*TL . 10 1 Bulletin of the British Museum (Natural History) Taxonomic studies in the Labiatae tribe Pogostemoneae J. R. Press Botany series Vol 10 No 1 30 September 1982 V The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales British Museum (Natural History), Cromwell Road, London SW75BD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.) Trustees of the British Museum (Natural History), 1982 The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Mr J. F. M. Cannon Editor of Bulletin : Mr P . W. James Assistant Editor: Mr J. R. Laundon ISSN 0068-2292 Botany series Vol 10 No 1 pp 1-89 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 30 September 1982 Taxonomic studies in the Labiatae tribe Pogostemoneae GENERAL 1 OCT 1982 jfr J. R. Press_ Department of Botany, British Museum (Natural History), Cromwell Road, London SW75BD Contents Synopsis 2 1 . Introduction 2 2. Historical review 4 Taxonomic position of the Pogostemoneae 4 Previous taxonomic treatments of the Pogostemoneae 5 3. Methods 8 Reconstitution of material 8 Data compilation 8 Measurement of characters 10 The programs 12 4. Character variation 12 Leaves 12 Heterophylly 12 Leaf size 12 Phyllotaxis 12 Petiole 12 Bracts 17 Braceteoles 17 Indumentum 17 Calyx 19 Corolla 19 Stamens 22 Style 22 Disc 22 Nutlets 22 Inflorescence 25 Pollen 25 5. The analyses 27 Principal co-ordinates analysis 27 Clustering to maximise within-group-mean-similarity 27 Single-linkage analysis 42 6. The taxa : discussion 43 Comanthosphace and Leucosceptrum 43 Rostrinucula 47 Elsholtzia 47 Elsholtzia sections Cyclostegia and Elsholtzia 49 Elsholtzia section Aphanochilus series Stenelasmeae 51 Elsholtzia section Aphanochilus series Platyelasmeae 53 Relocated OTUs 54 Elsholtzia beddomei and E. kachinensis 56 Elsholtzia penduliflora 56 Elsholtzia flava, E.fmticosa, andE.hunanensis 56 Elsholtzia concinna 56 Leucosceptrum plectranthoidewn 57 Bull. Br. Mus. not. Hist. (Bot.) 10 (1): 1-89 Issued 30 September 1982 J. R. PRESS Dysophylla mairei 57 Elsholtzia integrifolia 57 'Elsholtziajaponica' 57 Keiskea 58 Tetradenia 59 Eurysolen 60 Pogostemon and Dysophylla 62 Subgeneric divisions 64 Relocated OTUs 64 Elsholtzia aquatica 64 Distribution of Pogostemon 66 Colebrookea 66 7. Taxonomic conspectus 67 Key to genera and sections 67 Conspectus 68 Species excludendae 74 Appendix 1 . Specimens used for scoring and producing averaged data 76 Appendix 2. Coded data for 138 OTUs 78 Appendix 3. List showing the five nearest neighbours for each OTU with the similarities expressed as percentages 83 Appendix 4. Distribution of OTUs in the six and nine group schemes for clustering to maximise WGMS analysis 87 Acknowledgements 88 References . . , 88 Synopsis Relationships within the tribe Pogostemoneae (Benth. ex Endl.) Briq. of the family Labiatae are investigated. After a review of the taxonomic history of the group, the distribution of sixty-four morphological characters in 138 species from ten genera is discussed. The CLASP 202 computer program incorporating cluster analyses and ordination methods is used to analyse the morphological data. Discussion of the relationships of supra-specific taxa is followed by an enumeration of the species within each genus. Close similarities are established between Comanthosphace S. Moore, Leucosceptrum Smith, and Rostrinucula Kudo (formerly included in Elsholtzia Willd.). Further investigation into style and fruit characters is required to confirm the status of Leucosceptrum as a genus separate from Comanthosphace. Close similarities are also established between Elsholtzia, Keiskea Miq., and Tetradenia Benth. (formerly grouped with Colebrookea Smith, Dysophylla Blume and Pogostemon Desf.). Elsholtzia is divided into three sections, one (section Platyelasmd) being raised from the rank of series. Section Cyclostegia is reduced to a synonym of section Elsholtzia. The bract and inflorescence characters which are alleged to distinguish them form a transition series between the two sections. Five of the six species of Keiskea are compared for the first time, and additional distinguishing characters are given for the genus. Eurysolen Prain is included within the tribe Pogostemoneae on the basis of its fruit morphology but its relationships within the tribe remain in doubt. Dysophylla is reduced to a section, section Eusteralis, of Pogostemon. The distinguishing characters form a transition series between the genera. In addition a number of characters formerly used to separate the genera are now believed to be merely responses to aquatic environments. Two new species, Pogostemon glabratus and P. trinervis are formally described. Sixteen new combinations are made. 1. Introduction The Pogostemoneae is a tribe containing some 200 species divided among 10 genera. The group has a wide distribution through the warm temperate and tropical regions of the Old World (Fig. 1). Most species are herbs or suffruticose perennials, but some, e.g. species of Dysophylla TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE - possible relicts of cultivation Fig. 1 Distribution of the Pogostemoneae. Blume, are aquatic or amphibious; a few, e.g. Elsholtzia fruticosa (D. Don) Render and Colebrookea oppositifolia Smith, are shrubs. Several species, particularly of Elsholtzia Willd. and Pogostemon Desf . , have commercial and homeopathic uses. The most important are Pogostemon heynianus Benth., which provides the patchouli oil of perfumery (Day, 1979), and several species of Elsholtzia, e.g. E. ciliata (Thunb.) Hylander and E. rugulosa Hensley, which are used as stomachics and carminatives (Chopra, Nayar & Chopra, 1956; Keys, 1976; Yunnan Xian Weishenju, 1973). A number of other species are used as condiments and in folk-medicine, especially in eastern and south-eastern Asia. Pogostemon mutamba (Hiern) G. Taylor (Fig. 30) from southern tropical Africa produces starchy edible tubers (Taylor, 1931). Several species are valuable food sources for bees, e.g. Pogostemon parviflorus (Benth.) Benth., from which pangol honey is produced. This relatively small but quite well-known tribe has been rather neglected by recent workers in the Labiatae. Various genera have been studied in part or in whole and a number of changes suggested, although these sometimes contradict with each other. In particular the generic limits of Dysophylla, Pogostemon, Comanthosphace S. Moore, and Leucosceptrum Smith bear further investigation, as do the positions within the tribe of Keiskea Miq., Rostrinucula Kudo and Tetradenia Benth., and the possible inclusion of Eurysolen Prain. No recent worker has considered the tribe in its entirety and this is the aim of this study. To ensure a thorough analysis and detailed assessment of all the available data, numerical phenetic methods have been used to investigate the interrelationships and definitions of the different taxa. 4 J. R. PRESS 2. Historical review Taxonomic position of the Pogostemoneae The only complete monographic treatments of the Labiatae are those of Bentham (1832-36, 1848). In his earlier work Bentham (1832-36) divided the family into eleven tribes, and described a number of ad hoc groups within them. Endlicher (1838) accepted Bentham's arrangement and also assigned the rank of subtribe to Bentham's unnamed groups. Subtribes Pogostemeae Endl. (containing Dysophylla and Pogostemori) and Elsholtzieae Endl. (contain- ing Elsholtzia and Tetradenia) were placed in the tribe Menthoideae Benth. (see Table 1). In his second account Bentham (1848) rearranged his tribes, reduced the number to eight and recognized a number of subtribes. For example, subtribe Elsholtzieae (containing Colebrookea Smith, Dysophylla, Elsholtzia, Pogostemon, and Tetradenia} was placed in the tribe Satureieae Benth. (see Table 1). Later classifications were minor variations of that of Bentham until Briquet (1897) produced an apparently very different classification, with more subdivisions than in the earlier ones. He recognized eight subfamilies, 14 tribes and 11 subtribes (see Table 1). Although Briquet altered much of the order of Bentham's and Endlicher's systems, the essential Table 1 Summary of the classifications of the Labiatae by Endlicher (1838), Bentham (1848), and Briquet (1897). Endlicher, 1838 (based on Bentham, 1832-36) Tribes & subtribes Bentham, 1848 Tribes & subtribes Briquet, 1897 Subfamilies, tribes & subtribes I Ocimoideae I Ocimoideae I Ajugoideae 1. Moschosmeae 3 unnamed divisons 1. Ajugeae 2. Plectrantheae II Satureieae 2. Rosmarineae 3. Hyptideae 1. Elsholtzeieae II Prostantheroideae 4. Lavanduleae 2. Menthoideae III Prasioideae II Menthoideae 3. Thymeae IV Scutellarioideae 1. Pogostemeae 4. Melisseae V Lavanduloideae 2. Elsholtzieae 5. Genera anomala VI Stachyoideae 3. Mentheae III Monardeae 1. Marrubieae 4. Meriandreae IV Nepeteae 2. Perilomieae III Monardeae V Stachydeae 3. Nepeteae 1. Salvieae 1. Scutellarieae 4. Stachydeae 2. Rosmarineae 2. Melitteae a. Prunellinae 3. Hormineae 3. Marrubieae b. Melittinae IV Satureineae 4. Lamieae c. Laminae 1. Origaneae VI Prasieae 5. Glechoneae 2. Hyssopeae VII Prostanthereae 6. Salvieae 3. Cunileae VII Ajugeae 7. Meriandreae V Mellisinae 8. Monardeae VI Scutellarineae 9. Hormineae VII Prostanthereae 10. Lepechinieae VIII Nepeteae 11. Satureieae IX Stachydeae a. Melissinae 1. Melitteae b. Hyssopinae 1. Lamieae c. Thyminae 3. Marrubieae d. Menthinae 4. Balloteae e. Perillinae X Prasieae 12. Pogostemoneae XI Ajugoideae VII Ocimoideae a. Hyptidinae b. Plectranthinae c. Moschosminae VII Catopherioideae TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 5 difference between the classifications is one of rank, Briquet's divisions generally being made at one rank higher than those of the other authors. Although some authors (e.g. Erdtman, 1945; El-Gazzar & Watson, 1970) have expressed dissatisfaction with Briquet's classification, it is accepted by most modern authors (e.g. Keng, 1969) as being superior to those of earlier workers. Briquet regarded the Pogostemoneae as a tribe characterized particularly by the 4-5 lobed corolla being equal or weakly bilabiate, and the anther locules showing some degree effusion, and this concept of the group is followed here. A brief synopsis of the nomenclature is given below. Tribe Pogostemoneae (Benth. ex Endl.) Briq. in Engl. & Prantl, Natiirl. Pflanzenfam. 4 (3aV 326 (1897). Tribe Menthoideae subtribe Pogostemeae Benth. ex Endl., Gen. pi. : 612 (1838). Tribe Menthoideae subtribe Elsholtzieae Benth. ex Endl., Gen. pi. : 612 (1838). Tribe Menthoideae subtribe Mentheae Benth. ex Endl., Gen. pi. : 612 (1838) propane quoad Colebrookia sphalm. Tribe Satureieae subtribe Elsholtzieae Benth. in DC., Prodr. 12 : 149 (1848). Tribe Satureieae subtribe Pogostemoneae Benth. & Hook.f., Gen. pi. 2 (2) : 1162, 1164 (1876). Tribe Satureieae subtribe Menthoideae Benth. & Hook.f., Gen. pi. 2 (2) : 1162, 1164 (1876) pro parte quoad Elsholtzia et Keiskea. Tribe Satureieae subtribe Pogostemoninae Kudo in Mem. Fac. Sci. Agric. Taihoku imp. Univ. 2 (2): 45 (1929). Previous taxonomic treatments of the Pogostemoneae The Labiatae tribe Pogostemoneae comprises some 10 currently recognized genera, of which only three, Dysophylla, Elsholtzia, and Pogostemon, contain more than a dozen species. The taxonomic affinities of the genera within the Pogostemoneae and in related tribes have been subject to a number of different assessments and interpretations. The early literature in particular is fragmentary and confused. Few authors agree on the importance that should be attached to the various characters, or indeed which characters should be used in the considera- tion of the affinities and subsequent groupings. The results are at variance with each other and provide no clear picture as to the internal groupings of the tribe. The following account attempts to trace the history of the genera of the Pogostemoneae. The first currently recognized genus, Dysophylla Blume, had earlier been given the poly- nomial Veronica hirsuta latifolia Zeylanica aquatica by Hermann (1717). Linnaeus (1747) used the generic name Alopecuro-Veronica. Later he (1767) described M. auricularia L., a plant of remote affinity to the other species of Mentha; here he placed in synonymy the names Alopecuro-Veronica and Majorana foetida (a plant described by Rumphius, 1750, as Majana foetida). When Blume (1826) separated Dysophylla as a distinct genus based on Mentha auricularia he related it to, and placed it next to Mentha L., but considered it to differ by the closure of the fruiting calyx, the fleshy swelling of the disc and the distinctly bearded stamens. Under the only species, Dysophylla auricularia (L.) Blume, he cited Rumphius's illustration. Willdenow (1790) described a new genus, Elsholtzia, based on a Silesian plant, E. cristata Willd. Although he gave a clear description and distinguished his genus from other Labiatae on corolla, calyx, stamen and inflorescence characters, a number of confused associations were made between this and other plants by later authors. Lamarck (1789) described five species in Hyssopus L. , but his rather vague limits of the genus allowed for inclusion of two species of Elsholtzia, one of which was Willdenow's type species. Persoon (1806) amended Willdenow's genus description to add 'Flor. secundi, bracteati' and described two new species, E. paniculata (referable to Pogostemon) and E. ocymoides. Don (1825) considered Elsholtzia congeneric with Perilla L. and reduced the name to synonymy. The description differed from that of Perilla sensu Linnaeus (1764) on a number of points but closely matched that of Willdenow's Elsholtzia. Two new genera, Leucosceptrum and Colebrookea, were described by Smith (1805) based on 6 J. R. PRESS Nepalese specimens. Leucosceptrum, because of its habit, corolla and deeply lobed disc, was said to have affinities with the Verbenaceae, and because of the four-lobed corolla, exserted stamens and bilocular anthers, was considered close to Mentha. Smith described the corolla as having four unequal segments, the upper deeply emarginate, the lower large and entire. However, his illustration incorrectly shows the opposite of this, the lower segment being emarginate and the upper entire. The second monotypic genus, Colebrookea, was chiefly characterized by its fruit morphology. The distinctive hairy calyx with its plumose teeth acts as a pappus for the single-seeded, dry fruit. Roxburgh (1815) published an account of a second species of Colebrookea, C. ternifolia Roxb., based on plants collected in Mysore. All later authors, however, regard this merely as a variety of C. oppositifolia, leaving Colebrookea a monotypic genus. C. oppositifolia is a shrub and seems very different from the herbaceous species of Elsholtzia. In fact they do share a number of floral characters. Poiret (1817) produced a rather confusing situation when he published a new generic name Elshotzia, and put into it Colebrookea oppositifolia as a new combination. He even added the comment in the text 'ce genre est le meme que le Colebrookea de Smith. II fait y reunir le barbula de Loureiro'. The 'barbula' of Loureiro (1790) is an earlier name for Caryopteris Bunge (Verbenaceae), which is strikingly similar to the Pogostemoneae in calyx, corolla, and anther characters. Desfontaines (1815) described a new genus, Pogostemon, for a species with bearded stamens, P. plectranthoides Desf . (now = P. benghalensis (Burm. f .) Kuntze). Citing the shared similarity of calyx and corolla characters of Pogostemon and Hyssopus he claimed an affinity for his new genus; 'Le genre Pogostemon a de 1'affinite avec 1'Hyssope. La corolle renversee, les trois lobes de la levre superieure entiere et arrondis au sommet, les filets des etamines abaisses et barbus, sont les principaux characteres qui le distinguent'. Blume (1826) followed Desfontaines' description and published a second species from Java, P. menthoides, but which had naked filaments. The first comprehensive review of all genera was Bentham (1829). He considered the original descriptions of Dysophylla and Pogostemon unsatisfactory and suggested modifications for both. In Dysophylla he placed less emphasis on the connivence of the calyx teeth, since it is a feature not common to all species. In Pogostemon he regarded the declination of the stamens to be so slight as to be of little significance, and he questioned Blume's inclusion of P. menthoides, since it appeared anomalous with the absence of hairs on the stamens, hairy stamens being a constant feature of all the other species of both genera. The delimitation of Dysophylla was expanded by Bentham to include those species of Mentha described by Loureiro (1790) and Roxburgh (1814, 1832). Bentham reduced Elsholtzia to only one species, E. cristata. He created three new genera, all closely related to Elsholtzia: Aphanochilus, very similar to Elsholtzia, but differing in the exserted stamens with anther-locules confluent, the shrubby habit and non-secund inflorescence (the Perilla species described by Don (1825), although unknown to Bentham, belonged to Aphanochilus), the monotypic Cyclostegia, easily distinguished by its densely strobilate in- florescence, and Tetradenia, a Madagascan genus based on a plant labelled Mentha fruticosa in Hooker's herbarium; this differs from the other two genera in the irregular calyx and bright red, glandular swellings around the nutlets from which it is named. A year later Bentham (1830) wrote a second account of the Labiatae. The only change from the earlier work was for Dysophylla, where he divided the genus for two parts on the basis of phyllotaxy. The first was characterized by opposite, paired leaves, and agreed with Blume's original description; it contained three species, D. auricularia, D. myosuroides Benth., and D. strigosa Benth. The second contained all Dysophylla species with verticillate leaves. The first complete treatment of the Labiatae was by Bentham (1832-1836) in which he repeated his earlier descriptions for most of the genera and formally recognized his own divisions of Dysophylla (1830) as sections Oppositifoliae and Verticillatae. On the basis of inflorescence characters he divided Pogostemon into Paniculatae and Racemosae. Apha- nochilus Benth. and Cyclostegia Benth., however, were reduced to sections of Elsholtzia, the distinctions being based primarily on morphological differences of the inflorescence and bracts. Bentham considered Leucosceptrum to be a section of the unrelated genus Teucrium L. in his TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 7 tribe Ajugoideae, which it resembled in lacking the upper lip of the corolla, in the hairiness of its fruits, and in its semi-shrubby habit. Bentham's division of Dysophylla into sections Oppositifoliae and Verticillatae was taken further by Rafinesque-Schmaltz (1847) who published the genus Eusteralis based on Mentha pumila Graham and M. verticillata Roxburgh, which effectively raised section Verticillatae to generic rank. The problems posed by Dysophylla were also reflected by Hasskarl (1842), Miquel (1859), and Kuntze (1891), who were unable to distinguish it adequately from Pogostemon and included it as a section of that genus. Miquel (1865) described four new species of Elsholtzia, three of which he referred to Bentham's section Cydostegia. Bentham (in Bentham & Hooker, 1876) referred them to Pogostemon (see Hooker, 1896). This arrangement was based on the similarity of the anther structure, the anthers being subglobose, imperfectly two-celled and two-valved, as in Pogoste- mon, and not ovoid, distantly two-celled with the cells two-valved, as in Elsholtzia. One year later S. Moore (1877) removed Miquel's four species of Elsholtzia and placed them in the new genus Comanthosphace. Moore's view was that Comanthosphace approached Elsholtzia more nearly than Pogostemon, particularly in the two-lipped, five-lobed form of the corolla. This arrangement was fairly reliable, but Moore mistakenly described the verticillasters as being obscurely bracteate, an error pointed out by Hooker (1896). In fact the bracts are very large, another feature in common with Elsholtzia, but often caducous. In the same paper, Miquel (1865) proposed a new genus, Keiskea, to accommodate material collected in Japan. When considering its affinities he gave its position as close to Mentha, although the description suggested closer affinities to Elsholtzia in that the corolla was sub-bilabiate with an emarginate upper lip and three-lobed lower lip. The filaments were hairy at the base, forming an incomplete annulus. Miquel described one species, Keiskea japonica, and the genus remained monotypic until Diels (1924) described a second species from China, K. sinensis. A third species, K. elsholtzioides, was described by Merrill (1937). He had difficulty in determining the affinities of this species and commented: 'In making the preliminary examina- tions this was placed in Comanthosphace from which its calyx characters exclude it. In its characteristic persistent, broad bracts it resembles Elsholtzia but in spite of these, seems to belong in Keiskea. ' K. elsholtzioides Merrill differs markedly from the two previously known species of Keiskea which have narrower bracts and broadly campanulate calyces. Masamune (1940) described a Formosan species of Keiskea, K. macrobracteata. The conspicuous charac- ters, bilabiate calyx and broadly ovate bracts, which distinguish it from K. japonica Miq. and K. sinensis Diels, prompted him to propose dividing the genus into two sections, Macrobracteatae and Eukeiskea. The next major work on the Labiatae after Bentham was by Briquet (1897) although his account of the Pogostemoneae had no major changes from that of Bentham (1832-36). However, he described new subgeneric groupings in three genera. In Pogostemon he divided Racemosae Benth. into: A. Glabriuscula with naked filaments and B. Barbata with hairy filaments. Paniculatae Benth. was also divided into two groups designated only as A and B and distinguished by the density of the verticillasters (see Table 13). Dysophylla was divided into two sections: section Goniocalicinae, with strongly five-angled calyces, and section Rhabdocalici- nae, with cylindrical calyces. Section Rhabdocalicinae was further divided by annual or perennial habit (see Table 14). Two new series were described in Elsholtzia section Aphanochi- lus : series Platyelasmeae Briq. (containing only E. densa Benth. and E. eriostachya (Benth.) Benth. and distinguished by broad bracts and matt nutlets) and series Stenelasmeae Briq. (containing the remaining species and distinguished by narrow bracts and shiny nutlets). Another Elsholtzia species of dubious position was described by Rehder (1917) as E. dependens. This species seems not closely related to any other species of the genus. According to its broadly bracted spikes it ought to be placed in the group Platyelasmeae Briquet of the section Aphanochilus Bentham, but it differs from the species of this group as from those of the other groups in the entire upper lip of the corolla, in the irregular ring of hairs at the mouth of the corolla formed by hairy disc-like excrescences at the base of the filaments and in a hairy crescent-shaped crest below the base of the lower lip, and in the rostrate nutlets. The drooping g J. R. PRESS habit of the long and slender spikes is also very peculiar and, so far as I know, does not occur in any other Elsholtzia.' Rehder's views were supported by Kudo (1929) in his monograph of the Chinese Labiatae. However, he took it one stage further and removed it from Elsholtzia and established a new, monotypic genus, Rostrinucula. Kudo also revived Bentham'sAphanochilus, considering it to merit generic rank. This was not wholly accepted by other authors since many considered Aphanochilus and Elsholtzia to be congeneric, the calyx, anther, and bract characters being unreliable for separating the two. During the last 50 years, very few major systematic changes have been made within the Pogostemoneae. Kitamura & Murata (1962) proposed the union of Comanthosphace and Leucosceptrum, citing the shape of both calyx and corolla, exsertion of stamens, anther shape, aerole size, and presence of stellate hairs on the leaves, as shared characters. As the authors did not examine all the species of either genus their views have not been generally accepted. El-Gazzar & Watson (1967) made a study of Dysophylla and Pogostemon, using leaf characters, presence or absence of crystals in the calyx, and presence or absence of stem aerenchyma. They concluded that Bentham's Dysophylla section Oppositifoliae (containing four species, D. auricularia, D. myosuroides, D. rugosa Hook, f., and D. salicifolia Dalz, ex Hook, f.) should be sunk into Pogostemon since it shared the characters of opposite, broad, petiolate leaves, crystals present in the calyx, and stem-aerenchyma absent. Dysophylla section Verticillatae (containing all other Dysophylla species) was characterized by verticillate, linear, glabrous, sessile leaves, crystals absent from the calyx, and stem-aerenchyma present. Wu & Li (1975) also placed D. auricularia and D. falcata Wu in Pogostemon although they did not specify their reasons for so doing. These transfers of Dysophylla species, including the type species D. auricularia, to Pogostemon raised a nomenclatural problem for which a number of solutions were proposed and subsequently rejected. The latest and most satisfactory proposal, suggested by Bakhuizen van den Brink & van Steenis (1968), and also made by Panigrahi (1976), was to revive the name Eusteralis Rafin. for Dysophylla section Verticillatae. Keng (1978), however, combined Dysophylla and Pogostemon as congeneric taxa, and placed Dysophylla section Verticillatae in Pogostemon section Eusteralis. Wu & Huang in their precursor account for the Flora Reipublicae Popularis Sinicae (1974), and later in the flora itself (1977), gave a generic account of the Chinese species of Elsholtzia. This covered 33 species and produced two new subsections and eight new series. 