1710 /

(DlCI§CO f. PEDROCHE

Rep rint ed fro m R HOIlORI , Vol. 76, 008, 1074

INVESTIGATIONS OF NEW ENGLAND MARINE ALGAE VI: DISTRIBUTION OF MARINE ALGAE NEAR , MASSACHUSETTSl

DOUGLAS C. C OLEMAN AND ARTHUR C . MATHIESON

Cape Cod is a major phytogeographic boundary on the northeast coast of North America, delineating a distinctive northern and southern marine flora (Chapman, 1964; Humm, 1969; Setchell, 1922; Stephenson & Stephenson, 1949). Harvey (1852-1858) and Farlow (1870, 1882) first recognized its importance as a phytogeographic boundary. Additional information regarding the uniqueness of the Cape Cod flora was contributed by Collins (1900), Davis (1913a, b) and Taylor (1937, 1957) . Although th e marine flora of Cape Cod has received con­ siderable attention since the time of Harvey, no one has ever conducted simultaneous yea r round studies of the algae on both sides of the Cape. In addition, nothing is known about th e seasonal changes that occur in the Canal, which joins Cape Cod Bay and . Conover (1958) and Sears (1971 ) conducted seasonal st udies of the algae in southern Cape Cod, but they ga ve no consideration to the Canal. The purpose of the present investigation was to study the horizontal and vertical distribution of seaweeds at seven locations from Scituate to Woods Hole, Massachu­ setts, and reproduction including the Cap e Cod Canal. The seasonal occurrence of seaweeds at the same sites wiII be discussed in another paper. Monthly collections of all th e conspicuous algae at seven stations (Fig. 1) were made from J anuary to December, 1969. Severe winter conditions (heavy ice and surf) ex­ isted from December to March, and they restricted some

'Published wit h th e ap proval of the Dire ctor of th e Un iversity of New Ham pshire Agriculture Experiment Station as Scientific Con­ t ri bution Number 656.

G37 538 Rhodora [Vol. 76 collections. Specimens were collected on foot in the inter­ tidal zone and by SCUBA diving to a depth of 60 feet in the subtidal zone. Diving in the Canal was done at slack water (Anon. 1969a). The collections were brought to the Uni ­ versity of New Hampshire for identification and processing. Taylor (1937, 1957, 1960) was the primary source of identification and nomenclature. A number of additional r eferences were cons ulted for the identification of species and the determin ation of recent geogr aphical r ecords along the Atlantic Coast of North America subsequent to Taylor's (1960) summa ry. These include : Adey, 1964, 1965, 1966 ; Bell & McFarlane, 1933a,b; Blomquist & Humm, 1946; Cardinal, 1964, 1965, 1966, 1967a,b,c, 1968; Co]J ins, 1909 ; Edelstein, et al., 1967, 1969, 1970; Edelstein & McLachlan 1966, 1967a,b, 1968a,b, 1969; Fritsch, 1935, 1945; Hoek, 1964; Hoyt, 1920; Lamb & Zimmermann, 1964; Lee, 1968 , 1969; Lewis, 1914; MacFarlane & Bell, 1933; McFarlane & Milligan, 1965; Mathieson & Fuller, 1969; Mathieson, Dawes & Humm, 1969; Rhodes, 1970; Sou th & Cardinal, 1970; Stone, et al., 1970; Wilee, 1959, 1965; Willi ams, 1948, 1949; Wulf, et al., 1968 ; Zaneveld & Barne-s, 1965 ; Zaneveld, 1965 , 1966a,b, 1972. The nomenclature of Parke & Dix on (1968) was applied in most cases. Surface water temperature and salinity information was recorded wh en each station was investigated. Temperature wa s determined with a laboratory grade mercury thermom­ et er; salinit y wa s mea sured with a set of hydrometers (G. M. Mfg., Co.). In addition, hourly and daily surface water temperature data were obtained for each end of t he Canal from therrnographs monitored by the Fish eries Division of the State of . Additional (daily) temperature and salinity information was supplied by the Woods Hol e Oceanographic Institute, F almouth, Massachusetts (per­ sonal communication with Dean Bumpus and Jo seph Chase) . Figure 1 illustrates the location of th e seven study sites. One station (Scituate) wa s located north of the east end of the Canal, four were in th e Can al proper a nd t wo (Woods 1974] Marine Algae - Coleman and Mathieson 539

CAPE COD BAY

70'30' 70'10'

Figure 1. Map of Cape Cod, Massachusetts, showing' the seven stations. Hole and Wings Neck) were located south of the west end of the Canal. The land-cut of the Canal cuts across the Cape in an east-west direction from the town of Buzzards Bay (approximately 41°44' N Latitude, 70°37' W Longi­ tude) to Sandwich (approximately 41°46' N Latitude, 70°30' W Longitude). It is seven miles in length, with an additional seven miles of approach channels. Several differences are obvious between the two sides of the Cape. To the north, the substrate consists of granite outcrops, boulders and cobbles, while to the south the sub­ 540 Rhodora [Vol. 76

22 / /' \ / \ /' ". 20 / \ / .~ _ ...... \ 18 ,..;, '"./ \ \ / . ./ "~ \ / / -: \ \ 16 I I t7 \\ / / .' \ .\ I / ,/ . \ \ 14 / . .I' '- . \ u i/ ,I \'.\

V) 12 I I ,I '- \\ L.U /i/ '-\\ L.U I . • \ .\ 0<0 10 / /,1 \~~ C) L.U Cl 8 ~\\ t/'. / . 'i/ 6 1./ \ ~ . . / \. 4 .f / • . / STATION 1-­ / , STATION 3 .-._._ .. 2 <: I ,­ STATION 5 - '-'­ .':'""--:i/ , ...... STATIO N 7 ---­ 0 ::"'----'

