2010 NORTHEASTERN NATURALIST 16(Monograph 5):1–48
Floristic Studies of Seaweeds from Cobscook Bay, Maine
Arthur C. Mathieson1,2,*, Clinton J. Dawes3, Edward J. Hehre2,
and Larry G. Harris1
Abstract - The benthic marine algal flora of Cobscook Bay, ME, which is located
near the mouth of the Bay of Fundy and is contiguous with Passamaquoddy Bay, NB,
Canada, consists of 148 taxa, including 38 green, 46 brown, and 64 red algae. The
seaweed flora of Cobscook Bay contains 47% of the total taxa (315) known from the
Bay of Fundy and 37% of the 403 taxa recorded from the northeast coast of North
America. Cobscook Bay’s floristic affinities range from 63.8% (Bay of Fundy) to
74.3% (Casco Bay), with Passamaquoddy Bay having a 68.8% affinity. Of the 37
sites in Cobscook Bay, Eastport Harbor (70 taxa) and Reversing Falls Park (51 taxa)
had the highest number of species, and the lowest was found at one Lubec site near
Lead Mine Road (only “brackish” water taxa found). The Bay’s overall (R + C)/ P
floristic ratio ([red + green]/ brown algae) is 2.2, which indicates a cold-temperate
flora. Six introduced seaweeds were found in the Bay. The expanded vertical distribution
of intertidal (and subtidal) taxa within Cobscook Bay may be associated with
extreme tides and a high incidence of summer fog. Kelps in some Cobscook Bay
areas were larger than open coastal plants, reflecting the effect of protected embayments,
strong currents, and high levels of nutrients, as demonstrated by Saccharina
longicruris at Wilbur Neck, with an average length of 4.6 m. Saccharina latissima,
growing in areas with strong tidal currents at Reversing Falls Park, had elongate and
narrow blades similar to the narrow-bladed S. latissima f. angustissima known only
from mid-coastal Maine. Dwarf limicolous fucoids (i.e., Fucus species “muscoideslike”)
were found in some sandy salt marsh habitats similar to those described at a
few scattered Canadian Maritime and Gulf of Maine sites.
Introduction
Cobscook Bay, ME is located near the mouth of the Bay of Fundy on
the eastern boundary between the United States and Canada (Fig. 1). It
is a hydrographically and geologically complex estuary with high levels
of biodiversity and productivity (Brooks 2004, Kelley and Kelley 2004,
Larsen 2004a, Larsen and Campbell 2004). Observations on the region’s
fauna began in the 1840s, with Mighels’ (1843), Stimpson’s (1851, 1853),
and Fuller’s (1862a, b) descriptions of several invertebrate taxa. In the early
1870s, government-funded research, coordinated by US Fish Commissioner
Baird, addressed the decline of fisheries (Larsen 2004b, Verrill 1871), firmly
established the region’s species richness, and reported that damming of rivers
and streams by the timber industry was the cause of the mid-19th century
fisheries collapse. Most subsequent research in Cobscook Bay focused on
1Department of Biological Sciences University of New Hampshire, Durham, NH
03824. 2Jackson Estuarine Laboratory, University of New Hampshire, Durham, NH
03824. 3Department of Biology, University of South Florida, Tampa, fl33620. *Corresponding
author - arthur.mathieson@unh.edu.
2 Northeastern Naturalist Vol. 16, Monograph No. 5
impact analyses on fisheries and invertebrates associated with proposed tidal
power, oil facilities, and aquaculture (Larsen and Webb 1997). The faunal
studies emphasized the Bay’s species richness (Holmes 1905, Richardson
1905, Webster and Benedict 1887), its extraordinary natural productivity,
habitat diversity, occurrence of many typical subtidal species within the intertidal
zone (Fuller 1862a, b; Stimpson 1851), and the presence of gigantism
in invertebrates (Larsen 2004b, c). Trott and Larsen (2003) noted that the
extreme tides, upwelling, high incidence of summer fog which shields the
intertidal from solar radiation, and highly varied habitats might explain the
nearly 800 species of macroinvertebrates in Cobscook Bay (Larsen 2004c,
Trott 2004a)
Some of the earliest collections of seaweeds from Cobscook Bay were
made in Eastport, ME by Eaton (1873), who worked with the US Fish Commission,
and Farlow, who collected both independently (Farlow 1881) and
with several colleagues (Farlow et al. 1877–1889). Several collections of
seaweeds were made in eastern Canadian sites near Cobscook Bay, including
the Bay of Fundy (Hay 1886), Passamaquoddy Bay (Klugh 1916), and
the New Brunswick/Canadian Maritime area (Fowler 1901, 1902; Hay
1887a, b). Klugh (1916) recorded pronounced floristic differences in the
Figure 1. The Cobscook Bay, Maine area near the entrance to the Bay of Fundy.
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 3
Passamaquoddy Bay-Bay of Fundy region versus other Canadian Maritime
areas, which were confirmed by Bell and MacFarlane (1933), MacFarlane
and Milligan (1965), Colinvaux (1966), and Edelstein et al. (1970a). The
last study summarized monthly records of occurrence, vertical distribution,
and reproductive phenologies of seaweeds from Digby Neck, NS, Canada.
Analogous seasonal studies were made by Hehre et al. (1970) and Stone et
al. (1970) on South Wolf and Campobello Islands (New Brunswick), respectively,
near the mouth of the Bay of Fundy. Koetzner and Wood (1972)
described the summer flora of Kent Island, NB, Canada, which is southeast
of Grand Manan Island. Smith et al. (1978) surveyed seaweed populations at
13 sites in southwestern New Brunswick, detailing species composition and
distributional patterns. Wilson et al. (1979) produced a checklist of seaweeds
from the Bay of Fundy, including distributional maps of individual taxa
and a synopsis of earlier findings. The geographical limits of their checklist
extended from Grand Manan in the northwest to Cape Sable Island in southwestern
Nova Scotia (i.e., “Fundy approaches;” MacFarlane 1961). South
et al. (1988) surveyed the benthic marine algae at 50 sites in southwestern
New Brunswick, including the Passamaquoddy Bay area, which is one of the
richest marine environments in northeast America (Thomas 1983).
The effects of nori aquaculture on the species composition and abundance
of various Porphyra taxa were assessed at two nori aquaculture sites in Cobscook
Bay (Mathews Island, Eastport and Huckins Ledge, Lubec; Appendix
2, sites 7 and 19) during 1997 and 1998, using detailed transect and quadrat
analyses, plus molecular evaluations (Klein et al. 2003; Levine 1998; Watson
et al. 1998, 1999; Yarish et al. 1998). The occurrence and distribution
of fouling seaweeds at eight Cobscook Bay sites were also evaluated during
a rapid assessment survey (Mathieson et al. 2008b; Pederson et al. 2005),
which attempted to identify native, introduced, and cryptogenic species
(Carlton 1996) growing on floating docks and other artificial structures. The
distribution, ecology, and genetics of dwarf Fucus populations on sandy high
intertidal salt marshes at Reversing Falls (Appendix 2, site 28) were also
evaluated (Mathieson et al. 2006). Starting in 1994, the Nature Conservancy
fostered a decade of interdisciplinary research on Cobscook Bay, which attempted
to gather a broad base of scientific information that would aid in the
conservation and management of the Bay (Larsen 2004a, b). Five botanical
studies were conducted: phytoplankton productivity (Phinney et al. 2004);
biomass and productivity of intertidal rockweeds (Vadas et al. 2004c); and
growth and productivity of kelps (Vadas et al. 2004a), red and green algae
(Vadas et al. 2004b), and eelgrass (Beal et al. 2004).
The objectives of the present study were as follows: (1) to assess Cobscook
Bay’s benthic marine algal flora using a variety of historical and recent
collections; (2) to compare the species composition of seaweeds at 37 sites;
(3) to compare Cobscook Bay’s total marine flora with the Bay of Fundy, Passamaquoddy
Bay, and Casco Bay; (4) to summarize distributional patterns
of individual taxa within Cobscook Bay; (5) to document any new records
4 Northeastern Naturalist Vol. 16, Monograph No. 5
for the state of Maine, including introduced species; (6) to summarize zonation
patterns of several conspicuous benthic organisms extending from
“Downeast” Maine (i.e., from Mount Desert Island to the Bay of Fundy) to
New Hampshire; (7) to evaluate morphological variability of kelp populations
at four Downeast sites and one mid-coastal Maine location; and (8) to
evaluate the ecology and occurrence of the dwarf embedded (i.e., limicolous)
Fucus taxa in Cobscook Bay versus other Northwest Atlantic sites.
General Ecology of Cobscook Bay
Cobscook Bay, ME (Fig. 1) is located in northeastern North America
between Cutler, ME and Point Lepreau, NB, Canada, and from Grand
Manan Island to the tidal head (dam) on the St. Croix River and other minor
tributaries (“Quoddy area;” Larsen 2004b). Included within this region are
Passamaquoddy and Cobscook Bays, interconnected passages (e.g., Oak
Bay), Campobello Island, the Deer Island archipelago, and many smaller
islands. At high tide, Cobscook Bay’s surface area is approximately 110 km2
and has a convoluted shoreline of approximately 325 linear km; at low tide,
the Bay is about 74 km2 and has approximately 37 km2 of intertidal mudflats
(Larsen 2004b). Kelley and Kelley (2004) estimated that more than 70% of
Cobscook Bay’s bottom is covered by gravel and rock, while mud occurs in
scattered shallow-water coves and in two large deposits within the central
Bay. Larsen and Gilfilan (2004) emphasized that Cobscook Bay differs from
other Maine estuaries and embayments because of the coarse nature of its
bottom sediments, in contrast to the more typical mud and sand in other estuaries
(Larsen 1979, Shorey 1973).
Cobscook Bay has narrow openings to the sea at Head Harbor Passage
between Campobello Island and Eastport and near Lubec (Fig. 1). Strong
tidal currents (≈2 m/s) within the Bay maintain cold temperatures and effi-
cient exchange with offshore waters year-round (Brooks 2004, Brooks et al.
1999), which cause mixing of the water column and contribute to the Bay’s
unusually productive and diverse ecosystems (Larsen 2004a, Trott 2004a).
Typically, temperatures range from 4–14 °C (Sowles and Churchill 2004),
while salinities are >30‰, except at the tidal headwaters of the Dennys
and Pennamaquan Rivers (Hertzman 1992). The former river is the largest
stream entering the Bay, with an estimated maximum discharge of 8 m/s,
while the latter is about 45% the size of the Dennys River (Brooks et al.
1999). The annual ranges of temperatures and salinities within Cobscook
Bay are narrower than those found at many other areas on the east coast
of North America (Larsen 2004c, Mathieson et al. 1991). Turbidity is low
throughout most of the Bay, and the euphotic zone (i.e., 1% of surface irradiance)
reaches the bottom at all peripheral stations around the Bay (5.7 to
11.5 m; Phinney et al. 2004), while the Bay has mean and maximum depths
of 10 and 45 m, respectively.
Cobscook and Passamaquoddy Bays have similar climatic, geological,
and tidal settings, but the former is smaller, shallower (mean of 10 versus
25 m), has larger ratios of shoreline/area and intertidal/total area, and a
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 5
greater tidal prism (Larsen 2004c). The tides of Cobscook Bay are mixed
semi-diurnal, with a mean amplitude of 5.7 m at Eastport and a maximum of
7.6 m during extreme spring tides. Using modeling studies, Brooks (2004)
reported flushing times in the outer Bay of 1–2 days versus a week or more
within inner embayments that are repositories of particulate matter (Kelley
and Kelley 2004). Brooks (2004) also notes that about one half a cubic kilometer
of seawater enters and leaves the Bay on each flood and ebb tide. The
amount of freshwater entering the Bay via the Dennys and Pennamaquan
Rivers is negligible and less than 1% of the intertidal volume (0.5 km3) that
enters the Bay on each flood and leaves at ebb (Brooks 2004; Campbell
2004). The Bay’s hydrological patterns differ from many other macrotidal
estuaries in Maine, which may receive substantial river input (Gleizon et
al. 2003). Trott (2004b) also noted that all of the water entering the inner
Bay, which is formed by the confluence of Whiting and Dennys Bays, must
pass through a single narrow opening restricted by an island. Therefore, all
processes occurring in the inner Bay are dependent on mixing with offshore
waters. The occurrence of a narrow bedrock constriction near Eastport (Kelley
and Kelley 2004) produces two large gyres (back-eddies) in the Bay
(Brooks 2004, Brooks et al. 1999).
In discussing the effects of salmon aquaculture facilities in Cobscook
Bay, Kelley and Kelley (2004) noted that particulate organic materials can
accumulate within shallow muddy subtidal regions and intertidal mudflats
throughout the Bay. Garside and Garside (2004) considered Cobscook Bay
to be nutrient rich and a potentially eutrophic habitat. Their salinity-nitrate
plots showed that the most important source of these nutrients was the Gulf of
Maine as diminishing concentrations occurred inland. Despite high natural
nutrient loading, grazing of phytoplankton serves to limit their accumulation
and potential eutrophication; thus, man-made nutrient contributions (e.g.,
via salmon aquaculture) appear to have only local impacts on phytoplankton
(Sowles and Churchill 2004). Phinney et al. (2004) suggested that these nutrients
may be utilized by macrophytes to form “green tides” (Fletcher 1996)
rather than phytoplankton blooms (but see Sowles and Churchill 2004). For
example, Vadas et al. (2004c) reported increased seaweed cover and biomass
near salmon pens, and Larsen et al. (2004) recorded increased green algal
coverage (cf. Timson 1976, Vadas and Beal 1987). However, Sowles and
Churchill (2004) recorded variable occurrences of algal mats on mudflats
before the 1970s, prior to extensive marine salmon farming.
Since 1970, the intertidal fauna at several inner Cobscook Bay sites has
declined in species richness, with a shift from species typical of hard bottoms
to the formation of mussel beds (Trott 2004b). The primary cause of
this change appears to be increased sedimentation, possibly due to increased
commercial dragging. Thus, many kelp beds are no longer present at several
locations, and rocks, shells, and algae often are coated with a veneer of mud
that resists removal. Anoxic conditions were so extreme at some locations
(e.g., Wilbur Neck) that a black zone was evident just millimeters below the
surface of the sediment (Trott 2004b).
6 Northeastern Naturalist Vol. 16, Monograph No. 5
Methods
Collections of Cobscook Bay seaweeds, including historical specimens
dating back to the 1870s, are deposited in various herbaria, including the
Farlow Herbarium (FH), the New York Botanical Garden (NY), the D.C.
Eaton Algal Herbarium (YALE), the University of Michigan (MICH), the
Jepson Herbarium of the University of California, Berkeley (UC), and the
Brooklyn Botanical Garden (BKL). Most of these early collections were
made by D.C. Eaton (1873), W.G. Farlow (1881), and various colleagues
(Farlow et al. 1877–1889) during summer or fall in the Eastport, ME area.
In 1970, I.M. Lamb made some collections from Edmunds, ME that are in
FH, while G.R. South and his students collected in the Edmunds and Pembroke
areas during 1977, and their specimens are deposited in the Atlantic
Reference Center of the Huntsman Marine Science Center (St Andrews, NB,
Canada). All herbarium samples were evaluated to confirm their identifications,
locations, and habitat characterizations.
Our studies of Cobscook Bay seaweeds were conducted between 1966
and 2006 and included seasonal comparisons of 37 sites (Appendix 2), including
outer and inner embayments, tidal and non-tidal rapid sites, various
salt marsh habitats, tidal rivers (i.e., Dennys and Pennamaquan), and two
sites contiguous with nori aquaculture farms (Mathews Island and Huckins
Ledge). Approximately 1600 voucher samples resulting from these collections
(Appendix 2) are deposited in the Albion R. Hodgdon Herbarium at
the University of New Hampshire (NHA). Species composition, numbers
of taxa, and percent occurrence of each taxon/site were summarized. The
presence and origin of introduced species were documented using historical
and recent collections throughout the Northwest Atlantic (Hofmann et al., in
press; Mathieson et al. 2008a, b, c; Villalard-Bohnsack 2002).
Cobscook Bay’s seaweed flora was compared with the Bay of Fundy
(NB, NS, and ME; Wilson et al. 1979), Passamaquoddy Bay (NB; South et
al. 1988), and Casco Bay (ME; Mathieson et al 2008a). The numbers and
similarities of shared taxa from each of the four locations were documented,
with the latter summarized using Kulezynski’s coefficient as outlined by
Bray and Curtis (1957): C = 2W/(A +B), where C = their index of similarity,
W = the number of taxa in common to both areas, A = the number at one site,
and B = the number in the other location.
