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J. Linn. SOC. (Bot.), 60, 383, p. 261 With 6 plalea Printed i n Great Britain Februnry, 1988 25 1 Taxonomic and nomenclatural notes on the genus Cystoseira C.A& BYMARGARET ROBERTS, F.L.S. Department of Botany, University of Hull (Accepted for publication March, 1987) Taxonomic and nomenclatural confusions encountered in a study of the British species of Cyatoaeira are discussed; a number of non-British species, problems of whose synonymy have become involved with those of the British species, are also included. Fucw concatenutw, L. is shown to be a synonym of F. foeniculaceus L. The combination Cyatoeira nodkaulb (With.) M. Roberts replaces C. grnnulata (L.) Orev., as the name C. granulala is illegitimate and its type does not belong to this species. Fucua aelaginoides L. is the earliest description of Cyatoaeira tanlariscifolia (Huds.) Papenf., but the combination C. aelaginoides cannot here be used since Naccari had earlier employed the name to described another species. The identity of the type specimen of Fucw barbatw L. and other descriptions of this species are considered. Cyatoaeira firnbriata (Desf.) Bory is the correct name for the species often referred to &8 C. abrohnifoh C.Ag. The species currently known aa c. discore (L.) Ag. has no legithate name. Cyatoeedra wmeoidea (L.) M. Roberts is shown to be the correct name of the species known aa C. concalenala Ag. Cyatoaeira selaginoidea Nacc., C . aauvageauanu Hamel and C. hppG mnsu Valiante are briefly discussed; the last has no legitimate name. CONTENTS Introduction . Systematic descriptions . C'yatoaeirafoeniculacea (L.) Orev. . (lyatoaeiramym'ophylbides Sauv. . C'ystoaeira rwdicaulia (With.) M. Roberts Cyntoaeira tamariscifolia (Huds.) Papenf. Cyatoaeira barbata Ag. . Cyel~seirajmbrida (Desf.) Bory . cy8lo8eiTa di8COT8 WnSU SaUVagefLU. . Cyatoaeirausneeidea (L.) M. Roberts . Oyatoaeira aelaginuidea Nacc. . Cyatoeeira aauvagmwna Hamel. . Cyatoaeira hoppii sensu Valiante . Acknowledgements . . . References . . . PAQE . . . . 261 . . . . . 262 . 262 . 264 . 264 . 266 . 267 . 268 . 269 . 269 . 281 . 281 . . . 282 . . . 282 . . . . 282 INTRODUCTION These notes give an account of some of the taxonomic and nomenclatural confusions which have been encountered during the preparation of a series of papers on the British species of Cystoseira. Past taxonomic confusion in this genus has been due partly to the inability of early workers to recognize specific limits with any certainty, because of the great range of morphological form frequently seen within a single species, and partly to lack of information on valid criteria for specific separation. In addition there has been nomenclatural confusion because of incorrect application of specific epithets, very often resulting from failure to consult the original type specimensof the description concerned,

Taxonomic and nomenclatural notes on the genus Cystoseira C.Ag

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Page 1: Taxonomic and nomenclatural notes on the genus Cystoseira C.Ag

J . Linn. SOC. (Bot.), 60, 383, p. 261 With 6 plalea Printed in Great Britain Februnry, 1988

25 1

Taxonomic and nomenclatural notes on the genus Cystoseira C.A&

BY MARGARET ROBERTS, F.L.S.

Department of Botany, University of Hull (Accepted for publication March, 1987)

Taxonomic and nomenclatural confusions encountered in a study of the British species of Cyatoaeira are discussed; a number of non-British species, problems of whose synonymy have become involved with those of the British species, are also included. Fucw concatenutw, L. is shown to be a synonym of F. foeniculaceus L. The combination Cyatoeira nodkaulb (With.) M. Roberts replaces C. grnnulata (L.) Orev., as the name C. granulala is illegitimate and its type does not belong to this species. Fucua aelaginoides L. is the earliest description of Cyatoaeira tanlariscifolia (Huds.) Papenf., but the combination C. aelaginoides cannot here be used since Naccari had earlier employed the name to described another species. The identity of the type specimen of Fucw barbatw L. and other descriptions of this species are considered. Cyatoaeira firnbriata (Desf.) Bory is the correct name for the species often referred to &8 C. abrohnifoh C.Ag. The species currently known aa c. discore (L.) Ag. has no legi thate name. Cyatoeedra wmeoidea (L.) M. Roberts is shown to be the correct name of the species known aa C. concalenala Ag. Cyatoaeira selaginoidea Nacc., C. aauvageauanu Hamel and C. h p p G mnsu Valiante are briefly discussed; the last has no legitimate name.

CONTENTS

Introduction . Systematic descriptions .

C'yatoaeira foeniculacea (L.) Orev. . (lyatoaeira mym'ophylbides Sauv. . C'ystoaeira rwdicaulia (With.) M. Roberts Cyntoaeira tamariscifolia (Huds.) Papenf. Cyatoaeira barbata Ag. . Cyel~seirajmbrida (Desf.) Bory . cy8lo8eiTa di8COT8 WnSU SaUVagefLU. . Cyatoaeira usneeidea (L.) M. Roberts . Oyatoaeira aelaginuidea Nacc. . Cyatoeeira aauvagmwna Hamel. . Cyatoaeira hoppii sensu Valiante .

Acknowledgements . . . References . . .

PAQE . . . . 261 . . . . . 262 . 262 . 264 . 264 . 266 . 267 . 268 . 269 . 269 . 281 . 281 . . . 282 . . . 282 . . . . 282

INTRODUCTION

These notes give an account of some of the taxonomic and nomenclatural confusions which have been encountered during the preparation of a series of papers on the British species of Cystoseira. Past taxonomic confusion in this genus has been due partly to the inability of early workers to recognize specific limits with any certainty, because of the great range of morphological form frequently seen within a single species, and partly to lack of information on valid criteria for specific separation. I n addition there has been nomenclatural confusion because of incorrect application of specific epithets, very often resulting from failure to consult the original type specimens of the description concerned,

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262 M. ROBERTS

The extent of the confusion may be seen from a study of synonymy in many early accounts. Speciesrepresented in the British Isles are dealt with first: non-British species mentioned

in these sections are dealt with more briefly at the end.

