cape Cod Canal formed a cold-water flora, while those limited to the south of the Canal were somewhat less boreal. Most taxa from each state were annuals and exhibited cosmopolitan local distributional patterns, occurring in both open coastal and estuarine sites.
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A theory that introduced species can be identified from their evolutionary, ecological, and geographical attributes (criteria for introduced species) is corroborated by the accuracy of its predictions. A human-borne, global invasion of the Oriental isopod Synidotea laevidorsalis (Miers, 1881) that began over 100 years ago was discovered in a test of criteria for introduced species (Chapman and Carlton, 1991). Two explicit predictions from that work were(1) ''Many new records of endemic and introduced Synidotea populations around the world remain to be discovered,'' and (2) ''Synidotea laevidorsalis is established, for example, in Brisbane, or Sydney, Australia.'' Both predictions were correct. It appears that many estuarine and nearshore marine introductions have occurred that have not been discovered.
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