The nature of the seaweed floras at these same four sites, plus a composite
of the three northern embayments was assessed using Cheney’s (1977) floristic
ratio: (R + C)/P, where R = the number of Rhodophyceae, C = the number
of Chlorophyceae, and P = the number of Phaeophyceae. A value of <3.0 indicates
a temperate or cold-water flora, while values of >6.0 indicate a tropical
one; intermediate values represent a mixed (i.e., warm temperate) flora.
A variety of taxonomic references, including Taylor (1962), South and
Tittley (1986), Van Oppen et al. (1995), Sears (2002), Gabrielson et al.
(2006), Guiry and Guiry (2009), and Hommersand and Lin (2009), were
employed to identify Cobscook Bay seaweeds. Several other sources of
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 7
references are in Mathieson et al. (1998, 2001) and a review of seaweeds
from the northwest Atlantic (Mathieson and Dawes, in prep.). Nomenclature
primarily follows Silva et al. (1996) and Sears (2002), except for some recent
changes resulting from molecular studies of the Phyllophoraceae (Fredericq
and Ramírez 1996), Ulvales (Hayden et al. 2003), and Laminariales (Lane et
al. 2006).
Zonation patterns of several conspicuous benthic intertidal organisms
were determined at four macrotidal Downeast Maine sites during the fall of
1994: West Quoddy Head, Lubec (66°57'W, 44°49'N) and three Cobscook
Bay Pembroke sites (two at Reversing Falls and another at Wilbur Neck [Appendix
2, sites 28a, 28b, and 30, respectively]). A comparison of zonation
patterns at three southern Maine and New Hampshire open coastal sites with
reduced tidal amplitudes was made, including an exposed site at Bald Head
Cliff in York, ME (Femino and Mathieson 1980), a semi-exposed Fort Stark
site on Newcastle Island, NH (Mathieson et al. 1981a), and a sand-abraded
site at Bound Rock, in Seabrook, NH (Daly and Mathieson 1977). A line
level and surveying rod were used to make height measurements after pulling
a level line from a primary or secondary reference point (Dawes 1998).
Vertical heights were calculated above or below mean low water (MLW) by
plumbing down to low water from a reference point and using the Harbor
Master Computer Program (Version 3, Zihua Software, Marlboro, MA). The
accuracy of these elevational measurements is about ± 5.0 cm (Mathieson et
al. 1998).
Measurements of maximum thallus length of six kelps (Agarum clathratum,
Alaria esculenta, Laminaria digitata, Saccharina latissima, S.
longicruris, and Saccorhiza dermatodea) were made at the same four Cobscook
Bay sites outlined above. Depending upon the presence and abundance
of various kelps, 25–30 fronds of each taxon were collected and measured to
enumerate morphological differences and potential patterns of gigantism. We
also calculated the mean (± SD) frond widths and length/width ratios of 25–30
S. latissima plants at four distinct Maine sites, including exposed open coastal
sites at West Quoddy Head (Lubec) and Bald Head (Phippsburg), a composite
of six non-tidal rapids sites within Cobscook Bay, and an exposed tidal rapids
habitat at Reversing Falls Park (Pembroke).
Results
Species composition and historical floristic comparisons
The composite benthic marine algal flora of Cobscook Bay is based
upon 37 collection sites in Cobscook Bay and consists of 148 taxa (Appendix
1, Appendix 2), including 38 green, 46 brown, and 64 red algae.
Six heteromorphic life-history partners were recorded plus four taxa with
questionable affiliations—i.e., “Chlorochytrium inclusum”, “Codiolum
petrocelidis”, “Codiolum pusillum”, and “Ralfsia clavata”. Two other
gametophytic/ sporophytic pairs (i.e., Bonnemaisonia hamifera/“Trailliella
intricata” and Mastocarpus stellatus/“Petrocelis cruenta”) were identified
8 Northeastern Naturalist Vol. 16, Monograph No. 5
(Appendix 3), with their life-history partners sometimes being partially or
completely uncoupled.
Patterns of species richness at the 37 Cobscook Bay sites were highly
variable (Fig. 2), with the highest numbers of taxa occurring at Eastport Harbor
(site #1; 70 taxa), Reversing Falls Park (site #28; 51 taxa), and Wilbur
Neck and the area near the Friedman Marine Laboratory (sites #30 and #35;
43 taxa each). Intermediate numbers ranging from 29 to 18 taxa were found
at four sites (#4, 5, 17, and 24), while the remaining 29 sites varied from 11
taxa (site #10, Carlow Island, Eastport and site #29, Crows Neck, Trescott)
to 0 taxa (site #20, Lead Mine Road, Lubec).
Several freshwater algal taxa, not included in Appendix 1 or Appendix 3,
occurred at inner estuarine or ”brackish” water sites, including one green
(Zygnema sp. at site #33, August 1994) and two red algae (Audouinella hermanii
(Roth) Woelkerling and Lemanea fucina Bory de Saint-Vincent at sites
#27 and 33, August 1994). Audouinella hermanii was a specific epiphyte on
stunted plants of L. fucina. An undescribed yellow green alga (Tribonema
sp.) was found during spring runoff (March–April 1996) at three sites (#20,
23, 36). The marine macroscopic colonial diatom Berkeleya rutilans (Trentopohl)
Grünow was recorded once at site #5 during August 2005.
In comparing historical (1800s) and recent collections of seaweeds from
Cobscook Bay (Appendix 3), 67 taxa were recorded by Eaton (1873), Farlow
(1881), Farlow et al (1877–1889), and Verrill (1871), including 15 green, 23
brown, and 29 red algae, in contrast to the 148 taxa cited above. Such differential
numbers of taxa were probably due to limited seasonal and spatial sampling
of earlier collections. Nine historical records were not confirmed by recent
collections (i.e., Urospora penicilliformis, Ralfsia fungiformis, Sphacelaria
radicans, Stictyosiphon griffithsianus, Colaconema daviesii, Hydrolithon
Figure 2. Patterns of species richness at the 37 Cobscook Bay, ME sites.
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 9
farinosum, Lithothamnion ungeri, Scagelia americana, and Titanoderma pustulatum),
while 82 taxa were not found previously (Appendix 3).
Floristic affinities
A total of 316 taxa are known from the four Northwest Atlantic embayments,
including 81 green, 111 brown, and 124 red algae (Tables 1, 2). The
number (%) of taxa/area declined from the Bay of Fundy (250 taxa, 79.4%),
to Casco Bay (201 taxa, 64%), Passamaquoddy Bay (192 taxa, 61%), and
Cobscook Bay (148 taxa, 47%). Thus, Cobscook Bay had the lowest numbers
(%) of the region’s total seaweed flora. The patterns for mean number
(± SE) of shared taxa (Table 1) showed values ranging from 169.8 ± 27.9
(Bay of Fundy) to 152.3 ± 16.4 (Passamaquoddy Bay), 151.5 ± 16.9 (Casco
Bay), and 130.5 ± 6.5 taxa (Cobscook Bay). By contrast, the mean % similarity
(± SE) values varied from 78.0 ± 8.9 (Passamaquoddy Bay) to 76.7 ± 8.1
(Cobscook Bay), 76.1 ± 8.4 (Casco Bay), and 74.8 ± 8.9 (Bay of Fundy).
Using Cheney’s floristic ratio of (R + C)/P, the combined flora of 316 taxa
from the four embayments had a ratio of 1.9, which designated a cold-temperate
flora. The individual floristic ratios ranged from 1.8 (Bay of Fundy)
to 1.9 (Casco Bay), 2.2 (Cobscook Bay), and 2.3 (Passamaquoddy Bay).
These minor differences in floristic ratios primarily reflect variable acreages
(habitats), numbers of brown algae, and frequency of seaweed evaluations.
Distributional patterns of individual taxa
As Figure 3 shows, 38 of the 148 seaweed taxa recorded from Cobscook
Bay were restricted to a single location (2.7%), 62 were collected at
2 or 3 sites (5.4–8.1%), and 14 species were found at 8 to 13 sites (21.6 to
35.1%). The most circumscribed taxa (2.7% occurrence) included 11 green
(Acrochaete wittrockii, Bryopsis plumosa, “Chlorochytrium inclusum”,
Cladophora ruchingeri, Codium fragile subsp. fragile, Gomontia polyrhiza,
Table 1. Floristic affinities of seaweed populations from the Bay of Fundy (BF), Passamaquoddy
Bay (PB), Cobscook Bay (COB), and Casco Bay (CAB), expressed as the number and (percent)
of shared taxa including means, standard deviations (SD) and standard errors (SE).
BF PB COB CAB
BF 250
(100)
PB 164 192
(74.2) (100)
COB 127 117 148
(63.8) (68.8) (100)
CAB 138 136 130 201
(61.1) (69.0) (74.3) (100)
Mean 169.8 152.3 130.5 151.5
SD ±55.7 ±32.8 ±12.9 ±33.8
SE ±27.9 ±16.4 ±6.5 ±16.9
Percent 74.8 78.0 76.7 76.1
SD ±17.7 ±14.9 ±16.1 ±16.7
SE ±8.9 ±7.5 ±8.1 ±8.4
10 Northeastern Naturalist Vol. 16, Monograph No. 5
Pringsheimiella scutata, Spongomorpha aeruginosa, Ulva flexuosa, Ulva
torta, and Urospora penicilliformis), 10 brown (Asperococcus fistulosus,
Fucus distichus subsp. edentatus, Fucus sp. “muscoides-like”, Haplospora
globosa, Hincksia granulosa, Pseudolithoderma extensum, Punctaria
plantaginea, Ralfsia fungiformis, Sphacelaria radicans, and Stictyosiphon
griffithsianus), and 17 red algae (Acrochaetium alariae, Audouinella
polyidis, Bangia fuscopurpurea, Callithamnion tetragonum, Colaconema
dasyae, Colaconema daviesii, Gracilaria tikvahiae, Halosacciocolax kjellmanii,
Hydrolithon farinosum, Lithothamnion ungeri, Phyllophora truncata,
Phymatolithon lenormandii, Polysiphonia denudata, Porphyra yezoensis f.
narawaensis, Scagelia americana, Titanoderma pustulatum, and Turnerella
pennyi). The most cosmopolitan seaweeds, which were found at 8–13 sites
(21.6 to 35.1%), included 5 green (Acrosiphonia spinescens, Ulva intestinalis,
U. lactuca, U. linza, and U. prolifera), 6 brown (Ascophyllum nodosum,
Chordaria flagelliformis, Fucus vesiculosus, Pylayella littoralis, Saccharina
Table 2. Elevational comparisons of several conspicuous benthic organisms at seven sites ranging
from West Quoddy Head, Lubec, ME to Bound Rock Rock, Seabrook, NH expressed as
meters above or below mean low water (MLW). WQH= West Quoddy Head, Lubec, ME; RF
“#1” = Reversing Fall site ”#1”, Pembroke, ME; RF “#2”= Reversing Falls site “#2”, Pembroke,
ME; WN= Wilbur Neck, Pembroke, ME; BHC= Bald Head Cliff, York, ME; FS= Fort Stark
(Jaffrey Point), Newcastle, NH; Bound Rock, Seabrook, NH.
WQH RF “#1” RF “#2” WN BHC FS BR
Salt marsh - - 2.9–3.8 3.9–4.6 - 2.7–3.1 -
Chlorophyceae
Cladophora sericea - - 0.0–0.8 0.0–0.8 - 2.1–2.7 -
Ulva intestinalis/ - - 0.0–0.8 - 1.8–2.8 2.1–2.4 -0.5–+0.9
U. compressa
Ulva lactuca 0.0–0.9 0.0 0.0–0.8 0.0–0.8 0.1–2.0 -0.1 -0.5–+0.9
Phaeophyceae
Ascophyllum nodosum 1.3–6.0 0.9–3.5 0.2–2.7 0.0–3.5 1.5–2.0 0.0–2.0 -
Fucus distichus subsp. 0.0–0.9 - - - 1.2–1.7 0.0–0.6 0.02–0.5
edentatus
Fucus distichus subsp. - 0.0–1.4 0.0–0.6 - - 0.0–0.6 -
evanescens
Fucus spiralis 6.0–6.3 3.8–4.1 2.4–2.9 3.5–3.9 - 2.3–2.7 -
Fucus vesiculosus - 1.0–4.1 0.0–2.2 - 0.2–2.8 0.0–2.4 0.0–1.7
Saccharina latissima/ 0.0 0.0 0.0 0.0 - -0.1 -
S. longicruris
Pylaiella littoralis - - 0.0–0.8 - - 0.0–1.3 -
Rhodophyceae
Palmaria palmata 0.0–0.9 0.0 - 0.0–0.2 1.7–1.8 -0.1 -
Polysiphonia lanosa - 0.9–2.1 0.2–1.8 - - 0.2–2.0 -
Porphyra umbilicalis 0.0–2.2 0.0–1.4 0.0–1.2 0.0–0.4 1.7–1.8 0.0–2.5 1.0–1.8
Major invertebrates
Mytilus edulis - 0.0–1.7 - - 0.1–3.0 - 0.0–1.8
Semibalanus balanoides - 1.1–3.56 - - 1.8–3.4 0.0–2.7 0.4–1.8
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 11
latissima, and Scytosiphon lomentaria), and 3 red algae (Palmaria palmata,
Polysiphonia stricta, and Porphyra purpurea). Three seaweeds (Colaconema
dasyae, Lithothamnion ungeri, and P. yezoensis f. narawaensis), all members
of the Rhodophyceae, were restricted to Cobscook Bay and not found in
the Bay of Fundy, Passamaquoddy Bay, or Casco Bay (Appendix 1). By contrast,
the Bay of Fundy had 162 taxa that were not found in Cobscook Bay,
while Passamaquoddy and Casco Bays had 74 and 68 taxa, respectively, that
were not recorded in Cobscook Bay (Appendix 1). Three detached Fucus
taxa (i.e., F. spiralis ecad lutarius, F. sp. “muscoides-like”, and F. vesiculosus
ecad volubilis) were restricted to salt marsh habitats within Cobscook
and Casco Bays.
Introduced species
Six introduced seaweeds were recorded from Cobscook Bay (Appendix 1),
with four of these originating from Asia: Codium fragile subsp. fragile,
Figure 3. Distributional patterns of 148 seaweed taxa from Cobscook Bay expressed
as the number of red, brown, and green algal taxa/site at the 37 study sites.
12 Northeastern Naturalist Vol. 16, Monograph No. 5
Bonnemaisonia hamifera as its “Trailliella intricata” sporophyte stage,
Neosiphonia harveyi, and Porphyra yezoensis f. narawaensis. One taxon
was from the North Pacific (Melanosiphon intestinalis) and another from
Europe (Dumontia contorta). Codium and P. yezoensis f. narawaensis were
only collected once (2.7%), with the former occurring in drift at Wilbur Neck
(site #30) and the latter growing on Coastal Plantation’s nori cultivation nets
at Goose Island (site #8). “Trailliella intricata” and M. intestinalis were
found at two sites (5.4%), while D. contorta and N. harveyi were collected
at three locations (8.3%). Overall, introduced seaweeds were found at 10 of
the 37 sites (27.0%). A single introduced taxon occurred at sites #7, 8, 10,
14, 22, 28, 29, and 35, while two were collected at site #4 (M. intestinalis
and N. harveyi), and three at site #30 (C. fragile subsp. fragile, D. contorta,
and the “Trailliella intricata” phase of B. hamifera).
Five other introduced species (Colpomenia peregrina, Fucus serratus,
Ulonema rhizophorum, Lomentaria clavellosa, and L. orcadensis) were
found at one or more of the 3 other embayments (Appendix 1), with all of
these originating from Europe. Overall, a total of 8 introduced taxa were recorded
in the Bay of Fundy, 4 in Passamaquoddy Bay, and 8 in Casco Bay.