SYSTEMATIU DESCRIPTIONS

Cystoseira foeniculacea Cystoseira foeniculaceu (L.) Grev., AZg. Brit., 6 (1830) Savageau in Bull. Stat. Bio2.

d’drcachn, 14: 437 (1912). Fucus foeniculaceus L., Sp. PI., 2: 1161 (1753). Fucus abrotanifolius L., Sp. PI., 2 : 1161 (1753). Fucus concatenatus L., Sp. PI., 2: 1160 (1753). Fucus discors L., Syst. Nut. ed. 12,2: 717 (1767). Cystoseira abrotanifolia (L.) Ag., Sp. AZg., 1: 63 (1820). Cystoseiru concutenata (L.) Ag., Sp. AZg., 1 : 57 (1820) excl. descr. Cystoseira discors (L.) Ag., Sp. AZg., 1: 62 (1820).

‘ 17. FUCUS vesiculis ovetis pedunculatis, foliis multi- fidis linearibus. Fucus folliculaceus, foeniculi folio. Bauh. Pin. 365 Habitat in Oceano.’

Fucw foeniculaceus was described by Linnaeus (1753: 1161) as:

Linnaeus is known to have based his ideas on the species in Bauhin’s Pinaz very largely on the specimens in Burser’s Herbarium (of. Stearn, 1957: 116; Savage, 1935: 19). In Burser’s Herbarium, Vol. XX Sheet 93 is labelled ‘Fucus folliculaceus Foeniculi folio Bauh. Ad pagum Perault in mari’ (Juel & Svedelius, 1936 : 136) and in his manuscript list of determinations of the plants in Burser’s Herbarium Linnaeus identified this as ‘Fwus foeniculaceus’ (Savage, 1937 : 69) ; this specimen should thus be designated as holotype of this species. This appears to be a good specimen of the species Cystoseira foeniculaceu (L.) Grev., as now recognized.

Fucw abrotanifolius was also described by Linnaeus in 1753 (p. 1161) as: ‘ 18. FUCUS fronde bipinnata subfiliformi:

apicibus vesiculosis dilatatis. Loef. epist. 1.4. Habitat in m r i Anglico.’

This diagnosis was repeated in 1759 (p. 1345) and 1763 (p. 1629) without significant variation but in 1767 a (p. 716) it was expanded and concluded with the sentence. ‘Fucus pinnatus, ramis dichotomis: extremitatibus dilatato-vesiculosis. h e $ . it. 174’. This is a direct translation from Lofling’s Iter Hispanicum (1758), and Sheet 1274:95 in the Linnaean Herbarium, which is a rather denuded fertile plant of Cystoseira foeniculacea (L.) Grev., carries a similar form of words in Lofling’s hand. Linnaeus accepted Lofling’s manuscript diagnosis as the two had collaborated. This specimen is therefore the holotype of Fucus abrotanifolius L. ; i t is unquestionably native to Great Britain and is conspecific with F. foeniculaceus L.

In 1767a (p. 717) F. discms was described by Linnaeus for the f is t time as: ‘48.F. fronde tereti inermi aculeatissima, foliis distichis subpinnatis linearilanceolatis serratis. Cadis pal- mris , tererr, undique vestitwr ramentis s. mu- leis inemibus, confertissimis: Folk distich posi- ta, plana, laevia linmri-lanceolata, serrata & haec swpe parum pin&.’

The type of this description is Sheet 1274:21 in the Linnaean Herbarium on which are mounted two specimens. The left-hand one is a fragment of a flattened juvenile lateral

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Notes on Cystoseira C.Ag. 253

branch system of Cystoseira foeniculacea (L). Grev. of the form commonly found off the coast of Devon. The other is a member of the Rhodophyceae and is ringed round in pencil 80 that it is excluded from the part of the sheet which is labelled ‘discors’ in the hand of Lofling (Savage, 1945; 200).

All these descriptions thus refer to the same species, that currently known as C. foeniculacea .

After the publication of the Species Plantarum, Linnaeus added a specimen to the herbarium (1274: 14) and labelled it Fucus concutenatus. This specimen was collected in Spain by Alstromer, and probably reached Linneaus in about 1761 (Savage, 1945, Key to list of Specimens). This specimen was taken by Goodenough & Woodward (1797 : 135) to be the type of F. concutenatus L., and in this they were followed by Agardh (1820: 57). This specimen, however, belongs to species whose correct name appears to be Cystoseira usneoides (4.v.). The name Fucus concatenatus was also applied by various early British authors to Cystoseira nodicaulis (4.v.).

Pucus concutenatus however, was originally described by Linnaeus in 1753 (p. 1160) as: ‘ 10. FUCUS caule tereti ramosissimo, fructificationibus

oblongis concatenatis. Roy. Lugdb. 614. Habitat in Oceano.’

This description is taken almost verbatim from that of van Royen (1740: 514) with only the substitution of the term ‘wncdenatis’ for the ‘catenulatis’ of van Royen. Linnmus presumably accepted van Royen’s description because he had himself assisted in its preparation (Dixon, 1964 : 57).

The type of Linnaeus’s description is thus the specimen of van Royen (Pl. l), now in the Rijksherbarium, Leiden (Herb. Lugdb. 910. 153-1332), and this consists of two entire fertile primary branches of the alga currently known as Cystoseira foeniculacea (L.) Grev. (PI. 2). This specimen does not entirely agree with the published description, as its ultimate ramuli consist of concatenate vesicles terminated by small fusiform receptacles. Possibly the distinction between cryptostomata-bearing vesicles and the terminal receptacles was not realized at the time of Linnaeus’s description. LinnaeuR repeated this diagnosis in 1759 (p. 1344) and 1763 (p. 1628), but in 1767a (p. 717) gave an expanded description using only the term Lvesicle’ and omitting ‘fructification’. The specimen in Linnaeus’s own herbarium must be assumed to be a determination from memory and erroneous (cf. Dixon, 1964: 59).

