The distribution of Phyllophora in the North Atlantic and Arctic regions

P. R. NEWROTHI Department of Biology, University of New Brunswick, Fredericton, New Brunswick

Received January 5, 1971

NEWROTH, P. R. 1971. The distribution of Phyllophora in the North Atlantic and Arctic regions. Can. J. Bot. 49: 1017-1024.

The distributions of five species of Plzyllophora Greville (Order Gigartinales) were examined in the North Atlantic and Arctic regions. Less detailed distributional observations of Pf7yllophora were made in the Mediterranean and Black Seas. Distributional records, based on extant specimens from extensive personal field observations in the United Kingdom and eastern North America and from collections in various herbaria, are recorded in tables and on maps; these records are discussed.

Introduction

Phyllophora Greville (Order Gigartinales, Family Phyllophoraceae) is well represented in the North Atlantic. Five species of Plzyllophora were listed by Parke and Dixon (1968) for the British Isles. Nomenclatural studies (Newroth 1970; Newroth and Taylor 1971) have shown that three of these, P. crispa (Huds.) Dixon, P. palmettoides J. Ag., and P. traillii Holm. ex Batt. are legitimate names. However, two new com- binations were proposed: P. truncata (Pallas) Newroth et A. R. A. Taylor (= P. brodiaei (Turn.) Endl., = P. interrupts r rev.) J. Ag.) and P. pseudoceranoides (Gmelin) Newroth et A. R. A. Taylor ( =P. rnernbranfolia (Good. et Woodw.) Endl.).

The literature does not provide a summary of the distribution of Phyllophora in the North Atlantic and Arctic regions. Moreover, because of the difficulty in identifying several species, there are numerous cases of incorrect citations. In this paper, observations on the distributions of the five species listed by Parke and Dixon (1 968) are reported.

Materials and Methods In the period from 1965 to 1970 many sites on the east-

ern seaboard of North America and in Britain were ex- amined for Phyllopl7ora by the author. Eighty-four collections were made at 35 localities from Rhode Island to Newfoundland, and. in the British Isles, 142 collections were made at 53 localities. Single collections were made at Ple Tatihou, Cherbourg Peninsula. France, and Bayona, Spain. In 35 of these collections material was collected by diving, 28 using SCUBA.

After studying Phyllophora specimens in these collec- tions (included in Tables 1 and 4), the author examined

l ~ r i s e n t address: Department of Botany, University of British Colunlbia, Vancouver 8, B.C., Canada.

collections in 38 private and institutional herbaria in North America and Europe. Attempts were made to study all the collections of P/~yllophora from Arctic and North Atlantic areas.

Distribution lists (Tables 1 to 4) were compiled from specimens observed in herbaria or collected by the author. Specimens of doubtful attribution were not included. The location of a voucher specimen is indicated by the ac- cepted herbarium abbreviation (after Lanjouw and Stafleu 1964) or an unofficial abbreviation placed in parentheses. A key to the unofficial abbreviations has been provided in Table 5.

The maps in Figs. 1 and 2 illustrate the distribution of Phyllophora in eastern North America and in the British Isles.

Observations and Discussion EASTERN CANADA AND UNITED STATES

The distributional records of Plzy/lophora for this area are listed in Table 1 and shown on Fig. 1 .

Taylor (1957) outlined the North American ranges of P. pseudoceranoides (as P. rnernbranfo- lia), P. truncata (as P. brodiaei and P. interrupta), and P. traillii. More recent work has considerably expanded knowledge of the distribution of Phyl- lophora. Hehre and Mathieson (1970) reported their investigation of the marine flora of New Hampshire, while MacFarlane, Cardinal, and Edelstein et a[. have contributed much to knowl- edge of Phyllophora distributions in Eastern Canada.

Bell and MacFarlane (1933) and MacFarlane and Milligan (1965) reported the presence of P. pseudoceranoides (as P. ~nernbranfolia) and P. trurzcata (as P. brodiaei) in the Gulf of St. Law- rence, and P. pseudoceranoides (as P: rnernbrani- folia) in the Bay of Fundy and the Atlantic coast of Nova Scotia. Cardinal (1965, 1967) reported P. pseudoceranoides (as P. rnernbranfolia) and P. truncata (as P. brodiaei) from the Bay of Chaleur

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1018 CANADIAN JOURNAL OF BOTANY. VOL. 49. 1971

and GaspC. Edelstein and McLachlan (1966) re- corded these species from Halifax Co., N.S., and later (Edelstein and McLachlan 1968) reported P. traillii from Digby Co., N.S.