3. Methods Reconstitution of material All materials used in this study were obtained from herbarium specimens. Flowers and fruiting calyces were too brittle to be dissected without prior softening. The material was soaked in 4% sodium-orthophosphate solution for 12 hours, washed in distilled water, and stored in absolute alcohol. Data compilation The number of species and bulk of material available precluded the recording of every specimen for computation. Nevertheless, all available species were included in the sampling and all material was examined. To obtain full data for species (OTUs), three typical specimens were selected for scoring (Appendix 1) as a basis for each operational taxonomic unit. The species, their OTU numbers and acronyms are given in Table 2. The data were then averaged to produce single character statements for each species. Obscure or poorly understood taxa were scored wherever possible from authentic specimens. Data on pollen were obtained from the literature except for Eurysolen gracilis Prain and Leucosceptrum canum Smith. Pollen samples from these two species were acetolysed and examined by light microscopy. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE Table 2 The OTUs used in the analyses. EGRA 1. Eurysolen gracilis DFAL 58. falcata EJAP 2. 'Elsholtzia japonica' DSZE 59. szemacensis EFED 3. feddei DFAU 60. faurei EELE 4. 'elegans' DKOE 61. koehneana EPSE 5. pseudocristata DGRA 62. gracilis ENIP 6. nipponica DLYT 63. lythroides EOLD 7. oldhamii DPUM 64. pumila ECON 8. concinna DMAI 65. mairei ECIL 9. ciliata DAND 66. andersonii EKAC 10. kachinensis DGLA 67. 'glabrata' EPYG 11. pygmaea DAUR 68. auricularia ESOU 12. soulei DSAL 69. salicifolia EARG 13. argyi DRUG 70. rugosa ELUT 14. luteola DMYO 71. myosuroides EBOD 15. bodinieri PBEN 72. Pogostemon benghalensis EHET 16. heterophylla PPAR 73. parviflorus ESTR 17. strobilifera PPAN 74. paniculatus EHUN 18. hunanensis PTUB 75. tuberculosus EERI 19. eriostachya PGLA 76. glaber EDEN 20. densa PHEY 77. heynianus EMAN 21. manshurica PCAB 78. cablin EAQU 22. aquatica PELS 79. elsholtzioides EINT 23. integrifolia PAMA 80. amarantoides EBED 24. beddomei PFOR 81. formosanus EPEN 25. penduliflora PMEN 82. menthoides ERUG 26. rugulosa PFRA 83. fraternus ESTA 27. stauntonii PBRA 84. brachystachyus EFLA 28. flava PMIC 85. micangensis EFRU 29. fruticosa PMUT 86. mutamba EPUB 30. pubescens PNIG 87. nigrescens ECOM 31. communis PPHI 88. phillipensis EALO 32. alopecuroides PREF 89. reflexus EGRI 33. grifftthii PRET 90. reticulatus EGLA 34. 'glanduligera' PMOL 91. mollis EELA 35. elata PTRA 92. travancoricus EWIN 36. winitiana PATR 93. atropurpureus ESTC 37. stachyodea PSPE 94. speciosus EBLA 38. blanda PVEL 95. velatus EMYO 39. myosurus PWIL 96. williamsii EOCH 40. ochroleuca PPUR 97. purpurascens EPIL 41. pilosa PPAL 98. paludosus ECAP 42. capituligera PVIL 99. villosus DTRI 43. 'Dysophylla trinervis' PWIG 100. wightii DHEL 44. helferi PROT 101. rotundatus DTOM 45. tomentosa PSTR 102. strigosus OPEN 46. pentagona PHIS 103. hispidus DSTO 47. stocksii PPUB 104. pubescens DSAM 48. sampsonii PBRE 105. brevicorollus DPEG 49. peguana PNEL 106. nelsonii DGRI 50. grifftthii PBAT 107. battakianus DYAT 51. yatabeana PWAT 108. wattii DCRA 52. crassicaulis PHIR 109. hirsutus DSTE 53. stellata PRUP 110. rupestris DLIN 54. linearis PMAC 111. macgregorii DCRU 55. cruciata PGAR 112. gardnerii DQUA 56. quadrifolia PCHA 113. chaixii DTSI 57. tsiangii PDIE 114. dielsianus 10 J. R. PRESS PGRI 115. griffiithii LCAN 127. Leucosceptrum canum PNIL 116. nilagiricus LPLE 128. plectranthoideum PLIT 117. litigiosus TGOU 129. Tetradenia goudotii RDEP 118. Rostrinucula dependens THIL 130. hildebrandtii RSIN 119. sinensis TFRU 131. fruticosa CFOR 120. Comanthosphace formosana KJAP 132. Keiskea japonica CSTE 121. stellipila KELS 133. elsholtzioides CBAR 122. barbinervis KGLA 134. glandulosa CSUB 123. sublanceolata KSIN 135. sinensis CJAP 124. japonica KSZ 136. szechuanensis CNIN 125. ningpoensis CTER 137. Colebrookea ternifolia CNAN 126. nanchuanensis COPP 138. oppositifolia Measurement of characters The characters used in the study, their states and character type according to Gower's coefficient of similarity (Gower, 1971), are given in Table 3. Petiole length (character 2) was measured to the nearest millimetre. Bracteole and pedicel lengths (characters 15 and 64) were measured to the nearest 0-5 mm. The lengths of the longest and shortest calyx tooth, calyx tube at anthesis and in fruit, corolla tube and upper and lower corolla lips (characters 17, 18,21, 22, 30, 32 and 33 respectively) were all measured to the nearest 0-05 mm. The number of leaves per whorl (character 4) was recorded only for those species with more than 2 leaves at a node (character 3). Wherever possible the leaves were counted from whorls in the middle of the stem. Characters 6, 7 and 8 refer to overall shape, basal shape, and apical shape respectively in the leaves. Each of the character states represents a broad class of shape : e . g . , ovate . Each specimen was assigned to the most appropriate class and coded accordingly. A similar procedure was adopted with bract and bracteole shape (characters 11 and 14). The coded data for each OTU is given in Appendix 2. An asterisk denotes missing or non-applicable data. Table 3 The characters scored, their states, and character types according to Gower's coefficient of similarity. Characters Type 1. Leaves : type 1 homophyllus 1 heterophyllus 2. Leaves : petiole length 3 in millimetres 3. Leaves : phyllotaxis 2 opposite 1 verticillate 4. Leaves : number per whorl 3 5. Leaves : comparative size of 2 members of a pair or whorl equal 1 unequal 6. Leaves : shape 2 linear 1 ovate 2 orbicular 3 trifid 7. Leaves : basal shape 2 cuneate 1 attenuate Characters 2 rounded 3 truncate 8. Leaves : apical shape acute 1 acuminate 2 obtuse 9. Leaves : margin entire 1 dentate 2 doubly dentate 10. Bracts : type not membranous 1 membranous 11. Bracts : shape linear 1 ovate 2 at least as broad as long 12. Bracts : fusion Ofree 1 connate for at least part of their length Type 2 TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 11 Characters Type 13. Bracts : persistence 2 persistent 1 deciduous 14. Bracteoles 2 absent 1 linear 2 lanceolate 15. Bracteoles : length in tenths of 3 a millimetre 16. Calyx : symmetry of teeth 2 regular 1 irregular 17. Calyx : length of longest tooth 3 in hundredths of a millimetre 18. Calyx : length of shortest tooth 3 in hundredths of a millimetre 19. Calyx : annulus in throat 2 absent 1 present 20. Calyx : number of main veins 3 21 . Calyx : tube length at anthesis 3 in hundredths of a millimetre 22. Calyx : tube length in fruit 3 in hundredths of a millimetre 23. Calyx : tooth form in fruit 2 not plumose 1 plumose 24. Calyx : tooth position in fruit 2 incurved 1 erect 2 spreading 25. Calyx : teeth hairs 2 absent 1 present 26. Calyx : tube hairs 2 absent 1 present 27. Cah.v: angles 2 more or less terete 1 strongly angled 28. Corolla : symmetry 2 lower lip with 1 lobe 1 lower lip with 2 lobes 29. Corolla : division 2 upper lip not emarginate 1 upper lip emarginate Corolla : tube length 3 in hundredths of a millimetre Corolla : exsertion 2 exserted 1 not exserted 32. Corolla : length of upper lip 3 in hundredths of a millimetre 33. Corolla : length of lower lip 3 in hundredths of a millimetre 34. Corolla : colour 2 not white Characters Type 1 white 35. Corolla : colour not yellow 1 yellow 36. Corolla : colour not purple 1 purple 37. Corolla : annulus absent 1 complete ring of hairs 2 interrupted ring of hairs 38. Corolla : invagination invagination absent 1 invagination present 39. Corolla : shape corolla not gibbous above annulus 1 corolla gibbous above annulus 40. Stamens : exertion exserted 1 not exserted 41. Anthers : size all anthers equal 1 lower pair reduced 42. Filaments : length equal 1 lower pair longer 2 upper pair longer 43. Anthers : locule number unilocular 1 bilocular-fused 2 bilocular-free 44. Filaments : hairs glabrous 1 hairy towards base 2 hairy towards middle 45. Filaments : base not bulbous 1 bulbous 46. Style : type gynobasic 1 terminal 47. Style : lobes plain 1 clavate 48. Style : basal shape not bulbous 1 bulbous 49. Disc : number of tumescent glands Onone 1 one 2 four 50. Nutlets : hairs absent 1 present 51. Nutlets : surface not verrucose 1 verrucose 12 J. R. PRESS Characters Type 52. Nutlets : apex 2 not rostrate 1 rostrate 53. Nutlets : number at maturity 2 Gone Hour 54. Hairs : septate, eglandular 2 absent 1 present 55. Hairs : septate, glandular 2 absent 1 present 56. Hairs : branched, stalked 2 absent 1 present 57. Hairs : branched, sessile 2 absent 1 present 58. Indumentum : adaxial leaf surface 3 glabrous Characters Type 1 sparsely hairy 2 densely hairy 59. Indumentum : abaxial leaf surface 3 glabrous 1 sparsely hairy 2 densely hairy 60. Indumentum : stem 3 glabrous 1 sparsely hairy 2 densely hairy 61 . Inflorescence : number of 3 verticils per spike 62. Inflorescence : form of verticils 2 not secund 1 sub-secund 2 secund 63. Inflorescence : number of 3 flowers per verticil 64. Flowers : length of pedicel 3 in tenths of a millimetre The programs The coded data were used to construct a similarity matrix using the measure of similarity described by Gower (1971). This matrix became the basis for analysis using the programs of the CLASP package (Rothamstead Experimental Station). This includes single-linkage and prin- cipal co-ordinates analyses (for full explanation see Gower, 1967 and Sneath & Sokal, 1973), a nearest neighbours list, and a method for clustering to maximize within-group mean similarity (WGMS). In this latter program all OTUs are randomly assigned to a preselected number of groups. An OTU is then transferred from one group to another if by doing so the WGMS is increased (and conversely the between-group mean similarity (BGMS) is decreased). This process is continued until further transfers no longer increase the WGMS. Since the number of groups is preselected, and every OTU must be assigned to a group, one may encounter the 'rag-bag' effect in which a number of unrelated OTUs are clustered to give a group with a very low WGMS. 4. Character variation Most taxa in the Labiatae are very similar to each other in general appearance, and morpho- logical variation tends to occur in relatively few characters. Characters which have long been considered of taxonomic importance (see Bentham, 1832-36, 1848; Endlicher, 1838; Bentham & Hooker, 1876; Briquet, 1897) within the Pogostemoneae include calyx and corolla shape, arrangement of stamens, and anther-locule number. A survey of 64 characters (see Table 3), and descriptions of variation within those which appear to be of taxonomic significance in the Pogostemoneae, is given below. Leaves All taxa have dorso-ventrally flattened leaves. Three basic lamina shapes are found; linear, ovate to orbicular, and lobed. The most common is the ovate to orbicular type, found in seven of the 10 genera (Colebrookea, Fig. 2, Comanthosphace, Eurysolen, Fig. 29, Keiskea, Leucoscep- trum, Rostrinucula, Fig. 26, and Tetradenid). In addition most species of Elsholtzia (e.g. E. ciliata, Fig. 2) some species of Pogostemon (e.g. P. mollis Benth., Fig. 2) and P. trinervis Fig. 2 Leaf shapes, margins, and petioles in the Pogostemoneae. (a) Pogostemon mollis x 1. (b) 'Dysophylla trinervis' x 2. (c) Elsholtzia kachinensis x 1. (d) Pogostemon glaber x 1. (e) Pogostemon speciosus x 1. (f) Pogostemon paniculatus x 1. (g) Elsholtzia stachyodea x 1. (h) Colebrookea oppositifolia x 0.5. (i) Elsholtzia ciliata x 1. (j) Elsholtzia densa x 1. (k) Dysophylla stellata x 1. (1) Dysophylla quadrifolia x 1 . 14 J. R. PRESS Chermsirivathana ex Press (see p. 74, Figs 2 & 33, provisionally called 'Dysophylla trinervis' by Chermsirivathana, 1963) have leaves of this shape. The distinction between an ovate and an orbicular leaf is a fine one and the two characters tend to grade into each other. Linear leaves are found only in Dysophylla (see Fig. 2), Pogostemon nilagiricus Gamble, and Elsholtzia pyg- maea W. Smith, although the latter is somewhat dubious since the observation is based on one specimen the leaves of which may better be considered as narrowly ovate. Three-lobed leaves occur in Elsholtzia integrifolia Benth. Variation in the leaf apex shape is limited. Usually it is acute or acuminate, although obtuse apices are found in some species of Dysophylla (e.g. 'D. trinervis', Figs 2 & 33), Elsholtzia (e.g. E. katchinensis Prain, Fig. 2), Pogostemon (e.g. P. mollis, Fig. 2), and Tetradenia (e.g. T. fruticosa Benth.). The leaf base shape can be divided into four character states; attenuate, cuneate, rounded, and tuncate. Colebrookea (Fig. 2), Comanthosphace, Keiskea, and Rostrinucula (Fig. 26) are exclusively cuneate. All Tetradenia species have rounded leaf bases. Eurysolen gracilis and Leucosceptrum canum (Fig. 29) have attenuate leaf bases. Elsholtzia and Pogostemon usually have cuneate (e.g. E. heterophylla Diels, Fig. 3, P. glaber Benth. in Wallich, Fig. 2), or attenuate (e.g. E. stachyodea (Link) Raiz & Saxena, Fig. 2) leaf bases, although a few species have rounded leaf bases (e.g. E. kachinensis, Fig. 2, P. speciosus Benth., Fig. 2). Truncate leaf bases are confined to Dysophylla (e.g. D. stellata (Lour.) Benth., Fig. 2) although 'D. trinervis' (Figs 2 & 33) and D. koehneana Muschler in Fedde have cuneate leaf bases, and in D. quadrifolia (Roxb.) Benth. (Fig. 2) the leaf base is usually attenuate. The leaf margins show varying degrees of incision, which may be conveniently scored as entire, dentate, or doubly dentate. Colebrookea (Fig. 2), Comanthosphace, Keiskea, Rostrinu- cula (Fig. 26), Eurysolen gracilis and Leucosceptrum canum (Fig. 29) all have dentate margins. In Pogostemon and Tetradenia the margin may be dentate (e.g. Pogostemon mollis, Fig. 2, Tetradenia fruticosa} or doubly dentate (e.g. Pogostemon paniculatus (Willd.) Benth., Fig. 2, Tetradenia goudotiiBriq.). In Dysophylla the margins may be entire (e.g. Dysophylla tomentosa Dalz., Fig. 3) or dentate (e.g. Dysophylla stellata, Fig. 2). Elsholtzia exhibits the whole range from entire (e.g. Elsholtzia integrifolia) to dentate (e.g. Elsholtzia densa Figs 2 & 28) and doubly dentate (e.g. Elsholtzia ciliata, Fig. 2). Heterophylly Heterophylly is rare but is found in two species of Elsholtzia (E. bodinieri Vaniot and E. heterophylla, Fig. 3). Unlike most taxa these two species possess leaves on the stolons which are much smaller, usually broader, and much hairier than the cauline leaves. Leaf size The Labiatae normally have one pair of leaves at each node of the stem; each leaf of a pair is identical to the other. Pogostemon gardneri Hook, f . and P. paniculatus (Fig. 3) are unusual in having one leaf of a pair much smaller than the other. This feature is occasionally found in P. purpurascens Dalz. , but is absent from the remainder of the Pogostemoneae. Phyllotaxis The Pogostemoneae (excluding Dysophylla section Verticillatae) usually follow the general Labiatae pattern of opposite, decussate leaf pairs, Colebrookea ternifolia is supposed by some authors to have ternate leaves but I have never found this. Species of Dysophylla section Verticillatae Benth. (Fig. 3) are very distinct in having whorls of leaves at each node. The number of leaves per whorl varies between , but is usually constant within each species ; the most common numbers are three, four, and five, but D. stocksii Hook.f . may have 14 or more leaves per whorl. Petioles Petiole length is sometimes difficult to ascertain, especially in species with attenuate leaf bases, but when a distinct petiole is present it varies from 1 mm (e.g. Elsholtzia heterophylla, Fig. 3) to 80 mm (e.g. E. fruticosa) long. In most Dysophylla species the leaves are sessile, but D. quadrifolia (Fig. 2) and a few other species have short but quite distinct petioles. d u Fig. 3 Phyllotaxis in the Pogostemoneae. (a) Pogostemon paniculatus x 0.5. Leaves in opposite pairs. Note the disparity in size of the members of the pairs of leaves, (b) Dysophylla tomentosa x 1. Leaves in whorls of six. (c) Elsholtzia densa x 1. Leaves in opposite pairs, (d) Dysophylla linearis x 1. Leaves in whorls of four, (e & f ) Elsholtzia heterophylla x 1 . (e) Leaves in opposite pairs, (f ) Stolons bear smaller, more rounded leaves than those of the stems. 16 J. R. PRESS Fig. 4 Bract types in the Pogostemoneae. (a) Eurysolen gracilis x 5. (b) Elsholtzia luteola x 5. (c) Elsholtzia ciliata x 5. (d) Dysophylla auricularia x 10. (e) Elsholtzia kachinensis x 5. (f) Rostrinucula dependens x 5. (g) Elsholtzia flava x 5. (h) Elsholtzia densa x 5. (i) Pogostemon paniculatus x 5. (j) Tetradenia fruticosa x 10. (k) Comanthosphace japonica x 5. (1) Elsholtzia pilosa x 5. (m) Keiskea elsholtzioides x 5. (n) Leucosceptrum canum x 5. CENeRAi * Bulletin of the British Museum (Natural History) *. ^v f/f\ Botany series Vol 10 1982 British Museum (Natural History) London 1982 Dates of publication of the parts No 1 30 September 1982 No 2 28 October 1982 No 3 . . . 25 November 1982 23 December 1982 ISSN 0068-2292 Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset Contents Botany Volnme 10 No 1 Taxonomic studies in the Labiatae tribe Pogostemoneae J. R. Press No 2 The typification of Hudson's algae: a taxonomic and nomenclatural reappraisal L. M. Irvine & P. S. Dixon 91 No 3 Seaweeds of the Faroes . . . 107 No 4 The lichen genus Steinera A. M. Henssen & P. W. James . . 227 TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 17 Bracts The Pogostemoneae show considerable bract variation, particularly in colour, shape, fusion, and persistence. Six combinations of these four characters can occur: (1) Bracts brown, broader than long, free, deciduous; this combination is found in Com- anthosphace, Rostrinucula and Leucosceptrum canum (Figs 4, 26). The bracts often fall before the flower buds open. (2) Bracts brown, broader than long, free, persistent; this combination is found in Tetradenia and Elsholtzia section Elsholtzia and Elsholtzia section Aphanochilus series Platyelasmeae (Figs 4, 28). In Elsholtzia section Elsholtzia the bracts are membranous. (3) Bracts brown, broader than long, fused, persistent; in Elsholtzia section Cydostegia and E. luteola Diels (Figs 4, 27) the bracts are again membranous and each pair is connate at the margins, forming a cyathium. (4) Bracts green, broader than long, free, persistent; in Elsholtzia concinna Vaut. and E. kachinensis (Fig. 4) the bracts are green and never membranous. (5) Bracts green, ovate, free, persistent; this type includes the following (all shown in Fig. 4): some Dysophylla species (e.g. D. auricularia) , Elsholtzia flava (Benth.) Benth., Eurysolen gradlis, Keiskea, and some Pogostemon species (e.g. P. paniculatus). (6) Bracts green, linear, free, persistent; Colebrookea and the remaining species of Dysophyl- la (e.g. D. peguana Prain), Elsholtzia (e.g. E. pilosa (Benth.) Benth. Fig. 5), and Pogostemon (e.g. P. fraternus Miq.) all have this character combination. Bracteoles Bracteoles may be present or absent. When present they are usually shorter, narrower, and more hairy than the bracts. Bracteoles are absent in Keiskea, Tetradenia, Elsholtzia sections Elsholtzia and Cydostegia, and section Aphanochilus series Platyelasmeae. In Comanthos- phace, Rostrinucula, and Leucosceptrum canum the bracteoles are caducous, and, like the bracts, often fall before the flower buds open. Indumentum Four hair types (Fig. 5) are found: (1) Septate eglandular hairs; these are simple hairs each at least several cells long. The cells are laterally compressed and arranged so that the narrow sides alternate along the length of the hair, and the terminal cell is usually somewhat pointed. (2) Septate glandular hairs; these are structurally identical with the septate eglandular hairs except for the terminal cell, which is a globular, single-celled gland. (3) Branched stalked hairs; these are the 'stellate' hairs of earlier authors, each hair in fact having a central axis bearing irregular branches along its length. The hairs are multicellular, the septa clearly visible. (4) Branched sessile hairs; similar to branched stalked hairs, but distinguished by the lack of a central axis, the branches radiating irregularly from a central point. Most species have septate, eglandular hairs only. A number of species e.g. Dysophylla linearis Benth. and Pogostemon brachystachyus Benth. have a mixture of glandular and eglandular septate hairs. Branched stalked hairs are less common, being found in five genera. Comanthos- phace, Leucosceptrum, and Rostrinucula bear branched stalked hairs with a few septate eglandular hairs. Tetradenia and a number of Elsholtzia species, e.g. E. capituligera (Dunn) Y. C. Wu and E. stachyodea, have a mixture of branched stalked and septate eglandular hairs. Other species, e.g. E. eriostachya and E. fruticosa, have branched stalked, septate eglandular and septate glandular hairs. Branched sessile hairs are found in two species of Pogostemon: in P. tuberculosus Benth. the hairs have a central boss from which the short stiff branches radiate, whilst in P. velatus Benth. there is no central boss and the branches are long and flexuous. All species possess some hairs, although these may be restricted to the inflorescence. The abaxial leaf surface is always more hairy than the adaxial; the veins more hairy than the surface of the lamina. The density of hairs on the stems decreases with age, so that the lowest parts of the 18 J. R. PRESS Fig. 5 Hair types in the Pogostemoneae. (a) septate glandular hair, (b) septate eglandular hair, (c & d) branched stalked hairs, (e) branched sessile hair, the branches short and stiff, (f ) branched sessile hair, the branches long and flexuous. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 19 stem are frequently glabrous. The density of the indumentum varies between individuals as well as between species. Only Pogostemon glaber and some species of Dysophylla, e.g. D. yatabeana Makino, have glabrous stems and leaves. All other species have some indumentum here, ranging from very thin, e.g. Elsholtzia luteola, to very dense, e.g. Pogostemon mollis. Calyx The calyx is generally campanulate, five toothed, with the teeth sometimes much elongated. The teeth are unequal in Keiskea (Fig. 6), most Elsholtzia species (e.g. E. ciliata, Fig. 6 and E. densa, Figs 6, 28) and some species of Pogostemon (e.g. P. fraternus, Fig. 6). In Tetradenia (Fig. 6) the upper tooth is much the broadest and overlaps the lateral teeth. The calyx tube is strongly veined, the most prominent veins generally considered as the main veins. Three species, Comanthosphace ningpoensis (Hemsley) Hand.