Early accounts of this species from British waters include those of Stackhouse (1801, PIS. 14, 17), who identified two forms from British waters, and, whilst identifying them with the Linnaean species Fwus abrotanifolius and P. diswrs, suggested that F. diswrs might not be a good species, but merely a developmental stage of F. abrotanifolius Smith & Sowerby (1810a, pls 2130 and 2131) also recognized F. abrotunifolius and F. discors, but called the latter a ‘supposed species’ and quoted Mrs GriBth’s observations that it was merely a spring form.

Turner (1802) also recognized both F. discors and F. abrotunifolius as distinct species, but in 1819 (pl. 252) described the species as F. foeniculaceus with two varieties, abrotanifolius and diswrs, corresponding to the Linnaean species. This is the first treatment of the Limaean names as synonymns, and this choice of epithet should be followed. Greville (1830: 7) first made the correct combination Cystoseira foeniculacea, and included the C . diswrs and C . abrotanfolia of Agardh as synonyms.

Cystoseira discors (L.) Ag. was incorrect when published since Fucus foeniculacew is quoted as a synonym. It is, however, a legitimate name, since its basionym is legitimate (Lanjouw et al., 1961, Art. 63, Note). If its type specimen were considered to belong to a species other than that to which the types of F. foeniculaceus, F. abrotanifolius and P. concatenatus belong, then it would be the correct name for the species (seepp. 259,260).

Cystofleira abrotanifolia (L.) Ag. included both the Fwus abrotunifolius of Linnaeus and 17

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264 M. ROBEBTS

the F. $mbri&w of Desfontaines (1798-99 : 423), and was found in the eastern Atlantic south of Cadk and in the Mediterranean.

Greville’s inclusion of Agardh’s Cystoeeira abrotanifolia within C. foeniculaceu is now generally accepted, but with the exclusion of the Mediterranean element. This is believed (Sauvageau, 1912: 434) to be distinct, though related, species, to which the name F w w , fimbri&w of Desfontaines applies (sea p. 259).

Although Linnaeus’s own specimens are not the type specimens of Cystoseira foeniculacea (L.) Grev., a consideration of them seems appropriate since they have been consulted from time to time by botanists as a bash for their interpretation of the species.

Sheets 1274 : 4,5,6 and 7, in the Linnaaan Herbarium are all labelled Fucw foeniculacew. Sheet 7 is a single, fertile lateral branch system and shows the typical form of the fertile frond of Cystoseira foeniculacea (L.) Grev. with the proximal region lacking appendages and the distal region profusely branched with terminal receptacles, and is probably a good specimen of this species. Sheet 5, although purely vegetative and a small fragment, is also typical of this species. Sheet 4 is more doubtful; it could be a vegetative frond of a rather more flattened form than the preceding specimen, but it does not seems possible to determine this specimen without soaking out a small portion: it might not belong to the genus Cystoseira at all. On Sheet 0, the upper specimen carries the comment ‘ concatenatus of British Authors. D.T. t J.E.S.’ Although this may be a specimen of C. nodimdie, it seems more to resemble C. barbata (L.) C . Ag. both in the solitary aerocysts and in the very small, muoronate and pedicellate receptacles (of. Harvey, pl. ccclx). This specimen may have been responsible for Goodenough and Woodward’s misinterpretation of Fww foenicu2acew L. (see p. 8). The lower specimen, which is labelled ‘barbatus. Sp. Pl.’ in an unknown hand, does not appear to resemble Cystoeeira barbata Ag., but seems more like an evesiculate fertile frond of C . foeniculacea (L.) Grev. These sheets thus provide an illustration of the difficulty early botanists had in recognizing this species.

Cystoseira myriophylloides Cystoeeira myriophyl2oides Sauvageau in Bull. r3tat. Bbl. d ’ A r m h , 14: 466 (1912).

Cystoseira barbata sensu Desmaz., PI. Crypt. Fra.m, ed. 2: no. 23 (1826), non Ag. Cystoseira barbata var. turneri sensu Crouan frat., Alg. mar. FinistAre, no. 109 (1852),

This species has only recently been recorded from the British Isles (Blackler, 1961 : 75) at various localities in the Channel Isles. A specimen from this original collection is now in the Herbarium of the Marine Biological Association a t Plymouth. Since that date, several further collections have been made from the Devon coast and a number of specimens found by the present author in the collections of the British Museum (Natural History) and the Marine Biological Association, previously misidentified as other species.

The type material is the specimens studied by Sauvageau (1912 : 455) in preparing his account of this species. A specimen collected by Sauvageau on 2 March 1911 and now in the collection of the Museum National d’Hist.de Naturelle has been selected by Professor J. Feldmann as the lectotype (Pl. 3). This is a single plant with four main axes bearing laterals which are nearly all in the overwintering condition proximally but with fertile growth developing distally. One frond bears receptacles.

non J.G. Ag.

Cystoseira nodicaulis Cystoeeira mdimdia (With.) M. Roberts 1967.

F w w concatenatwr semu Lightf., Fl. h t . , 2: 923 (1777), non L.-Velley, Col. Fig.

Fww, nodicaulh With., Arrang. Brit. Pl., ed. 3,4: I11 (1796), excl. syn. Stackh. Bar. PI., 2 : fig. 1 (1796).-With., Arrang. Brit. PI., ed. 3,4: 89 (1796).

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Notee on Cystoseira C.Ag. 255

Fucw foeniculaceus sensu Gooden. and Woodw., Trans. Linn. Soc. Lond., 3 : 134

Fucus mucronatus Turn., Syn. Brit. Fuc., 1 : 73 (1802), m. mperjf. Putus granulatus sensu Smith in Sowerby, Engl. Bot., t. 2169 (1810), non L.-Turn.,

Cystoseira granulata sensu Grev., Alg. Brit., 5 , t. 2 (1830), non Ag.-Sauvageau in

(1797), non L.Stackh., Ner. Brit., Appendix: pag. ult., t. E. 5 (1801).

Puci, 4: 131, t. 251 (1819).