F ig-ure 2. Mea n m on thl y variati on of sur face w ater temperatures at sta tions 1, 3, 5 and 7 du ri ng 1969. strate is largely sand and mud with fewer boulders. The Cape is also the dividing lin e between the Labrador Current to the north and the Gulf Stream to the south, Thus, a marked difference in water temperatures occurs during the summer. The yearly temperature range south of the Cape was approximately 22°C., while that to the north was about 17°C. (Fig. 2) . The salinity ranges south of t he Cape were usually higher than those to the north, particularly during spring and summ er (Fig. 3) . Th e tidal amplitude is much greater on the north than the south side. Thus, a three foot tidal amplitude exists in Buzzards Bay, while it is approxi­ mately ten feet at Scituate (Anon. 1969a). The waters of Cape Cod Bay are over 100 fe et deep while those in Buz­ zards Bay are sh allower. 32.5~ .....,.----., \. -' , ~ \. /\ 32.0 '\. ..,..". -a:- - -e_ .~" , <, "",,'~ ,-­ ;I <. ~o .::,.c.;.-_~_ .0 ...... ,,_-, 0 _~,./.,' x'­,_.0 -, \ o ~ ...... _\- '---ti" 0 -.,;I '\. '.J' ~ 31 .5 /' ,". ~"b~ \ / \ I -"'0 >­ \ I ...... I­ . ' I ~ \ _0 / \ , Z 31 .0 ,I \ , ...... J V ~30.5 V> 30.0 STATION 1 STATION 3 -----­ 29.5 STATION 5 -'-'­ STATION 7 ---­ 29.0

F'igur e 3. Mean mo nth ly var iation of su r face wa ter sa linity at sta tions 1, 3, 5 and 7 during 1969, 542 Rhodora [Vol. 76

STATION 1 -- ~ STATION 3 .~_.._- -_. STATION 5 -._._. 25 STATION 7 ---­

•, ,I , 20 •l ·I , • • I, \ I t . \ • . I I I , ,\ ,I ,• I \ I I ! I, \ ,I , I I \ I \ i I \ 15 \ I I I \ I U , ·I \ \ I I \ I V) \ I \ ~ \ I, u.J I 1 \ I u.J , \ ell:: \ I (J 10 1 . u.J JULY 0

5 JANUARY

t

o t • T T % i ~ l j • LJ LA DAYS F igure 4. Mean dail y variation of sur face water temperatures at stations 1, 3, 5 and 7 during .January and July, 1969. 1974] Marine Algae - Coleman and Mathieson 543

The Canal has a limited amount of solid substrate, no wave action and a reduced intertidal zone. The seasonal temperature ranges are intermediate between the north and south sides of the Cape, but daily fluctuations during the summer are greater than on either side of the Cape (Fig. 4). The

60 NORTHERN SOUTHERN COSMOPOLITAN

50

a.. :::::> o 40 oCk: <, « x « I­ 30 ou.. Ck: UJ cO ~ 20 :::::> Z

10

,; .._.-._e_ ._._ ,; _...... O.l....-r------.----.-----.-----.---r------.--­.... 2 3 4 5 6 7 STATIONS Figure 5. Number of northern, southern an d cosmopolitan species at each station. 1974] Marine Algae - Coleman and Mathieson 545

A listing of the known ge ographical distribution of each taxon along the Atlantic Coast of North America is also designated numerically in Table r. The species ca n be grouped into three components (Le., northern, southern and cosmopolitan) according to their major centers of distribu­ ti on. The northern species are found m ost commonly north of the Cape, and they in clude groups 1, 2, 3, 4 and 15. The souther n species ar e more common south of the Cape; th ey include group s 7, 11, 12 and 13. Cosmopolitan speci es are found on each side of the Cape, and they include group s 5, 6,8, 9, 10, 14 and 16. Figure 5 sum mar izes the number of northern, s outher n and cosmopolitan species at eac h station. Most of the spe­ cies were cosmo polita n. Th e northern component declined towards the so uth, while the southern component incr eased towards the south, reaching a maximum at 'Wings Neck and Woods Hole. Cosmopolitan s pecies included Cha eiom orpha me lagonium , R hizoclonium tortuosum, Vlva laciuca, A sco­ phyllum nodosum , Petalonia f ascia, Ahnf eltia plicata and Ceramium rubrurn, while conspicuous northern components included Codiolum petrocelidis, Mono st roma spp., Chorda filum, Fucus spirulis, Ltimiruiria diqitato; Chor eocolax poly­ siphoniae, and Porphyra umbilicalis. Representative species more common to the south of th e Ca pe in cluded Codium fragile subsp . tomentosoides, S ar oceeum. filipendula, Agard­ hiella ienera, Cham pia parvula and Hypnea musciformis. Scituate had a species composition typical of northern Ne w England , while Wings Neck and Woods H ole were more typical of war me r water locations. Thus, Laminaria spp., Gigar tina stellata, Ralf sia fu ngiform is, Chaetomorpha atrovirens, Uros pora collabens, and Fucus ve siculosus were common at the former station, while S argassum filip endula, A gardhiella tenere, Calli th amnion baileyi, and Codium fra g­ ile s ubs p. tom eniosoules were common at the latter stations . The Canal stati ons, 3-5, included representatives from both sides of the Cape (e.g. Lam inaria saccharina, Chondrus crispus, Rhodymenia ptumat«, Agardhiella tenem, Codium fm gil e subsp. tomentosoides and Saroaesum. {ilipendula. 546 Rhodora [V ol. 76