-Mazz., Elsholtzia integri- folia, and ' Elsholtzia japonica' have 15 main veins; Comanthosphace (excluding C. ningpoensis) and Leucosceptrum canum (Fig. 6) have 12. The remaining taxa have five or 10 main veins, although in some species of Dysophylla (e.g. D. tomentosa) the number is difficult to determine. The main, central veins of the teeth are thickened to form five strong ribs in Colebrookea (Fig. 6), D. griffithii Hook.f., Dysophylla pentagona C.B. Clarke ex Hook.f., and D. stocksii. The calyx is usually hairy on the outer surface, and the teeth are fringed with cilia. Three species, Elsholtzia kachinensis, E. penduliflora W. Smith and Pogostemon glabratus Chermsiri- vathana ex Press (see p. 71, Fig. 32, provisionally called 'Dysophylla glabrata' by Chermsiri- vathana, 1963) have a glabrous calyx. Keiskea japonica (Fig. 6) has a hairy calyx-tube but glabrous teeth, while several species, including Pogostemon amarantoides Benth. and P. paludosus Benth. , have a glabrous calyx-tube and hairy teeth. The presence of an annulus of stiff hairs in the throat of the calyx is confined to Keiskea and some species of Pogostemon (e.g. P. fraternus and P. litigiosus Doan, both in Fig. 6). The calyx may be accrescent at the fruiting stage. In Elsholtzia series Platyelasmeae the increase in size is striking, the fruiting calyx (Figs 6, 28) being up to eight times larger than at anthesis. In Colebrookea the calyx teeth are plumose, becoming greatly elongated but not much widened in fruit. The calyx teeth are normally erect during fruiting, but in Dysophylla stellata and Pogostemon litigiosus they are spreading and in other species of Dysophylla (e.g. D. pentagona) and Pogostemon (e.g. P. nelsonii Doan) strongly incurved. Corolla The corolla is zygomorphic, although in some species of Dysophylla weakly so and not obviously bilabiate. Generally the corolla lobes are arranged to form an upper group of three lobes and a single lower lobe. Elsholtzia integrifolia is exceptional in having a bifid lower lobe. In Dysophylla, Pogostemon, and Rostrinucula (Fig. 26) the upper central lobe is entire; in the remaining six genera it is shallowly to deeply emarginate. The corolla tube shows little variation except in length. At maturity it may reach 9-5 mm in length (e.g. Elsholtzia bodinieri) or be as little as 0-5 mm long (e.g. Tetradenia hildebrandtii Briq.) and is usually rather slender at the base widening gradually to the throat. In Pogostemon the upper lip is longer than or equals the lower lip. In the other genera the lower lip is longer than or equals the upper lip. An annulus of hairs is found in the corolla throat of six genera. In Comanthosphace, Keiskea, and Tetradenia (Fig. 7) the annulus is a complete ring at, or slightly below, the level of insertion of the staminal filaments. In a number of Elsholtzia species, e.g. E. capituligera, E. hunanensis Hand.-Mazz., and E. stauntonii Benth. (Fig. 7), the annulus is an open ring, the lines of hairs projecting dorsally and ending under the upper lip. Rostrinucula (Figs 7, 26) has a partial annulus composed of clusters of hairs at the insertion point of each staminal filament. In addition to the hair-clusters there is an invagination on the ventral surface of the corolla forming a crescent-shaped papilla in the throat, itself also hairy and forming part of the annulus. In Eurysolen gracilis (Figs 7, 29) the corolla is gibbous slightly above the base. An invagina- 20 J. R. PRESS Fig. 6 Calyx shapes and veination in the Pogostemoneae. Only the main veins are shown, (a) 'Dysophylla trinervis' x 5. (b) Elsholtzia flava x 5. (c) Leucosceptrum canum x 5. (d) Pogostemon parviflorus x 5. (e) Elsholtzia densa (fruiting) x 5. (f) Colebrookea oppositifolia (fruiting) x 10. (g) Tetradenia goudotii x 5. (h) Elsholtzia ciliata x 5. (i) Pogostemon litigiosus x 5. (j) Pogostemon fraternus x 5. (k) Keiskea japonica x 5. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE Fig. 7 Corolla structures in the Pogostemoneae. (a) Keiskea japonica x 5. (b) Eurysolen gracilis x 5. (c) Elsholtzia stauntonii x 5. (d) Rostrinucula dependens x 5. (e) Tetradenia fruticosa x 5. (f) Comanthos- phace japonica x 5. (g) Colebrookea oppositifolia x 20. (h) Pogostemon tuberculosus x 5. (i) Leucosceptrum canum x 5. 22 J. R. PRESS tion similar to that in Rostrinucula occurs immediately below the gibbous curve forming a hairy papilla which all but closes the corolla tube. The corollas are usually yellow, white, and rose-pink to purple, the latter being most common. Stamens In common with most Labiatae, the Pogostemoneae have four stamens arranged into two pairs: an upper and a lower pair. The filaments may be equal in length or one pair longer than the other. In Colebrookea, Rostrinucula (Fig. 26) and Tetradenia the filaments are equal; in Eurysolen gracilis (Fig. 29) the upper filaments are longest; in Comanthosphace, Keiskea, and Leucosceptrum canum the lower filaments are longest. In the remaining genera all three conditions can occur. The anthers are almost always exserted, the only exceptions being Elsholtzia aquatica C.H. Wright and Elsholtzia series Platyelasmeae (Fig. 28). The filaments are inserted towards the top of the corolla tube. In Eurysolen gracilis (Figs 26, 29), Rostrinucula dependens (Rehder) Kudo, and R. sinensis (Hemsley) Y.C. Wu, each filament is marked by a bulbous swelling below the point of insertion, which bears annular hairs. Dysophylla, Pogostemon, and Leucosceptrum canum (Fig. 8) have hairs on the filaments. In the first two genera the hairs are long, thread-like, and usually purple due to cross-wall pigments. In Leucosceptrum canum the hairs are short and white. In Dysophylla and most species of Pogostemon the hairs are borne towards the middle of the filament; in Leucosceptrum canum (Fig. 8) and some species of Pogostemon (e.g. P. hispidus Prain and P. travancoricus Beddome, Fig. 8) the hairs are borne towards the base of the filament. The anthers are usually equal in size. However Elsholtzia pilosa has two smaller, lower anthers often reduced to half the size of the upper ones. Anthers are bilocular in Elsholtzia, Keiskea (Figs 8, 27, 28), and Tetradenia and unilocular in the remaining genera. Tetradenia and most species of Elsholtzia have locules confluent through partial fusion. Keiskea, Elsholtzia hunanensis, and 'E. japonica' have distinctive free locules. Style Two style types occur in the Labiatae; (1) gynobasic, when the style arises from the base of, and between, the deeply-divided lobes of the ovary, (2) terminal, when the ovary is shallowly-lobed and the style is not basal. Gynobasic styles are found throughout the family except for the subfamily Ajugoideae which have terminal styles. In Leucosceptrum canum (Fig. 9) the style usually appears terminal but in some specimens it more nearly approaches the gynobasic condition. The reverse seems true of Elsholtzia flav a (Fig. 9). All other Pogostemoneae have gynobasic styles. The style is typically long, slender and straight or slightly curved, with a bifid tip. The lobes at the tip are subulate and equal. In Elsholtzia the style may vary in base and lobe shape. In some species (e.g. E. ochroleuca Dunn, E. stachyodea, Fig. 8) the style bears a bulbous swelling at the base, just above the point of emergence from between the nutlets. In Elsholtzia series Platyelasmeae (Fig. 8) the style lobe tips are clavate, a condition found also in Pogostemon litigiosus. Disc The disc may be regular or may bear one to several tumescent lobes. In Elsholtzia and Keiskea (Figs 8, 27, 28) there is a single, somewhat elongated lobe on the posterior edge of the disc. In Tetradenia (Fig. 8) there are four lobes, spaced regularly around the disc. They are bright-red and quite large, overtopping the young nutlets. The other genera have regular discs. Nutlets In the Labiatae there are usually four nutlets, a feature generally true of the Pogostemoneae. Exceptions are Colebrookea oppositifolia (Fig. 10), Dysophylla stocksii, Esholtzia kachinensis o Fig. 8 Stamen types in the Pogostemoneae. All stamens x 10. All anther details x 20. (a) Keiskea japonica. (b) Colebrookea oppositifolia. (c) Comanthosphacesublanceolata. (d) Elsholtzia ciliata. (e) Leucoscep- trum canum. (f) Dysophylla linearis. (g) Pogostemonparviflorus. (h) Pogostemon atropurpureus . 24 J. R. PRESS Fig. 9 Style and disc types in the Pogostemoneae. All x 10. (a) Leucosceptrum canum. (b) Elsholtzia densa. (c) Elsholtzia stachyodea. (d) Elsholtzia fruticosa. (e) Comanthosphace japonica. (f) Pogostemon tuberculosus. (g) Keiskea japonica. (h) Tetradenia fruticosa. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 25 and Keiskea japonica (Fig. 10) with only one nutlet at maturity, the other three nutlets apparently undergoing early abortion. The nutlets are generally obovoid and blunt at the apex. However, Rostrinucula nutlets (Figs 10, 26) have a rostrate tip at maturity. Hairy nutlets are found in Colebrookea (Fig. 10), Rostrinucula (Figs 10, 26), all Comanthos- phace species (Figs 9, 10) except C. nanchuanensis Y.C.Wu & Li, and in Elsholtzia japonica where the hairs are confined to the apex of the nutlet. All other species have glabrous nutlets. Variation in nutlet ornamentation is limited. Most species have nutlets with a smooth to slightly rugulose surface. By contrast those of Elsholtzia series Platyelasmeae (Figs 10, 28) are distinctly verrucose towards the apex. In Keiskea japonica (Fig. 10) the nutlet has ridges which form an almost reticulate pattern. In Colebrookea the single ripe nutlet does not separate from the calyx, the two structures acting as a single dispersal unit. Inflorescence The inflorescence is usually racemose (paniculate in some species of Pogostemon). It varies greatly in length, and in the number of whorls of flowers, or verticils. The verticils are secund in Keiskea and many species of Elsholtzia (e.g. E. ciliata, E. luteola (Fig. 27) and E. stauntonii). In the paniculate species of Pogostemon the verticils are sub-secund, i.e. the flowers are secundly arranged in each verticil but the verticils themselves are not secund on the stem. The flowers in all genera may be pedicellate or sessile. Pollen There is no complete pollen survey of the Pogostemoneae. However, sufficient data is available to make some considerations. Labiatae pollen is remarkably uniform and only two features, the number of colpi and the number of nuclei, show taxonomically useful variation. The pollen grains are either bi-nucleate and tri-colpate or tri-nucleate and hexa-colpate (see Erdtman, 1945, 1952). A number of authors (e.g. Wunderlich, 1963; El-Gazzar & Watson, 1968) have advanced the view that these characters have a high taxonomic value and have used them to good effect in studies on relationships within the Labiatae and closely related groups. Pollen from eight Pogostemoneae genera has been examined by other workers. To provide a full sample I examined Eurysolen gracilis and Leucosceptrum canum pollen under the light microscope. The data are summarized in Table 4. Table 4 Numbers of nuclei and colpi in pollen grains of the Pogostemoneae. Genus Number of nuclei per cell Number of colpi per cell Elsholtzia 3 6 Keiskea 3 6 Tetradenia 3 6 Eurysolen - 3 Comanthosphace 2 3 Leucosceptrum - 3 Rostrinucula 2 3 Pogostemon 2 3 Dysophylla 2 3 Colebrookea 2 3 26 J. R. PRESS Fig. 10 Nutlet types in the Pogostemoneae. Profile and inner face of nutlet are shown. All x 10. (a) Keiskea japonica. (b) Colebrookea oppositifolia. (c) Eurysolen gracilis. (d) Dysophylla stellata. (e) Pogostemon mollis. (f) Leucosceptrum canum. (g) Comanthosphace barbinervis. (h) Elsholtzia flava. (i) Elsholtzia densa. (j) Rostrinucula dependens. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 27 5. The analyses Principal co-ordinates analysis A general view of overall similarities can be obtained by looking at two-dimensional plots using the first four eigenvectors of the principal co-ordinates analysis (Figs 11-13). These vectors account for 43% of the total variation and although low, this percentage is sufficiently high to provide a basis for comparing the different groups. The results show a clear division of the points into two major groups; one formed by Dysophylla and Pogostemon, the other containing the remaining taxa except Colebrookea and Rostrinucula which occupy intermediate positions between the two groups. Comanthosphace and Leucosceptrum canum (OTU 127) are rather peripheral members of the second group in Fig. 13, where they are quite distant from the main cluster. Two OTUs are outstanding in their positions. Dysophylla mairei Leveille (OTU 65) is clustered with Elsholtzia, while Elsholtzia aquatica (OTU 22) is clustered with Dysophylla and Pogostemon. This separation into two major groups provides a convenient division for further investiga- tion. Each group was used as a subset for which a second series of plots was prepared. Colebrookea and Rostrinucula were omitted from these plots. Figs 1416 show the plots for the Comanthosphace/ Elsholtzia subset. Comanthosphace and Leucosceptrum canum (OTU 127) are separated from the main cluster of Elsholtzia species (Fig. 14). Keiskea and Tetradenia are also separated from Elsholtzia but only when the third (Fig. 15) and fourth (Fig. 16) eigenvectors are used. Elsholtzia is divisible into two sub-groups, and this is more easily seen when only the species of Elsholtzia are marked on the plots (Figs 17-19). The first sub-group represents sections Cyclostegia and Elsholtzia and occupies that end of the cluster furthest away from Comanthos- phace. The species forming these two sections are quite intermixed. The second sub-group represents section Aphanochilus series Stenelasmeae and occupies that part of the main cluster nearest to Comanthosphace. The two sub-groups are separated in the original plots (Figs 14-r6) by Keiskea and Tetradenia. Figs 17-19 show an interesting distribution for the three species of Elsholtzia section Aphanochilus series Platyelasmeae. E. eriostachya (OTU 19) consistently segregates with section Aphanochilus series Stenelasmeae, whilst E. densa (OTU 20) segregates with sections Cyclostegia and Elsholtzia. The third species, E. manshurica (Kitagawa) Kitagawa (OTU 21), occupies an intermediate position between the sub-groups, although in Fig. 17 it is part of the sections Cyclostegia and Elsholtzia sub-group. Note also the position of OTU 8, E. concinna, in each of the plots. In all analyses Dysophylla mairei (OTU 65), Eurysolen gracilis (OTU 1) and Leucosceptrum plectranthoideum (Leveille) Marquand (OTU 128), remain within the sub- group formed by Elsholtzia section Aphanochilus series Stenelasmeae. The analysis of the Dysophylla/ Pogostemon subset is shown in Figs 20-22. These two genera form a single cluster within which is a marked distribution of points. Most Dysophylla species occupy one half of the cluster and Pogostemon occupies the other half. Elsholtzia aquatica (OTU 22) always clusters with Dysophylla. Clustering to maximize within-group-mean-similarity (WGMS) As an independant check to help decide which real groups may be present in the Pogostemoneae the data were analysed using a method for clustering to maximize WGMS. For this method the number of groups must be preselected (see p. 12) and here the similarity matrix was used as a guide. A pictorial representation of the similarity matrix was prepared by replacing the percentage similarities with appropriate symbols to make groups of similar OTUs more easily visible (Figs 23 and 24). There are two distinct patterns, one with six groups and a second with nine groups. Allocating the OTUs among six groups yields little information. The WGMS and BGMS values (Table 5) are so close that, with the possible exception of group four Comanthosphace, Rostrinucula, and Leucosceptrum canum (OTU 127), no discrete groups can be recognized. 28 J. R. PRESS vector 1 *'" A A ;:** si. AA A A A (I) A A (?) A A A A A A A A AA o AA A AA A A 65 o o o o o o O u o oo o o vector 2 Fig. 11 Two-dimensional plot of the Pogostemoneae using principal co-ordinates analysis. Key: Pogostemon, ; Dysophylla, O; Elsholtzia, A; Comanthosphace , ; Keiskea, A; Tetradenia, ; Leucosceptrum, ; Rostrinucula, Rostrinucula, ; Colebrookea, ; Eurysolen, . TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 29 vector 1 A A A A A A A A 65, V/ *$ A A 22 o O V o o 00 00 vector 3 Fig. 12 Two-dimensional plot of the Pogostemoneae using principal co-ordinates analysis. Key: Pogostemon, ; Dysophylla, O; Elsholtzia, A; Comanthosphace , ; Keiskea, A; Tetradenia, ; Leucosceptrum, ; Rostrinucula, ; Colebrookea, ; Eurysolen, . 30 J. R. PRESS vector 1 65 A * A A A A(r) AA A A A A A A A A A o O O O o o o o o o 22 vector Fig. 13 Two-dimensional plot of the Pogostemoneae using principal co-ordinates analysis. Key: Pogostemon, ; Dysophylla, O; Elsholtzia, A; Comanthosphace, ; Keiskea, A; Tetradenia, @; Leucosceptrum, ; Rostrinucula, ; Colebrookea, ; Eurysolen, . TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 31 vector 1 65 A A A A 128 A A A (J) A A 127 A A A A 126 vector 2 Fig. 14 Two-dimensional plot of the Comanthosphacel Elsholtzia subset using principal co-ordinates analysis. Key: Dysophylla, O; Elsholtzia, A; Comanthosphace, ; Keiskea, A; Tetradenia, ; Leucoscep- trum, ; Eurysolen, . 32 J. R. PRESS vector 1 127 128 126 65 O A A A A A A A 25 A AA vector 3 Fig. 15 Two-dimensional plot of the Comanthosphacel Elsholtzia subset using principal co-ordinates analysis. Key: Dysophylla, O; Elsholtzia, A; Comanthosphace , ; Keiskea, A; Tetradenia, ; Leucoscep- trum, ; Eurysolen, (j). TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 33 vector 1 IT) (T) 00 128 127 126 A A A A A A A A A A A A A A vector Fig. 16 Two-dimensional plot of the Comanthosphacel Elsholtzia subset using principal co-ordinates analysis. Key: Dysophylla, O; Elsholtzia, A; Comanthosphace, ; Keiskea, A; Tetradenia, ; Leucoscep- trum, ; Eurysolen, . 34 J. R. PRESS vector 1 A A A A A A A A A A A ,20 O vector 2 Fig. 17 Two-dimensional plot of the Comanthosphacel Elsholtzia subset showing the distribution of species of Elsholtzia only. Key: Section Elsholtzia ; Section Cydostegia O; Section Aphanochilus series Stenelasmeae A; Section Aphanochilus series Platyelasmeae A. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 35 vector 1 21 A A * A A A A A O vector Fig. 18 Two-dimensional plot of the Comanthosphacel Elsholtzia subset showing the distribution of species of Elsholtzia only. Key: Section Elsholtzia ; Section Cyclostegia O; Section Aphanochilus series Stenelasmeae A; Section Aphanochilus series Platyelasmeae A . 36 J. R. PRESS vector 1 19 A A A A A A vector Fig. 19 Two-dimensional plot of the ComanthosphacelElsholtzia showing the distribution of species of Elsholtzia only. Key: Section Elsholtzia ; Section Cyclostegia O; Section Aphanochilus series Stenelasmeae A; Section Aphanochilus series Platyelasmeae A . TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 3? vector 1 6 I O 7 o 22 O OOAO 61 00 00 O O I 7 , R o O \ \ 70 o ^ Q o b / 53 / v w Op O / ^^ 59 R P 69 o 58 K / o / / o // / 55 43 I % 109 / 86 66 / w 102 85 .' 82 vector 2 Fig. 20 Two-dimensional plot of the Dysophyllai Togostemon subset using principal co-ordinates analysis. Key : Pogostemon , ; Dysophylla , O ; Elsholtzia , A . 38 J. R. PRESS vector 1 o o o o 22. o O O 62 O O^" 54 O OO / 67 / O / 85 53 ' 59 O 43 \ 2 ^O / N _ 69 110 109 65 ' O * X 86 . / \ /O \ " 6 66 82 X 71 vector 3 Fig. 21 Two-dimensional plot of the DysophyllalPogostemon subset using principal co-ordinates analysis. Key: Pogostemon, ; Dysophylla, O; Elsholtzia; A . TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 39 vector 1 "f 56 \ A O 5 , 4 O * * O 66 - 00 T o \ o ' 58 O J 43 * QC. _^ 59 O X O 86 O O 7Q 68 - O ^O A - f O 58 67 f O O 61 109 vector Fig. 22 Two-dimensional plot of the Dysophylla/Pogostemon subset using principal co-ordinates analysis. Key: Pogostemon, : Dysophylla, O; Elsholtzia, A. 40 J. R. PRESS Colebrookea Tetradenia Comanthosphace , Rostrinucula, Leucosceptrum canum /?///// Pogostemon Dysophylla Fig. 23 Diagram showing six possible groups indicated in the similarity matrix. Allocating the OTUs among nine groups (Table 6) is more informative and confirms the general picture obtained from the principal co-ordinates analyses. Comanthosphace and Leucosceptrum canum (OTU 127) are confirmed as forming a group separate from other OTUs but here the group includes Rostrinucula. As in the principal co-ordinates analysis (Figs 11-13) the remaining taxa (except Colebrookea) are divided into two subsets. The first contains groups four, five, and seven which share low BGMS values with the remaining groups, but high BGMS values with each other. Dysophylla section Verticillatae is placed in group four, but species of section Oppositifoliae are divided between all three groups, D. salicifolia (OTU 69) in group four, D. myosuroides (OTU 71) and D. rugosa (OTU 70) in group five, and D. auricularia (OTU 68) in group seven. Elsholtzia aquatica (OTU 22) is placed with Dysophylla section Verticillatae in group four. Pogostemon is divided between groups five and seven. The second subset contains groups two, three, six, and nine, but groups two (Keiskea) and three (Tetradenia) are quite discrete. Not so Elsholtzia in groups six and nine, which share high TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 41 Colebrookea Tetradenia Comanthosphace , Rostrinucula , Leucosceptrum canura Pogostemon Elsholtzia section Aphanochilus series Stenelasmeae Elsholtzia sections Elsholtzia Cyclostegia Keiskea Fig. 24 Diagram showing nine possible groups indicated in the similarity matrix. BGMS values compared to their respective WGMS values. Group six contains Elsholtzia section Aphanochilus series Stenelasmeae together with E. eriostachya (OTU 19) from series Platyelasmeae and E. concinna (OTU 8). Once again Dysophylla mairei (OTU 65), Eurysolen gracilis (OTU 1), and Leucosceptrum plectranthoideum (OTU 128) are all included in this group. Group nine contains Elsholtzia sections Cyclostegia and Elsholtzia plus E. densa (OTU 20) and E. manshurica (OTU 21) of Elsholtzia section Aphanochilus series Platyelas- meae. Group eight is an anomaly, containing the unlikely combination of Colebrookea with one species each of Dysophylla and Pogostemon. The heterogeneity of this combination is reflected in the very low WGMS value of 79-3% and the obvious similarity with Dysophylla and Pogostemon in groups four, five, and seven. A slight rearrangement of the OTUs to remove the anomaly results in the similarities shown in Table 7. Here the species of Dysophylla and Pogostemon are removed, 'Dysophylla glabrata' (OTU 67) now placed in group four and 42 J. R. PRESS Table 5 WGMS and BGMS values for six groups in the cluster- ing to maximize WGMS analysis. 1 84-8 2 82-4 88-9 3 73-7 72-0 79-7 4 71-6 70-4 72-1 85-8 5 82-3 81-4 70-9 71-2 87-9 6 80-8 78-8 70-1 67-7 76-4 88-3 1 Key to groups: 1 Colebrookea, species of Pogostemon, and Dysophylla section Opposi- tifoliae. 2 species of Pogostemon. 3 Keiskea, Tetradenia, Elsholtzia and Eurysolen. 4 Comanthosphace, Rostrinucula and Leucosceptrum canum. 5 species of Pogostemon. 6 Dysophylla section Verticillatae. (For exact distribution of OTUs see Appendix 4.) Table 6 WGMS and BGMS values for nine groups in the clustering to maximize WGMS analysis. 1 2 3 85-8 67-9 70-3 90-8 70-2 94-7 4 67-9 64-2 65-8 88-8 5 71-8 69-3 70-4 80-0 85-8 6 72-6 74-8 74-0 71-8 74-3 83-1 7 70-7 70-0 68-0 78-9 83-0 72-2 89-6 8 67-2 65-8 66-7 75-4 73-8 71-3 76-5 79-3 9 72-8 77-3 76-4 70-8 72-2 78-3 73-0 68-5 86-7 1 Key to groups: 1 Comanthosphace, Rostrinucula, and Leucosceptrum canum. 2 Keiskea. 3 Tetradenia. 4 Dysophylla section Verticillatae. 5 species of Pogostemon and Dysophylla section Oppositifoliae. 6 Elsholtzia section Aphanochilus and Eurysolen. 7 species of Pogostemon. 8 Colebrookea, 'Dysophylla glabrata', and Pogostemon amarantoides. 