Bull. Stat. Biol. d’Aramhon, 14: 262 (1912).

This is the species commonly called Cystoseira granulata but, as pointed out below (Rub C. usneoides), that name is illegitimate and its true type does not belong to this species.

English botanists in the latter half of the eighteenth century confused this species with both Fucus concatenatus L. and F . foeniculaceus L. The descriptions by Lightfoot (1777 : 923), Velley (1795 : 281) and Withering (1796 : 89) of F. concatenatus all clearly refer to this species, as does the F. foeniculaceus of Goodenough & Woodward (1797: 134) and Stack- house (1801 : pag. ult.). The F. concatenatus of Hudson (1778: 574) may also refer to this species, but this cannot be determined with certainty. In his attempt to define the species Hudson quoted Linnaeus’s 1753 (p. 1628) and 1767a (p. 717) descriptionofF. concatenatus. He also quoted ‘FUCUB folio tenuissime diviso, siliquatus. R. syn. 48’. This specimen is in the Fielding Druce Herbarium, Oxford and was identified by Batters (in Druce & Vines, 1907: 26) as Cystoseira foeniculacea (L1) Grev. However, soaking out of the receptacles shows that it also belongs to the species under discussion. Hudson next quoted a Reaumur synonym (Reaum, act. gall. 1712: 31, t , 3, f, 5 ) , which is also rather a problem: the illustration is clearly the species under discussion, whilst the text refers to C. tamariscifolia (Huds.) Papenf. This is perhaps a further example of the difficulty these early botanists had in defining specific limits in this genus.

Withering (1796: 89) first described this species under Fucw concatenatus and then (1796: 111) as new species F. nodicaulis. This is its first description as an independent species, and the epithet of this name must be adopted. Withering’s description refers to the species under discussion, but the illustration of Stackhouse which he quotes (1797 : pl. 11) is Cyatoseira tamariscifolia. Withering obviously mistook the mounds of the clustered young laterals for tophules, a mistake which was also made by J. Agardh (1848 : 218) ; C. nodicaulis was not illustrated by Stackhouse until 1801 (Appendix. t. E. 5 ) , sub Fucus foeniculaceus.

Turner in 1802 (p. 73) described this species under F. mucrmtus, having concluded that it was neither F. concutenatus nor F . foeniculaceus. Turner quotes F. nodicuulis as a synonym, so his name is superfluous and illegitimate.

Smith (in Smith & Sowerby, 1810b, pl. 2169) was the first British author to apply the name F. granulatus to this species, and was followed in 1819 (p. 131) by Turner, who retained var. mucronatus and included F . usneoidea (i.e. F. granulatw L.) in his concept of the species.

C. Agardh (1820: 55) made the combination Cystoseira granulata basing his description on specimens provided by Cabrera and Heredia. These specimens were almost certainly not C. nodicuulit?, and the description omits any mention of the tophules already known from the illustrations of Stackhouse (1801: Appendix. t. E. 5; 1816: pl. 18), Smith (in Smith & Sowerby, 1810b: pl. 2169) and Turner (1819: pl. 251).

Greville (1830: 5, t . 2) gave a clear account of this species, illustrated by a good figure. Although he included C. Agardh’s C. granulatu in his synonymy, Qreville doubted whether Agardh’s description really referred to the British species since it lacked any mention of the tophules and because Agardh questioned whether Turner’s FWW granuldus (1819 : fig. 261) belonged to this species and placed it in Cystoseira barb&. Greville (p. 0) comments that Turner’s figure agrees well with the species, and is of a rather immature specimen.

Subsequent authors have all used the combination C. granulata to describe this species, usually referring i t to Linnaeus or Agardh. Sauvageau (1912: 262) uses the citation

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266 M. ROBEBTS

C. granulata Grev. All of these, however, are incorrect, and the correct name for the species is C. nodicaulis (Withering) M. Roberts.

The type of Withering’s description has not been l o c h d .

Cystoseira tamariscifolia Cystoseira tamriscijolia (Huds.) Papenf. in Hydrobiologia, 2: 185 (1950).

Fwus selaginoides L., Syst. Nut., ed. 10,2: 1346 (1759). Fww tamriscifolius Huds., F1. Angl., 469 (1762). Fww ericoides L., Sp. Pl., ed. 2, 2,: 1631 (1763). Cystoseira ericoides (L.) Ag., Sp. Alg., 1: 62 (1820).Sauvageau in Bull. Stat. Biol.

This species was first described by Linnaeus in 1759 (p. 1345) as Fucus selaginoides, with

d’Arcuchon, 14: 316 (1912).

the following description :

‘ F. filiformis teres, ramis fasciculatis, fructificatioriibus imbricatis sessilibus foliolo subulato terminatis.’

In the Linnaean Herbarium there are two sheets (1274: 12, 13) labelled F . selaginoides, possibly in the handwriting of Solander (Savage, 1946: 200). Of these, Sheet 12 is un- doubtedly the type of Linnaeus’s species as it has ‘D’ written on it by Linnaeus, the letter which he assigned to the new species F . selagimides in the System Naturae, edition 10 (p. 1345; 17673: 134).

In 1762 (p. 469) Hudson described a species to which he gave the name of F. tamarisci- folius. His description is based on specimens. The locality, however, is given as ‘ Yorkshire ’, where the species is not now found. Specimens cannot be found in any of the Herbaria and Collections examined, other than from Cornwall, Devon, Dorset, The Scilly Isles, the Channel Islands, Galway and Donegal : Stewart (1962) reports communities on the west coast of Barra in the Hebrides. It seems unlikely that 200 years ago the species extended as far north as Yorkshire on the east coast; any changes of range of algal species over this period which can be substantiated appear to be northward extensions rather than regreasions (Dixon, 1965). However, the description agrees quite well with the species which we now know as Cystoseira tamriscijolia (Huds.) Papenf. and there seems no reason to suppose that it refers to anything else. The record of Yorkshire must be regarded as a mistake, perhaps a confusion with another locality. As a synonym Hudson quotes ‘Fucus foliis Ericae seu Tamarisci. Ray, Syn. Meth. 8tirp. Brit., ed. 3: 49 (1724)’, of which the type is the specimen in the Fielding Druce Herbarium at Oxford labelled with Ray’s name and which is certainly this species. He also quotes Ray’s locality of Cornwall.