VERTICAL DISTRIBUTION Figure 6 summarizes the vertical distribution of the con­ spicuous species at each station. The distributions were recorded as follows: 1) species restricted to the intertidal zone; 2 ) species restricted to th e subtida l zone ; 3 ) species found in both th e intertidal and subtidal zones. Most of the plants at Scituate were collected from the intertidal and subt idal zones, including tide pools. Spe cies that were common to both zones included Chon drus crispus , Chord a spp.. E ctoctu-pus siliculosus, Petalonia fasc ia, Ceramium ru brum , Corollina offi cirui lis, Dum ontia in crasstiia, and Chae tomorpha linum. F ew species (e.g. Chaetom orpha atrovirens, Cladophora fiexuosa, Polyide s rotundus, A spero­ coccus echina ius, and Laminori o: spp.) were restricted to the subtidal zone or inter tidal zone (e.g. Codi olum spp., Enteromorpha intestinalis, Rhizoclonium iortuosum, Ulo­ thri x fiacca, Choreocolax polysiphon'iae, and F'ucus spi ralis ) , Sub tidal substrate was lacking at station 2, an d the lowest number of species was found here. The Canal stations (2-5) had the highest subtidal com­ ponent. Species restricted to th e subti dal zone in the Canal were also found in the intertidal zone at other stations. Th e most common subtidal s pecies in th e Canal included Lam i­ naria spp., Chon drus crispus, Aqardhieila tenere; Chorda spp., Chordar ia fiagelli formis, Ulua lcctuca, P etal oni a fas­ cia, and Scutosiphon. lomenuiriue. Speci es restricted to th e intertida l zone included Urospora p enicillif orm is, A scophyl­ lum nodosum , Fucus spiralis, and Choreocolax polysiphon­ iae; those common to both zones included E ctocarpus sili­ cuiosus, Fucus vesiculosus , and Sphacelaria cirrosa. Th e majority of species collected at Win gs Neck and Woods Hole was found in the subtida l zone an d the inter­ tidal-subtidal zones. Few species wer e restricted to th e intertidal zone. Species r estricted to th e subtidal zone in­ cluded A gardhiella tenere; Sarg assum filipendula, Callitham­ nion roseurn, Dosu o. pedicellate, Seirospora grifJithsiana, Leath esia difformis, Chortlaria fiagelliform1:S , and Clado­ 1974] Marine Algae - Coleman and Mathieson 547

IN TERTIDA L INTER-SUBTIDAL 90 • SUBTIDAL 80

V) 70 UJ u 60 UJ 0­ V) 50 u, o 40 a:::: LlJ co 30 ~ ::::::> Z 20

10

Figure 6. Ver tica l d ist r-ibution of species at each sta tion.

phora s pp. Species found in both zones in cluded Chondrus cris pus, Melobesia leioiisii, E ctocarpue siliculosus, Fucus oesiculosus va l'. spuerocarpus, and Chaetomorpha linum, 548 Rhodora [Vol. 76 while those restricted to the intertidal zone included Bangia [uscopurpurea; Porphyra umbilicolis, Fucus spiralis, and Ulothrix flacca,

DISCUSSION Setchell (1917) was one of the first workers to emphasize the role of temperature in determining the geographical distribution of seaweeds. He divided the oceans into 5°C. intervals or isotherms, according to the maximum summer water temperatures. Hutchins (1947) also confirmed that 5°C. intervals either favored or inhibited growth and re­ population of marine organisms. 'Williams (1948), Parr (1933), Well and Gray (1960), and Humm (1969) have reported a similar relationship between temperature and species composition. As suggested by Setchell (1917) Cape Cod is a dividing line between the 15° and 20°C. maximum summer isotherms. Hence, it is not surprising that it is a major phytogeographic boundary, with distinct floras oc­ curring north and south of the Cape. Several factors are responsible for the marked difference in summer temperatures between the north and south side of Cape Cod. A southern extension of the Labrador Current carries cold water as far south as the Cape, where it re­ mains throughout the year. In addition, the deep waters of Cape Cod Bay warm up slowly during the summer. Davis (1913a, b) described Cape Cod Bay as a "holding pocket" of cold water. He further described the islands of Martha's Vineyard and as barriers protecting Buzzards Bay from the intrusion of cold waters off Gay Head. The high surface water temperatures in Buzzards Bay result from a northern extension of the Gulf Stream into the shallow confines of Buzzards Bay. During the winter, cold air lowers the temperature on both sides of the Cape and the sharp differential of temperature disappears. Winter temperatures in Buzzards Bay, however, may be somewhat lower than in Cape Cod Bay, because of the shallowness of the water of the former location. 1974] Marine Algae - Coleman and Mathieson 549

The transitional nature of the Cape Cod Canal is docu­ mented by its species composition. The Canal is dominated by cosmopolitan· species; northern species increase toward the east end of the Canal, while southern components in­ crease toward the west end. Setchell (1922) also recognized a cosmopolitan group of plants common to both sides of the Cape - in contrast to more northern and southern elements. The completion of the Canal in 1914 presumably provided a direct route for spores to pass from one side to the other. Stephenson (1944) has shown a similar transition of flora and fauna near the Cape of Good Hope in Africa; on the west coast the waters are relatively cold, while on the east coast the shore is bathed by the warm waters of the Indian Ocean. Seven major components are evident near the Cape of Good Hope; the major components are the cosmopolitan, warm- and cold-water elements. Physical factors, such as tidal amplitude, wave action and availability of solid substrate, determine the local distribu­ tion of species at the seven sites. Salinity was not con­ sidered a major factor in the areas studied because of the small range recorded. Scituate had the highest number of species ; it is the most exposed site having ample substrate. The lack of wave action, lower tidal amplitude, and reduced substrate in the Canal were probably responsible for the low number of species at stations 2-5. Woods Hole had the second highest number of species, even though tidal ampli­ tude, wave action, and substrate were reduced south of the Cape. The abrasive action of sand at 'Wings Neck resulted in low species numbers. A variety of physical factors, such as temperature, salin­ ity, light intensity, substrate and exposure, determine the vertical distribution of seaweeds (Stephenson & Stephen­ son, 1949; Chapman, 1964). Most algae at Scituate were collected from the intertidal and subtidal zones, while in the Canal and south of the Cape the majority of species was restricted to the subtidal zone. The former location (Scitu­ ate) is characterized by greater tidal amplitude and wave action, both of which are necessary for the development of 550 Rhodora [V ol. 76 an exte nsive intertidal flora - ass uming substrate is not limiting. The vertical position of species var ied from sta­ tion to station. Species f ound in the intertidal and subti da l zones at Scituate were often restricted to the subt idal zone in the Canal or south of the Canal.