Zonation and ecology of selected seaweeds and invertebrates
Salt marsh communities were present at two sites within Cobscook Bay
and one in southern New Hampshire (Table 2); their vertical stratifications
and maximum heights at the more sheltered Reversing Falls site #2
(2.9–3.8 m) and Wilbur Neck (3.9–4.6 m) exceeded those at Fort Stark,
NH (2.7–3.1 m). Similar expansive patterns of four fucoid algae (i.e., Ascophyllum
nodosum, Fucus distichus subsp. evanescens, F. spiralis, and
F. vesiculosus) occurred at one or more Downeast sites versus those in
southern Maine and New Hampshire. By contrast, populations of Cladophora
sericea at Fort Stark and Ulva intestinalis/U. compressa at both Bald
Head Cliff and Fort Stark showed the opposite pattern and were restricted
to high tide pools—i.e., 2.1–2.7 and 2.1–2.4 m, respectively. Four of the
nine taxa found at the exposed Bald Head Cliff site exhibited the most expansive
distributional patterns versus the other six sites—i.e., Ulva lactuca
(0.1–2.0 m), Fucus distichus subsp. edentatus (1.2–1.7 m), Palmaria palmata
(1.7–1.8 m), and the mussel Mytilus edulis L. (0.1–3.0 m). The upper
distributional limits of Pylaiella littoralis and Porphyra umbilicalis were
highest at Fort Stark (i.e., 0.0–3.3 and 0.0–2.5 m, respectively), while the
sand-abraded Bound Rock site had the most circumscribed upper distributional
limits for the barnacle Semibalanus balanoides (L.) at the and the
most expansive lower zonation for U. intestinalis/U. compressa (+0.5 to
+0.9 m) and U. lactuca (-0.5 to + 0.9 m).
Figure 4 summarizes the mean frond length (± SD) of six different kelp
species at four Downeast Maine sites. Agarum clathratum, Alaria esculenta,
and Saccharina longicruris occurred at three of the four sites. Agarum
ranged from 55 ± 11 cm to 77 ± 5 cm at the Reversing Falls sites #1 and #2,
respectively, while Alaria varied from 31 ± 2.0 cm to 124.8 ± 34.8 cm at West
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 13
Quoddy Head and Wilbur Neck, respectively. Saccharina longicruris exhibited
a pronounced morphological pattern, being smallest at West Quoddy
Head (81.7 ± 62.0 cm) and the Reversing Falls site #1 (143.5 ± 30.7 cm) and
largest at Wilbur Neck (459.3 ± 91.6 cm). Laminaria digitata and Saccorhiza
dermatodea occurred at two sites; the former at West Quoddy Head (81 ±
62.0 cm) and Reversing Falls site #1 (137.7 ± 30.7 cm), and the latter at the
Reversing Falls site #2 (127.7 ±16.7 cm) and Wilbur Neck (165.8 ± 87.7 cm).
Only Saccharina latisssima was found at all four sites, with its smallest plants
at West Quoddy Head (62.5 ± 62 cm) and largest ones at Reversing Falls site
#2 (261.7 ± 115.7 cm).
Figure 5 illustrates morphological variability of Saccharina latissima
populations from four Maine sites, including West Quoddy Head (5a–c),
Cobscook Bay (Boot Cove; 5d), Reversing Falls Park (Pembroke; 5e–g),
and Bald Head (Phippsburg; 5h–j). A conspicuous morphological gradient
was evident between West Quoddy Head (Fig. 5a) and Bald Head (5j), with
Reversing Falls specimens (5e–g) being intermediate. A comparison of morphological
features (Fig. 6) shows that West Quoddy Head and Cobscook
Bay S. latissima had reduced length/width ratios of 5.0 ± 2.1 and 3.5 ± 0.6,
respectively and mean frond widths of 10.5 ± 2.0 cm and 16.2 ± 3.7 cm,
respectively; Reversing Falls and Bald Head had much higher length/width
Figure 4. Mean frond length (± SD) of six different kelp species at four Downeast Maine
sites. WQH = West Quoddy Head, RF = Reversing Falls, and WN = Wilbur Neck.
14 Northeastern Naturalist Vol. 16, Monograph No. 5
Figure 5. Morphological variability of seven Saccharina specimens from Downeast
Maine (A–G) plus three from mid-coastal Maine at Bald Head, Phippsburg (H–J):
(A) A broad open coastal specimen of S. latissima from West Quoddy Head, South
Lubec, 21 May 1966; (B and C) Two somewhat narrower and smaller open coast
plants of S. latissima from the same site, 13 February1967; (D) A somewhat narrow
open coastal specimen of S. latissima from Boot Cove, Lubec, 5 December 1997;
(E–G) Three very narrow and elongated plants of S. latissima from Reversing Falls
Park, Pembroke, ME, 16 May 1996; (H–J) Three very narrow and elongated specimens
of S. latissima f. angustissima from Bald Head, Phippsburg, Maine collected
on 10 June 1995 (H), 5 March 2000 (I), and 1 September 1996 (J). Note the morphological
similarity of Reversing Falls (E–G) and Bald Head specimens (H–J) and the
continuous morphological gradient between plants in A–G.
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 15
ratios (27.6 ± 2.1 and 52.5 ± 5.3, respectively) and lower frond widths (2.1
+0.17 and 1.3 ± 0.08 cm).
Annotated checklist
Appendix 2 summarizes the 37 study sites within the Cobscook Bay
area and the total number of voucher specimens per site. A synopsis of
all historical and recent collections of seaweeds from Cobscook Bay is
summarized in Appendix 1 and Appendix 3, with the different taxa listed alphabetically
within each class. Appendix 3 also lists the sites where seaweed
Figure 6. Mean length/
width ratios (± SE) and
frond widths of Saccharina
latissima plants
from West Quoddy Head
(Lubec), a composite of
six non-tidal rapids sites
from Cobscook Bay, Reversing
Falls Park (Pembroke),
and Bald Head
(Phippsburg).
16 Northeastern Naturalist Vol. 16, Monograph No. 5
samples were collected, numbers of voucher specimens, dates of collections,
collector(s), and percent occurrence of each taxon within Cobscook Bay.
At least one collection record for each taxon per site is included, although
several others were often made.
Discussion
The number of red, brown, and green marine benthic algae recorded
from Cobscook Bay (148 taxa) represents 37% of the total flora (ca. 403
taxa) recorded by Sears (2002) for the northeastern coast of North America
extending from the Strait of Belle Isle near Newfoundland to Long Island
Sound. The Bay’s flora also represent approximately 47% of the composite
seaweed biota (315 taxa) found in all four embayments ranging from
the Bay of Fundy to Casco Bay. Furthermore, Cobscook Bay’s floristic
diversity is lower than the Bay of Fundy (250 taxa), Casco Bay (201 taxa),
and Passamaquoddy Bay (192 taxa). A comparison of Cobscook Bay’s
flora with several other Northwest Atlantic sites also shows some interesting
patterns. For example, its flora exceeds that found in Penobscot Bay,
ME (145 taxa; Mathieson et al. 1996, 1998) and the York River Estuary
(ME) plus the adjacent open coast (131 taxa; Mathieson et al. 1993), while
it is the same as that found within Brave Boat Harbor, ME and coastal
environs (148 taxa; Mathieson et al. 2001), but considerably less than
Newfoundland’s flora (ca. 254 taxa; South 1983, South and Hooper 1980)
and the Great Bay Estuarine System of New Hampshire-Maine (216 taxa;
Mathieson and Hehre 1986). Several factors may contribute to these varying
patterns of species richness, including collecting efficiency, frequency
of observations, habitat size and diversity, hydrographic stability, pollution,
water temperatures, and availability of rocky substrata (Larsen 2004a;
Mathieson et al. 1991, 2008a).
Only two sites in Maine—Mount Desert Island (Mathieson et al. 1998)
and Casco Bay (Mathieson et al. 2008a)—have detailed historical records
that can be used to make floristic comparisons of the late 1800s/early
1900s and the present time. The similarity indices (historical versus recent)
for Mount Desert and Casco Bay’s floras were comparable, being 75 and
79.9, respectively. By contrast, lower values were evident for individual
Mount Desert sites, with these indices ranging from 43 at Seal Harbor to 68
at Otter Cliffs. The situation in Cobscook Bay was obviously different than
that of Mount Desert or Casco Bay, as the early floristic studies of Eaton
(1873), Farlow (1881), and colleagues (Farlow et al. 1877–1889) were
restricted seasonally and spatially. For example, Eaton (1873) wrote that
his collections from Eastport “were sought for only a few weeks in August
and September, and if this coast could be thoroughly explored at different
seasons, the list would doubtless be much extended.” Thus, the index of
similarity (≈56.0%) and number of taxa in common were low when the two
time periods were compared.
Following the initial botanical studies of seaweeds in Cobscook Bay,
there was a hiatus of such studies until the 1980s, in contrast to the extensive
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 17
floristic studies conducted during this period in the Canadian Maritime Provinces
and Newfoundland (e.g., Bird and McLachlan 1992, Cardinal 1968,
Edelstein et al. 1970a, South 1983, South and Hooper 1980, South et al.
1988, Wilson et al. 1979). Other than the present study, most recent investigations
of Cobscook Bay seaweeds have been associated with environmental
impact studies (Larsen 2004b, Timson 1976, Vadas et al. 2004c), autecological
investigations of individual seaweeds (Mathieson et al. 2006; Vadas and
Beal 1987; Watson et al. 1998, 1999), or projects associated with the Nature
Conservancy (Beal et al. 2004; Vadas et al. 2004a, b, c).
Timson (1976) estimated that approximately 1062 ha of tidal mudflats were covered with green algae based upon ground-truthing of Maine
Geological Survey maps, while Larsen et al. (2004) reported 1370 ha of
green-tide populations based upon Landsat-derived evaluations. Sowles and
Churchill (2004) interviewed many Cobscook Bay resident who recalled
variable patterns of green-tide populations, some of which covered entire
mudflats prior to extensive marine salmon farms (i.e., before the 1970s).
Recent rapid assessment studies in Cobscook Bay (Mathieson et al. 2008b)
have shown extensive green-tide populations at several sites in Eastport,
West Lubec, Lubec, Pembroke, Dennysville, Edmunds, Trescott, and Whiting
(Appendix 2), many of which were either near or contiguous to various
salmon farms. Increased seasonal and spatial sampling will no doubt enhance
our knowledge of green-tide blooms as well as the Bay’s total flora.
An evaluation of Cheney’s (1977) (R + C)/ P floristic ratios showed a
strongly cold-temperate flora (2.0) for the three Quoddy regions (Cobscook
Bay, Passamaquoddy Bay, and Bay of Fundy), while Cobscook Bay had a
slightly higher ratio of 2.3. Hence, the floristic patterns for Cobscook Bay
correspond with its moderate temperature range of 4–14 °C (Sowles and
Churchill (2004) versus more variable estuarine sites (-2.0 to 28 °C) like the
Great Bay Estuarine System (Mathieson and Hehre 1986). Values of <6.0 (=
“mixed” or “cold-water” floras) were recorded at outer and mid-estuarine
sites in southern Maine and New Hampshire, while inner riverine sites had
reduced numbers of brown algal taxa and ratios of 7–15 (Mathieson and
Penniman 1986a, b). Adjacent open coastal sites in southern Maine and New
Hampshire had an overall mean of 2.5 (Mathieson and Penniman 1986b),
while those from Massachusetts through New York ranged from 3.2 to 7.3
(A.C. Mathieson, unpubl. data). Thus, Cheney’s ratio is primarily useful
for open coastal populations, and this may explain some of the high ratios
recorded in southern New England and New York where reduced numbers
of cold-water brown algae occur (Chapman 1964; van den Hoek 1975, 1982;
Hooper et al. 2002; Humm 1969; Mathieson et al. 1991; Setchell 1922; Stephenson
and Stephenson 1949). The above floristic ratios are in contrast to
Florida (5.9) and Caribbean (7.1) reef algae (Dawes and Mathieson 2008).
Relatively few warm-water or estuarine disjunct seaweeds (Bousfield and
Thomas 1975, Mathieson and Hehre 1986, Pielou 1979, Vadas 1977) occur
within Cobscook Bay, except for a few populations of Bryopsis plumosa,
Gayralia oxysperma, Ceramium deslongchampii, Gracilaria tikvahiae, and
18 Northeastern Naturalist Vol. 16, Monograph No. 5
Porphyrostromium ciliare. By contrast, some areas to the north and south of
Cobscook Bay have a greater number of such taxa, presumably because
of unique environmental conditions (e.g., shallow topographies) and/or pronounced
hydrographic variability (Mathieson and Hehre 1986, Mathieson
and Penniman 1986a). For example, 12 warm-water taxa are recorded from
the Canadian Maritime Provinces (see Appendix 1), while Collins (1911) recorded
six from Casco Bay (i.e., Cladosiphon zosterae, Eudesme virescens,
Sphaerotrichia divaricata, G. tikvahiae, Polysiphonia fibrillosa, and Rhodophysema
georgei) and Mathieson et al. (2008a) noted an additional three
(Striaria attenuata, Chondria baileyana, and Polysiphonia denudata) from
the same embayment. Several of these southerly taxa have “modified” life
histories and extensive vegetative reproduction (Hehre and Mathieson 1970,
Mathieson and Dawes 1975). Wilkinson (1980) and Mathieson et al. (1993)
also describe varying ecological requirements between inner and outer estuarine
floras, with the former representing seasonally dynamic warm temperate
or “mixed” floras (i.e., annuals) with wide ecological tolerances, and the latter
being cold-temperate, perennial taxa with more limited tolerances.
The number of taxa/site within Cobscook Bay depended in part on accessibility.
While many collections were possible at the tidal headwaters of the
Pennaquan (5 visits) and Dennys Rivers (7 visits), only a few marine taxa
were present. Several authors (Doty and Newhouse 1954, Josselyn and West
1985, Ketchum 1983, Mathieson and Penniman 1986b, Mathieson et al.
1981b, Wilkinson 1980) have described similar reduction patterns and altered
species composition near the headwaters of temperate estuaries, presumably
because of wide-ranging hydrographic conditions (particularly salinity) and
limited substratum. The occurrence of the freshwater red algae Audouinella
hermanii and Lemanea fucina on the Pennaquan and Dennys Rivers confirms
the presence of very low salinities within these tidal headwaters (Mathieson
and Hehre 1986) as both taxa have limited “time-intensity tolerances” to
salinity at low tide (Wood and Straughhan 1953). The occurrence of depauperate
and/or green-algal dominated floras at other downstream sites also
indicates environmental stress, including hydrographic variability, high
sedimentation, or eutrophication (Clokie and Boney 1980, Cotton 1910,
Edwards 1972, Fritsch 1956, Round 1981, Sawyer 1965).
Attached intertidal fucoid algae (particularly Ascophyllum nodosum and
Fucus vesiculosus) dominate sites with stable salinities, large rocky substrata,
and limited sedimentation (Mathieson et al. 1991). By contrast, detached
or entangled masses of A. nodosum ecad scorpioides and F. vesiculosus
ecad volubilis often occur at scattered sites amongst Spartina alterniflora
Loiseleur, while dwarf limicolous populations of Fucus grow in high sandy
marsh habitats near S. patens (Aiton) Muhlenberg (Appendix 3).
Tidal rapids sites like Wilbur Neck (site #30) and Reversing Falls State
Park (site #28) with large numbers of seaweed taxa have also been described
for several areas throughout the world (Kitching and Ebling 1967; Lewis
1964, 1968; Mathieson et al. 1977, 1981b, 1983). The floras at Reversing
Falls and Wilbur Neck, ME (Fig. 1), plus Dover Point, NH also have strong
open coastal affinities. Lewis (1964) noted that organisms in the upper
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 19
shorelines of tidal rapids are more typical of sheltered locations, while lower
ones are more open coastal in character. The reduced quantities and stature
of A. nodosum at Reversing Falls can be explained by high currents (Mathieson
et al. 1983). Previous studies (Chapman 1964, Conover 1968, Lewis
1964, Kitching and Ebling 1967) have emphasized that strong tidal currents
can affect both the growth and size of seaweeds. Thus, some plants (e.g.,
Chaetomorpha picquotiana) can grow 3–5 m in length in strong currents,
while others (e.g., Fucus vesiculosus) can be very diminutive (Mathieson et
al. 1983).
As outlined by Mathieson et al. (2008c) and Hofmann et al. (in press),
21 introduced seaweeds are known from the Northwest Atlantic. Six of these
were recorded during recent collections from Cobscook Bay (Appendix 1
and Appendix 3), but none were known in the 1800s. In comparing spatial
patterns within Cobscook Bay, a single introduced taxon was found at 8
of the 37 sites (21.6%), while a maximum of three were found at Wilbur
Neck (Appendix 1 and Appendix 3). The Asiatic green alga Codium fragile
subsp. fragile was only found at one site in drift (Wilbur Neck, site #30).