9 Elsholtzia sections Cyclostegia and Elsholtzia. (For exact distribution of OTUs see Appendix 4.) Pogostemon amarantoides (OTU 80) in group seven. These transfers produce only small changes in the similarity values for these groups but the effect on group eight is striking. The WGMS rises to 95-2% and the links with groups four, five, and seven are greatly reduced. Single-linkage analysis In broad terms the single-linkage analysis reproduces the groupings given by the PCA and WGMS analyses but some minor deviations are of interest. Colebrookea, Keiskea, and Tetradenia form discrete groups in the dendrogram (Fig. 25). Elsholtzia can be divided into two subgroups, sections Cyclostegia/ Elsholtzia and section Aphanochilus series Stenelasmeae. However, section Aphanochilus series Platyelasmeae is noticeably separate from the remainder of Elsholtzia, and for the first time Eurysolen gracilis (OTU 1) is separated from Elsholtzia section Aphanochilus series Stenalasmeae. Dysophylla and Pogostemon are very similar to each other, linking at, or above, the 91% level; Dysophylla section Verticillatae forms a subgroup but TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 43 Table 7 Adjusted WGMS and BGMS values for nine groups in the clustering to maximize WGMS analysis. 1 2 3 88-5 69-5 71-3 90-8 70-2 94-7 4 68-1 64-2 65-8 88-8 5 68-9 69-3 70-4 80-0 85-8 6 74-0 74-8 74-0 71-8 74-3 83-1 7 67-4 70-0 68-0 78-9 83-0 72-2 89-6 8 68-5 65-4 70-3 65-4 71-6 72-2 69-4 95-2 9 73-9 77-3 76-4 70-8 72-2 78-3 73-0 67-1 86-7 1 Key to groups: 1 Comanthosphace and Leucosceptrum canum. 2Keiskea. 3 Tetradenia. 4 Dysophylla section Verticillatae. 5 species of Pogostemon and Dysophylla section Oppositifoliae. 6 Elsholtzia section Aphanochllus and Eurysolen. 1 species of Pogostemon. 8 Colebrookea. 9 Elsholtzia sections Cydostegia and Elsholtzia. species of section Oppositifoliae are scattered among the species of Pogostemon. Comanthos- phace, Leucosceptrum canum (OTU 127) and Rostrinucula form a group but with C. nan- chuanensis (OTU 126), L. canum and Rostrinucula linking at low similarities. Dysophylla mairei (OTU 65) and Leucosceptrum plectranthoideum (OTU 128) link with Elsholtzia at high similarity levels. Elsholtzia aquatica (OTU 22) links with Dysophylla and Pogostemon at a somewhat lower similarity. 6. The taxa: discussion Comanthosphace and Leucosceptrum Comanthosphace, Rostrinucula, and Leucosceptrum canum form a rather complex group. Rostrinucula is discussed separately on p. 47 and Leucosceptrum plectranthoideum (OTU 128) is discussed on p. 57. Six species of Comanthosphace are morphologically very similar and are always closely grouped together. The seventh species, C. nanchuanensis (OTU 126), is much less similar to the other six and less obviously a member of the group, especially as shown by the second principal co-ordinates analysis (Figs 14-16) and the single-linkage dendrogram (Fig. 25). Leucosceptrum canum (OTU 127) is most similar to species of Comanthosphace but, like C. nanchuanensis, tends to lie on the fringe of the main group. Comanthosphace can be recognized by a number of characters and character combinations. The most obvious and consistent are the broad, membranous, deciduous bracts and narrow, deciduous bracteoles, the closed annulus of hairs within the corolla, the hairy nutlets, and the dense pubescence of branched and unbranched hairs on all parts of the plant. Also, the flowers are large with relatively short, more or less equal calyx teeth, the upper lobe of the corolla is emarginate, and the anthers are unilocular. Comanthosphace nanchuanensis appears to lack bracteoles although this is uncertain since both bracts and bracteoles fall very early and are not present on every specimen in related species. Similarly, the glabrous appearance of the nutlets may be recorded erroneously since no fully mature nutlets were available. The annulus within the corolla is incomplete. All the other characters distinguishing Comanthosphace are shared by C. nanchuanensis and the inclusion of this species within the Comanthosphace group seems intrinsically correct. Certainly the differ- ence in overall similarity between Comanthosphace pro majore and C. nanchuanensis is not sufficiently great to justify the removal of the latter to a separate genus. 44 J. R. PRESS pco c\j cop p co cop pco CO cop o> pco o cnp pen O rvl o vO rvj LA o 00 rvj LA LA o rvj LA o \o OJ LA o u> OJ o OO OJ o o OJ s rA o o LA lA OJ LA o OJ lA J. IA J- OJ o R. 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PRESS 8 -R ft i a o o o IA o J- o LA o o o o LA o o o o LA o o o o ^0 ^ O LA O LA LA LA o o LA OJ o o o IA OJ OJ UA *" "* C^ 1 - rA OJ OJ o o OJ OJ r- ,- O o o o ,- OJ o o r- r- OJ OJ ,- - ,- T - O <- OJ ,- r- <- OJ r- CM OJ o OJ OJ o r- t- o I- OJ o - - OJ OJ r- T- OJ t- O <- OJ r- o 1- OJ I- OJ o O o o o o o O o i- o o o o o o O o O o o o o o o o o o O o o o o O o o o o o o O o o O o o o o o o o o o o o - *- o o O o o o o o o o o o o , o o o o o o o o O o o o o o o o o o o o o o o o o * o o o o o o o O o o o o o o o o o o o o o o o o o o o o I- o o o o o o o o o o o o o o o o o o o o o o o r- o o o o o o o o o o o o o o o o o o <- o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o OJ OJ OJ OJ OJ OJ OJ OJ o OJ OJ OJ OJ OJ OJ OJ OJ OJ OJ OJ OJ OJ OJ OJ OJ OJ OJ OJ o o o o o o o o 1- o o o o o o o o o o o o o o o o o o o o o o o o o * OJ r- o o o o o OJ o <- OJ * o v- o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o <- J n g o OJ o L^- EH o K> o oo p. w o J- ON z. n to o o o o OJ E o CSTE121 20 CBAR122 050 o o o rA OJ CP g o o o o -3- (V < 3 o IA o LA OJ z. h- 1 z o o LA o vO |H'I1 OTU dix 3 cont. 1 J. R. PRESS 2 3 4 5 OTU % OTU % OTU % t OTU % OTU % 106 107 90.9 112 89.5 113 88 .9 77 88.5 108 88.4 107 108 97.6 113 94.0 112 93 .3 77 92.4 86 9L8 108 107 97.6 113 95.5 112 i 95 .2 77 94.1 104 94.1 109 112 96.1 110 93.8 69 91 .6 111 9L2 82 90.2 110 102 93.8 108 93.8 81 91 .2 69 90.5 113 90.5 111 102 93.1 100 93.0 105 93 .0 108 92.6 103 92.3 112 77 95.2 108 95.2 113 94 .9 74 94.1 72 93.4 113 108 95-5 115 95.1 112 94 .9 77 94.4 107 94.0 114 75 93.4 79 92.5 77 92 .3 73 9L1 74 9LO 115 99 95.1 113 95.1 75 92 .2 79 92.2 77 92.0 116 86 92.4 91 91.1 70 90 .6 102 89.5 101 89.4 117 83 86.3 82 86.1 109 86 .0 102 84.4 84 84.0 118 119 94.8 126 83.6 125 83 .4 124 82.1 122 81.8 119 118 94.8 125 83.9 120 82 .7 126 81.6 124 81.4 120 125 93.5 122 89.4 124 89 .4 123 89.2 121 88.4 121 124 97.7 123 97.7 122 95 .7 125 92.3 120 88.4 122 123 96.1 124 95.9 121 95 .7 125 93.7 120 89.4 123 124 99.2 121 97.7 122 96 .1 125 93.4 120 89.2 124 123 99.2 121 97.9 122 95 .9 125 93.2 120 89.4 125 122 93.7 120 93.5 123 93 .4 124 93.2 121 92.3 126 123 85.1 120 85.1 118 83 .6 119 81.6 7 80.5 127 124 85.7 123 85.2 122 85 .0 121 84.3 120 84.2 128 42 87.7 29 84.1 24 84 .1 26 83.5 39 83.5 129 130 94.1 131 92.2 101 79 .6 13 79.4 12 79.3 130 131 97.7 129 94.1 24 79.9 13 79.1 40 79.1 131 130 97.7 129 92.2 13 81. .0 12 80.9 6 80.0 132 135 94.0 133 89.7 136 88 .7 134 83.0 25 80.1 133 136 96.0 135 93.7 134 93.3 132 89.7 7 83.0 134 133 93.3 136 89.5 135 87.7 5 86.4 7 85.7 135 132 94.0 133 93.7 136 92.7 134 87.7 8 82.1 136 133 96.0 135 92.7 134 89, .5 132 88.7 14 84.5 137 138 95.2 105 79.8 80 79.4 33 78.7 36 77.3 138 137 95.2 105 80.5 71 77.9 70 77.8 74 77.7 Appendix 4. Distribution of OTUs in the six and nine group schemes for clustering to maximize WG MS analysis Group 1. 66 : 70 : 71 : 86 : 87 : 91 : 103:105:109: 137:138. Group 2. 68: 72: 73: 77: 78: 79: 99:103:106: 113:114:115. Group 3. 1: 2: 7 13 19 26 32 38 3 9 15 21 28 34 40 14 20 27 33 39 128:129:130 134:135:136. Group 4. 118:119:120 124:125:126 Groups. 83 : 88 : 89 94 : 95 : 96 Six group scheme 82 : 84 : 85 : 98:101:102: 110:111:116: 74: 75: 76 80 : 81 : 97 107:108:112 4: 5: 6 10: 11: 12 16: 17: 18 23 : 24 : 25 29: 30: 31 35 : 36 : 37 41 : 42 : 65 131:132:133 121:122:123: 127. 90 : 92 : 93 : 100:117. Group 6. 22 : 43 : 44 : 45 : 46 : 47 : 48 : 49 : 50 : 51 : 52 : 53 : 54 : 55 : 56 : 57 : 58 : 59 : 60 : 61 : 62 : 63 : 64 : 67 : 69. Nine group scheme Group 1 . 118:119:120:121:122:123: 124:125:126:127. Group 2. 132:133:134:135:136. Group 3. 126:130:131. Group 4. 22: 43 44 45 46 47 48: 49 50 51 52 53 54: 55 56 57 58 59 60: 61 62 63 64 65 69. Group 5. 70: 71 82 83 84 85 86: 87 88 89 90 91 92: 93 94 : 95 96 98 100:101 102 :103 105 :109 110:111 116 :117. Group 6. 1: 2 8 19 23 24 25: 26 27 : 28 29 30 31: 32 33 : 34 35 36 37: 38 39 : 40 41 42 65:128. Group 7. 68: 72 73 : 74 75 76 77: 78 79 : 80 97 99 103 :106 :107 :108 :112 :113 114:115. Group 8. 67: 80 :137 :138. Group 9. 3: 4 : 5 : 6 7 9 10: 11 : 12 : 13 14 : 15 16: 17 : 18 : 20 21. Acknowledgements I would like to thank the directors and keepers of the following herbaria for providing facilities (asterisked) and for the loan of material: Edinburgh (E*), Geneve (G), Kew (K*), Kunming (KUN), Paris (P), Taipei (TAI), and Tokyo (TI). Mr A. Lauener came to my aid in locating Chinese material, especially from Leveille's herbarium. On the technical side I am grateful to Kay Shaw and Roger White for help with the computer programs. I would also like to thank Mr A. O. Chater and Mr I. Hedge for many useful discussions and comments. Miss Ann Lum kindly provided translations of Chinese literature, Dr N. K. B. Robson checked the latin diagnoses, and Miss Loveday Hosking typed the manuscript. My special thanks go to Dr C. J. Humphries for his help and encouragement throughout the compilation of this manuscript and for critically reading the result. References Bakhuizen van den Brink, R. C. & van Stennis, C. G. G. J. 1968. A note on Pogostemon Desf. and Dysophylla Blume. Taxon 17: 235-236. Bentham, G. 1829. Bot. Reg. 15: t. 1282 & t. 1300. - 1830. Synopsis of the genera and species of Indian Labiatae enumerated in the catalogue of the collections in Dr Wallich's charge. In N. Wallich, Plantae Asiaticae Rariores 1: 28-31. London. 1832-36. Labiatarum genera et species. London. - 1848. Labiatae. In A. de Candolle, Prodromus 12: 27-603. Paris. 1870. Flora Australiensis 5. London. 1876. Labiatae. In G. Bentham & J. D. Hooker, Genera Plantarum 2 (2): 1160-1223. London. Blume, C. L. 1826. 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Alabastra diversa. /. Bot., Lond. 15: 289-298. Mukerjee, S. K. 1940. A revision of the Labiatae of the Indian empire. Rec. hot. surv. Ind. 14 (1): 1-228. Ohwi, J. 1965. In F. G. Meyer & E. A. Walker (Eds), Flora of Japan. Washington, D.C. Panigrahi, G. 1976. Taxonomic notes on certain taxa of Asiatic angiosperms. Phytologia 32: 473-479. Persoon, C. H. 1806. Synopsis plantarum. 2. Paris. Poiret, J. L. M. 1817. Elsholtzia. In J. B. A. P. M. Lamarck, Encyclopedic methodique Botanique suppl. 5: 663. Paris. Prain, D. 1898. On three new genera of plants from the Kachin hills. Sclent. Mem. med. Offrs Army India H: 41^4. 1901. Eurysolen gracilis. In G. King, J. F. Duthie & D. Prain, A second century of new and rare Indian plants. Ann. R. hot. Gdn Calcutta 9 (1): 61-62. Rafinesque-Schmaltz, C. S. 1837. Flora Telluriana 2. Philadelphia. Rehder, A. 1917. Labiatae. In C. S. Sargent (Ed.) Plantae Wilsonianae 3: 380-384. Cambridge. Roxburgh, W. 1814. Hortus Bengalensis. Serampore. 1815. 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Actaphytotax. sin. 12: 337-346. 1977. Labiatae. In Flora Reipublicae Popularis Sinicae 66. Peking. & Li, H. W. 1975. Some changes of botanical name in Chinese Labiatae. Actaphytotax. sin. 13: 72-95. Wunderlich, R. 1963. The Pogostemoneae a debatable group of Labiatae (some remarks on their taxonomic position with regard to their pollen grains). /. Indian hot. Soc. 42A: 321-330. Yunnan Xian Weishenju (Ed.) 1973. Chinese herbal medicines of Yunnan: supplement. Kumming. British Museum (Natural History) Chance, change & challenge Two multi-author volumes from one of the foremost scientific institutions in the world . General Editor: P. H. Greenwood The Evolving Earth Editor: L. R. M. Cocks The Evolving Biosphere Editor: P. L. Forey In the first volume, The Evolving Earth, twenty scientists have been asked to review the present state of knowledge in their particular field, ranging from the origin of the Earth, through ocean sediments and soils to continental drift and palaeogeography. 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Press The typification of Hudson's algae: a taxonomic and nomenclatural reappraisal. By L. M. Irvine & P. S. Dixon Seaweeds of the Faroes (3 papers). ByD.E. G.Irvine; I. Tittley, W. F. Farnham & P. W. G. Gray; J. H. Price & W. F. Farnham The lichen genus Stelnera. By A. M. Henssen & P. W. James Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed by Henry Ling Ltd, Dorchester Bulletin of the British Museum (Natural History The typification of Hudson's algae: a taxonomic and nomenclatural reapprais Linda M. Irvine & Peter S. Dixon Botany series Vol 10 No 2 28 October 1982 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. 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Tustees of the British Museum (Natural History), 1982 The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Mr J. F. M. Cannon Editor of Bulletin: Mr P. W. James Assistant Editor: Mr J. R. Laundon ISSN 0068-2292 Botany series VollONo2pp91-105 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 28 October 1982 The typification of Hudson's algae: a taxonomic and nomenclatural reappraisal Linda M. Irvine Department of Botany, British Museum (Natural History), Cromwell Road, London SW7 5BD Peter S.Dixon Department of Ecology and Evolutionary Biology, University of California, Irvine, California 92717, U.S.A. Contents Synopsis 91 1. Introduction 92 2. Hudson's species concept 92 The algae of Flora anglica: first edition 92 The algae of Flora anglica: second edition 93 3. The typification process 94 Historical comments 94 Procedure adopted 95 4. Examples of the typification of Hudson's algal names .... 96 I. Holotype: a pre-Linnaean illustration 96 a. Fucus scorpioides 96 II. Lectotype: a pre-Linnaean illustration 96 a. Conferva imbricata 96 b. Fucus pinnatifidus 97 c. Fucus plumosus 98 III. Lectotype (provisional): a Hudson specimen 99 a. Conferva elongata .......... 99 b. Fucus plicatus 100 IV. Lectotype: a Hudson description 102 a. Conferva fucoides 102 b. Conferva nigra 102 V. Species inquirenda 103 a. Conferva fulva 103 5. Acknowledgements 104 6. References .... 104 Synopsis William Hudson (1734-1793) was the earliest English botanist to adopt the Linnaean systems of classification and nomenclature. The two editions of Flora anglica are of special significance to phycologists because both contain numerous descriptions of new species of algae; in other groups of plants (except lichens) these are very few. The algal descriptions given by Hudson are imprecise by modern standards, but his taxonomic views can often be interpreted by critical typification of the names of his new taxa. This requires an understanding both of the materials Hudson used and of his philosophy and method of work. Although these were not stated explicitly by Hudson, they can to some extent be deduced even 200 years after publication, as indicated here. A selection of nine species from amongst those described by Hudson provides a series of examples illustrating the problems to be dealt with in the typification of Hudson's algal names. Bull. Br. Mus. not. Hist. (Bot.) 10 (2): 91-105 Issued 28 October 1982 92 LINDA M. IRVINE & PETER S. DIXON 1. Introduction William Hudson (1734-1793) was the earliest English botanist to adopt the Linnaean systems of classification and nomenclature. The two editions of Flora anglica (Hudson, 1762; 1778) are of special significance to phycologists because both contain numerous new species of algae, whereas for other groups of plants (except lichens) these are relatively few. The posthumous, so-called third, edition (Hudson, 1798) is identical to the second edition, except for the correction of some printer's errors. In the course of preparing the various checklists of British marine algae (see Parke & Dixon, 1976) and Seaweeds of the British Isles (Dixon & Irvine, 19770), the significance of Hudson's works showed repeatedly. Thus, in the current Check-list of British marine algae third revision (Parke & Dixon, 1976), 37 species of Rhodophyta, Phaeophyta and Chlorophyta have Hudsonian basionyms. Some of them, and other names now reduced to synonymy, have already been considered (Dixon, 19596, 1960, 1962, 1963, 1967; Dixon & Irvine, 19770, b), and Laundon (1963, 1966, 1976) has typified three Hudson lichen names. We eventually realized, however, that this piecemeal approach was unsatisfactory, and that a more comprehensive study was desirable. The algal descriptions given by early authors such as Hudson are imprecise by modern standards, so that critical typification of the names of new taxa is imperative. This requires an understanding of the materials used and also of the author's philosophy and method of work which were not stated explicitly, but can to some extent be deduced even 200 years after publication. In the case of Hudson, a resume of his life and of the history of his personal herbarium has been previously presented (Dixon, 19590). Herbaria are cited by their official abbreviations: BAS: Botanisches Institut der Universitat Basel, Switzerland. BM: British Museum (Natural History), London. BM-K: Specimens formerly in the Herbarium of the Royal Botanic Gardens, Kew, and now at the British Museum (Natural History), London. BM-SL: The Sloane Herbarium at the British Museum (Natural History), London. K: The Herbarium, Royal Botanic Gardens, Kew. LINN: Linnean Society of London. NMW: National Museum of Wales, Cardiff. OXF: Fielding-Druce Herbaria, The University, Oxford. For details of the Morison, Ray and Dillenius herbaria, see Clokie (1964). 2. Hudson's species concept The algae of Flora anglica: first edition Hudson's species concept was obviously derived directly from that of Linnaeus , for which Stearn (1957) has given a detailed exposition. This close relationship means that the mechanics involved in the typification of Hudson's species are similar to the Linnaean examples discussed by Stearn. The similarity is particularly noteworthy in the first edition of Flora anglica where 49% of the algal binomials were taken from the first edition of Species plantarum (Linnaeus, 1753), with the remainder newly described. The sources which Hudson used in the preparation of his algal descriptions for the first edition of Flora anglica fall into three categories, used singly or in combination, as follows: 1. Herbarium specimens which were originally in his possession From the comments made and additional information provided by Hudson in his species' treatments, it is clear that he must have had specimens of his own, although it may be difficult to find these at the present time. It was common for botanists of that period to replace a specimen (even one which had been used for a description) with another which was considered to be more appropriate, but not necessarily conspecific by modern standards. Secondly, Hudson's posses- sions were largely destroyed by fire in 1783 and it is not known with any certainty what parts (if any) of his early herbarium survived. The dispersal of herbarium material which once belonged THE TYPIFICATION OF HUDSON'S AL'GAEI A TAXONOMIC AND NOMENCLATURAL REAPPRAISAL 93 to Hudson was chronicled by Dixon (1959a), who showed that about 200 specimens were then located at BM and K. The algal collections at the latter have now been transferred to the former (see Ross & Brenan, 1970). The algal herbarium of Hudson's contemporary, Lightfoot, which contains many specimens received from Sir Thomas Frankland said to have been identified by Hudson, was eventually found in an attic at Saffron Walden Museum (Dixon, 1959a). It is now on permanent loan to K and has been transferred with other algal material to BM. The folder, once the property of Frankland, containing algal specimens, mentioned by Dixon (19590) as being at LINN, has also been transferred to BM. Thus, at present, virtually all known Hudson material is preserved at BM but it is always possible that some authentic Hudson specimens, even types, remain undetected and may come to light in obscure places. 2. Other herbarium specimens quoted by Hudson Three collections of particular relevance to Hudson algal binomials are those of Buddie, Petiver and Plukenet. These are all now in the Sloane Herbarium (BM-SL), of which Dandy (1958) has given a detailed account, and were probably consulted by Hudson when he was employed as a resident sublibrarian at the British Museum between 1757 and 1758. Buddie and Petiver were two of the most famous plant collectors of the early 18th century who corresponded and exchanged specimens with many other botanists. Their herbaria were acquired by Sir Hans Sloane in 1715 and 1718, respectively, and became part of the national collections on the death of the latter in 1753. Although Hudson made direct reference to material from these collections only infrequently, the herbaria are of great importance because they had previously been used extensively by Ray (1724) and Dillenius (1742). Knowledge of these collections enabled Hudson to interpret the two pre-Linnaean authors better than any other botanist. 3. Previously-published descriptions and illustrations Unlike many of his contemporaries, Hudson obviously considered illustrations of particular importance when he was interpreting a previous description, so that these were quoted whenever possible. The illustrations cited most frequently in the first edition of Flora anglica (Hudson, 1762) are those by Morison (1680-99), Ray (1724), and Dillenius (1742), although there is no evidence that Hudson ever consulted their herbaria at Oxford (OXF). The third edition of Ray's Synopsis was prepared anonymously by Dillenius and published after Ray's death. Hudson's citation R. Syn. refers to this edition. The three genera Fucus, Viva and Conferva contain 91 algal species in Hudson's first edition. Of these, 45 are species described previously by Linnaeus, 13 are new species based entirely on his own herbarium material while 33 are new species based on specimens, illustrations or descriptions by pre-Linnaean authors, sometimes in combination with Hudson's own herbarium material and sometimes not. The algae of Flora anglica: second edition In the second edition of Flora anglica (Hudson, 1778), Linnaeus was still by far the most frequently-quoted author. Here the second edition of Species plantarum (Linnaeus, 1763) was a major source, although other Linnaean publications were also used (e.g. Linnaeus, 1753, 1767, 1771, 1774) and, in addition, Hudson occasionally quoted other phycological works (Ellis, 1767; Gmelin, 1768; Oeder, 1762-1883). Analysis of the second edition is more difficult as the species treated do not fit into categories as neatly as for the first edition, reflecting the increasing complexity of phycology between 1762 and 1778. In the second edition 141 algal species are treated in the same three genera mentioned previously, 72 species being carried over from the first edition. Of the 62 species which occur for the first time, eight are species described previously by Linnaeus (in 1753 or later), four are based on previous treatments by Gmelin (1768), and two are based on treatments by Ellis (1767). New species based entirely on Hudson's own material total 29, while 16 new species are based on entities described by pre-Linnean authors, sometimes in combination with his own 94 LINDA M. IRVINE & PETER S. DIXON material and sometimes not. A few species from the second edition which cannot be charac- terized are omitted from this analysis. The two editions differ, therefore, in the number of algal species treated, with a more than 50% increase in the second edition over the first. The actual treatment of a particular species in the two editions of Flora anglica may differ in various ways, often subtle. A point frequently overlooked by subsequent workers is that, for both Hudson and Linnaeus, the nomenclatural system which they used differed from current practice in two principal ways. The first difference is in relation to the use of binomials. These are usually regarded as a revolutionary advance made by Linnaeus in 1753, whereas, as Steam (1957) has indicated, they were introduced primarily as 'an indexer's paper-saving device'. Although the polynomials of previous authors were cumbersome, investigators still thought in such terms even after Lin- naeus's introduction of binomials. As a consequence, Hudson cites synonyms exclusively in polynomial form in the first edition. Ellis (1767) and Gmelin (1768) were the first phycologists to accentuate binomials, the latter probably having the greater influence through the typographic device of heading each species treatment with a binomial in bold type and relegating polynomial synonyms to a minor position. This is in marked contrast to the procedures of Linnaeus and Hudson, where the specific epithet is merely a marginal annotation to a polynomial beginning with the generic name. Thinking in terms of binomials probably had a great effect on stability; polynomials could be constantly modified, an open invitation to transfer of application. Binomial synonyms began to appear in Hudson's second edition, with references to Ellis (1767) and Gmelin (1768). Secondly, the basic principle two centuries ago was that the more recent version of a text was the more definitive. Because the later publication was considered to be superior to the earlier work ('altera, emendata et aucta'), the second edition rarely refers back to the first. In a very few cases species were placed in synonymy by citation of an additional marginal epithet, although a few more were given as page references to polynomials. It was much later that de Candolle (1867) first stressed the principle of priority, which led to the formulation of a nomenclatural code. Thus, changes introduced by Hudson in an effort to improve had a significance to him different from that of current nomenclatural practice. The changes between the two editions are of several kinds: 1 . modifications in the polynomial description of a species, including substitution of words or phrases; 2. deletion, addition or transfer of synonyms (particularly pre-Linnaean polynomials), always without explanation; 3. changes of epithet, usually without explanation, but apparently done in most cases because one epithet was considered more appropriate than another; 4. changes in the anglicized 'common name'. Casual inspection of the two editions did not reveal some of these changes; they became apparent only after making detailed comparisons of all the species treated. It is interesting to consider them in the light of the comments made above, for they were obviously regarded by Hudson as improvements. 