ACKNOWLED GEMENTS We would like to thank Dr . A. Hodgdon for his cr itical review of the manuscript. In addit ion we express our grati­ tude to Dr. R. Fralick, Dr. R. Turner, and Mr. J ohn Hans­ man for their assistan ce with diving and collection of specimens, to th e Army Cor ps of En gin eers at Cape Cod for diving access to the Cape Cod Canal, and to Dean Bumpus and J oseph Chase for th e use of unpublished tem­ per atu re and salinity data.

AP P ENDI X

DESCRIPT ION OF S TAT ION S Scit uate (statio n 1) is located about 30 miles north of the east end of the Canal at approximately 42°12' N latit ude and 70°43' W longitu de (Fig. 1) . It is a semi-exposed site. consisting of massive granite outcropping's. Th ere are several ti de pools and a well developed int ertidal zone. Th e substrate in the subtidal zone consists of large outcropping's, boulders, cobbles and sand. Th e surfa ce water temper atures ranged from approximately 2-16°C. The highest temper a­ tures occurred in August, while the lowest occurred in March. The daily temperature range was abo ut 2°C. dur ing th e summer and 3°C. durin g the winter . The ave rage monthly summer temp eratures at Scitu ate were the lowest of all the s tations. Sali nity values remained re latively con­ stant throughout the year, ranging fro m a low of 30.6% 0 in the summer to a high of 32.5% 0 during the winter. Sta tion 2 is located at t he east end of the Can al at th e Scusset Breakwat er (Fig. 2). Collections were res tr icted to the int ertidal zone because of the lack of subtidal sub­ strate. An extensive intertidal zone is present. Th e water 1974] Marine Algae - Coleman and Mathi eson 551 temp eratures ranged f rom 12°C. to 19°C. The tidal ampli­ tude is 10 feet (Anon. 1969a). Station 3 is located at the power plan t (F ig. 2). Rip-rap extends to about 6 feet below M.L.W., while a shelf extends out 30-40 f eet and to a depth of 15 feet. During operation, th e power plant discharges wate r of over 23.9°C. Substrate is limited and a small intertidal zone is present. Th e tidal ampl itude is approximate ly 10 feet. Stati on 4 is ap proximately 2.5 miles eas t of the west end of the Can al proper (Fig. 2). The st udy site was on the south bank of th e Cana l. Th e subst rate grades from rocks (boulders) in th e upper shor e to sand-silt in t he lower sub­ tidal zone. The tidal amplitude is about 6 feet (Anon. 1969a) . A limited intertidal zone is exposed du rin g low tide. Station 5 is located at the Engineer's station at the west end of the Canal. Collectio ns were made in the vicini ty of the pier. Rip-rap extends to about 12 feet below M.L.W., whil e a shelf extends out to about 75 feet and s lopes to about 20 feet below M.L.W. The tidal amplitude is about 4 feet (Anon . 1969a ) . Wings Neck (station 6) is located at appr oximately 41°31' N latitude and 70°40' W longitude in the tow n of Bour ne, Massachusetts (Fig. 1) . It is a semi-exposed area subject to southwest winds during the sum mer. A limited inter tidal zone is present; it is composed of small cobbles and a few large boulders. A fe w shallow tide pools are evident. Th e ti dal am plitude is about 4 feet (Anon . 1969a ) . Th e range of water te mperatures was 0.2-22.1°C. The mean monthly win ter temperat ure (2.2°C.) was lowest in Janu­ a ry. At that time the temperature "vas lower tha n at Scitu­ ate, the northernmost station. The area is sub ject to severe ice scour ing during the win ter . Th e sali nity remained rela­ tively constant throughout the yea r, with a range of about 0.9"/ 00 (Fig. 4). Woods Hole (station 7) is located at approxima tely 41°31' N lati tude an d 70°40' W longitude in the tow n of Falmouth, Massachusetts (Fig. 1) . Collections were mad e at the jetty behind the U. S. Fi sh eri es Building. The bottom 552 Rhodora [Vol. 76 is approximately 20 feet below M.L.W., at the deepest point, and the bottom substrate is composed of sand and silt. Some small rocks were present at one side of the jetty. The tidal amplitude is about 2 feet (Anon. 1969a). The temperature ranged from a low of 0.2°C. in January to 22.5 °C. in Au­ gust (Fig. 3). Daily temperature ranges were small during the year (Fig. 5). The salinity remained relatively con­ 0 stant throughout the year with a range of only 0.8 / 0 0 (Fig 4) .

F ootn otes to Table I -Geograph ical distribution zones relating to T abl e I . r igh t man d column. 1. Nor thern lVI assachusetts to Newfoundland and n orth. 2. North er n Massachu setts to Nova Scotia. 3. Southern Ne w England-Long Island Sound to Gas pe an d Labrador. 4. Southern New England-Long Island to Newfoundlan d an d north. 5. New Jersey-Mar yland to Newfound la nd and nor th. 6. New J er sey-Maiyland to Ga spe an d Labr ador . 7. New .Icrs ey-Mar ylan d to 'Cape Coo. 8. North Carolina to N ewfoundland and north. 9. North Ca rolina to Ga spe . 10. South Ca rolina to Newfoundland a nd nor th . 11. South Carolina to northern Ma ssa chuset ts. 12. Tr opics to southern New England-Cape Cod. 13. Tropic s t o nor ther n Massachusetts. 14. Tropics to Newfoundland and north. 15. Southern New England-Long Island Sound to northern New Eng­ land-Nova Scotia. 16. Tro pics to northern New England-Nova Scot ia .