Newroth and Taylor (1968) first cited P. trun- cata (as P. brodiaei) from the Bay of Fundy and also presented additional data on the distribution of this species and Ceratocolax hartzii Rosen- vinge. Additional new records of Phyllophora- ceae in Eastern Canada include P. traillii and Gymnogongrus norvegicus (Gunnerus) J. Ag. from Point Lepreau, St. John Co., N.B. The Nova Scotia record of P. traillii, cited by Edel- stein and McLachlan (1968) and later by Edel-

stein et al. (1970), appears to be incorrect. Examination of their specimen (also illustrated, Edelstein and McLachlan 1968) suggested that it may be a juvenile specimen of P. truncata with developing nemathecia.

The Delaware record of P. pseudoceranoides (Table 1) is the southernmost record of Phyllo- phora in the western Atlantic.

ARCTIC REGIONS

As indicated by Newroth and Taylor (1968), the distribution of P. trurzcata is truly circum- boreal. However, the known northern distribu- tion of the other species of Pi~yllophora is more

TABLE 1 The distribution of Pl~yl/op/zora in Eastern Canada and the United States.

(Letters following the specific epithets are used in Fig. 1 to designate the localities where they have been found)

Locality P. tr~tncata (a) P.psettdocerat~oides (b) P. traillii (c)

United States Maine NH A TCD New Hampshire (PN), NHA (PN), NHA Massachusetts (PN), US, UC (PN), US, UC NY

(PW, NHA, L

Rhode Island (PN), UC, BM (PN), CANA, (F) NY Connecticut New York UPS, NY 3 $4 (F)

New Jersey US US, UC, TCD Delaware NHA

Canada 1. New Brunswick

Charlotte Co. St. John Co.

(UNB)

Westrnorland Co. (PN), (UNB) (PN), (UNB) (PN)

Kent Co. (UNB) (UNB)

Gloucester Co. 2. Nova Scotia

(PN) (PN), (AC), (NSRF)

Curnberland Co. Kings Co. Annapolis Co. Digby Co. Shelburne Co. Lunenburg Co. Halifax Co. Victoria Co. Inverness Co. Antigonish Co. Pictou Co.

3. Prince Edward Island

4. Newfoundland and Labrador

West Coast East Coast, Avalon

Peninsula Labrador

5. Quebec Bay of Chaleur and

Bay of GaspC St. Lawrence Estuarv

(NRcH) (UNB), (MUN) (NSRF) NY CANA (PN), (NRCH) (NRCH) (PN) CANA

(PN)

(NRCH), (NSRF) (NRCH) (NSRF) (PN), (NRCH), (PN), (NRCH), (NSRF), (UNB) (UNB)

(AC) CANA, (F), NY (NSRF) (AC) Magdalen Islands

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NEWROTH: PHYLLOPHORA DISTRIBUTION 1019

limited (see Tables 2 and 3). Blargesen and northwest Newfoundland. Zinova (1955, 1957) J6nsson (1905), in summarizing many of the summarized the distribution of P. truncata (as P. earlier reports, gave a general listing of Phyllo- brodiaei and P. interrupta) and other Phyllopho- phora in the Arctic Sea and North Atlantic. raceae in the Soviet Arctic. Wilce (1959) reported the presence of P. truncata (as P. brodiaei and P. interrupta) from Labrador EUROPEAN COASTS

and Ungava Bay, and P. pseudoceranoides (as P. The distribution of Phyllophora has been membranifolia) from southern Labrador and studied extensively in Scandinavia. Rosenvinge

TABLE 2 The distribution of Phyllophora trlrncata in Arctic North America and Greenland

Location of Locality Coordinates specimens

A. Canada Alaska Harbour,

Hudson's Bay S.E. Foxe Basin

53.21 N, 79.01 W NY, CANA 66.46 N. 79.15 W US

Pingil-Kalik, Foxe Basin 68.00 N; 84.00 W US Fury and Hecla Strait 69.56 N, 84.00 W US Orfot Island, Hudson's Bay 63.55 N, 90.20 W UC Port Kennedy 72.40 N, 93.00 W BM Fort Churchill, Manitoba 58.46 N, 94.10 W CANA Coronation Gulf 69.10 N, 105.51 W Dolohins-Union Strait 69.05 N. 114.45 W

Cpw CANA

-~ - .~-- " -~ - ~ - - - - - .~~ -.-- - -

Eglington Island 78:48 N: 118.30 W CANAb Prince Patrick Island 76.06 N, 119.03 W CANAb Amundsen Gulf 69.49 N, 122.44 W CANA Sachs Harbour, Banks Island 71.59 N, 125.00 W CANAb

B. Alaska Camden Bay Point Barrow Port Clarence

C. Greenland Proven Anmagasslik

70.00 N, 145.00 W CAN A 71.22N, 156.30W US, UC 65.00 N, 166.00 W CANA

anm marks Havn 70.30 N; 26.20 W C, UC Brsnlunds Fjord 82 .30N,26 .00W C

.Specimen from Dr. J. G . Hunter, Arctic Biological Station, Fisheries Research Board of Canada. bspecimens from Dr. R. K. S. Lee, National Museum of Natural Sciences, Ottawa, Ontario, Canada.