Villalard-Bohnsack (2002), Mathieson et al. (2008c), and Hofmann et al. (in
press) summarized the sources, sites and dates of first observations, mode(s)
of transfer, and current distributions of 21 introduced taxa known from the
Northwest Atlantic. Two of the six taxa found in Cobscook Bay are closely
tied to the Quoddy area, with Melanosiphon intestinalis being initially reported
from the Bay of Fundy (Edelstein et al. 1970b), and the U-51 strain of
Porphyra yezoensis f. narawaensis cultivated in Cobscook Bay (Bray 2006,
Levine 1998, Yarish et al. 1998). Apparently, the latter has not become a
permanent member of Cobscook Bay’s flora (Watson et al. 1998, 1999). Bray
(2006) also noted that this strain of nori is not the source of disjunct southern
materials (e.g., Long Island Sound) that predates it and has different genetic
patterns. Four other introduced taxa were found within the Bay of Fundy
and Passamaquoddy Bay areas, with the highest numbers of introduced taxa
(8) occurring in the former and 5 in the latter. In comparing other geographies,
7 introduced seaweeds were recorded during rapid assessment studies
at 67 sites between Downeast Maine and Staten Island, NY (Mathieson et
al., 2008b); only one of these was not found in the present study—i.e., the
Asiatic red alga Grateloupia turuturu Yamada. A comparison of the seaweed
flora of Mount Desert Island, ME showed three introduced taxa (Mathieson
et al. 1998), while nine were recorded from Casco Bay, ME (Mathieson et
al. 2008a) and ten from the Great Bay Estuarine System of New Hampshire-
Maine (Mathieson and Hehre 1986; Hofmann et al., in press).
The zonation patterns for the seven sites ranging from Downeast Maine
to New Hampshire showed several conspicuous differences. Foremost, the
upper limits of salt marsh vegetation plus Ascophyllum nodosum, Fucus distichus
subsp. evanescens, F. spiralis, and F. vesiculosus were much higher
at one or more of the macrotidal Downeast sites versus more southerly
mesotidal locations. However, Cladophora sericea and Ulva intestinalis/U.
compressa were restricted to high tide pools at Bald Head Cliff and Fort
Stark, while in Downeast areas they occurred in the low intertidal zone.
20 Northeastern Naturalist Vol. 16, Monograph No. 5
Downeast Maine populations of Saccharina latissima and S. longicruris occurred
slightly higher (0.0 m) than those at Fort Stark (-0.1 m). Although not
summarized in Table 2, there was a conspicuous uplifting of several other
subtidal populations (Alaria esculenta, Laminaria digitata, and Devalarea
ramentacea) within the mid-intertidal zone in Cobscook Bay, while they
were restricted to the subtidal zone farther south. The uplifting of these and
other seaweeds is a characteristic of sites near the Bay of Fundy and is probably
due to the rapid and large tidal amplitude, frequent cool, overcast to
foggy weather, and tidal upwelling (Trott 2004a).
South et al. (1988) noted that Saccharina latissima (as Laminaria
saccharina) was the dominant kelp in sheltered subtidal sites within Passamaquoddy
Bay, with plants reaching 3 m in length and having delicate
blades. Vadas et al. (2004a) stated that most kelps within the Gulf of Maine
rarely exceeded 3 m in length, although some specimens of S. longicruris (as
L. longicruris) on Mt. Desert Island reached l0 m (Vadas and Steneck 1988).
By contrast, Taylor (1962) listed kelp lengths from New England as follows:
S. longicruris (as L. longicruris), 3–5 m and up to 12 m; Laminaria digitata,
to ca 1.0 m; Saccorhiza dermatodea, to 2.0 m; Saccharina latissima (as L.
saccharina), >2.0 m; Agarum clathratum [as A. cribrosum (Mertens) Bory],
to 1.5 m; and Alaria esculenta, to 1.5 m. In a year-long study at three sites
in Cobscook Bay, Vadas et al. (2004a) reported S. longicruris frond lengths
of up to 2.5 m at Bar Island (mean 1.7 m), to 1.9 m at Mahar Point (mean
1.3 m), and to 1.2 m at Garnet Point (mean 1.0 m). The last two sites were
high-flow areas, while Bar Island had lower tidal currents. Our measurements
of kelp lengths (Fig. 4) were similar to those of Vadas et al. (2004a),
except that shorter fronds occurred in areas of very high currents. For example,
fronds of S. longicruris at West Quoddy Head—a highly turbulent
exposed open coastal area—were about 0.8 m long, while those at Wilbur
Neck—an area of intermediate tidal currents upstream from Reversing Falls
(Fig. 1)—were 4.6 m. Similar patterns were noted for L. digitata and S. dermatodea,
with smaller plants at West Quoddy Head and the largest at Wilbur
Neck. Giant kelps in Cobscook Bay appear to be associated with areas of
intermediate water motion, which reduces physical damage but provides
ample nutrients. The damping of wave action within Cobscook Bay, the upward
expansion of kelps due to macrotidal patterns, frequent fog conditions,
and cooler temperatures may result in slower growth (cf. Vadas et al 2004a)
yet allow for larger plants. Temperature races of North Atlantic S. latissima
(as L. saccharina) and S. longicruris (as L. longicruris) reported by Egan
et al. (1990) support the concept that temperature could play a role in plant
size in Cobscook Bay. A similar pattern of large-sized invertebrates has been
described by Larsen (2004b) for Cobscook Bay.
Saccharina latissima found in strong tidal currents at Reversing Falls
were elongate and narrow (Fig. 5e–g) and very similar to the unique
narrow-bladed S. latissima f. angustissima known only from mid-coastal
Maine (Fig. 5h–i). Collins (1880) initially recorded the latter plants
from exposed habitats on Fox Island (Phippsburg) and Bailey Island
(Harpswell), which are located on the easternmost limits of Casco Bay and
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 21
the contiguous “indented” mid-coastal area of Maine (Kelley 1987, Kelley
et al. 1989), respectively. Collins (1880) initially described the narrowbladed
kelp as Laminaria longipes Bory or L. agardhii Kjellman, but later
(Collins 1906, 1911) considered it to be L. agardhii f. angustissima, which
is currently designated as S. latissima f. angustissima (Lane et al. 2006,
Mathieson et al. 2008a). We are aware of only five earlier accounts of this
unique narrow-bladed kelp (Collins 1895–1919, Colt 1999, Mathieson et
al. 2008a, Sears 2002, Taylor 1962), which is thought to be endemic and
restricted to mid-coastal Maine. Molecular (Lane et al. 2006) as well as
detailed anatomical, morphological, ecological, and transplant studies
are obviously needed to confirm if these Reversing Falls and mid-coastal
Maine (e.g., Bald Head, Phippsburg) specimens are genetically similar or
simply phenotypic variants due to extremes of environmental conditions
(Kain 1979; Sundene 1964; Wilce 1959, 1960, 1964). For example, Sundene
(1964) conducted reciprocal transplant experiments of two different
“forms” of L. digitata (Hudson) Lamouroux within the Oslofjord (i.e., L.
stenophylla (Kützing) J. Agardh with numerous long narrow blades, and
L. digitata f. typica with broader, shorter and less divided blades), finding
that they were environmentally induced. Similar phenotypic plasticity after
transplantation has also been described for several other kelp taxa (Kain
1979, Norton 1969, Svendsen and Kain 1971, Yamada 1974). The only other
kelp that approximates Saccharina latissima f. angustissima from Casco
Bay or the narrow, elongated Reversing Falls specimens is S. angustata
(Kjellman) C.E. Lane, C. Mayes, L. D. Druehl & G. E. Saunders subsp.
sibirica Petrov & Suchovejeva. All three taxa have similar habitat ecology,
growing near low water on rocks exposed to strong water motion.
Some unique dwarf populations of Fucus were also found at Reversing
Falls (Mathieson et al. 2006); these limicolous or embedded populations
grow in the high intertidal sandy salt marshes in Cobscook Bay (Larsen
1979) and at a few other scattered northwest Atlantic sites ranging from
Nova Scotia to Long Island Sound (Kucera and Saunders 2008, Mathieson
and Dawes 2001, Mathieson et al. 2006). Dwarf plants, which we
designated as Fucus sp. “muscoides-like” (Mathieson and Dawes 2001,
Mathieson et al. 2006), produce moss-like turfs that are dichotomously or
irregularly branched and about 17.4 mm long. These dwarf Fucus plants
may be hybrids of F. vesiculosus and F. spiralis, as shown for populations
from salt marshes in southern Maine (Wallace et al. 2004), Nova Scotia,
and New Brunswick (Kucera and Saunders 2008). Three other detached/
entangled salt marsh fucoid ecads (Mathieson and Dawes 2001, Norton and
Mathieson 1983) were also found with these dwarf populations, including
Ascophyllum nodosum ecad scorpioides, F. spiralis ecad lutarius, and F.
vesiculosus ecad volubilis.
Conclusions
In conclusion, Cobscook Bay has a limited benthic marine algal flora that
has a large vertical distribution and is impacted by strong tidal currents. The
22 Northeastern Naturalist Vol. 16, Monograph No. 5
148 taxa of seaweeds from Cobscook Bay, ME represent 47% of the total
flora (316 taxa) known from the four embayments ranging from the Bay of
Fundy to Casco Bay; it also represents 37% of the 403 taxa recorded from
the northeast coast of North America (Sears 2002). Cobscook Bay’s coldtemperate
seaweed flora is most similar to that of Casco Bay (74.3%) and
Passamaquoddy Bay (68.8%) and least similar to that of the Bay of Fundy
(63.8%). The lower diversity of Cobscook Bay’s flora (148 taxa) versus the
Bay of Fundy (250 taxa) may reflect it’s smaller area compared to the Bay
of Fundy; this pattern contrasts with the very high numbers of invertebrates
(>800 taxa) known from Cobscook Bay. The six introduced seaweeds recorded
from Cobscook Bay are less than those reported for Casco Bay (8 taxa).
The large vertical distribution of intertidal and subtidal benthic algae and the
large stature of kelps within Cobscook Bay parallels that reported for the Bay
of Fundy; it is probably due to a combination of large tidal amplitudes, frequent
cool, overcast to foggy weather, and tidal upwelling. The very elongate
and narrow blades of some kelp species growing at Reversing Falls State Park
are similar to the narrow-bladed Saccharina latissima f. angustissima known
only from high wave-impacted sites in mid-coastal Maine.
Acknowledgments
We thank several former undergraduate and graduate students at the University of
New Hampshire (UNH) for their help with various field studies in Cobscook Bay. Our
studies were supported by funds from Maine-New Hampshire Sea Grant (NOAA), the
New Hampshire Agricultural Experiment Station, and the Leslie Hubbard Marine Endowment
Fund. This paper is published as Scientific Contribution Number 2345 from
the New Hampshire Agriculture Experiment Station, as well as Contribution Number
452 from UNH Jackson Estuarine Laboratory and Center for Marine Biology. The first
two authors acknowledge the encouragement and help of their wives, Myla Mathieson
and Kathy Dawes, who were involved with several of the field studies.
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2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 31
Appendix 1. A comparison of the marine benthic algae from the Bay of Fundy (BF), Passamaquoddy
Bay (PB), Cobscook Bay (COB), and Casco Bay (CAB). Superscripts: 1 =
life-history phase or stage, 2 = introduced species. Question mark (?) refers to a questionable
taxonomic or geographic record.
Species BF PB COB CAB
Chlorophyceae
Acrochaete flustrae (Reinke) O’Kelly in Gabrielson et al. X X X X
Acrochaete ramosa (N.L. Gardner) O’Kelly X
Acrochaete repens N. Pringsheim X X
Acrochaete viridis (Reinke) R. Nielsen X X
Acrochaete wittrockii (Wille) R. Nielsen X X X X
Acrosiphonia arcta (Dillwyn) Gain X X X X
Acrosiphonia sonderi (Kützing) Kornmann X
Acrosiphonia spinescens (Kützing) Kjellman X X X X
Blidingia chadefaudii (J. Feldmann) Bliding X
Blidingia marginata (J. Agardh) P. Dangeard ex Bliding X X
Blidingia minima (Nägeli ex Kützing) Kylin X X X X
Blidingia ramifera (Bliding) Garbary et Barkhouse X
Bolbocoleon piliferum N. Pringsheim X X X
Bryopsis hypnoides J.V. Lamouroux X X X
Bryopsis plumosa (Hudson) C. Agardh X X
Capsosiphon fulvescens (C. Agardh) Setchell et Gardner X X X
Capsosiphon groenlandicus (J. Agardh) K.L. Vinogradova X X X
Chaetomorpha aerea (Dillwyn) Kützing X X X
Chaetomorpha brachygona Harvey X X
Chaetomorpha ligustica (Kützing) Kützing X X X X
Chaetomorpha linum (O.F. Müller) Kützing X X X X
Chaetomorpha melagonium (F. Weber et D. Mohr) Kützing X X X X
Chaetomorpha picquotiana Montagne ex Kützing X X X X
Chlorochytrium cohnii E.P. Wright ?X X
Chlorochytrium dermatocolax Reinke X
1“Chlorochytrium inclusum Kjellman” = phase of Acrosiphonia and X X
Spongomorpha
Chlorochytrium schmitzii Rosenvinge X
Chlorococcum endozoicum F.S. Collins X
Cladophora albida (Nees) Kützing X X X
Cladophora crystallina (Roth) Kützing X
Cladophora laetevirens (Dillwyn) Kützing X
Cladophora liniformis Kützing X
Cladophora ruchingeri (C. Agardh) Kützing X X
Cladophora rupestris (L.) Kützing X X X
Cladophora sericea (Hudson) Kützing X X X X
1“Codiolum gregarium A. Braun” = phase of Urospora X
penicilliformis
1“Codiolum petrocelidis Kuckuck” = phase of Acrosiphonia and X X X
Spongomorpha
1“Codiolum pusillum (Lyngbye) Kjellman” = phase of Urospora X X X
2 Codium fragile (Suringar) Hariot subsp. fragile X X
Derbesia marina (Lyngbye) Solier X X
1“Halicystis ovalis (Lyngbye) J. Areschoug” = phase of D. marina X
Epicladia perforans (Huber) R. Nielsen X
Eugomontia sacculata Kornmann X X
Gayralia oxysperma (Kützing) K.L. Vinogradova ex Scagel et al. X X X
Gloeocystis scopulorum Hansgirg X
Gomontia polyrhiza (Lagerheim) Bornet et Flahault X X X X
32 Northeastern Naturalist Vol. 16, Monograph No. 5
Species BF PB COB CAB
Halochlorococcum moorei (N.L. Gardner) Kornmann et Sahling ?X