2 = Neoagardhiella bailey'i (Harvey ex Kutzing) W ynne et Ta ylor. (W ynne an d Ta ylor, 1937). 3 = Phullophor« truncate (Pallas) Newroth et Taylor. (N ewroth and Taylor, 1971). • = Pnutiophora pseudo cerano ides (Gm elin) Newroth et Taylor. (Newroth an d Taylor, 1971) . TABLE 1. DISTRIBUTION OF SPECIES f--' Distribution eo -.J Chlorophyta Station -Geo- ~ Taxon 1 2 3 4 5 6 7 graphical Chaeiomorplui atrouirens Taylor X 4 ~ Chaeiomorplui linum. (MU lier ) Kiltzing X X X X X \I' X X 14 ""'l Chaeiomorpha m elaaonium. (Weber et Mohr) Kiitzing X X X X X 5 5' (D Cladophora fiexuosa (Mull er) Harvey X X XX X 14 Cladophora gracilis (Griffiths ex Harvey) KUtzing X X X 14 aq> \I' Cotliolscm oreqarium A. Braun X X 3 (tl Codiolum petroce lidis Ku ckuck X 2 I Cotiium /ra,qile (Sur.) Hariot s ubs p, tomentosoides () (van Goor) Sil va X X X X X 7 ro0 E ni eromorplui in iestinalis (L.) Link X X X X X X X 14 8 \I' E nt eromorpha linza (L. ) J, Aga rdh X X X X 14 ::l \I' Mon ostrom o. grevillei (Thuret) Whittrock X X X X <1 ::l Mon ostroma pulchrum F arlow X X X X 4 c, ~ Ii hizocioniurn. tor tuosum Kiitz.ing X X 8 \I' Sponqomor pha arc t a (Dillwyn) Ktitzing X X X X X 5 g; ~. (tl Sponaom orpha spinescens Kiit zing X X 1 U> 0 Ulot hri x iiacca (Dillwyn) Thuret X X X X 8 ::l Ulva lactuca L. X X X XXX X 12 Urospora collabens (C. Agardth ) Homes et Batters X 1 01 Urospora penicilli/ormis (Rot h ) Areschoug X X XX X 4 01 iXI Subtotal 18 13 4 6 9 11 13 T ABLE I cont. <:n <:n Distribution .;:.. Phaeophyta Sta tion "Geo- Taxon 1 2 3 4 5 6 7 graphical Ascophsjllum. nodosum (L.) Le Jolis X X X X X X X 8 Asperoco ccus echinaius (Mertens) Greville X X X X X X 4 Chorda filum. (L.) Stackhouse X X X X X X X 5 Chorda tomeniosa Lyngbye X X X X X X X 3 Chordaria flag clliforrnis (Muller) C. Agardh X X X X X X X 5 Desmarestia aculeata (L.) La mo uroux X X X X X 5 Desm arcstio. viridis ( MUller) Lamouroux X X X X X X 5 ~ ~ Desmotrichum bclticum. Kiitzing X X X 3 0 A. Desm otriclium: urululaium. (J. Agard h ) Reink e X 0 X 9 '"'l Dictyosiphon fo eniculaceus (Hu dso n) Greville X X X 5 lI' E ctocarpus silicu losus (Dillwyn) Lyngb ye X X X X X X X 14 E lachista [ucicola (Velley) Aresch oug X X X X X X 5 Fu cus disticliu» L. emend. su bsp. distichus (C. Agardh ) P owell X 1 Fucu s distichus L. emend. subsp. endentaius (C . Agardh ) P owell X X X 5 F1tCUS distichus L . eme nd . s ubsp. evan escen s ....., (C . Agardh) P owell X X X X X 4 -< Fucus spiralis L . X X X X X X X 4 ~ -'l Fucus »esiculosus L. X X X X 8 0> ...... TABLE I - Phaeophyta cont. so --;:r Distribution 01>­ '-' Stati on -Geo- Taxon 1 2 3 4 5 6 7 graphical ~ F'ucus ve siculosos val'. ephaerocarpus J, Agardh X X X 3 ~ .....'1 . F'ucus »esiculosus val'. spiralis Farlow X X 6 ~ Giffordia granulosa (Smith) Hamel X X X 5 (l) Giffordia secunda. (Kiitzing) Batters X 15 > ~ X ~ Laminaria diaitata (Hudson) La mou reux X 4 (l) Laminaria saccharina (L.) Lamouroux X X X X X X 5 I Leathesia dijf oTmis (L .) Areschoug X X X X X X 8 (') Myrionema eiranaulans Greville X X X X X X 14 ro0 Petaionia fascia (Muller) Kuntze X X X X X X X 14 8 ~ Pi lay ella liitorolis (L.) Kj eIIman X X X X X X X 5 ~ laii f oli a. ~ Punctaria. Gr eviIle X X X X 5 ~ Punctaria plamtaoinea (R oth) Greville X X 5 0. Ralfsia iunaiiormie (Gunner) Setchell et Gardner X X X ] ~ ~ Ralfs ia. VeTT'UCOSa (A reschoug ) J . Agardh X X X X X X X 5 ::re-t­ (l) Saroaesum. filipendula C. Agardh X X X X X 12 rn 0 S cutosipho« lomeniarius (Lyngbye) Link X X X X X X X I4 ~ Sphacelario: cirrosa (Roth ) C. Ag ardh X X X X X X X 4 Spluieroiri cia. divaricaia (C. Agardh) Kyl in X X X 5 01 Subtotal 26 18 23 2] 26 24 29 01 01 T ABLE I cont. ~ ~ Distribution 0') Rhodophyta Stati on 'Geo- Taxon 1 2 3 4 5 6 7 graphical Agardhiella tenere (J. Agardh) Schmitz" X X X X X X 12 Ahnfeltia plicata (Hudson) Fries X X X X X X X 5 Antit luimnion am ericanum (Harvey) F arlow X 5 B angia [uscopurpureo. (Dillwy n ) Lyngbye X X X X X 14 B onnem aisonia hami iera Hariot X 15 Callithamnion baileyi Harvey X X X 5 Callithamnion ros eum. (Ro th ) Lyngbye X X X X 16 ::0 Cercmium rubrum (Hudson) C. Agardh ::r X X X X X X X 14 0 Ceramium strictum H arvey X X X X 16 0...... 0 Champia parvula. (C. Agardh) Harvey X X X 13 ~ Chondria sediioiia Har vey X X 12 Chond rus crispus Stackhouse X X X X X X X 5 Choreocolax polysiphoniae Reinsch X X X X X 4 Claihromorphnim. circumscri ptum (Stromfelt) Foslie X 1 Corollina officinalis L. X X X X X X 14 Cu stoclonium. purpureum. (Hu dson ) Batters val'. cirrhosum Har vey X X X X X X 15 Dasya pedicellata (C. Agardh) C. Agardh X ....., X X X 11 <: D ermat olith on pustulatum (Lamoureux) Foslie X X X 15 ?-- Dumontia incrossat« (Muller) Lamouroux X X X X -l X X 3 0> ...... TABLE I- Rhodophyta cant. <0 -'I Distribution ~ St ation IGeo­ Taxon 1 2 3 4 5 6 7 graphical ~ II> Gigartina stellata (Stackhous e) Batters X 4 '"1 Gloiosiphonia capillaris (Hudson) 5' (l) Carmichael ex Berkeley X X X 4 Gonioiricnum. alsidii (Zanardini) H owe X X X X 4 oq> II> Griffithsia i enuis C, Agardh X 11 (l) Grinne llia am eric ana (C, Aga r dh) Harvey X X X X X 11 I Hild en bra ndia pro totupus Nardo X X X X X 14 Q Hupnea musciiormis (Wulfen) Lamoureux X X 12 ro0 Lith oph yllum corallinae (Cronan) H eydrich X 15 S II> Liiho thamnium glaciale Kj ellman X 1 ;::l II> Lom entaria baileyana (Harvey) F arlow X X X X 16 ;::l Lom entaria orcadensis (Harvey) Collins ex Taylor X X 15 P­ ~ Melobesia leio lisii Rosan off X X X 16 II> Petrocelis midderu iorfii (Ruprecht) Kj ellman X 15 ::rc-t­ ro' Pliullophora brodia ei (Turner) Endlick- X X X X 5 CIJ 0 Phyll ophora m embromifolia (G oodenough ex ;::l Wo odward) J. Agardh X X X X X X 5 Phym at oli thon lenormomdi (Arescho ug) Adey X 6 01 Plumaria elegans (Bonnemaison) Schmitz X X 5 01 -l <:II TABLE I - Rh odophyta cont, <:II Distribu tion 00 Station ' Geo- Taxon 1 2 i$ 4 5 6 7 graphica l Poly ides roiundus (Hud son ) Gr evill e X X X X X X 4 Polysiph onia deniulata (Dillwyn) Greville ex Harvey in Hooker X X X X X H j Polusiphonio. elon gata (Hudso n) Sprengel X X X 11 Polysiph onia har veyi Bailey X X X X X 10 Polysiphonia lan osa (L.) Tandy X X X X X X X 5 ~ Polysiphonia niqr escens (Hudson) Greville X X X X X X X 10 ::r 0 Poly si ph onia novae-angliae Taylor X X X X 5 c, 0 Polysiphonia urceolaio. (Lightfoot ex Dillwyn) Gr eville X X X X X X 5 '"1 ~ Porphsrra Ieuc osticta Thu ret X X X 9 Por phyra miniata (C. Agardh ) C. Agardh X X x 1 Por ph.ura umbilicalis (L.) J. Agard h X X X X X X X 10 Rhodochorton peniciiliiorm e (Lightfoot) Rosenvinge X X 4 Rhodom ela con/ ervoid es (Hudson) Silva X X X X X 5 Rhodym enia palm ata (L.) Gr eville X X X X X X 5 r- Seirospora grijfithsiana (H arvey) Dixon X I Trtiilliella intricata (J. Agardh ) Batters X X X X X 5 ~ <: Subtotal 37 14 20 32 31 30 38 ?­ TOTAL 81 45 47 59 66 65 80 -'l 0> 1974] Marine Algae - Coleman and Mathieson 559