3. The typification process Historical comments By definition (Stafleu etal., 1978), a nomenclatural type is that element to which the name of a taxon is permanently attached. A holotype is the original single specimen or other element used by the author, whereas the lectotype is a specimen or other element selected later from a group of original materials. For recently-described taxa, the type must be indicated for the name to be validly published. Prior to 1958 this requirement did not apply, and it is necessary to establish the types and their status. For earlier authors this is not always easy, and there are several serious problems involved with Hudson's algal names because of the changes in treatment between the editions. These may reflect improved knowledge or they may represent a real change in THE TYPIFICATION OF HUDSON'S ALGAE: A TAXONOMIC AND NOMENCLATURAL REAPPRAISAL 95 Hudson's taxonomic opinion and therefore in the application of an epithet. Of the materials which he used, specimens in the Sloane Herbarium (BM-SL) are still largely as they were when they were examined by Hudson. The situation with his own herbarium materials is much less satisfactory, however, and almost without exception specimens have been remounted and/or relabelled, probably eliminating much information which would be fundamental to the present study. Because of this, it is not clear whether any extant specimen was used as the basis for a diagnosis in either edition or represents a superior specimen, substituted later. Procedure adopted The basic procedure adopted in the typifications of the Hudson binomials presented here and elsewhere (e.g. Dixon, 1967) has been to analyse the descriptions and treatments of the two editions in their entirety and to identify the source material(s) used by Hudson for each entity. At the same time, complete searches of herbaria (BM, BM-K, BM-SL, etc.) were made to provide catalogues of all material now existing which was alleged to have been in Hudson's possession and to locate all materials to which he made reference, or which Frankland said Hudson had identified. Integrating these two aspects indicated whether the elements upon which a species had been based were homogeneous or heterogeneous. In addition, further analyses were conducted of other herbaria and publications (Dillwyn, 1802-9; Turner, 1802; Turner, 1808-19; Smith & Sowerby, 1790-1814) to obtain information on contemporary or near-contemporary opinion. Where it is clear that Hudson provided no information which might suggest possession of his own herbarium material and that his treatment was based entirely on a single pre-Linnaean illustration, this is the holotype, as in the case of Fucus scorpioides (Example la). In other cases one element was a pre-Linnaean specimen or illustration and the other personal herbarium material. Sometimes all pieces of evidence proved to be taxonomically homogeneous both by present-day and near-contemporary standards, so that there were no doubts as to the application of a name. In such cases the pre-Linnaean specimen or illustration was selected as lectotype because of the uncertainties surrounding the surviving Hudson herbarium specimens. Examples of this are Conferva imbricata (Example Ha), Fucus pinnatifidus (Example lib), and F. plumosus (Example He). The situation became complicated where the data proved to be heterogeneous or where the identity of the pre-Linnaean element was in doubt. The uncertainty over the dates of collection of the surviving Hudson specimens is such that to accept a Hudson specimen as unequivocal lectotype is unwarranted. For cases such as Conferva elongata (Example Ilia) and Fucus plicatus (Example Illb), an extant Hudson specimen was therefore assigned the status of provisional lectotype. Finally there remain those cases where Hudson described a species exclusively from his own material and where none of this can be located at the present time. Providing that there had been no disagreement about the application of a name by Hudson's contemporaries and no subse- quent deviation in usage, it seemed sensible to retain the epithet and select the original diagnosis as type in the absence of any material. Examples of this procedure are Conferva fucoides (Example IVa) and C. nigra (Example IVb). In cases where both contemporary and present-day opinion are uncertain about the attribution of a name, as in C. fulva (Example Va), typification is clearly not possible. The Code (Stafleu et al. , 1978) makes provision for the selection of neotypes in cases where the original specimens are missing. This procedure is frequently used by lichenologists, and Laundon (1976) adopted it for Caloplacaflavorubescens (Huds.) Laundon. Phycologists, on the other hand, seem less prepared to accept neotypes and we have not selected any here. Nevertheless, we would like to suggest that in future neotypes could be used in cases where this would expedite the clarification of the nomenclatural tangles which so often hinder the pursuit of taxonomy. 96 LINDA M. IRVINE & PETER S. DIXON 4. Examples of the typification of Hudson's algal names These are as follows: I. Holotype: a pre-Linnaean illustration a. Fucus scorpioides. II. Lectotype: a pre-Linnaean illustration a. Conferva imbricata, b. Fucus pinnatifidus, c. F. plumosus. III. Lectotype (provisional): a Hudson specimen a. Conferva elongata, b. Fucus plicatus. IV. Lectotype: a Hudson description a. Conferva fucoides, b. Conferva nigra. V. Species inquirenda a. Conferva fulva. I. Holotype: a pre-Linnaean illustration EXAMPLE a. Fucus scorpioides Huds. [Bostrychia scorpioides (Huds.) Mont.] The original description of Fucus scorpioides by Hudson (1762:471) is printed as follows: 23 . FUCUS caule tereti ramoso , ramis alternis ramo- scorpioides sissimis apice inflexis. Fucoides erectum fruticuli specie, summitatibus in- flexis. R. Syn. 38. t. 2. f. 6. Anglis, upright Fucus. Habitat in littore Sussexiano; at Selsey-Island plenti- fully. Dr. Dill. R. Syn. Hudson's treatment was evidently based entirely on the previous treatment and illustration by Dillenius in Ray (1724:38) because the only locality cited by Hudson was taken directly from the latter: 'In the Marshes at Selsey Island, Sussex, plentifully'. There is no evidence that Hudson ever examined the Oxford herbaria and he appears to have based his treatment on the Dillenius illustration (Ray, 1724: pi. 2, fig. 6). This illustration is somewhat stylized although there is little doubt that it refers to the alga known today as Bostrychia scorpioides (Huds.) Mont. The material on which the illustration was based is in the Synopsis herbarium (OXF), and is of that alga (see also Batters in Druce & Vines, 1907:21). The illustration is the holotype of Fucus scorpioides Huds. No material referred by Hudson to his Fucus scorpioides has been located in any herbaria. This may well be due to the changes which occurred between the two editions of Flora anglica. Gmelin (1768) described Fucus amphibius quoting as a synonym the Fucoides erectum fruticuli ... of Ray which Hudson had used as the basis for his Fucus scorpioides in the first edition. In the second edition Hudson (1778) referred to Gmelin, accepted Gmelin's binomial and quoted also the Fucoides erectum fruticuli ... of Ray, but made no reference to his own Fucus scorpioides. A 'Hudson Sale' specimen of Bostrychia scorpioides, which was acquired by BM through the Forster herbarium, is annotated Fucus amphibius. II. Lectotype: a pre-Linnaean illustration EXAMPLE a. Conferva imbricata Huds. [Halurus equisetifolius (Lightf.) Kiitz.j The original description of Conferva imbricata first appears in the second edition of Flora anglica (Hudson, 1778:603), printed as followed: 44. CONFERVAfilamentisgeniculatisramosissimis, imbricata. ramis acutis, ramulis verticillatis imbricatis dicho- tomis. Muscus marinus hirsutus, flagellis longioribus, rarius divisis, ruber. Hist. Ox. III. 650. s. 15. t. 9.f. 7. Anglis, imbricated Conferva. Habitat in rupibus saxis etfucis marinis. [perennial]. I-XII. THE TYPIFICATION OF HUDSON'S ALGAE! A TAXONOMIC AND NOMENCLATURAL REAPPRAISAL 97 The original description of C. imbricata was obviously based on the previously-published illustration by Morison (1699: pi. 9, fig. 7), for which Hudson gave a partially erroneous citation, and on any material which Hudson might have had in his possession at the time of publication. Whether Hudson had material of his own is not obvious, the habitat notes being too vague and imprecise for any definite conclusion on this point. The Morison illustration is somewhat stylized, but it would appear to be of the alga known currently as Halurus equisetifolius (Lightf.) Kiitz. , although the relevant material in the Morison Herbarium (OXF) consists of a herbarium sheet bearing two specimens, the larger of which is of that species, while the smaller is of the brown alga Cladostephus spongiosus (Huds.) C. Agardh. The discordant nature of the Morison material was first noted by Holmes (in Vines & Druce, 1914) and has been personally confirmed. Conferva imbricata Huds. (1778) is antedated by C. equisetifolia Lightfoot (1777) , the basionym of Halurus equisetifolius. In the appendix to the second edition of Flora anglica, Hudson (1778:663) cited the former as a synonym of C. imbricata, however. Four specimens of the taxon under discussion have been located at BM and all are specimens from the 'Hudson Sale', received through the Forster collection. The four specimens are named as C. equisetifolia, not C. imbricata, although the date of collection or acquisition by Hudson is not known. The Morison illustration is selected here as the lectotype of Conferva imbricata Huds. EXAMPLE b. Fucus pinnatifidus Huds. [Laurencia pinnatifida (Huds.) Lamouroux] The original description of Fucus pinnatifidus by Hudson (1762:473) is printed as follows: 32. FUCUS frondibusplanisramosis,ramisdentato- pinnatifi- pinnatifidis marginibus callosis. dus. Fucus dealensis pedicularis rubras folio. Mus. pet. 405. R. Syn. 48. Alga cervi cornu divisura. B. hist. III. 797. Anglis, jagged Fucus. Habitat in saxis et rupibus marinis. Found about Deal by Mr. Dandridge. R. Syn. in littore Devoniae/re- quens. This description was clearly based upon four elements: 1. the previous treatment by Petiver (1695-1703:39); 2. the previous treatment by Dillenius in Ray (1724:48); 3. the previous treatment by Bauhin (1651:797); 4. material collected in Devon as that county is not cited specifically in any of the preceding three elements. The synonyms cited by Hudson are taken almost directly from those quoted previously by Dillenius in Ray (1724:48), whose treatment is printed as follows: * 37. Fucus Dealensis Pedicularis rubrifolio Mus. Pet. 405. Alga Cervi cornu divisura /. B. III. 797. Found about Deal by Mr. Dandridge, Mr. Bonavert, and Mr. John Lufkin. This treatment by Ray was obviously based predominantly on the Petiver reference but with an additional citation of Bauhin (1651). The Petiver specimen is in the Sloane Herbarium (vol. 150 folio 25) (BM-SL) and consists of two fragments with a printed extract '405. Fucus Dealensis Pedicularis rubrae folio. My ingenious Friends Mr. Dandride, Mr. Bonavert, and Mr. John Lufkin, Apothecary at Colchester, have all observed this elegant Fucus about Deal'. Associated with it are two hand-written labels, in Petiver's hand and Buddie's hand respectively. The specimen is of the alga known today as Laurencia pinnatifida (Huds.) Lamouroux. The Alga cervi cornu divisura of Bauhin (1651) was not based upon an alga. Juel (1936: 136) has shown that the specimen in the Bauhin Herbarium at Basel (BAS) labelled 'Alga cornu cervi divisura Bauh. Dedit Bauhinus ab Imperato acceptem' is a lichen of the genus Ramalina. 98 LINDA M. IRVINE & PETER S. DIXON Two Hudson specimens are known to exist; one is a 'Hudson Sale' specimen received at BM through Forster and the other (in BM-K) is a specimen collected by Frankland at Scarborough. The former specimen bears no annotation indicating that it was collected in Devon, or that it was in Hudson's possession prior to the publication of the first edition of Flora anglica, so that neither appears to be directly relevant to the typification of Fucus pinnatifidus Huds. The Petiver specimen is the most appropriate selection as the lectotype of Fucus pinnatifidus Huds., supporting both old (i.e. near-contemporary) and modern opinion as to the application of the epithet. Bauhin's polynomial, the only discordant element, can be discounted as Hudson did not see the specimen on which it was based. EXAMPLE c. Fucus plumosus Huds. [Ptilota plumosa (Huds.) C. Agardh] The original description of Fucus plumosus given by Hudson (1762:473) is printed as follows: 35 . FUCUS caule teretiusculo compresso ramoso , ramis plumosus. duplicate pinnatis coloratis. Fucoides purpureum eleganter plumosum. R. Syn. 38. t. 2.f. 5. Anglis, Feathered Fucus. Habitat in littoribus marinis passim. This description was based on two elements: 1. an illustration given by Dillenius in Ray (1724: pi. 2, fig. 5); 2. material in Hudson's possession, suggested by the general statement of occurrence rather than the single specific locality (Dover, Kent), cited by Ray. Ray's illustration is somewhat stylized but is clearly of the alga known currently as Plumaria elegans (Bonnem.) Schmitz, not that known as Ptilota plumosa (Huds.) C. Agardh. The Dillenius in Ray (1724:38) treatment is as follows: * 2. Fucoides purpureum eleganter plumosum. Mus- cus marinus eleganter plumosus, obscure purpurascens Buddl. H. S. Vol. I. f. 29. A D. Rand prope Dover col- lectus. Figuram vid. Tab. 2. fig. 5. This shows that Fucoides purpureum eleganter plumosum was based on a specimen in the Buddie herbarium (BM-SL, vol. 114 folio 29) which is also of Plumaria elegans. Although not relevant to the typification of the Dillenius in Ray entity, it is interesting to note that the material of Fucoides purpureum ... in the Synopsis herbarium (OXF), collected at Llanfaethly (north Wales), is also of P. elegans (cf. Druce & Vines, 1907: 21). Three specimens referred to Fucus plumosus Huds. which are said to have been in Hudson's possession are known to exist. These are as follows: (a) a specimen from the Lambert herbarium, now in BM-K; (b) a specimen from the Forster herbarium, at BM; (c) a specimen from the Pulteney herbarium, said to be annotated in Hudson's hand, at BM. All are of the alga known currently as Ptilota plumosa, although it is not possible to ascertain the date at which Hudson referred these specimens to his Fucus plumosus. Thus, the Dillenius in Ray illustration and the Buddie specimen on which that was based are of one entity, whereas such Hudson material as has been located is of another. This discrepancy is curious as it is known that Hudson had access to the Sloane Herbarium, into which the Buddie herbarium has been incorporated in 1715 '. There appears to be no alternative to accepting the Ray illustration as lectotype of Fucus plumosus Huds. since the specimens said to have been identified by Hudson may not have been available to him in 1762. All specimens subsequently attributed to Fucus plumosus Huds., are referable to a species different from the lectotype selected above. The reason appears to lie in the change between the two editions of Flora anglica. The first edition treatment has been quoted above; that of the second edition is printed as follows (p. 587): THE TYPIFICATION OF HUDSON'S ALGAE: A TAXONOMIC AND NOMENCLATURAL REAPPRAISAL 99 47. FUCUSfrondecartilagineacompressaramosa,ra- plumosus. mis duplicato-pinnatis, fructificationibus peduncu- latis globosis radiatis. Fl. angl. 473. Gmel. fuc. 152. Fucus frondibus cartilagineis lanceolatis bipinnatis plumosis, caule filiform! compresso ramoso. Mant. 134. Syst. not. 718. FL dan. 350. Fucoides purpureum eleganter plumosum. R. syn. 38. r. 2./. 5. Anglis, plumous Fucus. Habitat in rupibus etsaxis submarinis. [perennial] VII-X. Although this is one of the few instances where Hudson's treatment of a species in the second edition refers back to the first edition (by his citation of 'Fl. Angl. 473.'), the descriptions in the two editions differ markedly. Hudson added in the second edition references to treatments of Fucus plumosus by Linnaeus (1767, 1774), Gmelin (1768) and Oeder (1767), the two latter being illustrated. Both illustrations (Gmelin, 1768: pi. 152 and Oeder, 1767: pi. 350) are of the alga to which the binomial Ptilota plumosa is currently applied rather than to Plumaria elegans, the alga of the lectotype. It would appear therefore that Hudson was influenced by the references and illustrations which he quoted in the treatment of Fucus plumosus in the second edition and his concept of the species changed completely. The specimens of F. plumosus which are said to have been in his possession represent material identified at a later date, after the change. It is surprising that the identity of the alga described and illustrated by Dillenius in Ray and quoted by Hudson has not been questioned previously, particularly as the epithet plumosus is apparently derived from the Ray polynomial. The alga known currently as Plumaria elegans, to which the lectotype is referable, is plumose and usually purple-coloured, whereas the alga known today as Ptilota plumosa is not obviously plumose; it is bipinnate and carmine-red. Secondly, the alga known currently as Ptilota plumosa is of northern distribution in the British Isles, not occurring in an attached state south of Yorkshire on the east coast and Caernarvon on the west. Dover, the specific locality cited by Ray, is some 250 miles south of the southern limit of Ptilota plumosa. The nomenclature of the genera Plumaria and Ptilota is already confused (cf . Silva, 1952) and the present discovery makes that situation worse. Full discussion and resolution of these problems are beyond the scope of the present paper, but a full morphological and nomenclatural study of the North Atlantic Ptiloteae is now approaching completion, as part of the preparatory work for Seaweeds of the British Isles, Volume 1, Part 3. III. Lectotype (provisional): a Hudson specimen EXAMPLE a. Conferva elongata Huds. [Polysiphonia elongata (Huds.) Sprengel] The original description of Conferva elongata Hudson (1762:484) is printed as follows: elongata. 25. CONFERVA filamentis geniculatis ramosissimis, ramulis longissimis distantibus acutis. Conferva marina geniculata ramosissima lubrica, lon- gis sparsisve ramulis. R. Syn. 61. Dill. muse. 35. t. 6./. 38. Anglis, Pointed Conferva. Habitat in littoribus marinisfrequens. The description was based on three elements: 1. the previous treatment by Dillenius in Ray (1724:61); 2. the previous treatment and illustration by Dillenius (1742); 3. possibly material in his own possession, although the description and distribution are too imprecise for definite proof. The treatment by Dillenius (1742:34) was clearly based directly on the earlier Ray (1724) account by Dillenius. The material now in the Historia Muscorum herbarium (OXF) preserved under the Dillenian name consists, as was shown by Batters (in Druce & Vines, 1907: 190), of 100 LINDA M. IRVINE & PETER S. DIXON four specimens. Three of these are referable to the genus Ceramium, while the fourth is of the alga known today as Polysiphonia nigrescens. The illustration given by Dillenius (1742: pi. 6, fig. 38) is too imprecise to allow specific identification, although it most likely refers to a species of Ceramium. The previous treatment by Dillenius in Ray (1724:61) is printed as follows: * 23. Conferva marina geniculata ramosissima lubri- ca, longis sparsisve ramulis. Muscus marinus capillaris rubens geniculatus ramosissimus Buddie Hort. Sice. Sent to him by Mr. Stevens from Cornwal. It is else com- mon enough at Cockbush, Sussex, and about Sheerness. The Ray description was based principally on material in the Buddie herbarium, with additional material from Kent and Sussex. There are now in BM-SL vol. 114 folio 30, two specimens labelled 'Muscus marinus capillaris rubens geniculatis ramosissimis Buddie N.D. sent by Mr Stevens from Cornwall'. These are of species of Ceramium', one specimen is dark red while the other is bleached. Dillenius (1742) comments on this alga 'color rubicundiis, interdum arenaceiis'. The specimens currently in the Synopsis herbarium (OXF) consist of a mass of material largely collected by Brewer in north Wales, with no trace of any collections from Cornwall, Kent or Sussex. The indications are therefore that the original treatment of Conferva elongata Huds. referred to material of species of both Ceramium and Polysiphonia. The treatment in the second edition of Hudson (1778:599) is very different, however: 27. CONFERVA filamentis geniculatis ramosis,ramis elongata. dichotomis longis setaceis, articulis brevissimis. Anglis, pointed Conferva. Habitat in saxis et rupibus submarinis, in Devonia, Cornubia, Sussexia, et in insula Mona, passim, [annual]. IV-X. Desc. Fila dodrantalia etpedalia, crassitie fili emporetici tenuioris, articulata, Icevia, fusco-purpurea, basi ra- mosa; Rami dichotomi, longissimi, setacei; articulis brevissimis. The two diagnoses differ, two localities were added (Devon, Anglesey), one of the original localities (Kent) was deleted, and a considerable amount of descriptive material was included in the second treatment. Most importantly, the Ray and Dillenian synonyms of the first treatment are transferred to the synonymy of Conferva rubra, the basionym of Ceramium rubrum. Furthermore, contemporary views on Conferva elongata Huds., such as Dillwyn (1803) and Smith & Sowerby (1790-1814) accord with current application of the epithet to a species of Polysiphonia. There is a specimen in BM, received through the Forster herbarium from the 'Hudson Sale' and labelled 'Conferva elongata', which is correctly identified as Polysiphonia elongata (Huds.) Sprengel (as currently understood) and filed in a type folder. The sheet also bears a determinavit label 'Polysiphonia elongata (Huds.) Grev. ex Harv. in Hook = Conferva elongata Hudson probably in sense of Ed. II 1778 p. 599 ... A. R. A. Taylor 1.IX.64', in A. R. A. Taylor's hand. Although the initial treatment of Conferva elongata Huds. was based on material which was heterogeneous, the treatment in the second edition was not; near-contemporary and current usage is in complete agreement as to the application of the epithet, supported by a specimen of the entity originating from Hudson. It can be postulated that Hudson eventually appreciated that his original description was based on several discordant elements and that the 'improved' text of the later treatment in the second edition represents a careful selection from that original miscellany, a procedure which is common nomenclatural practice nowadays. The specimen in BM has therefore been selected as provisional lectotype of Conferva elongata Huds. EXAMPLE b. Fucus plicatus Huds. [Ahnfeltia plicata (Huds.) Fr.] The original description of Fucus plicatus by Hudson (1762:470) is printed as follows: THE TYPIFICATION OF HUDSON'S ALGAE! A TAXONOMIC AND NOMENCLATURAL REAPPRAISAL 101 plicatus. 