In 1763 (p. 1631) Linnaeus described a further species Fwus ericoides baaed, not on a specimen but on the Ray synonym quoted on the Fielding Druce Herbarium specimen.

Thus, as these three descriptions, all of which refer to the same species, have different types, all three names are legitimate.

The name F . selaginoides was not generally used by early botanists. Withering (1796 : 86) and Goodenough & Woodward (1797 : 130) recognized a species F . selaginoides L. which they separated from F . tamariscijolius Hudson, considering it not to be a native of the British Isles. Some (Turner, 1811 : 132; C. Agardh, 1820: 132) recognized a variety or other unspecified taxon selaginoides, whilst others (Hudson, 1778: 676; Greville, 1830: 4) quote F . selaginoides L. as synonymous with F. tamrisc(fo1iuS and Cystoseira ericoides (L.) C. Ag. respectively. ’

Bory (1832: 319) used the combination C. selugin0ide-s (L.) Bory based on Fwua selaginoides L., but this is probably a misidentification, since Linnmus’s species does not extend to the Pelop6nnisos; Sauvageau (1912: 285) believes it to refer either to Cystoseira stricta Sauv. or to C. mediterranea Sauv. As the name C. selagimides Naccari had been

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Notes on Cystoseira C.Ag. 257

previously published (Naccari, 1828a: 96; 1828b: 86), Bory’s combination is, in any cam, a later homonym and so illegitimate. Consequently, although the earliest name for this species is Fucus selaginoides L., it cannot be called Cystoseira selagirwides. Thus Hudson’s Fucus tamriscifolius provides the earliest available epithet in Cystoseira for the species to which the types of Fucus selagimides L., F . ericoides L. and F . tamariscifolius Huds. all belong, and Cystoseira tamriscifolia (Hudson) Papenfuss is its correct name. The type specimen is lost (see Dixon, 1959) and the type locality cannot be determined.

Cystoseira barbata Cystoseira barbata Ag., Sp. A&., 1 : 57 (1820)-Sauvageau in Bull. Stat. Bid. d’Armhon,

14: 392 (1912). Fucus foeniculaceus sensu Gmel., Hist. Fuc., 86, t . 2A fig. 2 (1768), non L. Fucus barbatus Gooden. & Woodw., Tram. Linn. SOC. Lond., 3: 128 (1797), non

Cystoseira h p i i Ag., sp. A&., 1: 59 (1820).

‘ 16. FUCUS vesiculis oblongis pedunculatis : terminatis foliolis linearibus.

Fucus barbatus L.

Linnaeus first described Pucus barbatus in 1753 (p. 1161) as:

Fucus folliculaceus, foliis abrotani. Bauh. Pin. 365? Fucus maritimus, foliis tumidis barbatis. Bauh. Pin. 365 Quercus maritima barbata. Bauh. Prodr. 154. Habitat in Oceano.’

Like Fucus foeniculaceus, this appears to be a species based on Burser’s Herbarium, and in Burser’s Herbarium, Vol. XX sheet 94 is labelled ‘Fucus folliculaceus foliis Abrotani Bauh. Congularis marinae Imperati nomine dedit Bauhinus’. This specimen, which agrees with the modern concept of C. barbata, was determined by Linnaeus as ‘Fucus foeniculaceus’ (Savage, 1937 : 69), and this determination was before the Species Plantarum (1753) description of F . barbatus was published. The first and queried synonym* ‘Fucus folliculaceus, foliis abrotani. Bauh. Pin. 365? ’ is omitted from Linnaeus’s later descriptions of the species (1763 : 1629), and in 1767a (p. 717) barbatus appears as a subdivision within foeniculaceus thus :

‘20. F. fronde filiformi ramosissima, vesiculis ovatis ter- minatus foliolis multipartitis obtusis apice fructi- ficantibus. Fucus barbatus.’

This Burser specimen may well be the type of F . barbatus L., as it is quite possible that Linnaeus drew up his diagnosis on the basis of it, later deciding that it belonged to the same species as the type of his F . foeniculaceus and inserting the query after the reference but without altering the diagnosis. Unfortunately, the first copy of the manuscript of the Species Plantarum lacks this section and in the second there is no evidence of any insertion of a query a.t any later date than the writing of the diagnosis.

If this specimen is not the type, then the second Bauhin synonym is the type. There is no specimen in Burser’s Herbarium carrying the second Bauhin synonym cited by Linnaeus; the identity of the specimen to which this description refers is therefore problematic as Linnaeus never saw Bauhin’s actual specimens (Savage, 1936: 23)) but it must probably be sought in the Bauhin herbarium a t Basle. However, as the combination Cystoseira barbata is preoccupied, its identity is irrelevant provided i t applied to a Cystoseira.

Fucus barbatus Goodenough and Woodward must be regarded as a new species, a later homonym of F . barbatus L. since (1797: 136) under F . foeniculaceus they say ‘Whether barbatus Sp. P1. foeniculaceus Syst. Nat. be the plant we have called barbatus, as from the reference to Gmehi might reasonably be supposed, or whether it be some varied appearance

*Reappears in 1771 (Mentisea), p. 509 under Fucw, aeluginoides.

Page 8: Taxonomic and nomenclatural notes on the genus Cystoseira C.Ag

268 M. ROBERTS of this plant, it is impossible from the imperfection and uncertainty of the specimens so named in the herbarium to asoertain.’ F w foenieulacewr sensu Gmelin is quoted as a synonym, so there is no doubt concerning the species intended, but as the account is based primarily on specimens, one of these must be taken as its type. These have not been located.