REFEREN CES

ADEY, W. H. 1964. Th e genus Phymatolithon in th e Gulf of Maine. Hydrobiologia, 24: 377-420. 1965. The genus Clathromorpku ra (Cor allinaceae) in the Gulf of Ma ine. Ibid. 26: 539-573. - - - -. 1966 . The genus Lithoihamnion, L eptopliutum. (nov. gen.) and Phymatolitho n in th e Gulf of Mai ne. I bid . 28: 321­ 370. ANONYMOUS. 1969a. Tide tables, hi gh and low wa ter predictions for 1969, No r th and Sou th America includ ing Gr eenl and . En­ vironm en tal Scien ces Services Adm inistration, Coa st and Geo­ detic Survey. 280 pp. 1969b. Tide tables, Cape Cod Can al. U. S. Army E ngi ne er Di vision, Ne w England Corps of Engineers. 12 pp . BELL, H . P., & C. MACFARLANE. 1933a . The marine algae of the maritime provinc es of Can ada, I. Li st of species with their distribution and pr evalence. Can . J our. Res. 9: 265-279 . - - --. 1933b. The mar ine al gae of the maritime provinces of Canada, II. A study of their ecology. Ibid. 9 : 280-293. BLOMQUIST, H. L., & H. J . HUMM. 194 6. Som e marine algae new to Be aufo rt, N OI'th Carolina. J our. Elisha Mit chell Sc i. Soc. 62: 1-8. CARDINAL, A. 1964. Etude sur les Ectocarpacees de la Manche. Beihefte zur Nova Hedwi gia, 15. 86 pp . . 1965 . Liste pr eliminaire des algues benthiq ues de la bai e des Chaleurs. Rapp. Ann. 1964 , Sta. Bio!. mar. Grande Riviere : 41-51. 1966. Additions a la liste benthiques de la baie de s Chaleurs. Ra pp. Ann. 1965, Sta . B ioI. m ar. Grande Ri viere : 35-43. 1967a . I nventaire d es a lg-ues m arin es benth iques de la ba ie des Chaleurs etc de la ba ie de Gaspe (Qu ebe c), 1. Pheo­ phyc ees. Nat. Ca n. 94: 233-271. J967b . I bid. II. Chlorophycees. Ibid. 94 : 447-46 9. 1967c. Ibid. III. Rhodophycees, Ibid. 94 : 735 -7 60. _ _ --c-. 191~& . RAl~.~~'tl\;l' '' dC' /' n l "" , ~~ ,,,,n.. : ~ hO ' 1-.." ,·11, :, ...... 1.. " ... . Liu Canada. Cah. nf., ~t. HlO l. mar. ~ r ande Riviere, Que . No. 48. 213 pp . CHAPMAN, V. J . 1964. The Algae. Macmill an and Company Ltd., London. 472 pp. COLLINS, F . S. 1900. P reliminar y lists of New England pla nts V. Ma rine algae . Rhodora 2: 41-52. 1909. The green algae of North Ame r ica. Tufts Col­ lege Stud ies. (Sci . Ser. ) 2 : 79-480. 560 Rhodora [Vol. 76