19. FUCUS capillaris uniformis ramosissimus implicatus subdiaphanus. Fucus trichoides nostras aurei coloris, ramulorum a- picibus furcatis. Pluk. ph. t. 184. /. 2. R. Syn. 45. Fucus coralloides erectus. R. Syn. 51. Anglis, Matted Fucus. Habitat in littoribus marinis. This was based on: 1. an illustration by Plukenet (1696: pi. 184, fig. 2); 2. the previous treatment by Ray (1724:45). The general statement of localities provided by Hudson is too imprecise to be certain that he had his own specimens at the time of publication of the original description, but his extended description suggests that he did. The reference to the Plukenet illustration was taken from the previous treatment by Ray (1724:45), which is as follows: 26. Fucus trichoides nostras aurei coloris, ramulorum apicibus furcatis Pluk. Aim. 160. T. 184. f.2. Algaexi- gua dichotomos arenacei coloris Syn. II. 4. 10. Fucus ceranoides ramosus tenuissime divisus Dood. Syn. II. App. 329. Palmaris est, corneus tenax, albus, per siccitatem rigidus, ubique ejusdem fere crassitudinis, qua? filum parvum superat. In littore Essexiano, Sussexiano & alibi. Both Dillenius in Ray (1724) and Hudson (1762) refer to an illustration by Plukenet (1696: pi. 184, fig. 2). This illustration is poor and it is not possible to state categorically that it represents the alga known currently as Ahnfeltia plicata. Plukenet's herbarium has been incorporated into BM-SL and the reverse of Folio 80 of Volume 84 bears a specimen labelled 'Alga marina trichodes lutea. S. Trichoman. ceranoides marin. [deleted] aurea & ramosa, ramulos apicis furcatis' in Plukenet's hand. This specimen could possibly have formed the basis for the Plukenet illustration which it resembles in shape, proportions and branching pattern, and it is also of the alga known currently as Ahnfeltia plicata. A second specimen of A. plicata bearing a different annotation is mounted on folio 81. The specimens in this volume of the Sloane Herbarium are arranged in alphabetical order, and the two specimens in question occur among species of Fucus, the genus to which Plukenet referred his entity in his publication (Plukenet, 1696). Although it is known that Hudson had access to the Sloane Herbarium, there is no annotation or other indication that he actually examined these specimens. In other cases where Hudson had examined material in the Sloane Herbarium, as with Fucus plumosus (Example He) and Fucus pinnatifidus (Example lib), he made reference to the actual specimen, but he did not do so in this case. As noted above, Hudson may have had material of his own to hand when he drew up the original description of Fucus plicatus . A specimen labelled Fucus plicatus at BM, received from the Forster herbarium and annotated as having been obtained from the 'Hudson Sale', is of the alga known today as Ahnfeltia plicata. In view of the uncertainties surrounding the Ray and Plukenet synonyms, this specimen has been designated as the provisional lectotype of Fucus plicatus (see Dixon & Irvine, 1977). The infraspectific entity in the original treatment of Fucus plicatus is based directly on the Fucus coralloides erectus of Dillenius in Ray (1724) which is a repetition of an entry in an earlier Ray (1704) publication. As Hudson gave no locality presumably he had no material of his own. The material in the Synopsis herbarium (OXF) referred to this entity was tentatively identified by Holmes (in Druce & Vines, 1907:26) as the freshwater red alga Lemanea fluviatilis (L.) C. Agardh. It consists only of a few scraps, and is difficult to confirm from personal examination, but Holmes's identification was probably correct. Thus, the infraspecific entity described by Hudson has no connection with Fucus plicatus Huds. 102 LINDA M. IRVINE & PETER S. DIXON IV. Lectotype: a Hudson description EXAMPLE a. Conferva fucoides Huds. [Polysiphonia nigrescens (Huds.) Grev.] The original description of C. fucoides (Hudson, 1762:485) is printed as follows: 31. CONFER VAfilamentisgeniculatisramosissimisra- fucoides. mulis multifidis fasciculatisque. Anglis, Branched Conferva. Habitat in littore Eboracensi. This description was obviously based on Hudson's own material. The treatment in the second edition (Hudson, 1778:603) differs in various ways, and is printed so: 42. CONFERVA filamentisgeniculatisramosissimis fucoides. ramulis multifidis, inferioribus fasciculatis fructi- feris. Anglis, fucus Conferva. Habitat in rupibus, saxis et fuels marinis passim, [perennial]. I-XII. Desc. Filapedalia, geniculata, Icevia atro-rubescentia, ramosissima, Rami alterni; ramuli multifidi, subdicho- tomi, inferior es fasciculati, fructiferi. Fructifica- tiones terminales, radiates, parvce. There are no extant specimens of Hudson's referred to this entity although several contempor- ary specimens have been located which are relevant, the most important of these being specimens originating from Frankland. One of these is in the collection which was at LINN (Dixon, 1959a) and is now at BM; this is said to be illustrative of Hudson's application of the name. The other Frankland specimen occurs in the Lightfoot herbarium (now at BM) and is annotated 'named repeatedly by Hudson C. fucoides.' Both Frankland specimens are of the alga known currently as Polysiphonia nigrescens. Dillwyn (1802-09 fasc. 10: pi. 75), in his treatment of Conferva fucoides, comments The [taxonomic] difficulty in the present species has been removed by the kindness of my friends the Rev. Hugh Davies and Archibald Menzies, who, from among some authentic specimens which they fortunately possess, have obligingly spared me two pieces marked "C. fucoides" exactly corresponding with the plant here figured . . .'. Unfortunately, no trace of these can be found at present among Dillwyn's herbarium materials, which are now much scattered (Dixon, 1966), neither can any relevant Hudson material once belonging to either Menzies or Davies be traced. There are, however, several specimens which were once in Davies's possession, and referred by him to C. fucoides, in BM-K; these are all of the alga known currently as Polysiphonia nigrescens. Later Dillwyn added an Introduction and Synopsis to the fascicles of plates of British Confervae (1802-09). In the Synopsis (p. 81) he was still not convinced that C. fucoides was distinct from C. nigrescens. Although no authentic herbarium material can be located at the present time, near-contemporary opinion was unanimous in referring the entity to the alga known today as Polysiphonia nigrescens, and provides a basis for accepting Conferva fucoides, lectotypified by the original description, as a synonym of Polysiphonia nigrescens. EXAMPLE b. Conferva nigra Huds. [Polysiphonia nigra (Huds.) Batters] The original description (Hudson, 1762:481) is brief, printed as follows: 12. CONFERVA filamentisaequalibusramosis,ramis nigra. faciculatis brevissimis. Anglis, black Conferva. Habitat in littore Eboracensi copiose. The later treatment (Hudson, 1778:595) is more detailed, but no synonyms or specimens are added: THE TYPIFICATION OF HUDSON'S ALGAE! A TAXONOMIC AND NOMENCLATURAL REAPPRAISAL 103 15 . CONFERVA filamentis aequalibus ramosis Ion- nigra. gissimis, ramis alternis multifidis brevissimus. Fl. angl. 481. Anglis, black Conferva. Habitat in littore Eboracensi pass/m. [annual]. V-X. Desc. Fila quinque pollicaria, cequalia, rigidiuscula, ni- gra, ramosa, ramis alternis brevissimis multifidis, fas- ciculatis. This indicates that the species was one of those described by Hudson on the basis of his own material. Contemporary workers referred the entity to the alga to which the epithet is now applied. For example, Dillwyn (1802-09 fasc. 10: pi. 70) had described C. atro-rubescens but stated later (Dillwyn 1802-09 fasc. 15: pi. 101) that 'Authentic specimens with which I have been favored by Sir Thomas Frankland and the Rev. Hugh Davies, prove that Hudson's Conferva nigra, respecting which I had previously been accustomed to yield to the generally received opinion of its being the same as Fucus fruticulosus, is in reality the C. atro mbescens of this work'. There is no trace at the present time of any such relevant material in BM, BM-K, or in the Lightfoot herbarium at BM-K. The volume of Dillwyn specimens at NMW contains several specimens on p. 202, with a label in which atro-rubescens is crossed through and replaced by nigra. One curious piece of information is provided by Batters (1902:81) in making the combination Polysiphonia nigra (Huds.) Batters. Here he stated that his transfer of Conferva nigra is based 'e spec. auth. in Herb. Brit. Mus.' but there is no 'authentic specimen' of Polysiphonia nigra at the present time in BM. Since there is no evidence suggesting any misapplication of the epithet, it can be typified by the description given by Hudson (1762:481). V. Species inquirenda EXAMPLE a. Conferva fulva Huds. [unassigned] The original description of Conferva fulva (Hudson, 1762:484) is extremely brief: fulva. 26. CONFERVA filamentis geniculatis ramosis, ramis ramulisque brevissimis alternis. Anglis, short Conferva. Habitat in littore Eboracensi. The treatment in the second edition (Hudson, 1778:602) is slightly changed and expanded. It is printed as follows: fulva. 39. CONFERVA filamentis geniculatis ramosis, ramis ramulisque brevissimis fulvis. Anglis, tawny Conferva. Habitat in saxis etfucis marinis, in littore Eboracensi. [annual]. V-IX. No synonyms are cited in either treatment, so that the species must have been described entirely from Hudson's own material. The entity has been a matter of conjecture for many years. Dillwyn (1802-09 fasc. 2: pi. 18) in his original description of Conferva repens (= Spermothamn- ion repens) stated: 'May not the present be Hudson's C. fulva, the description of which, in the Flora Anglica, is unfortunately so short, that unless any authentic specimen of it exists, which I believe there does not, it will always be impossible to tell what he meant by that name'. Later Dillwyn (1802-09 Introduction: 34), in an analysis of species of Conferva treated by Hudson, commented with respect to C. fulva 'I suspect that C. repens, T. 18, is the plant here designed, but proof is wanting'. Thus, Conferva fulva Huds., is an entity of unknown assignment at the present time and one about which near-contemporary authors were uncertain. In both editions of Flora anglica the treatments are very brief and equally referable to many red or brown algae. There is thus good reason for accepting that C. fulva is a species which cannot yet be typified. 104 LINDA M. IRVINE & PETER S. DIXON 5. Acknowledgements Many colleagues have contributed to our discussions and we would particularly like to thank Mr J. R. Laundon for his valuable comments. We also wish to thank the keepers and curators of the Department of Botany, British Museum (Natural History), the Royal Botanic Gardens at Kew, the Fielding-Druce herbarium at Oxford, and the National Museum of Wales at Cardiff for providing facilities to examine material. We are grateful for financial support of the research which led to this paper to the Joint Committee on Research of the University of Liverpool, the then Nature Conservancy, the National Science Foundation, and the Faculty Research and Travel Fund of the University of California. 6. References Batters, E. A. L. 1902. A catalogue of the British marine algae. /. Bot., Land. 40 (suppl. 2): 1-107. Bauhin, J. 1651. Historiaplantarumuniversalis. 3: 1-866+12. Ebroduni. Clokie, H. N. 1964. An account of the herbaria of the Department of Botany in the University of Oxford. viii+280pp. Oxford. Dandy, J. E. 1958. The Sloane herbarium. 246 pp. London. De Candolle, A. L. P. P. 1867. Lois de la nomenclature botanique ... 60 pp. Paris. Dillenius, J. J. 1742 ['1741']. Historia muscorum. xvi+576 pp. Oxonii. Dillwyn, L. W. 1802-09. British Confervae. 16 fasc., Introduction, Synopsis. 109+ A-G pis, 94 pp. London. Dixon, P. S. 1959a. Notes on two important algal herbaria. Br. phycol. Bull. 1 (7): 35^42. 19596. Taxonomic and nomenclatural notes on the Florideae, 1. Bot. Notiser 112: 339-352. 1960. Taxonomic and nomenclatural notes on the Florideae, II. Bot. Notiser 113: 295-319. 1962. Taxonomic and nomenclatural notes on the Florideae, III. Bot. Notiser 115: 245-260. - 1963. Further comments on the typification of Hudson's algae. Br. phycol. Bull. 2: 265-266. - 1966. Notes on important algal herbaria, IV. The herbarium of Lewis Weston Dillwyn (1778-1855). Br. phycol. Bull. 3: 19-22. 1967. The typification oiFucus cartilagineus L. and F. corneus Huds. Blumea 15: 55-62. & Irvine, L. M. 19770. Seaweeds of the British Isles. 1. Rhodophyta (1). Introduction, Nemaliales, Gigartinales. xi+252 pp. London. & - - 1911 b. Miscellaneous notes on algal taxonomy and nomenclature IV. Bot. Notiser 130: 137-141. Druce, G. C. & Vines, S. H. 1907. The Dillenian herbaria, cxii+258 pp. Oxford. Ellis, J. 1767. On the animal nature of the genus of zoophytes, called Corallina. Phil. Trans. R. Soc. 57: 404-^27. Gmelin, S. G. 1768. Historia fucorum. [viii]+vi+[iv] 239 pp. Petropoli. Hudson, W. 1762. Flora anglica. vii+[8]+506+[23] pp. London. 1778. Flora anglica. 2nd ed. xxxviii+690 pp. London. - 1798. Flora anglica. 3rd ed. [4]+iv+xxxii+688 pp. London. Juel, H. 0. 1936. Joachim Burser's Hortus Siccus mit erklarungen heraus gegeben. Symb. hot. upsal. 2 (1): i-v+ 1-187. Laundon, J. R. 1963. The taxonomy of sterile crustaceous lichens in the British Isles. 2. Corticolous and lignicolous species. Lichenologist 2: 101-151. 1966. Hudson's Lichen siliquosus from Wiltshire. Lichenologist 3: 236-241. - 1976. Lichens new to the British flora: 5. Lichenologist %: 139-180. Lightfoot, J. 1777. Flora scotica 2. [4]+531-1151 + [24] pp. London. Linnaeus, C. 1753. Species plantarum. 2: 561-1200+31. Holmiae. 1763. Species plantarum. 2nd ed. 2: 782-1684. Holmiae. - 1767. Systema natura. 12th ed. 1 (2): 533-1327. Holmiae. - 1771. Mantissa plantarum altera. iv+ 143-510 pp. Holmiae. - 1774. Systema vegetabilium. 13th ed. [iv]+844 pp. Gottingae & Gothae. Morison, R. 1680-99. Plantarum historiae universalis oxoniensis 2, 3. Oxonii. Oeder, G. C. 1762-1883 ['1761-1883']. Icones plantarum sponte nascentium in regnis daniae et norvegiae . . . Florae danicae nomine inscriptum. 17 vol. + Suppl. Hafniae. Parke, M. & Dixon, P. S. 1976. Check-list of British marine algae third revision. /. mar. biol. Ass. U.K. 56: 527-594. Petiver, J. 1695-1703. Musei Petiveriani. 96 pp. London. THE TYPIFICATION OF HUDSON'S ALGAE: A TAXONOMIC AND NOMENCLATURAL REAPPRAISAL 105 Plukenet, L. 1696. Almagestum botanicum . . . [ii]+402+[2] pp. London. Ray, J. 1704. Historia plantarum 3. London. 1724. Synopsis methodica stirpium britannicarum. 3rd ed. [xiii]+482+[30]. Londini. Ross, R. & Brenan, J. P. M. 1970. Cryptogamic collections at Kew and the British Museum. Taxon 19: 136. Silva, P. C. 1952. A review of nomenclatural conservation in the algae from the point of view of the type method. Univ. Calif. Publs Bot. 25: 241-324. Smith, J. E. & Sowerby, J. 1790-1814. English botany. 36 vol. London. Stafleu, F. A. et al. (Ed.) 1978. International code of botanical nomenclature, xiv+457 pp. [Regnum veg. 97]. Utrecht. Stearn, W. T. 1957. An introduction to the Species plantarum and cognate botanical works of Carl Linnaeus. In C. Linnaeus, Species plantarum. A facsimile of the first edition 1753. 1: v-xiv+ 1-176. London. Turner, D. 1802. A synopsis of the British Fuci. 2 vol. xlvi+400 pp. London. 1808-19. Fuci, sive plantarum Fucorum generi ... 4 vol. London. Vines, S. H & Druce, G. C. 1914. An account of the Morisonian herbariumi lxviii+350 pp. Oxford. British Museum (Natural History) Seaweeds of the British Isles Volume 1 Rhodophyta Part 1 Introduction, Nemaliales, Gigartinales P S Dixon & L M Irvine Seaweeds of the British Isles The result of many year's research carried out by the British Museum (Natural History) and the British Phycological Society, this is the first of a series of books which will be published under this title covering all the British and the majority of northern Atlantic seaweeds. Volume 1 Rhodophyta Part 1 Introduction, Nemaliales, Gigartinales In this, the first of three parts comprising Volume 1, a general introduction to the Rhodophyta - dealing with such topics as morphology, reproduction and economic utilization - is followed by treatment of the orders Nemaliales and Gigartinales. Each species is described and illustrated and notes on the ecology and distribution are given. Keys to aid identification are also included. As with all the books in the series, this title will provide a standard work of reference in a field where for too long nothing up-to-date has been available. 264 pp, 90 figs ISBN 565 00781 5 1977 Approx. 12. Titles to be published in Volume 10 Taxonomic studies in the Labiatae tribe Pogostemoneae. ByJ. R. Press The typification of Hudson's algae: a taxonomic and nomenclatural reappraisal By L. M. Irvine & P. S. Dixon Seaweeds of the Faroes (3 papers) . By D. E. G. Irvine; I. Tittley, W. F. Farnham & P. W. G. Gray; J. H. Price & W. F. Farnham The lichen genus Steinera. By A. M. Henssen & P. W. James Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed by Henry Ling Ltd, Dorchester. Bulletin of the British Museum (Natural History) Seaweeds of the Faroes Botany series Vol 10 No 3 25 November 1982 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready; each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW75BD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.) Tustees of the British Museum (Natural History), 1982 The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Mr J. F. M. Cannon Editor of Bulletin: Mr P. W. James Assistant Editor: Mr J. R. Laundon ISSN 0068-2292 Botany series Vol 10 No 3 pp 107-225 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 25 November 1982 Seaweed^of the Faroes Contents 1: The flora. By D. E.G. Irvine 2: Sheltered fjords and sounds. By I. Tittley, W. F. Farnham and P. W. G. Gray 3: Open shores. By J. H. Price and W.F. Farnham S\SH Jf 2s Nov V /92 . 109 . 133 153 Seaweeds of the Faroes 1: The flora David Edward Guthrie Irvine Polytechnic of North London, Holloway Road, London N7 8DB Contents Synopsis ............. 109 1. Introduction ............ 109 2. Methodology 110 3. Species list ............ HI Cyanophyta ........... 112 Rhodophyta 112 Chrysophyta 120 Phaeophyta 120 Chlorophyta 126 4. Conclusions ............ 129 5. Acknowledgements ........... 130 6. References ............ 130 Synopsis A brief description of the Faroes is followed by an account of earlier phycological investigations and the reasons for carrying out a general survey of the marine algae at the present time. A list of the known seaweed flora is given, including the records of previous investigators, with brief habitat and distribution notes, and records of occurrence in the Orkneys, the Shetland Isles, and Iceland. The flora is shown to be similar to that of the Shetland Isles, though much poorer in species, but with a few subarctic species in the north which have not so far been found in the British Isles. It shows remarkably few changes from that described by B0rgesen and his colleagues around the end of the 19th century. Some species not recorded by us, e.g. Porphyra linearis, may have been present earlier in the season. If dubious records are excluded, 9 Cyanophyta, 95 Rhodophyta, 74 Phaeophyta, 44 Chlorophyta, and 1 marine Vaucheria, 223 spp. in all are recorded, whereas 301 spp. have been found in the neighbouring Shetland Isles. 1. Introduction The Faroes (Faer0erne; Faeroes; F0royar) are a group of 18 islands and innumerable rocks, stacks and skerries, with an area of 1325 sq. km (Fig. 1). The islands lie mainly in the Gulf Stream, but a cold north-westerly current impinges on the northernmost islands of the group (Fig. 2). They lie about halfway between the Shetland Isles and Iceland, and are under the protection of Denmark, but have been self-governing since 1948. The population is about 50 000, mostly forming scattered communities fringing the deeply indented coastline. The capital, Torshavn, is a thriving port in the centre of the group, and the other main population centre is the fishing village of Klaksvik in the northern group of islands. To the west and the north the shores tend to be precipitous with only a minimal littoral, and access from the land is not practicable over large stretches. The islands are deeply penetrated by narrow fjords and separated by equally narrow sounds. The tidal range is small, but the littoral is effectively greatly extended by wave surge and spray. The climate is mild, with an annual rainfall Bull. Br. Mus. nat. Hist. (Bot.) 10 (3): 109-131 Issued 25 November 1982 110 SEAWEEDS OF THE FAROES KUNOY STRE YM Y KALSOY(\ A \|V I 00 Y O VINOY S DOUR OY Fig. 1 Map of the Faroes showing the main islands of about 1600 mm; conditions are generally cloudy and fogs are frequent in the summer months, while gales may occur at any season. The seaweeds of the Faroes have been investigated at various times, notably by H. C. Lyngbye (1819), E. Rostrup (1870) and H. G. Simmons (1897), and a definitive survey was carried out by F. B0rgesen and various colleagues and published in a series of papers from 1895 to 1905. B0rgesen was able to make several visits to the islands, and with the co-operation of the Danish Marine Department was able to spend long periods on a fisheries protection vessel from which he was able to visit many of the otherwise inaccessible shores and to have dredging facilities. Although he only visited the islands in spring and summer, H. J0nsson provided him with an extensive collection made in autumn and early winter. The contributions of other workers are listed in detail in the introduction to B0rgesen's account of the marine algae (1902). Since the papers of B0rgesen, surprisingly little has been published concerning the marine algae of the Faroes, although their flora is of obvious interest to phycologists working in the North Atlantic area. B. Rex carried out some studies of vegetation profiles on several shores in 1970. His most important find was that Dilsea carnosa is common in the subtidal. It had been recorded earlier by P. A. Holm (1855), but as later investigators, including B0rgesen, had not found it, this record has been discarded by B0rgesen. In 1975 G. Holt compared the seaweed floras (including blue-green algae and diatoms) of eight widely differing sites, using a large number of different stations at each site. His work, however, did not produce many additions to the known seaweed flora. 