TABLE 3 The distribution of PlzyllopRor.a in the U.S.S.R., Iceland, Spitzbergen, The Faroes, and Continental Europe

Locality P. trlmcata P. pseudocerar~oides P. traillii P. crispa P. palmettoides

White Sea, U.S.S.R. Novaya Semla, U.S.S.R. Kara Sea, U.S.S.R. Iceland Spitzbergen The Faroes Norway Sweden Denmark Germany Helgoland France (Atlantic)

UC, L UC L C UC. UPS L, c UPS UC, UPS n L

UC, BM, US TCD

UPS, C L, UPS, C

Spain Mediterranean Sea Black Sea

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1020 CANADIAN JOURNAL OF BOTANY. VOL. 49, 1971

(1 931) gave excellent discussions of the Phyllo- Rosenvinge have not been located in Copenhagen phoraceae in Danish waters, and established the presence of P. truncata (as P. brodiaei), P. pseudo- ceranoides (as P. membranfolia), and P. crispa (as P. epipllylla). He also claimed the presence of P. traillii, and illustrated plants which appear to be this species, but are much smaller than those nor- mally encountered by the author in other areas. Specimens of P. traillii froin the locality cited by

or other major algal herbaria. Until further col- lections can be made, this record must remain in doubt. Because the author did not collect Plzyllo- phora in the Baltic Sea, no further comments on its distribution in this area are presented.

Levring (1937) and Jaasund (1965) reported P. tnuzcata (as P. brodiaei) from the Norwegian coast. In addition to this species, herbarium

TABLE 4 The distribution of Plzyllophora in the British Isles. (The letters beneath the specific epithets are used in Fig. 2 to

designate the areas where the species have been found)

P. trrrncatn P. pse~rdoceranoicles P. traillii P. crispn P. palmettoides County (0) (b) (c) (4 ( 4

A. England Cornwall Devon Dorset Durham Essex Hampshire Kent Northumberland Suffolk Sussex Yorkshire Isle of Man Channel Islands Scilly Isles

B. Wales Anglesey Caernarvon Cardigan Pembroke

C. Scotland Aberdeen Angus Argyll A Y ~ Berwick Bute Dumfries East Lothian Fife Inverness Midlothian Moray Nairn Orkney Wigtown Zetland

D. Ireland Antrim Clare Cork Donegal Dublin

(PN), BM BM, (MBA)

BM BM, E BM

BM, LIVU LIVU BM, TCD ( P N ) O (PN)

CpN), LIVU (PN)

E STA BM. TCD

E BM, STA

E (PN), BM BM. E

BM, E E

(MV) GL STA, (OM) E

Galway Kerry BM

(MV) GL, (MV)

Mayo BM BM BM, (My) Sligo Wexford (OM)

(MV) . ,

.Specimen collected by W. Farnharn.

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NEWROTH: PHYLLOPHORA DISTRIBUTION 1021

specimens of P. pseudoceranoides and P. crispa from that area were observed by the author. J6nsson (1901) claimed that specimens of P. crispa from Iceland, housed in the Botanical Institute, Copenhagen were doubtful. Perhaps these and other specimens in Leiden and Upp- sala were mislabeled.

The southern range of Phyllophora on the European coast has not been sharply defined in relation to the Mediterranean flora. Table 3 indicates that P. pseudoceranoides, P. traillii, P. crispa, and P. palmettoides were present on the Atlantic coast of France. Phyllophora heredia (Clemente) Endlicher, not examined in detail in this research, is common in the Mediterranean and also occurs on the southern Atlantic coast of France (Schotter 1968).

In the Mediterranean, Plzylloplzora appears to be restricted to P. heredia, P. crispa forma nervosa, and P. palinettoides. Two additional species of Phyllophora, P. jiinbriata ErcegoviC (1949) and P. aegei Giaccone (1968), were re- ported from the Adriatic Sea. As they are poorly known and their systematic position is uncertain, their distributions are not discussed here.

TABLE 5 List of herbarium abbreviations (unofficial)

Institutional UNB: University of New Brunswick, Fredericton,

N.R. MUN: Memorial University of Newfoundland, St.

John's, Newfoundland. NRCH: Atlantic Regional Laboratory, National Re-

search Council of Canada, Halifax, N.S. NSRF: Nova Scotia Research Foundation, Halifax,

N.S. F: ailo ow Reference Library and Herbarium of

Cryptogamic Botany, Harvard University, Cambridge, Mass., U.S.A.

MBA : Marine Biological Association of the United Kingdom, Plymouth.