CONOVER, J. T. 1958. Seasonal gr owth of benth ic marine pl ants as related to enviro nm en tal f actors in an es tua r y. Publ. Inst. Mar. Sci. 5: 97-147. DAVIS, B. M. 1913a. General characteristics of the algal vegeta­ tion of Buzzards Bay and Vineyard Sound in th e vicinity of Woo ds Hol e. Dept. Corum. and Labor, Bull. (U.S.) Bur. Fish eri es 31: 443-544. 1913b. A ca talogue of the marine flora of 'Woods H ole and vicinit y. Dept. Comm. a nd Labor, Bull. (U.S.) Bu r . Fish eri es 31: 795-833. EDELSTEI N, T., L. CHEN, & .J. McLACHLAN. 1970. Investigations of the marine algae of No va Scotia , VIII. T he flor a of Digby N eck Peninsula , Bay of Fundy. Can. Jour. Bot. 48 : 621-629. EDELSTEIN, T., & J . McLACHLAN. 1966. Investigations of the mar ine algae of Nova Scotia 1. Winter flora of the A tl antic Coast, Can. .Tour . Bot. 44: 1035-1055. & ----- . 1967 a. Tbid. , III. Species of Phaeoph y­ ceae new or r are to Nova Scotia . I bid. 45: 203-210. - - - - - & 1967 b. l bid., IV. Species of Chlorophy­ ceae ne w or rar e to Nova Scotia. Ib id. 45: 211-214. --- - & 1968 a . lbid., V. Ad ditional species new or rare to Nova Scot ia. [bid. 46: 993-1003. ----- & 1968b . Ibid., VI. Some species new t o North Amer ica. Ibid. 47: 555-560. - - - - & 1969. Petroderma macu liforme on th e coas t of Nova Scotia. Ib id. 47: 56]-563. - -- -, ---- -, & J. S. CRAIGE. 1967. I nvestigations of the marine algae of No va Scotia II. Sp ecies of Rhodophyceae new or r a re to Nova Scotia. Ibid. 45: 193-202. - - - - - , , & 1969. Prelim in ary su rvey of the su bli ttor al flora of Halifax County. J. F ish. Re s. Br. Canad a 26: 2703-27 13. FARLOW, W. G. 1870 . XVII. List of sea-weeds or ma r ine al gae of the south coast of N ew England pp. 281-29 4. In: U. S. Corum. of F ish and F isheries. Commissioner's Rep ort 1871-1872. W ash­ in gton . Government P rinting Office. 1882 . 1. The marine algae of New England. Ib ul., report 1879. pp. 1-21 0. FRITSCH, F. E. 1935. Th e str uctur e an d repro duc tion of the algae, 1. xviii + 791 pp. Cambridge Univ. P r ess. 1945. Ibid.. II. xiv + 939 pp . Cambridge Univ. Press. H ARVEY, W. H. 1852-58. Nereis Boreal i-Americana. 1. Melano­ sp ermae . Smi thson i nn Contri b. Kn oiol., 3: 1-150, pl. 1-12. 1852; II. Rhodosnerm ae. tua .. 5: 1-258, p L 1.1-,'1 (;. 1853 ; III. Chlorospormae. Ibid., 10 ~ ii +- 1-140, pl. 37-50. 1858. 1974] Marine Algae - Coleman and Mathieson 561