Kornmannia leptoderma (Kjellman) Bliding X X X
Monostroma grevillei (Thuret) Wittrock X X X X
Ochlochaete hystrix Thwaites ex Harvey var. ferox (Huber) R. X X
Nielsen
Palmellococcus marinus F.S. Collins X
Percursaria percursa (C. Agardh) Rosenvinge X X X X
Prasinocladus lubricus Kucuck X
Prasiola stipitata Suhr ex Jessen X X X X
Pringshemiella scutata (Reinke) Marchewianka X X X X
Protomonostroma undulatum (Wittrock) K.L. Vinogradova f. X X X
pulchrum (Farlow) M.J. Wynne
Pseudendoclonium fucicola (Rosenvinge) R. Nielsen X
Pseudendoclonium submarinum Wille X X X X
Pseudopringsheimia confluens (Rosenvinge) Wille X
Rhizoclonium riparium (Roth) Kützing ex Harvey X X X X
Rosenvingiella polyrhiza (Rosenvinge) P.C. Silva X
Spongomorpha aeruginosa (L.) van den Hoek X X X X
Stichococcus marinus (Wille) Hazen X X
Tellamia contorta Batters X
Ulothrix flacca (Dillwyn) Thuret in Le Jolis X X X X
Ulothrix laetevirens (Kützing) F.S. Collins X X
Ulothrix speciosa (Carmichael ex Harvey in W.J. Hooker) Kützing X X X X
Ulva clathrata (Roth) C. Agardh X X X X
Ulva compressa L. X X X X
Ulva flexuosa Wulfen X X
Ulva flexuosa subsp. paradoxa (C. Agardh) M.J. Wynne X X X X
Ulva intestinalis L X X X X
Ulva lactuca L. X X X X
Ulva linza L. X X X X
Ulva prolifera O.F. Müller X X X X
Ulva rigida C. Agardh X X
Ulva torta (Mertens) Trevisan X X
Ulvaria obscura (Kützing) P. Gayral et Bliding X X X X
Uronema ?curvata Printz X
Urospora penicilliformis (Roth) J.E. Aeschoug X X X X
Urospora wormskjoldii (Mertens ex Hornemann) Rosenvinge X X X
Total Chlorophyceae: 81 62 58 38 54
Phaeophyceae
Acrothrix gracilis Kylin X
Agarum clathratum Dumortier X X X X
Alaria esculenta (L.) Greville X X X X
Ascophyllum nodosum (L.) Le Jolis X X X X
Ascophyllum nodosum ecad scorpioides (Hornemann) Reinke X X X
Asperococcus fistulosus (Hudson) W.J. Hooker X X X X
1“Chilionema reptans (P.L. Crouan et H.M. Crouan) Sauvageau” ?X
= phase of A. fistulosus
Chorda filum (L.) Stackhouse X X X X
Chordaria flagelliformis (O.F. Müller) C. Agardh X X X X
1“Streblonema chordariae (Farlow) De Toni” = phase of C. X X
flagelliformis
Cladosiphon zosterae (J. Agardh) Kylin X X
Coilodesme bulligera Strömfelt X
2Colpomenia peregina Sauvageau X
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 33
Species BF PB COB CAB
Delamarea attenuata (Kjellman) Rosenvinge X X
Desmarestia aculeata (L.) J.V. Lamouroux X X X X
Desmarestia viridis (O.F. Müller) J.V. Lamouroux X X X X
Dictyosiphon chordaria J.E. Areschoug X X
Dictyosiphon ekmanii J.E. Areschoug X X
Dictyosiphon foeniculaceus (Hudson) Greville X X X X
Dictyosiphon macounii Farlow X X
Ectocarpus fasciculatus Harvey X X X X
Ectocarpus siliculosus (Dillwyn) Lyngbye X X X X
Elachista chondrii J.E. Areschoug X
Elachista fucicola (Velley) J.E. Areschoug X X X X
Entonema alariae Jaasund X
Entonema polycladum (Jaassund) Jaasund X
Eudesme virescens (Carmichael ex Berkeley in Hooker) J. Agardh X X X
Fucus distichus L. emend Powell subsp anceps (Harvey et X
Ward ex Caruthers) Powell
Fucus distichus subsp. distichus Powell X X X X
Fucus distichus subsp. edentatus (Bachelot de la Pylaie) Powell X X X
Fucus distichus subsp. edentatus f. abbreviatus (Gardner) X
Hollenberg et Abbott
Fucus distichus subsp. evanescens (C. Agardh) Powell X X X X
2Fucus serratus L. X
Fucus spiralis L. X X X X
Fucus spiralis ecad lutarius (Kützing) Sauvageau X X
Fucus sp. “muscoides-like” X X
Fucus vesiculosus L. X X X X
Fucus vesiculosus f. mytilii (Nienburg) Nienhuis X
Fucus vesiculosus ecad volubilis (Hudson) Turner X X
Halosiphon tomentosus (Lyngbye) Jaasund X X X X
Halothrix lumbricalis (Kützing) Reinke X
Haplospora globosa Kjellman X X
Halothrix lumbricalis (Kützing) Reinke X
1“Hecatonema terminale (Kützing) Kylin” = a possible phase of X X
several brown algae
Hincksia granulosa (J.E. Smith) P. C. Silva in Silva, Meñez et Moe X X X
Hincksia ovata (Kjellman) P. C. Silva in Silva, Meñez et Moe X X
Hummia onusta (Kützing) J. Fiore X
Isthmoplea sphaerophora (Carmichael) Kjellman X X
Laminaria digitata (Hudson) J.V. Lamouroux X X X X
Laminariocolax aecidioides (Rosenvinge) A.F. Peters in Burkhardt X X
et Peters
Laminariocolax tomentosoides (Farlow) Kylin X X
Leathesia marina (Lyngbye) Decaise X X X
Leptonematella fasciculata (Reinke) P.C. Silva X X
Litosiphon laminariae (Lyngbye) Harvey X X X
2Melanosiphon intestinalis (D.A. Saunders) M.J. Wynne X X X X
Microspongium globosum Reinke = possible phases of Petalonia X
fascia and Scytosiphon lomentaria
Microspongium immersum (Levring) P.M. Pedersen X
Mikrosyphar polysiphoniae Kuckuck X
Mikrosyphar porphyrae Kuckuck X
Myriocladia lovenii J. Agardh X
Myrionema corunnae Sauvageau X X X
Myrionema magnusii (Sauvageau) Loiseaux X X
Myrionema strangulans Greville X X X X
Myriotrichia clavaeformis Harvey X
34 Northeastern Naturalist Vol. 16, Monograph No. 5
Species BF PB COB CAB
Petalonia fascia (O.F. Müller) Kuntze X X X X
Petalonia zosterifolia (Reinke) Kuntze X
Petroderma maculiforme (Wollny) Kuckuck X X X
?Pilinia lunatiae F.S. Collins X
Pilinia rimosa Kutzing X
Pogotrichum filiformis Reinke X X
Pseudolithoderma extensum (P.L. Crouan et H.M. Crouan) S. Lund X X X X
Punctaria crispata (Kützing) Batters X
Punctaria latifolia Greville X X X
Punctaria plantaginea (Roth) Greville X X X
Punctaria tenuissima (C. Agardh) Greville X X X
1“Streblonema effusum Kylin” = phase of P. tenuissima ?X X
Pylaiella littoralis (L.) Kjellman X X X X
1“Ralfsia bornetii Kuckuck” = phase of Petalonia and Scytosiphon X X
1“Ralfsia clavata (Harvey in W.J. Hooker) P.L. Crouan et H.M. X X X
Crouan” = phase of Petalonia & Scytosiphon
Ralfsia fungiformis (Gunnerus) Setchell et Gardner X X X
Ralfsia pusilla (Strömfelt) Batters X X
Ralfsia verrucosa (J.E. Areschoug) J.E. Areschough X X X X
Saccharina latissima (L.) Lane et al. X X X X
Saccharina latissima f. angustissima (F.S. Collins) Mathieson in X
Mathieson et al.
Saccharina longicruris (Bachelot de la Pylaie) Kuntze X X X X
Saccorhiza dermatodea (Bachelot de la Pylaie) J.E. Areschoug X X X X
Scytosiphon complanatus (Rosenvinge) Doty X
Scytosiphon dotyi M.J. Wynne X X
Scytosiphon lomentaria (Lyngbye) Link X X X X
Sorapion kjellmanii (Wille) Rosenvinge X X
Sphacelaria arctica Harvey X
Sphacelaria caespitula Lyngbye X
Sphacelaria cirrosa (Roth) C. Agardh X X X X
Sphacelaria fusca (Hudson) S.F. Gray X X
Sphacelaria nana Nägeli ex Kützing X X
Sphacelaria plumosa Lyngbye X
Sphacelaria radicans (Dillwyn) C. Agardh X X X X
Sphacelaria rigidula Kützing X X
Sphaerotrichia divaricata (C. Agardh) Kylin X X X
Spongonema tomentosum (Hudson) Kützing X X X X
Stictyosiphon griffithsianus (Le Jolis) Holmes et Batters X X X X
Stictyosiphon soriferus (Reinke) Rosenvinge X
Stictyosiphon tortilis (Ruprecht) Reinke X
Stilophora tenella (Esper) P.C. Silva in Silva, Basson et Moe X
Streblonema infestans (H. Gran) Batters X X
Streblonema fasciculatum Thuret in Le Jolis X
Streblonema parasiticum (Sauvageau) De Toni X X
Striaria attenuata (Greville) Greville X
Stypocaulon scoparium (L.) Kützing X
Tilopteris mertensii (Turner in J.E. Smith) Kützing X
2Ulonema rhizophorum Foslie ?X X X
Total Phaeophyceae: 111 89 59 46 70
Rhodophyceae
Acrochaetium alariae (Jónsson) Bornet X X X
Acrochaetium collopodum (Rosenvinge) G. Hamel X
Acrochaetium savianum (Meneghini) Nägeli X X
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 35
Species BF PB COB CAB
Acrochaetium secundatum (Lyngbye) Nägeli X X X X
Aglaothamnion halliae (F.S. Collins) N.E. Aponte, D.L. Ballantine X
et J.N. Norris
Agardhiella subulata (C. Agardh) Kraft et M.J. Wynne ?X
Ahnfeltia plicata (Hudson) Fries X X X X
1“Porphyrodiscus simulans Batters” = phase of A. plicata X
Antithamnion cruciatum (C. Agardh) Nägeli X
Antithamnionella floccosa (O.F. Müller) Whittick X X X X
1“Audouinella polyidis (Rosenvinge) Woelkerling et Womersley” ?X X
= phase of Helminthora divaricata (C. Agardh) J. Agardh
Bangia fuscopurpurea (Dillwyn) Lyngbye X X X X
2Bonnemaisonia hamifera Hariot X X
1“Trailliella intricata Batters” = phase of B. hamifera X X X
Callithamnion corymbosum (J.E. Smith) Lyngbye X X
Callithamnion tetragonum (Withering) S.F. Gray X X X
Callocolax neglectus F. Schmitz ex Batters X
Ceramium cimbricum H.E. Petersen in Rosenvinge X
Ceramium deslongchampsii Chauvin ex Duby X X X X
Ceramium diaphanum (Lightfoot) Roth var. elegans (Roth) Roth X X
Ceramium virgatum Roth X X X X
Ceratocolax hartzii Rosenvinge X X
Champia parvula (C. Agardh) Harvey X
Chondria baileyana (Montagne) Harvey X
Chondrus crispus Stackhouse X X X X
Choreocolax polysiphoniae Reinsch X X X X
Choreocolax rabenhorstii Reinsch ?X
Chroodactylon ornatum (C. Agardh) Basson X
Clathromorphum circumscriptum (Strömfelt) Foslie X X X X
Clathromorphum compactum (Kjellman) Foslie X X X
Colaconema dasyae (F.S. Collins) Stegenga et al. X
Colaconema daviesii (Dillwyn) Stegenga X X X X
Colaconema membranaceum (Magnus) Woelkerling X X X X
1“Conchocelis rosea Batters” = phase of Porphyra and Bangia X X
Corallina officinalis L. X X X X
Cystoclonium purpureum (Hudson) Batters X X X X
Dasya baillouviana (S.G. Gmelin) Montagne in Barker-Webb & X
Berthelot
Devaleraea ramentacea (L.) Guiry X X X X
2Dumontia contorta (S.G. Gmelin) Ruprecht X X X X
Erythrodermis traillii (Holmes ex Batters) Guiry et Garbary X X X
Erythrotrichia carnea (Dillwyn) J. Agardh X X X X
Euthora cristata (Turner) J. Agardh X X X X
Fimbrifolium dichotomum (Lepechin) G.I. Hansen X X X X
Gelidium crinale (Turner) Lamouroux ?X
Gloiosiphonia capillaris (Hudson) Carmichael in Berkeley X X X
1“Cruoriopsis ensis C.-C. Jao” = phase of G. capillaris X X
Gracilaria tikvahiae McLachlan ?X X
Gymnogongrus crenulatus (Turner) J. Agardh X X
Haemescharia hennedyi (Harvey) K. Vinogradova et Yacovleva ?X
Halosacciocolax kjellmani S. Lund X X X
Harveyella mirabilis (Reinsch) F. Schmitz et Reinke in Reinke X X X
Hildenbrandia crouaniorum J. Agardh X
Hildenbrandia rubra (Sommerfelt) Meneghini X X X X
Hydrolithon farinosum (J.V. Lamouroux) Penrose et Y.M. X X
Chamberlain
36 Northeastern Naturalist Vol. 16, Monograph No. 5
Species BF PB COB CAB
Kvaleya epilaeve W.H. Adey et Speradini X
Leptophytum foecundum (Kjellman) W.H. Adey X
Leptophytum laeve (Strömfelt) W.H. Adey X X X
Lithophyllum orbiculatum (Foslie) Foslie X
Lithothamnion glaciale Kjellman X X X X
Lithothamnion lemoineae W.H. Adey X
Lithothamnion ungeri Kjellman X
2Lomentaria clavellosa (Turner) Gaillon X
2Lomentaria orcadensis (Harvey) F.S. Collins ex W.R. Taylor X X
Mastocarpus stellatus (Stackhouse in Withering) Guiry in Guiry X X X X
et al.
1“Petrocelis cruenta J. Agardh” = phase of M. stellatus X X X X
Meiodiscus spetsbergensis (Kjellman) G.W. Saunders et McLachlan X X
Melobesia membranacea (Esper) J.V. Lamouroux X
Membranoptera spinulosa (Ruprecht) Kuntze X X X X
2Neosiphonia harveyi (J. Bailey) M.-S. Kim et al. X X X X
Odonthalia dentata (L.) Lyngbye X
Palmaria palmata (L.) Kuntze X X X X
Pantoneura fabriciana (Lyngbye) M.J. Wynne X X
Peyssonnelia rosenvingei F. Schmitz in Rosenvinge X X X X
Phycodrys fimbriata (Bachelot De La Pylaie ex J. Agardh) Kylin X X X
Phyllophora crispa (Hudson) P. Dixon X
Phyllophora pseudoceranoides (S.G. Gmelin) Newroth et A.R.A. X X X X
Taylor
Phyllophora truncata (Pallas) A.D. Zinova X X X
Phymatolithon laevigatum (Foslie) Foslie X X X X
Phymatolithon lamii (Lemoine) Y.M. Chamberlain X X X
Phymatolithon lenormandii (J.E. Areschoug in J. Agardh) W.H. Adey X X X X
Plagiospora gracilis Kuckuck X
Tsengia bairdii (Farlow) K.C. Fan & Y.P. Fan X X
Plumaria plumosa (Hudson) Kuntze X X X X
Pneophyllum fragile Kützing X X
Polyides rotundus (Hudson) Gaillon X X X X
Polysiphonia arctica J. Agardh X
Polysiphonia brodiei (Dillwyn) Sprengel X
Polysiphonia denudata Dillwyn) Greville ex Harvey X X
Polysiphonia elongata (Hudson) Sprengel X X
Polysiphonia fibrillosa (Dillwyn) Sprengel X X X
Polysiphonia flexicaulis (Harvey) F.S. Collins X X X X
Polysiphonia fucoides (Hudson) Greville X X X X
Polysiphonia lanosa (L.) Tandy X X X X
Polysiphonia nigra (Hudson) Batters X X
Polysiphonia stricta (Dillwyn) Greville X X X X
Porphyra amplissima (Kjellman) Setchell et Hus ex Hus X
Porphyra birdiae Neefus et Mathieson X X
Porphyra leucosticta Thuret in Le Jolis X X X X
Porphyra linearis Greville X X
Porphyra miniata (C. Agardh) C. Agardh X X X X
Porphyra purpurea (Roth) C. Agardh X X X X
Porphyra umbilicalis Kützing X X X X
2Porphyra yezoensis Ueda f. yezoensis Ueda X
2Porphyra yezoensis f. narawaensis Miura (strains U51 and H25) X
Porphyropsis coccinea (J. Agardh ex J.E. Areschoug) Rosenvinge X
Porphyrostromium ciliare (Carmichael ex Harvey in W.J. Hooker) X X X X
M.J. Wynne
Pterothamnion plumula (J. Ellis) Nägeli in Nägeli & Cramer X
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 37
Species BF PB COB CAB
Ptilota serrata Kützing X X X X
Rhodochorton purpureum (Lightfoot) Rosenvinge X X X X
Rhodomela confervoides (Hudson) P. C. Silva X X X X
Rhodomela lycopodioides (L.) C. Agardh ?X
Rhodophysema elegans (P.L. Crouan et H.M. Crouan ex J. Agardh) X X X X
P.S. Dixon
Rhodophysema georgii Batters X X
Scagelia americana (Harvey) Athanasiadis X X
Scagelia pylaisaei (Montagne) M. J. Wynne X X X X
Scinaia furcellata (Turner) J. Agardh X
Spermothamnion repens (Dillwyn) Rosenvinge X
Spyridia filamentosa (Wulfen) Harvey in W.J. Hooker X X
Stylonema alsidii (Zanardini) K.M. Drew X X
Titanoderma corallinae (P.L. Crouan et H.M. Crouan) Woelkerling, X X
Y.M. Chamberlain et P. C. Silva
Titanoderma pustulatum (J.V. Lamouroux) Nägeli in Nägeli et X X X X
Cramer
Turnerella pennyi (Harvey) F. Schmitz X X
1“Cruoria arctica Schmitz” = phase of T. pennyi X
Total Rhodophyceae: 124 99 75 64 77
Grand Total: 316 250 192 148 201
38 Northeastern Naturalist Vol. 16, Monograph No. 5
Appendix 2. Thirty-seven seaweed collection sites in Cobscook Bay, including collection
dates, and number of voucher specimens [in brackets].