C. Agardh first published the name Cyetoeeira barbata (1820; 67), basing his account on specimens from Schouw, Cabrera and Mehns. As synonyms he quoted :

‘F. foeniculaceus, Gm. t. 2. A. f. 2. .. F. Crinitus, Desf. El. Atl. p. 424... F. barbatus, Linn. Tr. 111. p. 128 ... F. Granulatus, Turn. t. 261. f. a.? F. abies p. Bertol. am. p. 287. t. 4. f. 2. b? ’

None of the three unqueried synonyms refers to a legitimate name for the species since the first is a mistlpplication by Gmelin, and the other two are later homonyms. Cyebeira barbata Ag. must therefore be treated aa a new name and its lectotype chosen from among the specimens studied by Agardh.

According to Sauvageau (1912: 394), C. hqwpii Ag. is probably the same species as C. barbata Ag. The two were first combined under the name C. barbata by J. G. Agardh (1842: 60), who modified the species to include two varieties, viz. : a Turneri, in which he placed the F w w barbatus of Turner (1819: pl. 260), Stackhouse (1801: pl. 14), and of Sowerby and Smith (1810b: pl. 2170), all of which were based on the ‘Fww barbatw’ of Goodenough and Woodward: and H q p i i , with elongated receptacles and numerous concatenate vesicles, in which he placed the Cyetmeira Hoppii of C . Agardh (1820: 69). These varieties are considered by Sauvageau to represent s e ~ o m l modifications of C. barbata Ag., and that included under p Hoppii is the form illustrated by Harvey (1861 : pl. 360) and by Hauck (1883-86: fig. 124), the latter illustration being a copy of Kutzing’s (1860: t. 46) C. hppii. This is also the form of the Burser specimen, Vol. XX Sheet 94 (Pl. 4), which is the possible type of F w w barbatus L.

Cyeheira barbata Ag. has been included from time to time in floras of the British Isles, usually M a rarity washed up after storms, but it is certainly not native to these shores. It seems quite likely, as suggested by Sauvageau (1912 : 393) that Hudson’s original record of this species (1762 : 469) was based on a misidentification and this waa repeated by subsequent authors without reference to the actual specimen. Hudson himself later (1778 : 676) included it in his later interpretation of F w w foeniculaceus, thus following Linnaeus (1769: 717). Turner (1819: 124) and Harvey (1861: pl. 360) both consider this account to refer to Cyetoeeira barbatia but this may not be the case. Greville (1830: 3) includes F w w foev&icdamw Hudson (1778: 676) as a synonym of his Cyetoeeira foeniculacea.

A further source of error may have been confusion with the species now known as C . rnyriuphyhides Sauvageau. At least one specimen of this species has been found in the collection of the National History Museum, originally identified as C. barbata (cf. Desmezidres, 1836: No. 23).

The only authentic specimen of C. barbata which has been located is the small fragment provided by Woodward for the illustration of Smith & Sowerby (1810b: pl. 2170), but in the letter describing the specimen, no mention of its locality is made.

Cystoseira fimbriata

Cyeb8eira$mbr&z (Desf.) Bory, E q e d . Sc9ci. Morde, Sect. Sci. Phye., 3 (2) : 318 (1832).- Hamel, Phbphyc. Fr., 418 (1939).

Fww$mbriatw Desf., R. At&&., 2: 423 (July 1799). F w w cornpaw Esper, Ic. Fw. , 162, t. 77 (1799).

Page 9: Taxonomic and nomenclatural notes on the genus Cystoseira C.Ag

Notes on Cystoseira C.Ag. 269

Cystoseira abrotanifolia sensu J. Ag., Alg. Mar. Medit. Adr., 52 (1842), non (L.) Ag.-

Cystoseirajilicina Bory, Exped. Sci. Morke, Sect. Sci. Phys., 3 (2): 318 (1832). Sauvageau in Bull. Stat. Biol. d’drmhon, 14: 473 (1912).

The type material of FucusJimbriatus consists of the specimens on which Desfontaines based his original descriptions : these are probably in the Musee d’Histoire Naturelle, Paris. The type of Fucus mpressus is the material from the Adriatic sent to Esper by Wulfen, and is presumably a t Erlangen. If these two are conspecific, the correct name for this species depends on the relative dates of publication of Desfontaines description, issued in July 1799 (cf. Steam, 1938 : 148) and Esper’s, not dated more precisely than 1799. Until this can be established with certainty i t is best to follow Hamel (1939, p. 418) in using Cystoseira Jimbriata (Desf.) Bory.

This species was included by C. Agardh (1820 : 63) in his C. abrotanifolia. J. G. Agardh (1842, p. 52) first recognised it as distinct from the Linnaean species, and transferred the name C. ahrotanifolia Ag. to describe this Mediterranean and Adriatic species. In following Agardh in this name, Sauvageau (1912: 474) commented that a situation had arisen whereby ‘ le nom spdcifique crkk par Linnk pour une plante anglaise s’applique actuellement A une plante mediterrankenne.’ Both Funk (1955) and Ercegovic (1952) use C. abrotanifolia Ag., but more recent authors (Guern, 1962; Seaone-Camba, 1965) have used C. jimbriata (Desf.) Bory.

C.$licina Bory (1832: 318,1838: 74) is considered by Sauvageau (1912: 475) to be a growth form of this species.

Cystoseira discors

Cystoseira discors sensu Sauvageau in Bull. Stat. Biol. d’Arcachon, 14: 287 (1912), non (I,.)

The name Cystoseira diswrs was first used by C. Agardh (1820: 62) and applies to C. foenicukzcea (L.) Grev. (cf. p. 253).

Sauvageau (1912 : 415) pointed out that under the name C. diswrs Ag. three species had hitherto been confused. After comparing complete plants a t different seasons from Banyuls and the Gulf of Gascony, he concluded that the Mediterranean and Atlantic plants described under this name were sufficiently different to warrant separation into distinct species, whilst their resemblances were sufficient to suggest a common phyletic origin. He stated the principal difference between the two to lie in the manner of insertion of the secondary laterals, those of the Atlantic plants being inserted in a regular distichous fashion and those of the Mediterranean plants being rarely distichous. For the Mediter- ranean species he suggested the retention of C. diswrs Ag. This name was not referred by Sauvageau to the Linnaean species, as he considered that Linnaeus’s specimen was not identifiable with certainty and the description not precise as to location. Although Agardh’s description gives C . discm9 as purely Atlantic, Sauvageau (1912: 418) pointed out that Agardh (1820: 62) quoted de Candolle (1805) and Lamouroux (1813), both of whom listed the species as purely Mediterranean. Sauvageau also favoured retention of this name since Mediterranean algologists had always used the name C. discms C. Ag. for their species.