2. Methodology It was hoped that an intensive study of the intertidal and subtidal vegetation of the Faroes, using boats and modern diving equipment, would show the Faroese algal flora to be much richer in THE FLORA 111 Fig. 2 Map showing position of the Faroes in relation to main ocean currents and neighbouring land masses (after Ryder in B0rgesen 1905: 813). species than the findings of the earlier workers had indicated; this had been the case in a similar investigation carried out in the Shetland Isles (Irvine, 1974). Accordingly, a rather flexible expedition was arranged for 1980, based on two laboratories in Torshavn kindly lent by the Academia Faeroensis and formed of a number of semi-independent groups and individual scientists from various countries who stayed for limited periods. Although they largely pursued their own special interests, their contributions to this species list were of considerable import- ance. The main group was based on Torshavn for two weeks in early July 1980, then a week at Vibareibi in the northern island of Viboy, followed by a week in Torshavn. Most of the major islands, apart from Suburoy, were examined both intertidally and subtidally (using inflatables and scuba equipment), and particular attention was given to Skalafj0rbur, a large fjord in Eysturoy, not unlike Sullom Voe in the Shetland Isles in some respects. Extensive collections were made, and the species list which follows is an amalgam of earlier records and of the observations and collections of all the members of this expedition. 3. Species list The nomenclature chiefly follows Parke & Dixon (1976) for taxa which also occur in the British Isles, although many abbreviations of authors' names are revised to accord with editorial requirements. 112 SEAWEEDS OF THE FAROES Abbreviations: (!) Recorded on the 1980 expedition. New to the Faroes. 0. Recorded from the Orkneys. S. Recorded from the Shetland Isles (Irvine, 1980). 1. Recorded from Iceland (Caram & Jonsson, 1972). [ ] Name used by B0rgesen (1902). Note that B0rgesen's place names have been changed to Faroese. CYANOPHYTA Calothrix Crustacea Thuret ex Bornet & Flah. [C. aeruginea, C. scopulorum, Rivularia atra] Near high tide level. Widely distributed. (!) O. S. I. Entophysalis conferta (Kiitz.) Drouet & Daily [Dermocarpa farlowi, D. violacea, Pleurocapsa amethystea] Epiphytic on various intertidal algae, notably Polysiphonia lanosa. Common, widely distri- buted. (!) O. S. I. E. deusta (Menegh.) Drouet & Daily [Hyella caespitosa, H. endophytica} In shells of molluscs, subtidally to 40 m. Found on Streymoy by B0rgesen. O. S. I. Microcoleus lyngbyaceus (Kutz.) P. Crouan & H. Crouan ex Gomont Listed by Holt from Saksun under the names Hydrocoleum glutinosum, Lyngbya semiplena, Merismopedia elegans, and Oscillatoria tenuis. O. S. M. vaginatus (Vaucher) Gomont ex Gomont [Phormidium autumnale] Near high tide level, on wet rocks and in pools. Found by B0rgesen on Kalsoy: Husar, and on Streymoy: Tinganes. I. Oscillatoria lutea Agardh ex Gomont [Lyngbya lutea] Intertidally on somewhat exposed shores. Recorded by B0rgesen from Viboy: Viovik, and Streymoy: Torshavn. O. Schizothrix calcicola (Agardh) Gomont Listed by Holt from Torshavn (as Phormidium fragile) . O. S. I. S. tenerrima (Gomont) Drouet [Microcoleus tenerrimus] Found once by H. J0nsson at Hvannasund. Spirulina subsalsa Oersted ex Gomont In rock pools on moderately exposed shores. Recorded by B0rgesen from near Torshavn and by Holt (as S. subtilissima} from Saksun. O. S. I. RHODOPHYTA Actinococcus subcutaneus The sporophyte phase of Phyllophora truncata (q.v.) parasitic on the gametophyte. Ahnfeltia plicata (Huds.) Fr. [including Sterrocolax decipiens] Lower intertidal and shallow subtidal, in sheltered fjords and bays. Locally common. (!) O.S.I. Antithamnion boreale (Gobi) Kjellman [A. plumula var. boreale] Subtidally to 50 m, mainly on sheltered shores. O. I. A. floccosum (Mull.) Kleen Intertidal and shallow subtidal, mainly on sheltered shores. Occasional, widely distributed. (!)O. S. I. THE FLORA 113 A. plumula (Ellis) Thuret Subtidal on rocks, shells, and epiphytic, exposed and sheltered shores. Widely but sparingly distributed. (!)O.S. (I?). Audouinella alariae (Jonsson) Woelk. [Chantransia Alariae] Shallow subtidal on Alaria fronds, especially on exposed shores. Widely distributed and generally abundant, although B0rgesen recorded it only from Streymoy: near Torshavn. (!) O. S. A. daviesii (Dillwyn) Woelk. [Chantransia Daviesii] Subtidal on various algae. Widely distributed. (!) O. S. A. efflorescens (J. Agardh) Papenf. [Chantransia efflorescens] On sheltered coasts, subtidally, epiphytic on various algae. Eysturoy. A. membranacea (Magnus) Papenf. [Rhodochorton membranaceum] Intertidally and subtidally, to 40 m, sheltered and exposed coasts, on Bryozoans, often with A. spetsbergensis. Widely distributed, probably common but overlooked. I. A. purpurea (Lightf.) Woelk. [Rhodochorton Rothii] On rocks in caves, fissures, and waterfalls, from the upper spray zone downwards intertidally, and epiphytic on stipes of Laminaria hyperborea subtidally to about 20 m, commonly on exposed shores. Abundant and widely distributed. (!) O. S. I. A. secundata (Lyngbye) P. Dixon [Chantransia secundata] Epiphytic on various algae, intertidal on exposed and sheltered shores. Widely distributed, common. (!) O. S. I. A. seiriolana (Harvey-Gibson) P. Dixon [Rhodochorton seiriolanum] Recorded by B0rgesen as epiphytic on Ceramium shuttleworthianum on an exposed coast, near high-water mark. Streymoy: Velbastabur. Otherwise known only from the type locality [Anglesey (Wales): Puffin Island]. A. spetsbergensis (Kjellman) Woelk. [Rhodochorton penicilliforme] Intertidally and subtidally to 40 m, on Bryozoans. Widely distributed but sparsely recorded; probably overlooked. (!) S. I. A. virgatula (Harvey) P. Dixon [Chantransia virgatula. Probably Lyngbye's Callithamnion lanuginosum belongs here, fide B0rgesen, though Rostrup identifies it with A. daviesii.} Epiphytic on various algae intertidally and subtidally, on sheltered and exposed shores. Common and widely distributed. (!) O. S. I. Bangia atropurpurea (Roth) Agardh [B. fuscopurpurea] On rocks intertidally from upper spray zone downwards, especially on exposed shores, often associated with Urospora and Ulothrix spp. Scattered filaments occurred intermingled with larger algae. Common and widely distributed. See Conchocelis. (!) O. S. I. Bonnemaisonia hamifera Hariot See Trailliella. *Brongniartella byssoides (Gooden. & Woodw.) Schmitz On rocks subtidally on moderately exposed shores. Rare, widely distributed. (!) O. S. Callithamnion corymbosum (Sm.) Lyngbye (as var. amphicarpa} Epiphytic subtidally at about 16 m. Eysturoy: near Oyri, a few fronds only. O. S. *C. decompositum J. Agardh Epiphytic subtidally. Occasional. (!) S. [C. granulatum (Ducluz.) Agardh Recorded by B0rgesen, but almost certainly in error. All Faroese specimens distributed in herbaria under this name appear to be referable to C. hookeri (Dixon & Price, 1981) and no specimens of C. granulatum could be found at any of the sites listed for it by B0rgesen. O. S.] 114 SEAWEEDS OF THE FAROES C. hookeri (Dillwyn) Gray [C. polyspermum, C. scopulorum] On rocks and epiphytic; in caves, fissures, and on open rock faces at about midtide level on exposed coasts. Common and widely distributed. (!) O. S. I. C. sepositum (Gunnerus) P. Dixon & J. Price [C. arbuscula] On rocks, barnacles, and Mytilus at midtide level on exposed coasts, especially on steeply sloping faces. Abundant, widely distributed. (!) O. S. I. Callocolax neglectus Schmitz ex Batters Parasitic on Callophyllis laciniata subtidally to 50 m. Generally distributed with the host and often common, but not observed intertidally. (!) O. S. I. Callophyllis cristata (Agardh) Kiitz. [Euthora cristata] Epiphytic on haptera of Laminaria hyperborea, more rarely on rocks and shells, lower intertidal (rarely) and subtidal to 50 m, exposed to moderately sheltered coasts. Very varied in growth form, intertidal plants tending to be much branched and bushy. Widely distributed, common. (!) O. S. I. C. laciniata (Huds.) Kiitz. On stones and shells and epiphytic, especially on the lower stipe and haptera of Laminaria hyperborea, in shady pools and subtidally to 50 m, in exposed to moderately sheltered localities. Common and widespread. (!) O. S. Ceramium diaphanum (Lightf.) Roth Intertidal and shallow subtidal, a few scattered fronds probably referable to this species. Recorded by Lyngbye (1819:119, pi. 37) from Torshavn and Eibi, but these records dismissed by B0rgesen as referring to C. rubrum. (!) O. S. C. rubrum (Huds.) Agardh Intertidal and subtidal, on rocks and epiphytic, on exposed to sheltered coasts. Widespread and abundant. Very variable in size and growth form. (!) O. S. I. [C. secundatum J. Agardh Recorded by Lyngbye (1819) from between Torshavn and Hoyvik. Record dismissed by B0rgesen as referring to a form of C. rubrum]. C. shuttleworthianum (Kiitz.) Rabenh. [C. acanthonotum] On rocks, barnacles, and Mytilus, in spongy tufts about midtide on exposed shores. Abun- dant, widely distributed. (!) O. S. I. C. strictum Harvey Recorded by Holt from Sandoy. S. Chondrus crispus Stackh. On rocks, midtide to shallow subtidal, exposed to sheltered shores, and in rock pools, often in dense stands. Neither Lyngbye nor B0rgesen considered it common, but it is generally distributed and at least locally abundant. (!) O. S. I. Choreocolax polysiphoniae Reinsch Parasitic on Polysiphonia lanosa. Recorded only sparsely, but probably occurring generally where the host is present. First recorded by B0rgesen (1905: 773). (!) O. S. Clathromorphum circumscriptum (Stromf.) Foslie Found once by B0rgesen on Streymoy: in Sundini between Hosvik and Hvalvik. Later placed by Foslie in Phymatolithon compactum (B0rgesen 1905: 773) but probably belongs to Clathromorphum. I. Conchocelis (phase in the life history of spp. of Bangia and Porphyra) Subtidal to 50 m, sheltered and exposed shores, and in shells. Common, widely distributed. (!) O.S.I. THE FLORA 115 Corallina officinal is L. On rocks intertidally, in pools, and in shallow subtidal situations, on exposed to fairly sheltered shores, often in dense patches. Common, widely distributed. (!) O. S. I. Cruoria pellita (Lyngbye) Fr. Encrusting rocks, shells, and stones intertidally and subtidally, and epiphytic on the stipes and holdfasts of Laminaria hyperborea on both exposed and sheltered shores. Common and widely distributed. (!) O. S. I. Cryptopleura ramosa (Huds.) Kylin ex Newton [Nitophyllum laceratum} On rocks and amongst Corallina subtidally, as small prostrate specimens. Widely dispersed but not common. (!) O. S. Cystoclonium purpureum (Huds.) Batters [C. purpurascens] On rocks and stones in shallow subtidal, mainly on sheltered coasts. Lyngbye considered it rare, and B0rgesen remarks This species does not appear to be widely distributed along the shores of the Faeroes'. However, we found it frequently, especially just below low water on sheltered shores. (!) O. S. I. Delesseria sanguinea (Huds.) Lamouroux On rocks, lower intertidal (in pools, caves, and deep crevices) and subtidally to 50 m, on exposed to sheltered shores. Common and widespread. (!) O. S. I. Dermatolithon corallinae (P. Crouan & H. Crouan) Foslie Epiphytic on Corallina. Common and widespread with its host. (!) S. I. D. crouanii (Foslie) Lemoine [Lithophyllum Crouani] Epiphytic on stipes and holdfasts of Laminaria hyperborea, associated with D. pustulatum. Eysturoy: Gjogv. Suburoy: Lobra. I. D. hapalidioides (P. Crouan & H. Crouan) Foslie Noted by Ostenfeld on a Patella shell in Suburoy: Hvannhagi. O. D. pustulatum (Lamouroux) Foslie [D. macrocarpum] Epiphytic on various algae (notably Furcellaria, Gigartina, and Laminaria hyperborea), intertidally and subtidally, exposed to sheltered shores. Abundant and widespread. (!)O. S. I. Devaleraea ramentacea (L.) Guiry [Halosaccion ramentaceum] On loose stones about low water mark and immediately below, on very sheltered shores; locally abundant in a few northern fjords over a limited area. Borboy: Haraldssund (!) and near Klaksvik. Viboy: Hvannasund (!). Streymoy: Vestmanna. It has also been recorded by J0nsson from Svinoy: near the Havn, on a fairly exposed shore on rocks near low water, but a close search of this very atypical site failed to reveal any specimens. There were, however, some stands of Dumontia with a growth form very similar to that of Devaleraea, and it seems likely that the original record was a misidentification. (!) I. Dilsea carnosa (Schmidel) Kuntze Recorded by Holm (1855) but not found by later investigators, and the record therefore dismissed by both Rostrup (1870) and B0rgesen (1903). Nevertheless, it was found by both Rex (1970) and Holt (1975), and we found it to be abundant and generally distributed, growing on rocks in intertidal pools, and subtidally to relatively shallow depths. Discoloured specimens were widespread and prominent in the drift, and living specimens were large and healthy. Had it occurred like this in earlier times it could scarcely have been missed or confused with other species. It seems probable, therefore, that Dilsea has only entered the Faroes during this century, some time before 1970. (!) O. S. I. Dumontia contorta (S. Gmelin) Rupr. [D.filiformis] On rocks and stones, intertidally and in the shallow subtidal, on exposed and sheltered shores. Abundant, widely distributed. (!) O. S. I. 116 SEAWEEDS OF THE FAROES *Erythrotrichia boryana (Mont.) Berthold Eysturoy: Skalafj0rbur, Skalabotnur. (!) O. *E. carnea (Dillwyn) J. Agardh [E. ceramicola] Intertidally and in the shallow subtidal as isolated filaments interwoven with other algae, such as Ceramium shuttleworthianum and Corallina, mainly on sheltered shores. Probably widely distributed but never abundant; easily overlooked. (!) O. S. Fimbrifolium dichotomum (Lepechin) G. Hansen [Rhodophyllis dichotoma] On stones and shells, and especially on stipes and haptera ofLaminaria hyperborea, subtidally to about 50 m, on exposed to moderately sheltered shores in the northern islands. (!) I. [Fig- 3] Fig. 3 Two specimens of Fimbrifolium dichotomum (Lepechin) G. Hansen growing with Delesseria sanguined on Odonthalia dentata. The scale shows 1 cm. [This species has not so far been recorded from the British Isles, but should be looked for in north-east Scotland.] THE FLORA 117 Furcellaria I urn brie-alls (Huds.) Lamouroux [F. fastigiatus] On rocks, lower intertidal (in pools) and subtidally to 20 m, on moderately exposed to sheltered shores, in dense but scattered patches. Frequent and widely distributed. (!) O. S. I. Gigartina stellata (Stackh.) Batters [G. mamillosa] On rocks, intertidal and immediate subtidal, on exposed coasts, often forming zones or large patches near low water on very exposed rock faces. Abundant and widely distributed. (!)O. S. I. (ion lot rich u in alsidii (Zanard.) M. Howe Recorded by Holt from Torshavn as G. elegans. Not recorded from Orkney, Shetland or Iceland. Griffithsia corallinoides (L.) Batters [Conferva (Griffithsia) corallina] Reported by Landt (1800: 233) as occurring in the Faroes, but not found since. Possibly an error, but in Shetland it occurs on moderately sheltered coasts subtidally to 20 m. O. S. G. flosculosa (Ellis) Batters [G. setacea] On rocks, subtidally to 20 m, apparently very rare. Found by B0rgesen as a single small specimen (Suburoy: Trongisvagsfj0rbur) and during the present expedition by P. A. Asen as a single small tuft. (!) O. S. Harveyella mirabilis (Reinsch) Reinke Parasitic on Rhodomela confervoides . Apparently rare. Kunoy (J0nsson). Suburoy: Trongis- vagur (B0rgesen). A single specimen found by us. (!) I. Hildenbrandia rubra (Sommerf.) Menegh. [H. rosea] Intertidally and to the top of the spray zone, on open rocks and stones and in pools, on sheltered and exposed shores. Abundant and widespread. (!) O. S. I. Laurencia pinnatifida (Huds.) Lamouroux Intertidally on rocks and large stones on exposed and sheltered coasts, rare. Streymoy: between Torshavn and Hoyvik (Rostrup). Suburoy: Tv0royri and 0ravik (Ostenfeld). Searched for carefully, including the Streymoy site, but not found on the present expedition. Original specimens correctly identified (J. H. Price, pers. com.). O. S. Leptophytum laeve (Stromf.) Adey [Lithothamnion laeve] In deep water, on sheltered and exposed coasts. Found by B0rgesen on Bordoy: Haraldssund; Streymoy: Argir; Suduroy: Trongisvagsfj0rbur and Lopra. (!) S. I. Lithophyllum incrustans P. Philippi Intertidal, on Phymatolithon polymorphism. Recorded by Jonsson on Viboy: Vibareibi, and by B0rgesen on Suburoy: Hvalba. O. S. Lithothamnion glaciale Kjellmun On rocks, stones and shells, subtidal, exposed coasts. Frequent and widely distributed (!) S. I. Lomentaria articulata (Huds.) Lyngbye On rocks and stones in the lower intertidal, often accompanying Corallina, especially abundant on exposed coasts, where it often occurs in dense mats. Abundant and generally distributed. (!)O.S. L. clavellosa (Turner) Gaillon On rocks, stones, and larger algae, especially on the stipes of Laminaria hyperborea, lower intertidal and subtidal to 30 m, on very exposed to moderately sheltered shores. Very variable in growth form. Common and widely distributed. (!) O. S. I. L. orcadensis (Harvey) F. Collins ex Taylor [L. rosea] Near low water and subtidally to 30 m, on rocks and holdfasts of Laminaria hyperborea. Widely distributed, frequent on exposed shores. (!) O. S. I. 118 SEAWEEDS OF THE FAROES Membranoptera alata (Huds.) Stackh. [Delesseria alata] Intertidally in clefts, caves and pools, and subtidally on rocks, stones, and epiphytic on Laminaria hyperborea, on both sheltered and exposed shores. Common and generally distributed. (!) O. S. I. Odonthalia dentata (L.) Lyngbye On rocks, intertidally in shady pools, and subtidally on rocks, stones, and stipes of Laminaria hyperborea; on sheltered and exposed shores. Common, widely distributed. (!) O. S. I. Palmaria palmata (L.) Kuntze [Rhodymenia palmata] On rocks and epiphytic on stipes of Laminaria hyperborea, intertidally and in the shallow subtidal, on exposed and sheltered shores. Often forming compact tufted mats intertidally to well above high tide level where streams cascade over a rocky shore. Abundant and widespread. (!) O. S. I. Peyssonnelia dubyi P. Crouan & H. Crouan On rocks, stones, shells, and stipes of Laminaria hyperborea, subtidally to 20 m, on sheltered to exposed shores. Frequent, widely distributed. (!) O. S. Phycodrys rubens (L.) Batters [Delesseria sinuosa] Lower subtidal on rocks in shady pools and subtidally to 50 m, on rocks and stones and shells, and epiphytic on stipes of Laminaria hyperborea, on sheltered to exposed coasts. In very sheltered conditions occurs as the forma lingulata, with narrow spiky fronds drawn out into twisted tendrils, looking quite unlike the typical form. Widespread and abundant. (!) O. S. I. Phyllophora crispa (Huds.) P. Dixon [P. rubens] On rocks in subtidal, on sheltered and exposed shores. Occasional, in scattered localities, but well grown. Simmons (1896: 266) reported this species from Borboy: near Klaksvik, but B0rgesen was unable to find specimens in this locality, identified Simmons' specimens as young P. truncata and hence removed the species from the Faroese flora list, to which it should now be restored. (!) O. S. I. P. pseudoceranoides (S. Gmelin) Newroth & A. Taylor [P. membranifolia] On rocks intertidally in caves, and in the shallow subtidal on somewhat exposed shores. There is much confusion between young immature specimens of this species and of P. traillii. O. S. I. *P. traillii Holmes & Batters In caves and undercuts of cliffs, pools, and shallow subtidal, exposed shores. Occasional. (!) O. P. truncata (Pallas) Zinova [P. Brodiaei] On stones in shallow subtidal, especially in sheltered conditions. Fairly frequent in widely scattered localities. See Actinococcus. (!) O. S. I. Phymatolithon laevigatum (Foslie) Foslie On stones and shells subtidally to 20 m, on both sheltered and exposed coasts. Widely distributed. O. S. I. P. lenormandii (Aresch.) Adey [Lithothamnion lenormandi] Intertidally in pools on both exposed and sheltered shores. Recorded by B0rgesen from only two sites, but apparently common and widely distributed. (!) O. S. I. P. polymorphum (L.) Foslie On rocks near low water and in shallow subtidal to 20 m. According to B0rgesen it occurs luxuriantly in caves to well above sea level. Abundant and widely distributed. (!) O. S. I. Plocamium cartilagineum (L.) P. Dixon [P. coccineum] On rocks in pools in the lower intertidal, and subtidally on rocks and on stipes and holdfasts of Laminaria hyperborea. On exposed to sheltered coasts. Abundant and widely distributed. (!) O.S.I. THE FLORA 119 Plumaria elegans (Bonnem.) Schmitz On rocks in the lower intertidal on exposed shores, typically as a mat on shady vertical rock faces, more rarely in pools. Occasional, widely distributed. (!) O. S. I. Poly ides rotundus (Huds.) Grev. On stones subtidally to 20 m, on sheltered coasts. Locally abundant, widely distributed (!) O.S.I. Polysiphonia brodiaei (Dillwyn) Sprengel On rock intertidally near low water, usually in pools, mainly on exposed shores. Common, widely distributed. (!) O. S. P. elongata (Huds.) Sprengel On stones, shells, and other algae in the shallow subtidal to 20 m, on both sheltered and exposed coasts, occasional. Locally abundant, widely distributed. (!) O. S. I. *P. fibrata (Dillwyn) Harvey Lower intertidal amongst Corallina in pools on exposed shores. Locally abundant, widely dispersed. (!) O. S. *P. fruticulosa (Wulfen) Sprengel On rocks on exposed shores, and intertidally in pools. Occasional. (!) S. P. lanosa (L.) Tandy [P. fastigiata] Parasitic on Ascophyllum. Widespread and abundant with the host, but absent where moderate wave exposure occurs. (!) O. S. I. [P. lepadicola (Lyngbye) J. Agardh Recorded by Lyngbye (as Hutchinsia lepadicola), but his specimens were found by B0rgesen to be creeping filaments of P. urceolata]. P. nigra (Huds.) Batters [P. atrorubescens] A few specimens collected by Ostenfeld from 6-8 m subtidally. Suburoy: Trongisvagsfj0rbur. Recorded by Lyngbye as occurring in the Faroes, but no specimens were found in his herbarium. O. S. P. nigrescens (Huds.) Grev. On rocks and stones in shallow subtidal, on both sheltered and exposed shores. Occasional. (!) O.S.I. P. urceolata (Lightf. ex Dillwyn) Grev. On rocks intertidally near low water mark, especially on exposed shores, and subtidally on stipes of Laminaria hyperborea, to 20 m. Abundant and generally distributed. (!) O. S. I. P. violacea (Roth) Sprengel Epiphytic on Laminaria, shallow subtidal, rare, Suburoy: Trongisvagsfj0rbur (Ostenfeld and Simmons). Eysturoy: Torshavn (Holt). O. S. Porphyra leucosticta Thuret On rocks and stones at low water level, and in shallow subtidal, on exposed and sheltered shores. Common, widely distributed. (!) O. S. P. linearis Grev. [P. umbilicalis f. linearis] According to B0rgesen (1903) found by Lyngbye to be abundant on Suburoy: Hvalbiar- fj0rbur. Not found by us, but to be expected in winter and spring. O.S.I. P. miniata (Agardh) Agardh Free-floating in inner parts of fjords, on rocks and stones and epiphytic, subtidally to 30 m, on exposed and sheltered coasts. Common, widely distributed. (!) O. S. I. P. purpurea (Roth) Agardh [P. umbilicalis f . laciniata] Epiphytic on fucoids intertidally, and in shallow subtidal, on sheltered coasts. Common, widely distributed. (!) O. S. I. 120 SEAWEEDS OF THE FAROES P. u m hi lk-:il is (L.) J. Agardh On rocks intertidally, especially abundant in the swash zone on exposed coasts. Widely distributed. (!) O. S. I. Porphyropsis coccinea (J. Agardh ex Aresch.) Rosenv. [Porphyra cocdnea] Subtidally, epiphytic, mainly on Desmarestia aculeata, on exposed and sheltered shores. Common, widely distributed. (!) O. S. I. Pterosiphonia parasitica (Huds.) Falkenb. On rocks and shells in shallow subtidal on exposed shores. Common, widely distributed. (!) O.S.I. Ptilota plumosa (Huds.) Agardh On rocks and more usually epiphytic on Laminaria stipes and holdfasts, subtidal on exposed to relatively sheltered shores to 50 m. Common and widespread. (!) O. S. I. This species and P. serrata are often not easily separable on morphological grounds, and are possibly conspecific. P. serrata Kiitz. [P. pectinata] On rocks and epiphytic on Laminaria stipes and holdfasts, subtidal