Site
# Description
(1) Eastport Harbor (44°54.0'N, 66°59'W): 10/1875, 10/7/1995, 6/30/1998,
8/7/2005, [39]
(2) Eastport Harbor (44°54.5'N, 66°59'W): 10/1872, 10/8/1994, 10/7/1995–
10/9/1995, 3/11/1996, 11/14/1999, 8/6/2005, [58]
(3) Treat Island, Eastport (44°52.5'N, 66°59.5'W): 10/7/1995, [2]
(4) Estes Head, Cargo terminal at Eastport (44°53.5'N, 66°59.8'W): 8/8/2005,
[31]
(5) Harris Point (44°55'N, 67°00'W): 8/8/2005, [25]
(6) Deep Cove, Eastport (44°53.5'N, 66°59.5'W): 9/21/1997, 8/20/1998, [27]
(7) Mathews Island, Eastport, (44°55'N, 67°01.5'W): 5/21/1996, 9/11/1996,
9/21/1997, 2/26/1998, 4/10/1998. 5/21/1998, 6/30/1998 8/21/1998,
9/24/1998, 10/31/1998, 11/30/1998, 2/25/1999, 4/1/1999, 6/15/1999, [233]
(8) Goose Island, on aquaculture nets, Eastport (44°55.5'N, 67°02.5'W): 7/8/1995 ,
[1]
(9) Causeway off Rt. 190, Eastport (44°56.5'N, 67°02'W): 3/11/1996, [29]
(10) Carlow Island at cove, Eastport (44°56.5'N, 67°02'W): 7/7/1995, [22]
(11) Carrying Place Cove, Eastport (44°55.5'N, 67°01.5'W): 8/21/1998, 8/8/2005,
[13]
(12) Prince Cove Salmon Farm, Eastport (44°55'N, 67°02.5'W): 8/21/1998, [18]
(13) Lubec Neck at International Bridge (44°51.5'N, 66°59'W): 10/7/1995,
2/23/1996, 6/13/1997, [28]
(14) Town Landing, Lubec (44°51.5'N, 66°59.5'W): 2/23/1996, [1]
(15) Pirates Creek, Lubec (44°51.5'N, 67°01.5'W): 5/21/1966, 7/21/1966, [42]
(16) Seward Neck, Lubec (44°53'N, 67°01'W): 3/11/1996, [23]
(17) Comstock Point, North Lubec (44°53'N, 67°01'W): 8/8/2005, [18]
(18) Gove Point, North Lubec (44°54'N, 67°03.8'W): 6/7/1995, 7/8/1995, 7/1/1998,
8/21/1998, [69]
(19) Huckins Ledge, Lubec (44°52.5'N, 67°03): 9/21/1997, 11/21/1997, 2/26/1998,
4/10/1998, 5/21/1998, 6/29/1998, 8/21/1998, 9/24/1998, 10/30/1998,
11/30/1998, 2/25/1999, 4/1/1999, 6/15/1999, [151]
(20) Lead Mine Road, Lubec (44°49'N, 67°03.5'W): 3/24/1996, [20]
(21) Federal Harbor, West Lubec (44°51'N, 67°04.0'W): 3/11/1996, [2]
(22) Offshore urchin farm, West Lubec (44°51'N, 67°04.1'W): 8/8/2005, [14]
(23) Pennamaquan River mouth, Pembroke (44°55.3'N, 67°06.5'W): 8/19/1994,
4/19/1996, [20]
(24) Pennamaquan River off Hersey Neck Rd., Pembroke (44°55.5'N, 67°07.5'W):
4/19/1996, 8/6/ 2005, [10]
(25) Pennamaquan River, opposite Hersey Pt., Pembroke (44°56.5'N, 67°10'W):
8/19/1994, [1]
(26) Pennamaquan River, @ Hersey Pt., Pembroke (44°56'N, 67°09'W): 8/19/1994,
4/19/1996, [2]
(27) Pennamaquan River, @ headwaters, Pembroke (44°57.6'N, 67°10'W):
8/19/1994, [5]
(28) Reversing Falls Park, opposite Falls Island, Pembroke (44°53'N, 67°08'W):
8/19/1994, 10/8/1994, 7/9/1995, 10/7/1995, 5/16/1996, 10/26/1996,
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 39
2/16/1998, 5/20/1998, 5/21/1998, 6/30/1998, 10/30/1998, 10/31/1999,
8/2/2000, 4/12/2003, [313].
(29) Crows Neck, Trescott (44°52.5'N, 67°08'W): 5/21/66, 5/22/66, 5/16/1996,
9/1/1971, 7/2/1977, 5/16/1996, 6/16/1999, 10/31/1999 [65]
(30) Wilbur Neck, Dennys Bay, Pembroke (44°54'N, 67°09.5'W): 10/9/1994,
10/7/1995, 10/8/1995, 10/9/1995, 3/11/1996, 10/26/1996, [226]
(31) Dennys River @Rt. 1 bridge, Dennysville (44°54.5'N, 67°13.3'W): 8/20/1994,
4/19/1996, [3]
(32) Dennys River, Dennysville Station (44°54.6'N, 67°12.5'W): 8/20/1994, [7]
(33) Dennysville Station @ Rt. 86 (44°54.7'N, 67°13.3'W): 8/20/1994, [3]
(34) Cove opposite William Island on Rt. 1, Edmunds (44°52.5'N, 67°10'W):
3/24/1996, [9]
(35) Friedman Marine Laboratory, Edmunds (44°49.5'N, 67°09.5'W): 7/7/1995,
3/24/1996, [58]
(36) Cobscook Bay State Park, Edmunds (44°49'N, 67°09.5'W): 3/11/1996, [18]
(37) Leighton Cove, along Rt. 1, Whiting (44°48'N, 67°05'W): 4/17/1995,
3/24/1996
40 Northeastern Naturalist Vol. 16, Monograph No. 5
Appendix 3. An annotated Checklist of the seaweeds in Cobscook Bay including
“name”, sites (see Appendix 2), partners in heteromorphic life histories, and percent
occurrence (%) All accession numbers lacking a collector name are Mathieson, while
all collections lacking a herbarium listing are NHA. * signifies introduced taxa.
Chlorophyceae
Acrochaete flustrae: Site 30, 10/9/1994, leg. Mathieson & Hehre (#52729); Site 37,
3/24/1996, leg. (#62863), 5.4%.
Acrochaete wittrockii: Site 37, 4/17/1995, leg. (#56958), 2.7%.
Acrosiphonia arcta: Site 1, 8/1872, leg. Eaton (Eaton 1873), as Cladophora arcta
Dillwyn; Site 29, 7/9/1995, leg. Hehre (#58557); Site 35, 7/7/1995, leg. Hehre
(#58358), 8.1%.
Acrosiphonia spinescens: Site 1, 8/7/2005, leg. (#78377); Site 4, 8/8/2005, leg.
(#78325, 78339); Site 5, 8/8/2005, leg. (#78356); Site 12, 8/21/1998, leg. Mathieson
& Yarish (#67389); Site 17, 8/8/2005, leg. (#78461); Site 22, 8/8/2005,
leg. (#78441); Site 24, 8/6/2005, leg. (#78421); Site 30, 10/9/1994, leg. Mathieson
& Hehre (#53279), 21.6%.
Blidingia minima: Site 1, 3/11/1996, leg. (#63663) and 8/7/2005, leg. (#78383); Site
4, 8/8/2005, leg. (#78329); Site 5, 8/8/2005, leg. (#78365); Site 25, 8/19/1994,
leg. (#51432); Site 33, 8/20/l994, leg. (#51617), 16.2%.
Bryopsis plumosa: Site 28, 10/8/1994. leg. Mathieson & Hehre (#53095), 2.7%.
Chaetomorpha ligustica: Site 1, 8/1872, leg. Easton (Eaton, 1873), as Chaetomorpha
tortuosa Dillwyn; all others as Rhizoclonium tortuosum (Dillwyn) Kützing,
Site 1, 10/8/1994, leg. Harris (#53256); Site 1, 8/7/2005, leg. (#78381); Site 4,
8/8/2005, leg. (#78331); Site 5, 8/8/2005, leg. (#78349, 78370); Site 17, 8/8/2005,
leg. (#78455, 78460); Site 30, 10/9/1994, leg. Mathieson & Hehre (#52730),
13.5%.
Chaetomorpha linum: Site 13, 6/13/1997, leg. Jones (#64580); Site 28, 10/8/1994,
leg. Mathieson & Hehre (#53096), 5.4%.
Chaetomorpha melagonium: Site 1, 8/1872, leg. Eaton (Eaton, 1873); Site 2,
8/6/2005, leg. Harris (#78397, 78401); Site 30, 10/9/1994, leg. Mathieson &
Hehre (#52724), 8.1%.
Chaetomorpha picquotiana: Site 7, 9/21/1997, leg. Mathieson & Wilkes (#12636);
Site 30, 10/9/1994, leg. Mathieson & Hehre (#52726), 5.4%.
“Chlorochytrium inclusum:” Site 30, 10/9/1994, leg. Mathieson & Hehre (#52943),
2.7%.
Cladophora ruchingeri: Site 30, 10/8/1995, leg. Harrington (#59830), 2.7%.
Cladophora sericea: Site 1, 8/1872, leg. Eaton (Eaton, 1873), as Cladophora flexuosa
Griffiths; Site 1, 8/8/2005, leg. (#78387); Site 10, 7/7/1995, leg. Hehre (#58496);
Site 15, 5/21/1966, leg. Hehre, as Rhizoclonium tortuosum (#3684); Site 24,
8/19/1994, leg. (#51274); Site 35, 7/7/1995, leg. Hehre (#58348), 16.2%.
“Codiolum petrocelidis:” Site 1, 9/1877, leg. Farlow (FH); Site 30, 10/9/1994, leg.
Mathieson & Hehre (#53286), 5.4%.
“Codiolum pusillum:” Site 1, (Farlow, 1881); Site 30, 10/9/1994, leg. Mathieson &
Hehre (#53283), 5.4%.
*Codium fragile subsp. fragile: Site 30 (drift), 10/8/1995, leg. True & Carpenter
(#63966), 2.7%.
Gayralia oxysperma: Site 1, 8/1872, leg. Eaton, as Monostroma latissima Wittrock
(Eaton, 1873); Site 28, 5/21/1998, leg. Mathieson & Dawes (#67548); Site 33,
8/20/1994, leg. (#51615), as Monostroma oxyspermum (Kützing) Doty, 8.1%.
Gomontia polyrhiza: Site 35, 3/24/1996, leg. [#12781; (rock #2085)], 2.7%.
Monostroma grevillei: Site 10, 7/7/1995, leg. Hehre (#58493); Site 28, 5/16/1996,
leg. (#62086); Site 35, 3/24/1996, leg. (#62644), 8.1%.
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 41
Percursaria percursa: Site 17, 8/8/2005, leg. (#78460); Site 23, 4/19/1996, leg.
(#63548); Site 35, 7/7/1995, leg. Hehre (#58350); Site 37, 4/17/1995, leg.
(#56963), 8.1%.
Prasiola stipitata: Site 6, 8/20/1998, leg. (#67503); Site 28, 8/2/2000, leg. Mathieson
& Dawes (#74885), 5.4%.
Pringsheimiella scutata: Site 35, 3/24/1996, leg. (#69522), 2.7%.
Pseudendoclonium submarinum: Site 7, 2/26/1998, leg. [#68937 (rock #2656)]; Site
14, 2/23/1996, leg. [#12777; (rock #2082)], 5.4%.
Rhizoclonium riparium: Site 1, (Farlow, 1881); Site 1, 8/7/2005, leg. (#78375,
78387); Site 5, 8/8/2005, leg. (#78370); Site 11, 8/8/2005 Aug. 2005, leg. S.
Crawford (#78425); Site 15, 5/21/1966, leg. Hehre, as R. tortuosum (#3683);
Site 24, 8/6/2005, leg. (unnumbered); Site 28, 8/19/1994, leg. (#51427); Site 35,
7/7/1995, leg. Hehre (#58342), 18.9%.
Spongomorpha aeruginosa: Site 30, 10/9/1994, leg. Mathieson & Hehre (#55042,
55043), 2.7%.
Ulothrix flacca: Site 1, 1877, leg. Farlow (FH); Site 15, 5/21/1966, leg. Scribner
(#6708), 5.4%.
Ulothrix speciosa: Site 1, 8/1872, leg. Eaton (Eaton, 1873), as “Hormotrichum
speciosum Carmichael,” Site 9, 3/11/1996, leg. (#61385); Site 23, 4/19/1996, as
Urospora penicilliformis, leg. (#61426); Site 35, 3/24/1996, as Urospora penicilliformis
leg. (#62643), 10.8%.
Ulva clathrata: Site 10, 7/7/1995, leg. Hehre (#58485); Site 15, 5/21/1966, leg.
(#3120); Site 30, 10/8/1995, leg. (#59711); Site 35, 7/7/1995, leg. Hehre
(#58349), 10.8%.
Ulva compressa: Site 1, 8/1872, leg. Eaton, as Enteromorpha compressa (L.) Nees
(YALE); Site 28, 10/7/1995, as E. compressa leg. (#59803); Site 29, 10/31/1999,
as E. intestinalis subsp. compressa (L.) M. W. R. N. de Silva et E. M. Burrows,
leg. (#71305); Site 34, 3/24/1996, as E. compressa, leg. (#62875), 10.8%.
Ulva flexuosa: Site 11, 8/8/ 2005, leg. (#78424, 78425), 2.7%.
Ulva flexuosa subsp. paradoxa: Site 15, 5/21/1966, as Rhizoclonium tortuosum leg.
Hehre (#3687); Site 26, 4/19/1996, leg. (#62282), 5.4%.
Ulva intestinalis: Site 1, 8/8/2005, leg. (unnumbered); Site 4, 8/8/2005, leg. (#78321,
78322); Site 5, 8/8/2005, leg. (#78369); Site 10, 7/7/1995, as E. intestinalis,leg.
Hehre (#58484); Site 11, 8/8/2005, leg. S. Crawford (#78423–78825); Site 11,
8/8/2005, leg. (unnumbered); Site 14, 5/21/1966, as E. intestinalis leg. Searles
(#4600); Site 24, 8/6/2005, leg. (#78411); Site 26, 8/19/1994, as E. intestinalis
leg. (#51431); Site 29, 5/22/1966, as E. intestinalis leg. (#3226); Site 33,
8/20/1994, as E. intestinalis leg. (#51265); Site 35, 7/7/1995, as E. intestinalis,
leg. Hehre (#58360); Site 37, 4/17/1995, as E. intestinalis leg. (#56955), 35.1%.
Ulva lactuca: Site 1, 8/1872, leg. Eaton (Eaton, 1873), as Ulva latissima; Site
1, 10/8/1994, leg. Harris (#53255); Site 1, 8/7/2005, leg. (#78382); Site 4,
8/8/2005, leg. (#78331, 78346); Site 5, 8/8/2005, leg. (#78357); Site 11, 8/8/2005,
leg. (unnumbered); Site 17, 8/8/2005, leg. (#78454); Site 22, 8/8/2005, leg.
(#78427, 78432, 78433); Site 24, 8/6/2005, leg. (#78416); Site 28, 8/19/1994,
leg. (#51424); Site 31, 8/20/1994, leg. (#51616); Site 35, 7/7/1995, leg. Hehre
(#58362), 29.7%.
Ulva linza: Site 1, 8/8/2005, leg. (unnumbered); Site 4, 8/8/2005, leg. (#78328); Site
5, 8/8/2005, leg. (#78358); Site 10, 7/7/1995, as E. linza, leg. Hehre (#58491);
Site 17, 8/8/2005, leg. (unnumbered); Site 21, 8/8/2005, leg. (#78446); Site 26,
8/19/1994, as E. linza leg. (#51429); Site 35, 7/7/1995, as E. linza leg. Hehre
(#58338), 21.6%.
42 Northeastern Naturalist Vol. 16, Monograph No. 5
Ulva prolifera: Site 1, 8/1872, leg. Eaton, as E. prolifera f. tubulosa (Kützing)
Batters (FH); Site 1, 8/7/2005, leg. (#78384); Site 4, 8/8/2005, leg. (#78336);
Site 11, 8/8/2005, leg. S. Crawford (#78423, 78423); Site 15, 5/21/1966, as E.
intestinalis leg. Searles (#3234); Site 22, 8/8/2005, leg. (#78433, 78442); Site 24,
8/6/2005, leg. (#78418); Site 27, 8/19/1994, as E. prolifera leg. (#51270); Site 29,
5/22/1996, as E. intestinalis leg. (#3228), 24.3%.
