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Floristic Studies of Seaweeds from Cobscook Bay, Maine
Arthur C. Mathieson, Clinton J. Dawes, Edward J. Hehre, and Larry G. Harris

Northeastern Naturalist, Volume 16, Monograph 4 (2009): 1–48

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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. 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Klein, C.D. Neefus, G.G. Mitman, and I. Levine. 1998. Domestication of indigenous Porphyra (nori) species for commercial cultivation in Northeastern America. World Aquaculture 29:26–29. 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%.