HOEK, C. VAN DEN. 1963 . Revision of the European species of Cladophora. xi + 248 pp . Le iden . HOYT, 'Y\T . D. 1920. Marine algae of Beaufort. N. C. and adjacen t regions. Bull. U. S. Bur. Fish . 36: 367-556. HUMM, H. J . 1969. Distribution of marine algae along the Atlan­ tic Coa st of North America. Ph ycologia 7: 43-53. HUTCHINS, L. W. 1947. The bases for 'temper atur e zonation in geographical distribution. Ec oJ. Monog raph s 17: 325-335. LAMB, M., & H. ZIMMERMANN. 1964. Marine vegetation of , E ssex County, Ma ssachusetts. Rhodora 66: 217-254. LEE, R. K. S. ] 968. A collection of marine alga e from Newfound­ land, I. Introduction and Phaeophyta. Nat. Can. 95: 957-978 . 1969. Ibid ., II . Chlorophyta and Rhodoph yta. Ibid. 96: 123-145. LEWIS, I. F . 1914. Th e seasona l life-cycle of some red algae at Woods H ole. Pl an t World 17 : 31-35. MACFARLANE, C., & H. P. BELL. 1933. Observa tion of th e seasonal chang-es in t he marine algae in th e vicinity of Halifax with particular r eference to winter cond itions. Pro c. Nov a Scotian In st, Sci. 18: 134-176 . ---- , & G. M. MILLIGAN . 1965. Marine algae of th e Maritime Provinces of Canada. A prelimina rv chec klist. Nova Scotia Res. F ound., Seaw eed division , Halifax. Mimeographed, 24 pp. MATHIESON, A. C., & S. F ULLER . 1969. A preliminary inves t igation of the benthonic marine al gae of the Chesap eake Ba y Regi on. Rhodora 71: 524-534. - - --, C. DAWES, & H . J . H UMM . 1969. Contributi ons to the marine algae of Newf oundland. Rhodora 71: 110-159. NEWR OTH , P. R., & A. R. A. TAYLOR . 1971. The nom enclature of the North Atlantic species of Ptiullophorc: Gre ville, Ph ycologia 10: 93-97. PARKE, M., & P . S. DIXON . 1968. Checklist of British marine algae - second r evision. J. Mar. B ioI. Assoc. U. K. 48: 783-832, PARR, A. E. 1933, A ge ograph ic-ecological ana lysi s of the sea­ sonal changes in t emper ature conditions in the shallow wa ter al ong th e Atl an ti c Coast of the U . S. Bull. Bing-ham Ocean ogr, CoIl., 4, 90 pp. RHODES, R. G. 1970 . Sea sonal occurrence of marine algae on an oyster reef in Burston's Bay, Virginia, Che sapeake Sci. 2: 61-63. SEARS, J, R. 1971. Morphology, systematics and descri ptive ecology of the sublittor al benthic mar ine a lgae of southern Cap e Cod and adjacent islands. Ph.D. Th esis, 295 pp. University of Massa­ chusetts. 562 Rhodora [Vol. 76

SETCHELL, W. A. 1917. Geographical distr ibution of the marine a lgae. Science 4!l: 197-204. 1920. Stenothermy and zone-inva sion . Am. N a t. 54 : 385-397. 1922. Ca pe Cod in its relat ion to the marine flora of N ew E ngland. Rhodor a 24: 1-11. SOUTH, G. R., & A. CARDINA L, 1970. A checklist of m arine algae of ea stern Canada. Can . .J. Bo t. 48: 2077-2095. STEPHENSON , T. A. 1944. The cons t itu tion of the inter ti dal fauna an d flor a of South Afr ica. P ar t II. Ann. N atal. Mus. 10: 261-358. -----, & A. S TEP HEN~ON . 1949. The un iver sal featu res of zona tion betwee n tide-mark s on rocky coa sts. J our . Eco!. 37 : 289-305. STONE, R. A., E. H EHRE, .J. CON WAY, & A. C. MAT HIESON. 1970. A preliminary checklist of the marine algae of Campobello I sland. N ew Brunswick , Canada. Rhodor a 72: 314-338 . TAYLOR, W. R. 1937. The marine algae of th e nor theaster n coas t of N or th America. 405 pp. U niv, of Michigan Press, Ann Arbor. 1957. Marine algae of the northeast coast of Nor th Ameri ca . 509 pp. An n Arbor Press . 1960. Ma r ine al gae of 'the ea stern t r opical and sub­ tr opical coasts of the Amer icas. 870 pr. Ann Arbor P r ess. WELL, H. W., & I. E. GRAY. 1960. Summer up wellin g off the northeast coast of Nor th Ca rolina. Limno!. Oceanogr . 5 : 433­ 468. WI LeE, R. T . 1959 . The marine algae of th e Labrador Pe ninsula and northwest N ewf oundland (ecol ogy and distribution) . Nat . Mu s. Canada 158: 1-103. 1965. Studies in th e genus L am ina ria III. A revision of the North Atlantic species of the sim plices section of Lami­ n ar ia. Bota nica Gothobur gensia III. pp. 247-256. WILLIAM S, L. G. 1948. Seasonal alternat ion of marine flor as at Cape Lookout, No rth Ca rolina. Am. Jour, Bot. 35: 682-695. ----. 1949. Mar ine algal ecology at Cap e Lookout, Nor th Carolina. Bull. Furma n Uni v. 31 : 1-21. \VULFF, R. L ., E. M. T. WULFF, B. H . ROB INSON, .J. K. LoWRY, & H . J . HUMM. 1968. Summer marine a lgae of th e j ett y at Ocean City. Ma ryla nd. Chesapeake Sci. 9 : 56-60. WYNNE, M. .J., & W. R. TAYL OR , 1973. The status of Agm'dhielLa tenera and AgardhielLa boil euo. (Rhodoph yta, Gigart inales ). H ydr obiologica -13: 93-107 , ZANEVELD, .J. S., & W. D. BPRNES. 1965. Reproductive periodici ties of some benthic algae in lower Ches apeake Bay. Chesapeake Sci. 6: 17-32. 1974J Marine Algae - Coleman and Mathieson 563

Z ANE VELD, J. S. 1965. Th e benthic marine alga e of Virginia. Vir­ gi nia J our. Sci. 1965: 346. - ---. 1966a. T he mari ne a lgae of the American coast be­ tween Cape Ma y, N . J. and Cape H at teras, N . C., 1. The Cya no­ ph yta. Bot. Mar. 9: 101-128. - -- - . 1966b. The benthic algae of Delaw are. A pre liminary checklist. 35 pp. Scientific Series, Pub!. no. 2, Inst it, of Ocean­ ogr., Old Domi nion College, Norfolk, Vir ginia. --- - . 1972. Th e benthic marine alg ae of Delaware, U.S .A. Chesapeak e Sci. 13: 120-138.

JACK SON ESTUARINE LABORATORY UNIVERSITY OF NEW HAMPSHIRE DURHAM, NEW HAMPSHIRE 03824