Ulva torta (Mertens) Reinbold: Site 37, 4/17/1995, as Enteromorpha torta (Mertens)
Reinbold, leg. (#56961), 2.7%.
Ulvaria obscura: Site 3, 1877, leg. Farlow, as Monostroma fuscum (Postels et Ruprecht)
Wittrock (FH); Site 22, 8/8/2005 (unnumbered); Site 30, 7/9/1995, leg.
Hehre (#58552), 8.1%.
Urospora penicilliformis: Site 1, 8/1872, Verrill, as Hormiscia penicilliformis (Roth)
Areschoug (FH), 2.7%.
Phaeophyceae
Agarum clathratum: Site 1, Leg. Farlow, Algae Exsic. Am. Bor. #112, as A. cribrosum
Bory de Saint-Vincent (FH); Site 5, 8/8/2005, leg. (#78347); Site 28, 10/8/1994,
as A. cribrosum leg. Mathieson & Hehre (#53119), 8.1%.
Alaria esculenta: Site 1, 8/1872, leg. Eaton (FH); Site 1, 8/7/2005, leg. (#78372); Site
4, 8/8/2005, leg. (#78343, 78348); Site 5, 8/8/2005, leg. (unnumbered); Site 17,
8/8/2005, leg. (#78451); Site 28, 8/19/1994, leg. (#51397), 16.2%.
Ascophyllum nodosum: Site 1, 10/8/1994, leg. Harris (#53245); Site 1, 8/7/2005, leg.
(#78381, 78386, 78390); Site 4, 8/8/2005, leg. (#78326, 78333); Site 5, 8/8/2005,
leg. (#78363); Site 17, 8/8/2005, leg. (#78455, 78459); Site 24, 8/6/2005, leg.
(#78410, 78415); Site 28, 8/19/1994, leg. (#51400); Site 32, 8/20/1994, leg.
(#51613); Site 35, 8/7/1995, leg. Hehre (#58340), 24.3%.
Ascophyllum nodosum ecad scorpioides: Site 4, 8/8/2005, leg. (#78331); Site 5,
8/8/2005, leg. (#78370); Site 9, 3/11/1996, leg. (#61370); Site 14, 4/21/1966,
leg. Hehre (#1729); Site 27, 8/19/1994, leg (#51817); Site 32, 8/20/1994, leg.
(#51611); Site 35, 8/7/1995, leg. Hehre (#58337), 16.2%.
Asperococcus fistulosus: Site 19, 3/21/1998, leg. (#66965), 2.7%.
Chorda filum: Site 1, 8/1872, leg. Eaton (Eaton, 1873); Site 4, 8/8/2005, leg.
(#78318); Site 17, 8/8/2005, leg. (#78448); Site 18, 8/21/1998, leg (#67482); Site
34, 8/16/1977, leg. Weiss #1748 (Huntsman Lab, G.R. South #5131), 13.5%.
Chordaria flagelliformis: Site 1, 9/1877, leg. Farlow (FH); Site 1, 8/7/2005, leg.
(#78378); Site 4, 8/8/2005, leg. (#78330); Site 5, 8/8/2005, leg. (#78354, 78355,
78364); Site 17, 8/8/2005, leg. (unnumbered); Site 21, 8/8/2005, leg. (#78428,
78444); Site 24, 8/6/2005, leg. (#78422); Site 28, 8/19/1994, leg. (#51416); Site
35, 7/7/1995, leg. Hehre (#58357), 24.3%.
Desmarestia aculeata: Site 1, 8/1872, leg Eaton (Eaton, 1873); Site 2, 8/7/2005,
leg. (#78371, 78376); Site 17, 8/8/2005, leg. (#78449); Site 27, 8/19/1994, leg.
(#51275); Site 36, 6/17/1977, leg. Moskowitz #174 (Huntsman Lab, G.R. South
#5129), 13.5%.
Desmarestia viridis: Site 1, 8/1872, leg. Eaton (FH); Site 1, 8/8/2005, leg. (#78404);
Site 28, 7/9/1995, leg. Hehre (#58550), 8.1%.
Dictyosiphon foeniculaceus: Site 1, 9/1877, leg. Farlow in Farlow, Anderson & Eaton
Algae Exsic. Am. Bor. #95, as D. hippuroides (Lyngbye) Kützing (FH); Site 5,
8/8/2005, leg. (#78367); Site 17, 8/8/2005, leg. (#78458); Site 30, 10/8/1995, leg.
(#59692); Site 35, 7/7/1995, leg. Hehre (#58359), 13.5%.
Dictyosiphon macounii: Site 28, 8/19/1994, leg. (#51412); Site 35, 7/7/1995, leg.
Hehre (#58353), 5.4%.
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 43
Ectocarpus fasciculatus: Site 4, 8/8/2005, leg. (#78335); Site 5, 8/8/2005, leg.
(unnumbered); Site 7, 9/21/1997, leg. Mathieson & Wilkes (#12645); Site 17,
8/8/2005, leg. (unnumbered); Site 22, 8/8/2005, leg. (#78428, 78439, 78440); Site
30, 10/9/1994, leg. Mathieson & Hehre (#52728), 16.2%.
Ectocarpus siliculosus: Site 1, 8/1872 (Eaton, 1873); Site 1, 8/7/2005, leg. (#78376);
Site 4, 8/8/2005, leg. (unnumbered); Site 5, 8/8/2005, leg. (#78355); Site 22,
8/8/2005, leg. (#78435); Site 24, 8/6/2005, leg. (#8415); Site 28, 8/19/1994, leg.
(#51403),16.2%.
Elachista fucicola: Site 1, 10/1875, leg. Farlow (FH); Site 4, 8/8/2005, leg. (#78317,
78345); Site 5, 8/8/2005, leg. (#78370); Site 22, 8/8/2005, leg. (#78433); Site 28,
8/19/1994, leg. (#51421), 13.5%.
Fucus distichus subsp. distichus: Site 1, leg. Averill, as F. distichus (NY); Site 30,
10/8/1995, leg. True & Carpenter (#63942), 5.4%.
Fucus distichus subsp. edentatus: Site 28, 10/7/1995, leg. (#59805), 2.7%.
Fucus distichus subsp. evanescens: Site 1, leg. Farlow in Farlow, Anderson &
Eaton Algae Exsic. Am. Bor. no 107 (FH); Site 4, 8/8/2005, leg. (#8332); Site 5,
8/8/2005, leg. (#8366); Site 22, 8/8/2005, leg. (#78433); Site 28, 8/19/1994, leg.
(#51401); Site 35, 7/7/1995, leg. Hehre (#58341), 16.2%.
Fucus spiralis: Site 1, 6/30/1998, leg. (#67545); Site 1, 8/8/2005, leg. (#78394);
Site 4, 8/8/2005, leg. (unnumbered); Site 28, 8/19/1994, leg. (#51414); Site 35,
7/7/1995, leg. Hehre (#58333), 13.5%.
Fucus spiralis ecad lutarius: Site 11, 2/24/1996, as F. vesiculosus L. ecad limicola F.S.
Collins leg. (#60883); Site 28, 5/20/1998, leg. Mathieson & Dawes (#69068), 5.4%.
Fucus sp. “muscoides-like:” Site 28, 10/28/1996, as “F. vesiculosus L. ecad muscoides
Cotton” leg. Mathieson & Dawes (#62359), 2.7%.
Fucus vesiculosus: Site 1, leg. Farlow in Farlow, Anderson & Eaton, Algae Exsic.
Am. Bor. #109 (FH); Site 1, 8/7/2005, leg. (#78393); Site 4, 8/8/2005, leg.
(#78334); Site 5, 8/8/2005, leg. (#78359); Site 17, 8/8/2005, leg. (#78457); Site
22, 8/8/2005, leg. (#78430); Site 24, 8/6/2005, leg. (#78413); Site 28, 8/19/1994,
leg. (#51278); Site 31, 8/10/1994, leg. (#51264); Site 35, 7/7/1995, leg. Hehre
(#58335), 27.0%.
Fucus vesiculosus ecad volubilis: Site 16, 3/11/1996, as F. vesiculosus ecad limicola
leg. (#62926); Site 28, 2/16/1998, leg. (#66939); Site 35, 3/24/1996, as F. vesiculosus
ecad limicola leg. (#62649), 8.1%.
Halosiphon tomentosus: Site 10, 7/7/1995, leg. Hehre (#58494); Site 18, 7/8/1995,
leg. Mathieson & Hehre (#58170), 5.4%.
Haplospora globosa: Site 13, 6/13/1997, leg. Jones (#64569), 2.7%.
Hincksia granulosa: Site 28, 5/16/1996, leg. (#62070), 2.7%.
Laminaria digitata: Site 1, leg. Farlow in Farlow, Anderson & Eaton, Algae Exsic.
Am. Bor. #119, as L. flexicaulis (FH); Site 1, 8/7/2005, leg. (#78393); Site 28,
8/2/2000, leg. Mathieson & Dawes (#74895), 8.1%.
*Melanosiphon intestinalis: Site 4, 8/8/2005, leg. (#78327) Site 14, 2/23/1996, leg.
(#60790), 5.4%.
Myrionema corunnae: Site 6, 8/20/1998, leg. Mathieson, Yarish, Chopin, & Wilkes
(#67495); Site 29, 5/16/1996, leg. (#62466), 5.4%.
Myrionema strangulans: Site 18, 7/8/1995, leg. Mathieson & Hehre (#58184);
Site 28, 7/9/1995, leg. Hehre (#58551); Site 35, 7/7/1995, leg. Hehre (#58343),
8.1%.
Petalonia fascia: Site 1, 10/1875, leg. Farlow, as Phyllitis fascia (O.F. Müller) Kützing
(FH); Site 1, 8/7/2005, leg. (#78376); Site 4, 8/8/2005, leg. (#78342); Site 5,
8/8/2005, leg. (#78350); Site 22, 8/8/2005, leg. (#78431); Site 30, 10/9/1994, leg.
Mathieson & Hehre (#52969); Site 35, 3/24/1996, leg. (#62645), 18.9%.
44 Northeastern Naturalist Vol. 16, Monograph No. 5
Petroderma maculiforme: Site 30, 10/9/1994, leg. Mathieson & Hehre [#54520 (rock
#1193)]; Site 32, 8/20/1994, leg. Mathieson [#54595 (rock #1265)]; Site 35,
3/24/1996, leg. Mathieson [#12779 (rock #2084)], 8.1%.
Pseudolithoderma extensum: Site 23, 8/19/1994, leg. [#54573 (rock #1245)], 2.7%.
Punctaria latifolia: Site 7, 9/21/1997, leg. Mathieson & Wilkes (#12620); Site 18,
8/21/1998, leg. (#67473); Site 30, 10/9/1994, leg. Mathieson & Hehre (#53289);
Site 30, 7/7/1995, leg. Hehre (#58356), 10.8%.
Punctaria plantaginea: Site 24, 8/6/2005, leg. (#78419), 2.7%.
Pylaiella littoralis: Site 1, 10/1875, leg. Farlow (FH); Site 1, 8/6/2005, leg. Harris
(#78405); Site 4, 8/8/2005, leg. (#78331, 78333, 78340); Site 5, 8/8/2005, leg.
(#78352, 78362, 78370); Site 17, 8/8/2005, leg. (#78456); Site 22, 8/8/2005,
leg. Mathieson (#78429); Site 24, 8/6/2005, leg. (#78417); Site 30, 10/9/1994,
leg. Mathieson & Hehre (#53275); Site 31, 8/20/1994, leg. (#51266); Site 35,
7/7/1995, leg. Hehre (#58355), 24.3%.
“Ralfsia clavata”: Site 1, 10/1875, leg. Farlow (FH); Site 14, 2/23/1996, leg. [#12777
(rock #2082)]; Site 23, 8/19/1994, leg. [#54571 (rock #1243)]; Site 31, 8/20/1994,
leg. [#54594 (rock #1264)]; Site 35, 3/24/1996, leg. [#12780 (rock #2084)],
13.5%.
Ralfsia fungiformis: Site 1 (Farlow, 1881), 2.7%.
Ralfsia verrucosa: Site 1 (SCUBA), 11/13/1999, leg. L. Harris; Site 10, 7/7/1995,
leg. Hehre [#57465 (rock #1770)]; Site 23, 8/19/1994, leg.[#54572 (rock #1244)],
5.4%.
Saccharina latissima: Site 1, 9/71, leg. Short & Lee, as Laminaria sp. (#48865); Site
1, 8/7/2005, leg. (#78373, 78402); Site 4, 8/8/2005, leg. (#78319, 78320); Site
22, 8/8/2005, leg. (#78426, 78434, 78437,); Site 23, 4/19/1996, leg. (#61432);
Site 24, 8/6/2005, leg. (#78414); Site 34, 7/15/1977, leg. Weiss #1751 (Huntsman
Lab., G.R. South #5122); Site 35, 7/7/1995, leg. Hehre (#58334), 21.6%.
Saccharina longicruris: Site 1, 10/1875, leg. Farlow in Farlow, Anderson & Eaton
Algae Exsic. Am. Bor. #117 (FH); Site 1, 8/7/2005, leg. (#78391); Site 4,
8/8/2005, leg. (#78344); Site 5, 8/8/2005, leg. (#78368); Site 17, 8/8/2005, leg.
(#78450); Site 22, 8/8/2005, leg. (#78445); Site 28, 10/8/1994, leg. Mathieson &
Hehre (#53115), 18.9%.
Saccorhiza dermatodea: Site 1, 10/1875, leg. Farlow in Farlow, Anderson & Eaton
Algae Exsic. Am. Bor. #120 (FH); Site 1, 8/7/2005, leg. (#78394); Site 17,
8/8/2005, leg. (#78447); Site 28, 8/19/1994, leg. (#51277); site 35, 7/7/1995, leg.
Hehre (#58336), 13.5%.
Scytosiphon lomentaria: Site 1, 9/1877, leg. Farlow (FH); Site 4, 8/8/2005, leg.
(#78341); Site 5, 8/8/2005, leg. (#78351); Site 17, 8/8/2005, leg. (#78462); Site
22, 8/8/2005, leg. (#78436); Site 24, 8/6/2005, leg. (#78412); Site 30, 10/9/1994,
leg. Mathieson & Hehre (#52721); Site 36, 3/11/1996, leg. (#63781), 21.6%.
Sphacelaria cirrosa: Site 30, 10/9/1994, leg. Mathieson & Hehre (#53281); Site 35,
3/24/1996, leg. (#62634), 5.4%.
Sphacelaria radicans: Site 1, 10/1875, leg. Farlow (NY), 2.7%.
Spongonema tomentosum: Site 1, 8/1872, leg. Eaton, as Ectocarpus tomentosus (Eaton,
1873); Site 13, 6/13/1997, leg. Jones (#64562), 5.4%.
Stictyosiphon griffithsianus: Site 1, 8/1872, leg. Eaton, as Ectocarpus brachiatus
Harvey (Eaton, 1873), 2.7%.
Rhodophyceae
Acrochaetium alariae: Site 28, 8/19/1994, leg. (#51407), 2.7%.
Acrochaetium secundatum: Site 10, 7/7/1995, leg. Hehre (#58487); Site 22, 8/8/2005,
leg. (#78439); Site 24, 8/6/2005, leg. (#78417); Site 30, 10/9/1994, leg. Mathieson
& Hehre (#52723), all as Audouinella secundata, 10.8%.
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 45
Ahnfeltia plicata: Site 30, 10/8/1995, leg. True & Carpenter (#63941); Site 31,
8/20/1994, leg. (#51614), 5.4%.
Antithamnionella floccosa: Site 1, 8/1872, leg. Verrill, as Callithamnion floccosum
(O. F. Müller) C. Agardh (FH); Site 1 (SCUBA), 11/13/1999, leg. L. Harris;
ibid. 8/6/2005 and 8/7/2005, leg. (#78374, 78376, 78389, 78400, 78403, 78406,
78409); Site 4, 8/8/2005, leg. (#8316); Site 22, 8/8/2005, leg. (#78435); Site 28,
10/8/1994, leg. Mathieson & Hehre (#53097), 13.5%.
Audouinella polyidis: Site 30, 10/9/1994, leg. Hehre (#52936 & slide #38), 2.7%.
Bangia fuscopurpurea: Site 1, 3/11/1996, leg. (#63664), 2.7%.