However, this usage of the name is incorrect and the species appears to have no legitimate name. No name is proposed as the author is not sufficiently familiar with the species to suggest one.

Ag.

Cystoseira usneoides

Cystoseira usneoidm (L.) M. Roberts. comb. nov. Fucw usneoides L., Syst. Nat., ed. 10,2: 1346 (1759). Fucw granulatus L., 8p. PI., ed. 2,2: 1629 (1763) m. supegl.

Page 10: Taxonomic and nomenclatural notes on the genus Cystoseira C.Ag

260 M. ROBERTS

FwusmcatenutussensuGooden. & Woodw.inTram. Linn.Soc., 3: 135 (1797),nonL. Cystoseira granulata Ag., Sp. Alg., 1: 55 (1820) nom. super$., excl. descr. Cystoseira concatenuta Ag., Sp. Alg., 1 : 57 (1820) pro pa&, non Fwus concatenatus

L.-Mont. in Durieu de Maisonneuve, FZ. Alg., 1: 15, t.6 (1846).-Sauvageau in Bull. Stat. Biol. d’drcachon, 14: 304 (1912).

This is the species to which the name C. wncatenata was applied by C. Agardh in 1820 and by almost all authors since. The reason for this is because Linnaeus added a specimen ofFwuswmeoides tohis herbariumafter 1753andlabelleditF. concatenatus(cf.p.253).The type of that name, and hence also the name Cystoseira concatenata (L.) Ag. is a van Royen specimen which belongs to the species C. foeniculaceu (L.) Grev.

In 1759 (p. 1345) Linnaeus described a new species, Fucw usneoidm, in the following terms :

‘E. F. filiformes teres, ramis geminis ramosissimis, fructi- ficationibus innatis, concatenatis, foliolis acerosis.’,

and in 1763 (p. 1629) he described F. granulatus as:

‘ 18. E’UCXJS fronde varicosa: ramis filiformibus ramulosis, vesiculosis ovtltis nodulosis mucronatis. Habitat in Oceano Indico Fwus spithameus. Caulis erectis, crassiusculus, inaequalis. Rarni ramulique numerosi, alterni, vane subramosi, filiformes, inaequalis, terminati. Vesiculis subovatis, nodulosis, terminatis mucrone.’

In the Linnaean Herbarium there are three sheets (1274 : 9,10,11) labelled F. usneoides, which in each case has been altered by Linnaeus to ‘F. granularis’. These are certainly the type specimens of F. usneoSe-9, and are also generally accepted as the type specimens of Linnaeus’s description (1763 : 1629) of F. granulatus. The difference in spelling between the epithet on the sheet and the published one seems no reason for rejecting this view. Sheet 1274: 11 (PI. 5 ) carries the comment in the hand of Sir James Edward Smith ‘certainly the species described in Sp. P1. Ed. 2 No. 18’ (viz. the description of F. granulatus quoted above), and is the one of the three which agrees best with the description. However, this specimen is mounted in a clump, and soaking out and remounting the entire specimen would be needed to see whether the axis bears tophules. Soaking out of a small portion of the tip of a branch shows that it possesses slender receptacles of the type seen in the species described by C. Agardh as Cystoseira concatenata. Moreover, the basal region of the specimen bears a number of flattened, foliose branches whose persistence in the basal regions of the mature plant is one of the main features which separates this species from the British C. nodicaulis (With.) M. Roberts.

The name Fww usneoicEes does not appear to have been used at all except by Hudson who did not apply it correctly. In 1762 (p. 470) he described a plant which he assigned with a query to F. usneoiCEe-9 (1759 : 1346) of Linnaeus, but his synonyms show that the plant in question is Desmarestia muleatu (L.) Lamour. In 1778 (p. 585) he correctly assigned the plant to the Fww muleatus of Linneaus (1767a: 717).

Cystoseira nodicaulis (With.) M. Roberts has hitherto usually been referred to c. Agardh’s (1820: 55) C. granulata, but there has always been doubt as to whether Agardh’s description really concerned this species (cf. Greville, 1830: 5 ; Montagne, 1838: 340; Sauvageau, 1912: 262) and it appears to refer, at least in part, to C. usneoides (L.) M. Roberts. Agard’s var. 8 setaceus from the English Channel may well refer to C. nodicaulis (With.) M. Roberts, although it does exclude Skckhouse’s illustration of this species, and

Page 11: Taxonomic and nomenclatural notes on the genus Cystoseira C.Ag

Notes on Cystoseira C.Ag. 261

may be based, as suggested by Montagne (loc. cit.), on a specimen lacking tophules. A study of C. Agardh’s material is necessary to resolve this point.

Turner’s (1819: pl. 251f) Fucus granulatus var. concutenatus also seems to refer to this species. He gave its distribution as ‘very common in the Adriatic. Wulfen.’, but none of Wulfen’s specimens have proved to be Cystoseira concatenata (Sauvageau, 1912: 281 and 307) and it is not reported from the Adriatic by Ercegovi6 (1952) or from Naples by Funk (1955). Little is known of the distribution of this species. Attached plants have been reported from very few localities-only from Spain (Miranda, 1934; Feldmann, 1937 ; Seaone-Camba, 1965) and from Algeria (Feldmann, 1931). Unattached specimens have been recorded from a variety of localities including Madeira and Brazil but, as the plant is capable of floating for long distances, this probably does not represent its true distribution. Linnaeus’s quotation of the Indian Ocean must have been an error, since this species of Cystoseira has not been recorded from that locality.

Sauvageau (1912 : 304) uses the citation C. concatenata (?L.) Ag. for this species whilst Hamel (1939: 407) just refers the species to C. Agardh. The correct name is, however, C. usneoides (L.) M. Roberts.