UCNW: University College of North Wales, Bangor. Private

PN: Peter R. Newroth, Botany Department. Uni- versity of British Columbia, Vancouver, B.C.

AC: Andrt Cardinal, Departement de Biologie, UniversitC Laval, Quebec, Que.

MM: Margaret T. Martin, Botany Department, University College of North Wales, Bangor, Wales.

MV: Mairin de Valera, Botany Department, Uni- versity College, Galway, Eire.

OM: Osborne Morton, School of Botany, Trinity College, Dublin, Eire.

Dl: David E. G. Irvine, Northern Polytechnic, London.

FM: Francis Magne, Laboratoire de Botanique, UniversitC de Rennes, Rennes, France.

The Black Sea flora includes P. truncata and P. crispa, as reported by Mikhoulov and Mash- takova (1966), and Zinova (1967) also records P. membranifolia (=P. pseudoceranoides). How- ever, this is perhaps erroneous, as her figure (p. 261, Fig. 156) of this species has cystocarps labeled by her as "nemathecia." It is possible that a misidentification was made. Furthermore, no specimens of P. pseudoceranoides have been seen from the Black Sea.

BRITISH ISLES

Although the algae of this area have received considerable study, no overall survey of Phyl- loplzora has been made. Evaluation of cited literature records not substantiated bv extant specimens is difficult, and it appears that many workers have misidentified Phyllophora.

The citation, in numerous publications, of P. truncata from the south coast of England, is a good example of the persistence of incorrect old records. After examining many herbarium speci- mens, and collecting on the south coast, the author agrees with Cotton (1912) that P. truizcata occurs only in more northern waters (see Fig. 2). The recent citation of P. truncata (as P. brodiaei) from the Scilly Isles (Russell 1968) appears to be erroneous, based on a mis- identification of Rhodymenia pseudopalnzata (Lamour.) Silva (personal communication from Mrs. L. M. Irvine, British Museum (N.H.)). Cystocarpic specimens of this species closely resemble P. truncata. The author found many specimens of Rhodymenia pseudopalmata and P. palinettoides in algal herbaria were mis- identified as P. truncata.

Conclusions Although the range of distribution of the five

species of Phyllophora recorded here will no doubt be extended by future research, these species appear to occur in distinct areas. Only P. truncata shows a marked discontinuity in its distribution. I t is circumboreal in the Arctic and Atlantic Oceans, extending south to New Jersey in the western Atlantic (Table 1, Fig. 1) and to Counties Wexford and Kerry, Eire, and Caernar- vonshire, Wales, in the eastern Atlantic (Table 4, Fig. 2). This species also occurs in the Black Sea (Table 3), although there is no evidence that it is present in the Mediterranean Sea.

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1022 CANADIAN JOURNAL OF BOTANY. VOL. 49. 1971

FIG. 1. The distribution of Plzyllophora in eastern Canada and the United States.

P. pseudoceranoides is amphi-Atlantic, with its northern limit in Newfoundland and its southern limit in Delaware (Table 1, Fig. I), in the western Atlantic. In the eastern Atlantic, it occurs as far north as Iceland and as far south as the coast of France (Table 3).

P. traillii has been found in the western Atlantic from New Bruilswick south to Con- necticut (Table 1, Fig. 1) and in the eastern Atlantic from the Faroes to the south coast of England and the coast of France (Tables 3 and 4, Fig. 2).

The distribution of P. palnzettoides appears to be restricted to the southern part of Britain

(the most northerly record is from Wales, Table 4, Fig. 2), the French Atlantic coast, and the Mediterranean Sea (Table 3).

P. crispa is widespread in the eastern Atlantic, from its northern limit in Iceland (Table 3) to the southern limit in North Africa. This species is also present in the Mediterranean and Black Seas.

Acknowledgments

I am indebted to Dr. A. R. A. Taylor for his guidance during this research, and to Dr. K. Cole for her assistance in the preparation of the manuscript. Thanks are also due to the numerous

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NEWROTH: PHYLLOPHORA DISTRlBUTlON

FIG. 2. The distribution of Phyllophorrr in the British Isles. KEY TO SPECIES: a, P. fruncrrfa; b, P. pseudo- cerartoicies; c, P. fraillii; ci, P. crispa; e , P. pab?zetfoicies.

persons who assisted me in making field col- lections and to Mrs. Jana Bance for her help in the translation of papers written in Russian.

Loans of herbarium specimens and permis- sion to examine specimens in many herbaria are acknowledged with thanks. Thanks are also due to colleagues who permitted examination of personal herbaria, and to Professor D. J. Crisp and Dr. Eifion Jones for making available the facilities of the Marine Science Laboratories, University College of North Wales, Menai Bridge, Anglesey, Wales, U.K.

A National Research Council of Canada Scholarship provided financial support during this research.

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