*Bonnemaisonia hamifera, collected as the tetrasporic “Trailliella intricata Batters
stage: Site 28, 10/8/1994, leg. Mathieson & Hehre (#53094); Site 30, 3/11/1996,
leg. (#63610), 5.4%.
Callithamnion tetragonum: site 22, 8/8/2005, leg. (#78445), 2.7%.
Ceramium deslongchampsii: Site 1, 8/1872, leg. Verril (Eaton 1873), as Ceramium
hooperi Harvey; Site 1, 10/26/1875, leg. Farlow (FH); Site 30, 10/9/1994, leg.
Mathieson & Hehre (#52732, 53278 in part as C. strictum), 5.4%.
Ceramium virgatum: Site 7, 9/21/1997, leg. Mathieson & Wilkes (#12613); Site
22, 8/8/2005, leg. (#78435, 78445); Site 24, 8/6/2005, leg. (#78417); Site 28,
8/19/1994, leg. (#51422); Site 35, 7/7/1995, leg. Hehre (#58345), 14.4%.
Chondrus crispus: Site 1, 10/8/1994, leg. Harris (#53257); ibid. 8/7/2007, leg. Harris
(#78400); Site 17, 8/8/2005, leg. (#78460); Site 28, 8/19/1994, leg. (#51423),
8.1%.
Choreocolax polysiphoniae: Site 30, 10/9/1994, leg. Mathieson & Hehre (#53328);
Site 35, 3/24/1996, leg. (#68893), 5.4%.
Clathromorphum circumscriptum: Site 1, 10/1875, leg. Farlow, as Lithothamnion compactum
Kjellman (FH); Site 30, 10/9/1994, leg. Mathieson & Hehre [#54521 (rock
#1194)]; Site 35, 7/7/1995, leg. Hehre [#57466 (rock #1771)], 8.1%.
Colaconema dasyae: Site 30, 10/9/1994, leg. Hehre (#52967 and slide #46), 2.7%.
Colaconema daviesii: Site 1, 1893, leg. Averill, as Callithamnion daviesii (Dillwyn)
Lyngbye (NY), 2.7%.
Colaconema membranacea: Site 1, 10/7/1995, leg. Harris (59558); Site 28,
10/8/1994, leg. Mathieson & Hehre (#53112), 5.4%.
Corallina officinalis: Site 1, 8/1872, leg. Eaton (FH); Site 30, 10/8/1995, leg. True
& Carpenter (#63949); Site 30, 7/1/1977, leg, Moskowitz #853, filed under Ulva
lactuca (Huntsman Lab., G.R. South #5163), 8.1%.
Cystoclonium purpureum: Site 1, 8/1872, leg. Eaton, as C. purpurascens (Hudson)
Kützing (FH); Site 28, 8/19/1994, leg. (#51972); Site 30, 7/7/1995, leg. Hehre
(#58346), 8.1%.
Devaleraea ramentacea: Site 1, 4/1878, leg. Farlow, as Halosaccion ramentaceum
(L.) J. Agardh in Farlow, Anderson & Eaton Algae Exsic. Am. Bor. #78 (FH); Site
28, 8/19/1994, leg. (#51419); Site 30, 3/24/1996, leg. (#62641), 8.1%.
*Dumontia contorta: Site 10, 7/7/1995, leg. Hehre (#58486); Site 30, 3/11/1996. leg.
(#63156); Site 35, 7/7/1995, leg. Hehre (#58361), 5.4%.
Erythrotrichia carnea: Site 1, 8/7/2005, leg. (#78387); Site 19, 9/21/1997, leg. Mathieson
& Wilkes (#12668); Site 24, 8/6/2005, leg. (#78417); Site 30, 10/9/1994,
leg. Mathieson & Hehre (#53285), 5.4%.
Euthora cristata: Site 1, 8/1872, leg. Eaton (NY); Site 1 (SCUBA), 11/13/1999, leg.
Harris; ibid., 8/6/2005, leg. Harris (#78395, 78403, 78407); Site 28, 10/8/1994,
leg. Mathieson & Hehre, as Callophyllis cristata (C. Agardh) Kützing (#53098),
8.1%.
46 Northeastern Naturalist Vol. 16, Monograph No. 5
Fimbrifolium dichotomum: Site 1, 8/1872, leg. Eaton, as Calliblepharis ciliata Kützing
(Hudson) Kützing (Eaton, 1873); Site 29, 7/2/1977, leg. Churchill, as Rhodophyllis
dichotoma (Lepechin) Gobi (Huntsman Lab., G.R. South #5112); Site 30,
3/11/1996, leg. (#63159), 8.1%.
Gracilaria tikvahiae: Site 35, 9/17/1970, leg. Lamb (#A-1092), (FH), 2.7%.
Halosacciocolax kjellmanii: Site 7, 2/25/1999, leg. (#69526), 2.7%.
Hildenbrandia rubra: Site 10, 7/7/1995, leg. Hehre [#57465 (rock #1770)]; Site 30,
10/9/1994, leg. Mathieson & Hehre [#54522 (rock #1195)]; Site 31, 8/20/1994,
leg. [#543595 (rock #1265)]; Site 35, 7/7/1995, leg. Hehre [#57466 (rock
#1771)], 10.8%.
Hydrolithon farinosum: Site 1, as Melobesia farinosa J.V. Lamouroux (BKL #31158),
2.7%.
Lithothamnion glaciale: Site 1, 9/1877, leg. Farlow, as L. fasciculatum (Lamarck)
Areschoug (FH); Site 28, 10/8/1994, leg. Mathieson & Hehre [#54592 (rock
#1262)], 5.4%.
Lithothamnion ungeri: Site 1, 9/1877, leg. Farlow, as L. fruticulosum (Kützing) Foslie
(FH), 2.7%.
Mastocarpus stellatus [including its tetrasporic “Petrocelis cruenta” stage, cf. Guiry
and West 1983]: Site 1, 8/1872, leg. Eaton, as Gigartina mamillosa (Goodenough
et Woodward) J. Agrdh (FH); Site 1, 9/1877, leg. Farlow, as “Petrocelis cruenta
J. Agardh”, mixed with & filed under “Codiolum petrocelidis Kuckuck” (FH);
Site 30, 10/9/1994, leg. Mathieson & Hehre (#53331); Site 30, 10/9/1994, leg.
Harris, as “Petrocelis cruenta”, [#54586 (rock #1256)]. Site 35, 7/2/1977, leg.
Moskowitz #1745 (Huntsman Lab, G.R. South #5174), 10.8%.
Membranoptera spinulosa: Site 1, 8/1872, leg. Eaton, as Delesseria alata (Hudson)
J.V. Lamouroux (Eaton, 1873); Site 2, 8/6/2005, leg. Harris (#78400, 78403,
78405), as Membranoptera alata (Hudson) Stackhouse; Site 28, 10/8/1994, leg.
Mathieson & Hehre (#53107), as M. alata; 8.1%.
*Neosiphonia harveyi: Site 7, 9/21/1997, leg. Mathieson & Wilkes, as Polysiphonia
harveyi (#12641); Site 22, 8/8/2005, leg. (#78443); Site 29, 10/31/1999, leg. Mathieson,
as P. harveyi J. Bailey (#71336), 8.1%.
Palmaria palmata: Site 1, 8/1872, leg. Eaton, as Rhodymenia palmata (L.) Greville
(FH); Site 1, 8/7/2005, leg. (#78379, 78380); Site 4, 8/8/2005, leg. (#78338);
Site 5, 8/8/2005, leg. (#78361, 78362); Site 17, 8/8/2005, leg. (#78452); Site 22,
8/8/2005, leg. (#78438); Site 28, 8/19/1994, leg. (#51967); Site 35, 7/7/1995, leg.
Hehre (#58364), 21.6%.
Peyssonnelia rosenvingei: Site 1, 10/1875 leg. Farlow, as P. dubyi P.L. Crouan et
H. M. Crouan (FH); Site 28, 10/8/1994, leg. Mathieson & Hehre [#54593 (1263
rock)]; Site 35, 3/24/1996, leg. Mathieson [#12767 (rock #2074)], 8.1%.
Phycodrys fimbriata: Site 1, 8/1872, leg. Eaton, as Delesseria sinuosa (FH); Site 2,
8/6/2005, leg. Harris (#78397, 78403, 78408), as Phycodrys rubens (L.) Batters;
Site 28, 10/8/1994, leg. Mathieson & Hehre (#53114), as P. rubens; 8.1%.
Phyllophora pseudoceranoides: Site 1, 10/8/1994, leg. Harris (#53253); Site 30,
10/8/1995, leg. True & Carpenter (#63963), 5.4%.
Phyllophora truncata: Site 2, 8/7/2005, leg. Harris (#78396, 78401), 2.7%.
Phymatolithon laevigatum: Site 2, 10/7/1995, leg. Harris [#12977 (rock #2275)]; Site
2, 8/6/2005, leg. Harris (#78401); Site 11, 12/5/1997, leg. Mathieson [#12857
(rock #2156)]; Site 29, 5/16/1996, leg. Mathieson [#12820 (rock #2119)],
10.8%.
2010 A.C. Mathieson, C.J. Dawes, E.J. Hehre, and L.G. Harris 47
Phymatolithon lamii: Site 1, 1872, leg. Farlow, as Lithothamnion polymorphum (L.)
J.E. Areschoug (FH); Site 30, 10/9/1994, leg. Mathieson & Hehre [#54519 (rock
#1192)], 5.4%.
Phymatolithon lenormandii: Site 1, 10/8/1994, leg. Harris [#54555 (rock #1227)],
2.7%.
Plumaria plumosa: Site 1, 8/1872, leg. Eaton, as Ptilota elegans (Bonnemaison)
Schmitz (FH); Site 28, 8/19/1994, leg. (#51971), 5.4%.
Polyides rotundus: Site 1, 8/1872, leg. Eaton (#26553); Site 30, 10/9/1994, leg. Hehre
(#52944), 5.4%.
Polysiphonia denudata: Site 1, 8/6/2005, leg. Harris (#78397), 2.7%.
Polysiphonia flexicaulis: Site 1, 8/1872, leg. Prudden, as P. violacea (Roth) Sprengel
(FH); Site 4, 8/8/2005, leg. (#78315, 78323); Site 17, 8/8/2005, leg. (#78453);
Site 22, 8/8/2005, leg. unnumbered); Site 28, 10/8/1994, leg. Mathieson & Hehre
(#53106), 13.5%.
Polysiphonia fucoides: Site 2, 11/14/1999, leg. Harris (#70734); Site 3, 8/1872, leg.
Prudden, as P. violacea (Roth) Sprengel; Site 30, 3/11/1996, leg. (#63165); Site
36, 3/11/1996, leg. (#63790), 10.8%.
Polysiphonia lanosa: Site 1, 8/1872, leg. Eaton, as P. fastigiata (Roth) Greville
(BKL #30637); Site 1, 8/8/2005, leg. (#78390); Site 4, 8/8/2005, leg. (unnumbered);
Site 5, 8/8/2005, leg. (#78363); Site 17, 8/8/2005, leg. (#78455); Site 22,
8/19/1994, leg. (#51970); Site 30, 3/24/1996, leg. (#62647), 16.2%.
Polysiphonia nigra: Site 28, 10/8/1994, leg. Mathieson & Hehre (#53111); Site 29,
10/31/1999, leg. (#71300), 5.4%.
Polysiphonia stricta: Site 1, 8/1872, leg. Eaton, as P. urceolata (Lightfoot ex Dilwyn)
Greville (BKL #30678); Site 1 (SCUBA), 11/13/1999, leg. L. Harris; ibid.,
8/6/2005, leg. Harris (#78403); Site 4, 8/8/2005, leg. (#78324); Site 5, 8/8/2005,
leg. (#78353); Site 18, 7/8/1995, leg. Mathieson & Hehre, as P. urceolata
(#58175); Site 22, 10/8/1994, leg. Mathieson & Hehre, as P. urceolata (#53110);
Site 29, 5/16/1996, leg. (#62469); Site 35, 7/7/1995, leg. Hehre, as P. urceolata
(#58344), 21.6%.
Porphyra amplissima: Site 10, 7/7/1995, leg. Hehre (#58495); Site 18, 7/8/1995, leg.
Mathieson & Hehre (#58171); Site 28, 7/9/1995, leg. Hehre (#58558); Site 29,
5/16/1996, leg. (#62472); Site 35, 6/13/1977, leg. S. Weiss #730, as P. miniata
(C. Agardh) C. Agardh (Huntsman Lab, G.R. South #5169); Site 35, 7/7/1995,
leg. Hehre (#58352); 16.2%.
Porphyra birdiae: Site 4, 8/8/2005, leg. (unnumbered); Site 5, 8/8/2005, leg. (unnumbered),
5.4%.
Porphyra leucosticta: Site 1 (SCUBA), 10/8/1994, leg. Harris, as P. umbilicalis Kützing
(#53254); Site 28, 8/19/1994, leg. (unnumbered), as P. umbilicalis (#51404);
Site 35, 7/7/1995, leg. Hehre, as P. umbilicalis (#58351), 8.1%.
Porphyra miniata: Site 1, 8/1872, leg. Eaton, as P. vulgaris C. Agardh (YALE); Site
9, 3/11/1996, leg. (unnumbered), as P. leucosticta (#61386), 5.4%.
Porphyra purpurea: Site 4, 8/8/2005, leg. (unnumbered); Site 5, 8/8/2005, leg. (unnumbered);
Site 11, 12/5/1997, leg. (#66081); Site 17, 8/8/2005, leg. (unnumbered);
Site 19, 11/21/1997, leg. (#66060); Site 24, 8/6/2005, leg. (unnumbered);
Site 28, 10/7/1995, leg. X. Fei (#65279); Site 35, 3/24/1996 leg. (unnumbered),
as P. umbilicalis (#62629), 21.6%.
Porphyra umbilicalis: Site 3, 10/7/1997, leg. Yarish & Fei (#59566); Site 4, 8/8/2005,
leg. (#78337); Site 30, 10/9/1994, leg. Mathieson & Hehre (#53313), 8.1%.
48 Northeastern Naturalist Vol. 16, Monograph No. 5
*Porphyra yezoensis f. narawaensis (strains U51 & H25): Site 8 (Aquaculture nets),
7/8/1995, leg. Hehre (#58332), 2.7%.
Porphyrostromium ciliare: Site 19, 11/21/1997, leg. as Erythrotrichopeltis ciliaris
(Carmichael) Kornmann (#66069); Site 30, 10/9/1994, leg. Mathieson & Hehre,
as E. ciliaris (#52722), 5.4%.
Ptilota serrata: Site 1, 8/1872, leg. Pruden (Eaton, 1873); Site 1, leg. Farlow, as
P. plumosa C. Agardh in Farlow, Anderson & Eaton Algae Exsic. Am. Bor. #85
(FH); Site 2, 8/6/2005, leg. Harris (#78398, 78409); Site 28, 10/8/1994, leg. Mathieson
& Hehre (#53105), 10.8%.
Rhodochorton purpurem: Site 1, 8/1872, leg. Verrill and Pruden (Eaton, 1873), as
Callithamnion rothii (Turton) Lyngbye; Site 1, 10/1875, leg. Farlow, as Rhodochorton
rothii (Turton) Nägeli (FH.); Site 28, 7/9/1995, leg. Hehre (#58553),
5.4%.
Rhodomela confervoides: Site 2, 8/6/2005, leg. Harris (#78399); Site 27, 8/19/1994,
leg. (#51408); Site 35, 3/24/1996, leg. (#62637), 8.1%.
Rhodophysema elegans: Site 2, 10/7/1995, leg. Harris [#12977 (rock #2275)]; Site 2,
8/6/2005, leg. Harris (#78401); Site 29, 5/16/1996, leg. Mathieson [#12820 (rock
#2119)], 8.1%.
Scagelia americana: Site 1, 8/1872, leg. Eaton (Eaton, 1873), as Callithamnion
americanum Harvey; 2.7%.
Scagelia pylaisaei: Site 1, 8/1872, leg. Pruden (Eaton, 1873); Site 1, 9/1877, leg.
Farlow, as Antithamnion boreale (Gobi) Kjellman (FH); Site 28, 7/9/1995, leg.
Hehre (#58555), 8.1%.
Titanoderma pustulatum: Site 1, as Melobesia pustulatum J.V. Lamouroux (BKL
#31160), 2.7%.
Turnerella pennyi: Site 1, 10/7/1995, leg. Harris (#59560), 2.7%.