Cystoseira selaginoides

Fwus selqinoides sensu Wulf. in Jacq., Coll. Austr. Bot., 1 : 356 (1787) non L. Fucus aculeatus sensu Esper, Ic. Fuc., 1 : 72, t. 33 (1798) non L. Fucus abies-marina sensu Turner, Fuci, 4: 122, t. 249 (1819) non Gmel.

Cysto8eira seluginoides Nacc., F1. Venet., 6 : 96 (1828).

Naccari (1828~: 96) described specimens from the northern Adriatic, and gave the following synonymy :

‘ Cystoseira selaginoides. Nacc. Alg. Adr. mss. Fucus seluginoides. Wulf. (ma non Linneo)-Bertol.--Pollin. Fwus muleatus t . 33. Fucus Abies-marina Turner. 3. t. 249 (ma non Gmel.).’

The first two names quoted in this synonymy are referred by their authors to Linnaeus, and F. dies-marina is referred by Turner to Gmelin, so all the references quoted were to what Naccari believed to be misidentifhtions. Consequently, Cystoseira selugimides Naccari is to be regarded as a new species typified by one of the specimens used by Naccari in compiling the description. According to Sauvageau (1912: 283) an isotope in the Herbarium of Thuret is not identifiable and the species must be studied in the type locality. He applied the name in the sense of Valiante (1883 : 399) but this is not the correct application of the name. Hamel (1939 : 399) considered Valiante’s species to be different again from that of Naccari and proposed a further new name for it, ‘C. sauvageauiana’. On morphological grounds, Sauvageau (1912 : 282-83) considered the FWW dies-marina of Turner not to refer to Gnielin’s F. abies-marina, but to Wulfen’s F. eelaginoides.

Cystoseira sauvageauana

Cystoseira sauvaeauana Hamel, Phkophyc. Pr., 399 (1939) ‘sauvageauiana’. Cystoseira selaginoides sensu Valiante in Faun. PI. Golf Neapel, 7 : 19 (1883) [reimpr. in

Atti R. A d . Lincei. ser. 3, Mem. 15 : 399 (1883)], non Nacc.-Sauvageau in Bull. Stut. Biol. d’Armhon, 14: 283 (1912).

According to Hamel (1939 : 399) Valiante’s and Sauvageau’s description of C. selaginoides Nacc. refers to another hitherto undescribed species. The type of this species to which Hamel gave the name C. sauvageauuna is one of the specimens he described when compiling his description. Probably the best lectotype would be the Sauvageau specimen cited from Port-Vendres.

The spelling of the epithet must be changed from Lsauvageauiuna’ to Lsauvageawlna’

Page 12: Taxonomic and nomenclatural notes on the genus Cystoseira C.Ag

262 M. ROBERTS in accordance with the International Code of Botanical Nomenclature (Lanjouw et ad., 1961, Rec. 73C).

Cystoseira hoppii Cystoseira hoppii sensu Valiante in Faun. Fl. Gblj Neupel, 7 : 16, t. 7 (1883) (reimpr. in

Atti R. A d . Lincei, ser. 3, Mem. 15: 395 (1883)), non Ag.Sauvageau in Bull. Stat. Biol. d’drcuchm, 14: 412 (1912).

Sauvageau (1912 : 412) considered the species described by Valiante (1883 : 16) under C. b p p i i to be distinct from C. barbata under which it was included by de Toni (1895 : 170). This species appears to have no legitimate name ; C. hoppii C.Ag. is a synonym of C. bar@ C.Ag.

AOENOWLEDOEMENTS

I wish to thank Dr P. S. Dixon for help in planning this paper and for advice at all stages of the work and Mr R. Ross for help with all the details of synonymy involved. I am indebted to the Directors and Curators of the following Herbaria for permission to examine specimens and to photograph and soak out portions where necessary :

The Rijksherbarium, Leiden; The Musee d’Histoire Naturelle, Paris; The British Museum (Natural History), London; The Linnean Society, London; The Marine Biological Association Plymouth. The photograph of Linnaeus’s type specimen of F w w wneoides in Plate 6 and of Sheet 94, Vol. XX, of Burser’s Herbarium in Plate 4 were supplied by courtesy of the International Documentation Company, Zug, Switzerland ; the remaining plates were prepared by Mr P. E. Smith, Department of Botany, University of Hull. I also wish to thank Mr J. H. Price and the Secretary and Librarian of the Linnean Society for very considerable help with the references, and Professor J. Feldmann for examining the collections of Sauvageau in Paris, and selecting the lectotype of C. myrio- phylloides.

I am indebted to the International Federation of University Women and to the Botanical Research Fund for the Grants which enabled me to carry out this work.

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EXPLANATION OF PLATES

PLATE 1 The type specimen of Fucua co?~catenatua, collected by van Royen and now in the Rijksherbarium. Leiden. (Herb. Ludgb. 910. 163-1332). Photograph by Mr P. E. Smith.

PLATE 2 (a). A small portion of the receptacle of van Royen’s specimen of Fucus concatenutue soaked out in water and photographed in a glass dish, submerged in water. (b). A portion receptable of Cyetoseira joeniculacea (L). Grev., collected by M. de Val6ra from the West Coast of Ireland. Photographed in water. (c). A portion of receptacle of Cyetoaeira usneoidee (L.) M. Roberts (= C. concatenata As.) collected by Sauvageau from Guethary in July, 1909, and soaked out and photographed in water. Scale= x 3. Photographs by Mr P. E. Smith.

PLATE 3. The type specimen of Gystoaeira myriophylbidea Sauv., collected by Sauvageau and now in the collection of the M d e National d’Histoire Npturelle, Paris. Photograph by M i P. E. Smith.

P L A ~ 4. Sheet 94, Vol. XX in the Herbarium of Burser which is possibly the type Linnaeus’s Fucua barbatua. Photograph supplied by the International Documentation Company.

PLATE 6. Sheet 1274: 11 in the Herbarium of Linnaeus, which is the type of Cyatoeira ueneoidea (L.) M. Roberts. Photograph supplied by the International Documentation Company.

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