on exposed to relatively sheltered coasts. Occasional, widely distributed. (!) I. Rhodomela confervoides (Huds.) P. C. Silva [R. subfusca] On stones and shells intertidally, and in shallow subtidal, on sheltered coasts. Occasional. (!) O.S.I. R. lycopodioides (L.) Agardh On rocks and stones intertidally, and in shallow subtidal, on sheltered and exposed shores. Common, widely distributed. (!) O. S. I. This species and R. confervoides appear to intergrade morphologically, and are possibly conspecific. Rhodophysema elegans (P. Crouan & H. Crouan ex J. Agardh) P. Dixon [Rhododermis elegans] On stones and shells in subtidal, occasional. Found by J0nsson on Streymoy: Kaldbaksfj0rbur and by B0rgesen on Suburoy: Trongisvagsfj0rbur. I. Sterrocolax decipiens SeeAhnfeltia. *Trailliella tetrasporangial phase of Bonnemaisonia hamifera Epiphytic on various small algae, intertidally in pools, rare. Streymoy: Hoyvik, Eibi. Presumably a relatively recent introduction to the islands, as it appears to be very localized. Known from the Shetland Isles since 1949 (personal record), where it is now abundant and widely distributed. (!) O. S. CHRYSOPHYTA Vaucheria coronata Nordst. In muddy turf, upper edge of sheltered beach, Streymoy: near Hvalvik (det. T. Christensen). Also in mud-filled rock crevices, upper spray zone of moderately exposed shore, Streymoy: near Glyvursnes (B0rgesen). (!) O. PHAEOPHYTA * Aglaozonia phase of Cutleria multifida Epiphytic on stipes and haptera of Laminaria hyperborea, subtidal to 10 m. Locally common on Eysturoy: Skalafj0rbur near Strendur and Kumlavik. (!) S. THE FLORA 121 Alaria esculenta (L.) Grev. On rocks, lower intertidal to shallow subtidal, mainly on exposed shores but extending some distance into fjords. Abundant and widely distributed. In more sheltered localities the blade is broader and less tattered, approaching the growth-form attributed to A. pylaii. (!) O. S. I. A. pylaii (Bory) J. Agardh On rocks with moderate shelter, lower intertidal and shallow subtidal. Recorded sparsely from several of the southern islands, but seems doubtfully distinct from A esculenta. Looked for but not found with any certainty by our expedition. I. Ascophyllum nodosum (L.) Le Jol. On rocks, midtide level, abundant in sheltered localities, but also found where there was moderate exposure, sometimes even accompanying Alaria. Widespread. (!) O. S. I. Asperococcus fistulosus (Huds.) Hook. [A echinatus] Found by Simmons epiphytic on Corallina on Suburoy : Hvalbiarf j0rbur. Epiphytic on fucoids on inner face of harbour wall, Streymoy: Kollarfj0rbur Harbour, and at Hoyvik in a pool at low water. (!) O.S.I. *A. turneri (Sm.) Hook. On stones and shells, subtidal to 7 m in a sheltered fjord. Eysturoy: Skalafj0rbur, locally frequent. (!) O. S. Chorda filum (L.) Stackh. On pebbles and shells, shallow subtidal to 4 m in sheltered localities, especially near heads of fjords. Widespread and locally abundant. (!) O. S. I. Chordaria flagelliformis (Mull.) Agardh On stones and epiphytic, in lower intertidal pools and shallow subtidal to 10 m on sheltered to moderately exposed shores. Common and widespread. (!) O. S. I. Cladostephus spongiosus (Huds.) Agardh Found by B0rgesen only in lower intertidal rock pools on an exposed rock platform on Eysturoy: near Gjogv. O.S.I. Cutleria multifida (Sm.) Grev. See Aglaozonia. Desmarestia aculeata (L.) Lamouroux On rocks and stones, more rarely epiphytic, on exposed and sheltered coasts, subtidal to 20 m. Often in large detached masses on sand or mud in very sheltered localities. Widespread and abundant. (!) O. S. I. D. ligulata (Lightf.) Lamouroux On rocks and epiphytic on Laminaria stipes, rarely encountered attached, occasional as drift specimens. Our expedition found only one small tattered specimen, growing attached in the shallow subtidal. Recorded from Eysturoy, Suburoy and Streymoy. Rex (1970) records it as occasional on the stipes of Laminaria hyperborea on Streymoy: Hoyvik. (!) O. S. I. D. viridis (Mull.) Lamouroux On stones and rocks, more rarely epiphytic, on exposed to moderately sheltered shores, subtidal to 20 m. Abundant and widespread. (!) O. S. I. Desmotrichum undulatum (J. Agardh) Reinke Found once by B0rgesen in shallow water along with Cystodonium purpureum at Streymoy: Sundini. S. *Dictyosiphon chordaria Aresch. A specimen possibly belonging to this species was collected in shallow non-tidal water on Eysturoy: at the head of Skalafj0rbur. (!) O. S. I. 122 SEAWEEDS OF THE FAROES D. ekmanii Aresch. [D. Ekmani] Epiphytic on Scytosiphon, shallow subtidal on sheltered shore, found only on Borooy: Klaksvik by B0rgesen. I. D. foeniculaceus (Huds.) Grev. (includes D. hippuroides) Epiphytic on Chordaria flagelliformis in pools and shallow subtidal on sheltered shores. Common, widely distributed. (!) O. S. I. Ectocarpus fasciculatus Harvey On rocks and epiphytic on various algae, intertidal and shallow subtidal, on exposed and sheltered shores. Common and widespread. (!) O. S. I. E. siliculosus (Dillwyn) Lyngbye [Includes E. confervoides, E. dasycarpus] On rocks and shells and epiphytic on various algae, intertidal and subtidal to 10 m, on exposed and sheltered shores. Common and widespread. (!) O. S. I. [Elachista flaccid a (Dillwyn) Aresch. Recorded by Lyngbye as rather rare on Fucus vesiculosus, but his herbarium specimens (fide B0rgesen) are E. fucicola. Recorded by Simmons on Himanthalia elongata near Torshavn (Streymoy) and on Fucus vesiculosus in Hvalbiarfj0rbur (Sufouroy). It seems unlikely that any of these are reliable records. O. S.j E. fucicola (Velley) Aresch. Epiphytic on intertidal fucoids on exposed and sheltered shores. Very common and wide- spread. (!) O. S. I. E. scutulata (Sm.) Aresch. On receptacles of Himanthalia. Common and widely distributed. (!) O. S. Eudesme virescens (Carmich. ex Harvey) J. Agardh [Castagnea virescens] Found by B0rgesen on stones in shallow subtidal in sheltered locality, Streymoy: between Hosvik and Hvalvik, and by us in sheltered lower intertidal pools and shallow subtidal on Eysturoy and Streymoy. (!) O. S. I. [Fucus ceranoides L. Recorded by Lyngbye, but no Faroese specimens are to be found in his herbarium (fide B0rgesen). No other phycologist has reported finding this species in the Faroes. O. S. I.] F. distichus L. subsp. distichus [F. inflatus f. linearis] On rocks in high level pools on exposed coasts. Found by B0rgesen only on Suouroy: near Famjin. S. F. distichus subsp. anceps (Harvey & N. Ward ex Carruth.) H. Powell [F. inflatus f. disticha] On rocks intertidally, often well above sea-level on exposed shores. Widely distributed. (!) O.S.I. F. distichus subsp. edentatus (Bach. Pyl.) H. Powell [F. inflatus f. edentata] On rocks, shallow subtidal, on sheltered shores, often with reduced salinity. Very variable in size and growth form. Abundant and widespread. (!) O. S. I. [F. serratus L. Reported by Landt as widespread at the base of cliffs, but no other investigator records finding this species, so the record would seem to be an error. O. S. L] F. spiralis L. On rocks, intertidally, on sheltered to exposed shores. With increased exposure plants become dwarfed (f. nana Kjellman), and can grow up to 5 m above high water mark (fide B0rgesen) and may also occur in high level pools. Abundant and widespread. (!) O. S. I. F. vesiculosus L. On rocks and stones at midtide level to shallow subtidal, on sheltered shores, often almost to the heads of fjords in brackish water. Very common and widespread. (!) O. S. I. THE FLORA 123 Giffordia granulosa (Sm.) Hamel [Ectocarpus granulosus] Epiphytic on Laminaria blades, subtidal at 4-6 m on sheltered shores. Viboy: near Hvanna- sund (J0nsson). Suburoy: Trongisvagur (Ostenfeld). Eysturoy: Eibi. (!) O. S. I. G. hincksiae (Harvey) Hamel [Ectocarpus Hincksiae] Epiphytic on larger algae, intertidal and subtidal to 20 m, typically on exposed coasts. Common and widespread. (!) O. S. I. Halidrys siliquosa (L.) Lyngbye Found attached in shallow subtidal (2 m) in Eysturoy: Skalafj0rbur near Glyvrar, and in drift in several localities, but would seem to be uncommon and very localized. Our expedition searched for it fruitlessly in the recorded sites. O. S. Halosiphon tomentosus (Lyngbye) Jaasund [Chorda tomentosa] On rocks, pebbles and shells, subtidal to 10-12 m, associated with Alaria, Laminaria spp. and other large seaweeds. Borboy: Anir. Eysturoy: Molin and Eibi. Streymoy: Sundini (Rex). I. Hecatonema foecundum (Stromf.) Lois. Forming dense mat on blade of Laminaria hyperborea; found by B0rgesen at Torshavn. I. Herponema velutinum (Grev.) J. Agardh [Ectocarpus velutinus] Reported by Simmons on Himanthalia in Hvalbiarfj0rbur (Suburoy), but not found by B0rgesen on two original Himanthalia specimens of Simmons which did, however, bear Elachista scutulata. S. Himanthalia elongata (L.) Gray On rocks at low tide level on exposed shores or in strong currents; on very exposed shores often well above low tide level. Common and widespread. (!) O. S. Isthmoplea sphaerophora (Carmich. ex Harvey) Kjellman Epiphytic on various intertidal algae on exposed shores. Common and widespread. (!) O. S. I. Laminaria digitata (Huds.) Lamouroux On rocks, intertidal to subtidal, on sheltered to exposed coasts. Common and widespread. (!) O.S.I. L. faeroensis B0rg. (=L. sacchdrina forma?) On stones, rocks, and shells in subtidal to 20 m, in very sheltered localities such as harbours and heads of fjords. Abundant and widespread. For taxonomic status see Kain (1976). (!) S. L. hyperborea (Gunnerus) Foslie On rocks, subtidal to 40 m, exposed to moderately sheltered shores, forming dense forests. Widespread and abundant. (!) O. S. I. L. saccharina (L.) Lamouroux On rocks and stones or epiphytic on large algae, in pools in lower intertidal and in subtidal to 20 m; on exposed to sheltered coasts. Common and widespread. (!) O. S. I. Laminariocolax tomentosoides (Farlow) Kylin [Ectocarpus tomentosoides] Partly endophytic in stipes and blades of Laminaria spp., forming short matted growth. Common and fairly widespread. O. S. Leathesia difformis (L.) Aresch. On rocks or epiphytic on Corallina, intertidal, mainly in sheltered situations. Common and widespread. (!) O. S. I. Leptonematella fasciculata (Reinke) P. C. Silva [Leptonema fasciculata] Found by B0rgesen on Eysturoy: Glyvrar and Fuglafj0rbur. O.S.I. Litosiphon filiformis (Reinke) Batters [Pogotrichum filiforme] Epiphytic on blades of Laminaria saccharina, often forming dense mats. Found by B0rgesen on Streymoy: Torshavn and Vagar: Mibvagur, and by us on Eysturoy: Skalafj0rbur. (!) S. I. 124 SEAWEEDS OF THE FAROES L. laminariae (Lyngbye) Harvey Epiphytic on Alaria blades, often forming a dense mat. Frequent, widely distributed. (!) O. S. Microspongium globosum Reinke [Myrionema globosum] Epiphytic on Himanthalia and other large algae, forming small cushions or mats, intertidal and shallow subtidal. Found by B0rgesen on Streymoy and Suburoy. O. Mikrosyphar polysiphoniae Kuck. Endophytic in Polysiphonia urceolata and Callithamnion hookeri, intertidal and shallow subtidal. Found by B0rgesen on Eysturoy: Oyri and Streymoy: Hoyvik and Kvivik. S.I. M. zosterae Kuck. Epiphytic on Zoster a marina, often forming rather large pseudoparenchymatous patches. Found by B0rgesen on Suburoy: Vagsfj0rbur. Myrionema aecidioides (Rosenv.) Sauvageau On blades of 'Laminaria faeroensis'. Found at Eysturoy: Skalafj0rbur, and Streymoy: Hvalvik by H. J0nsson. S.I. M. corunnae Sauvageau Forming short dense cushions on Laminaria digitata blades. Found by B0rgesen on Suburoy: Famjin. I. M. faeroense B0rg. On Palmaria palmata, in small dense mats, often with Microspongium globosum, on Streymoy: near Torshavn, the type locality. Not found or investigated since first described by B0rgesen. M. speciosum B0rg. On conceptacles of Himanthalia, forming short dense mats, often with Microspongium globosum, on exposed coasts on Suburoy: Vagseibi, the type locality. Not found or investi- gated since first described. M. strangulans Grev. [M. vulgar e\ Epiphytic on Monostroma fuscum, and various spp. of the Ulvaceae. Streymoy: between Hosviic and Hvalvik. Eysturoy: Skalafj0rbur near Strendur (!) and Kumlavik (!). (!) O. S. I. *Myriotrichia clavaeformis Harvey Epiphytic on Scytosiphon in lower intertidal on Streymoy. (!) O. S. Pelvetia canaliculata (L.) Dene & Thuret On rocks about high tide level in sheltered situations. Common but rather sporadic, often of minute size. (!) O. S. I. Petalonia fascia (Mull.) Kuntze [Phyllitis fascia] Intertidal on rocks and stones and epiphytic, often in pools. On exposed and sheltered shores. Common. (!) O. S. I. P. zosterifolia (Reinke) Kuntze [Phyllitis zosterifolia] On rocks intertidally on exposed to fairly sheltered coasts. Rather rare and local. I. Petroderma maculiforme (Wollny) Kuck. Forming small brown patches on smooth rock surfaces near high tide level. Streymoy: Saksun (B0rgesen). I. S. Phaeostroma parasiticum B0rg. Epiphytic on blade of 'Laminaria faeroensis' in shallow sheltered subtidal. Streymoy: between Hosvik and Hvalvik (B0rgesen). Pilayella littoralis (L.) Kjellman [Ectocarpus littoralis] On rocks and epiphytic, intertidal and shallow subtidal, growing in damp clefts to 2 m or more above high tide level. Abundant and widespread. (!) O. S. I. THE FLORA 125 Pleurocladia lacustris Braun [Pilinia.maritima] Found by B0rgesen on rocks at high tide level, intermingled with crustose blue-green algae, on Sandoy: near Sandur. Pseudolithoderma extensum (P. Crouan & H. Crouan) S. Lund [Lithoderma fatiscens] On stones and shells subtidally, 1-40 m, on exposed and sheltered shores. Fairly widely distributed. (!) S.I. Punctaria latifolia Grev. On rocks and epiphytic on large algae in the lower intertidal and shallow subtidal on sheltered coasts. Fairly widely distributed. (!) P. plantaginea (Roth) Grev. On stones in shallow subtidal in very sheltered situations. (!) O. S. I. Ralfsia clavata (Harvey) P. Crouan & H. Crouan On rocks about 3 m above high water mark on Suburoy: near Famjin; a small quantity of an alga presumed by B0rgesen to belong to this species. O.S.I. R. verrucosa (Aresch.) J. Agardh On rocks intertidally and in shallow subtidal, especially in pools, on sheltered and exposed coasts. Abundant and widely distributed. (!) O. S. I. [Saccorhiza polyschides (Lightf.) Batters Recorded from the Faroes by Landt, but not found by B0rgesen or other investigating phycologists. It is, however, common in the Shetland Isles. O. S.] Scytosiphon lomentaria (Lyngbye) Link [5. lomentarius] On rock intertidally and in shallow subtidal, exposed to sheltered shores, able to withstand low salinities. Abundant and widespread. (!) O. S. I. Sorapion kjellmanii (Wille) Rosenv. Epiphytic on Chaetomorpha melagonium. Found by B0rgesen on Streymoy: near Torshavn. Sphacelaria caespitula Lyngbye On stipes of Laminaria, rare. Eysturoy: Nes (Lyngbye). Streymoy: Kvivik (B0rgesen). I. S. cirrosa (Roth) Agardh [S. cirrhosa] Epiphytic on Desmarestia aculeata and Chaetomorpha melagonium on exposed and sheltered shores. On Svinoy and on Suburoy: Vagsfj0rbur (Jonsson). On Cladophora rupestris, in midtide pools, on Streymoy: Hoyvik (!) , Torshavn (!). (!) O. S. [I?]. S. nana Naeg. ex Kiitz. [S. britannica] On damp rocks around high tide level, especially in caves and clefts, forming a dark brown mat, on fairly exposed shores. Found by B0rgesen on Streymoy: Kvivik and Glyvursnes. O. S. [I?]. S. plumosa Lyngbye [Chaetopteris plumosa] On cobbles, subtidal 4-6 m, Eysturoy: Anir (!), Skalafj0rbur (!). Specimens in Rostrup's herbarium said to have been found on Faroes by Mr Randtopp of Torshavn (fide B0rgesen). (!) O.S.I. S. rigidula Kiitz. [5. furcigera] On a Laminaria stipe, subtidal, 6-8 m, on fairly exp9sed shore. Found by B0rgesen at Torshavn, del. as S. furcigera by Sauvageau, Eysturoy: Anir (!), Skalabotnur (!). (!) O. S. Spongonema tomentosum (Huds.) Kiitz. [Ectocarpus tomentosus] Epiphytic on the larger algae, intertidal on both exposed and sheltered coasts. Common and widespread. (!) O. S. I. *Stictyosiphon griffithsianus (Le Jolis) Holmes & Batters Epiphytic on Palmaria palmata, Streymoy: Kirkjub0ur. (!) S. 126 SEAWEEDS OF THE FAROES S. tortilis (Rupr.) Reinke On stones and epiphytic on Chordariaflagelliformis, in shallow subtidal on sheltered shores, often where salinity is reduced. Found by B0rgesen on Eysturoy and Suburoy, but probably much more widely distributed. O.S.I. Streblonema stilophorae (P. Crouan & H. Crouan) Hamel [Ectocarpus Stilophorae] Found by B0rgesen in the sorus of a Laminaria plant in Torshavn. I. Ulonema rhizophorum Foslie Epiphytic on Dumontia contorta at mid to lower shore levels. Esturoy: near the inner end of Skalafj0rbur. (!) O.S. Waerniella lucifuga (Kuck.) Kylin [Ectocarpus lucifugus] On rocks in cracks and fissures near high tide level, intermixed with Rhizoclonium. Found at Viboy: Vibvik, and Borboy: Klaksvik, both, by J0nsson. Viboy (!).(!) S. CHLOROPHYTA Acrochaete repens Pringsh. Found in old fronds of Chorda filum by H. J0nsson on Eysturoy: Skalafj0rbur. Streymoy: Hvalvik. I. Blidingia minima (Naeg. ex Kiitz.) Kylin [Enteromorpha intestinalis var. minima] On exposed shores in spray zone. Abundant and widespread. Eysturoy: Gjogv (det. Burrows) (!).(!) O.S. I. Bolbocoleon piliferum Pringsh. Found by B0rgesen between cortical cells of Petalonia fascia and Scytosiphon lomentaria on Mykines and Streymoy: Sundini. O. S. I. Bryopsis plumosa (Huds.) Agardh On rocks in deep pools and in shallow subtidal. Widely distributed, but rare and sporadic (!). Svinoy: Svinoyareibi (!). Streymoy: Torshavn to Hoyvik (!).(!)O.S.I. Capsosiphon fulvescens (Agardh) Setch. & N. Gardner Amongst vascular plants at high tide level at the head of a fjord. Locally common. Eysturoy: Skalafjorbur. (!)O. S. I. Chaetomorpha capillaris (Kiitz.) B0rg. [C. tortuosa] On rocks or loose lying or entangled amongst other algae, in pools at high tide level. In sheltered habitats. Widely distributed (!). Streymoy: Hedfjordur, Kirkjub0ur, Argir (det. Burrows) (!). (!) O. S. [I?] C. melagonium (F. Weber & Mohr) Kiitz. On rocks and epiphytic, in intertidal pools and subtidal, exposed to sheltered shores. Very common (!). Streymoy: Argir (!). (!) O. S. I. *Chlorochytrium cohnii E. Wright Eysturoy: Skalafj0rbur, subtidal in 'Schizonema' tubes (det. Burrows) (!). (!) O. S. I. C. inclusum Kjellman non-specific phase in life-history of Spongomorpha spp. Endophytic in various red algae. Widely distributed. (!) O. S. I. [Cladophora fracta (Mull, ex Vahl) Kiitz. Recorded by Rostrup as a common Faroese species, but B0rgesen considered this a case of misidentifica- tion, and no reliable herbarium material of this species from the Faroes exists. It was not found by either B0rgesen or us. S.] C. rupestris (L.) Kiitz. On rocks and stones intertidally in pools and in shallow subtidal, on exposed and sheltered coasts. Abundant and widespread (!). Streymoy: Argir (det. Burrows) (!). Eysturoy: Skala- fj0rbur (det. Burrows) (!). (!) O. S. I. THE FLORA 127 C. sericea (Huds.) Kiitz. [C. gracilis] In midtide to high tide level rock pools on moderately exposed to sheltered shores. Frequent and widespread. Eysturoy: Skalafj0rbur, Raktangi (del. Burrows) (!), Strendur. Streymoy: Argir (det. Burrows) (!). Viboy: Hvannasund (det. Burrows) (!). (!) O. S. I. Codiolum gregarium phase in life-history of Ulotrichales spp. On rocks near high tide level on sheltered and exposed coasts. Frequent and widespread. I. C. pu sill u m phase in life-history of Ulotrichales spp. Found by Lyngbye on rocks at high tide level on Suouroy: Hvalba. Eysturoy: Elduvik. I. Derbesia marina (Lyngbye) Solier On rocks, Corallina, Laminaria stipes and worm-tubes, subtidally from extreme low tide level to 20 m. Not uncommon, but easily overlooked. See Halicystis. (!) S. I. Enteromorpha clathrata (Roth) Grev. In sheltered shallow subtidal locations, often free-floating in large tangled masses. Locally common. O. S. I. E. compressa (L.) Grev. [E. intestinalis var. compressa} On rocks or epiphytic, intertidal, often in pools. Common and generally distributed. (!) O.S.I. E. intestinalis (L.) Link On rocks and epiphytic, intertidal and above, and in shallow subtidal, on exposed to fully sheltered shores, from fully saline to virtually freshwater habitats. Abundant and widespread. (!). Streymoy: Argir (det. Burrows) (!), Kaldbaksfj0rbur (det. Burrows) (!). Eysturoy: Raktangi (det. Burrows) (!). (!) O. S. I. E. linza (L.) J. Agardh On rocks and stones on open shores near low tide level, in pools and in shallow subtidal. Common and widespread (!). Viboy: Hvannasund, near causeway (!). Eysturoy: Skala- fj0rbur, Raktangi (det. Burrows) (!), Strendur. (!) O. S. I. E. prolifera (Mull.) J. Agardh [E. intestinalis var. prolifera] On rocks or detached, intertidal in pools and subtidal, in sheltered localities such as the heads of fjords, often growing in virtually freshwater. Locally common. Eysturoy: Gjogv (det. Burrows) (!). (!) O. S. I. *E. torta (Mert.) Reinb. In shallow subtidal, 3-4 m, on holdfast of ' Laminaria faeroensis' ', with Rhizoclonium ripar- ium, Erythrotrichia carnea, andPolysiphoniaurceolata. Eysturoy: Skalafj0rbur, Strendur. (!) O. S. Gomontia polyrhiza (Lagerh.) Bornet & Flah. Endophytic in mollusc shells, subtidal to 30 m. Common and widespread. I. Halicystis ovalis haploid phase of Derbesia marina. [Valonia ovalis] On rocks and shells, extreme low tide level and subtidal. Found at various sites by Lyngbye. Sandoy: Tr011konufinger (!). S. Monostroma fuscum (Postels & Rupr.) Wittr. On rocks and stones and epiphytic, in runnels and subtidally to 20 m. Common and widespread (!). Streymoy: Argir (det. Burrows) (!). (!) O. S. I. M. grevillei (Thuret) Wittr. Epiphytic, especially on Corallina, or on rocks, in shallow pools on sheltered to moderately exposed shores. Common and widespread. (!) O. S. I. M. undulatum Wittr. Epiphytic, especially on Corallina, or on rocks, lower intertidal and shallow subtidal. Widely distributed. O. I. 128 SEAWEEDS OF THE FAROES Ostreobium queckettii Bornet & Flah. Endophytic in mollusc shells in subtidal to 50 m. Common and widely distributed. O. S. Percursaria percursa (Agardh) Rosenv. On flat sheltered shores at high tide level, with Vaucheria. Found by B0rgesen on Streymoy: Sundini. O. S. I. Phaeophila wittrockii (Wille) R. Nielsen [Endoderma Wittrockii] In cell walls of various brown algae. Widely distributed. S. I. Prasiola crispa subsp. marina B0rg. On steep rocks on exposed coasts. Common. S.I. P. crispa subsp. terrestris (Roth) B0rg. On rocks and shady soil just above high tide level in the spray zone. P. furfuracea (Mert.) Menegh. On rocks in surge zone of exposed coasts. Found by B0rgesen on Streymoy: Tinganes. I. P. stipitata Suhr On rocks and stones at high tide level and above, especially with high nitrate levels from bird droppings, etc. Abundant and widespread. (!) O. S. I. Pringsheimiella scutata (Reinke) Marchew. Found by B0rgesen epiphytic on various algae and on Zostera marina, intertidally and in shallow subtidal, on exposed and sheltered shores. Widely distributed. O.S.I. Pseudopringsheimia confluens (Rosenv.) Wille [Ulvella confluens] Found by B0rgesen epiphytic on Gigartina intertidally on exposed shores on Streymoy: Velbastabur. P. fucicola (Rosenv.) Wille [Ulvella fucicola] Found by B0rgesen on Fucus distichus on Suburoy: Tv0royri. S.I. Rhizoclonium riparium (Roth) Harvey On rocks near high tide level and in the spray zone, forming large thick mats, often associated with percolating freshwater. Common and widely distributed. Eysturoy: Skalafj0rbur, subtidal (del. Burrows) (!). (!) O. S. I. Spongomorpha aeruginosa (L.) Hoek [S. lanosa] On Cladophora rupestris in intertidal pools and shallow subtidal. Found sparsely by B0rgesen on Streymoy: between Hosvik and Hvalvik (!) and on Eysturoy: Kumlavik (!), Raktangi (det. Burrows) (!).(!) O.S.I. S. arcta (Dillwyn) Kiitz. [Acrosiphonia albescens. Probably includes A. binderi, A. flagellata, and A incurva] On rocks and stones and epiphytic, near low tide level and in shallow subtidal, forming dense mats, on exposed and sheltered shores, or occurring in sheltered areas as large detached masses. Common and widely distributed. Eysturoy: Raktangi (det. Burrows) (!), head of Skalafj0rbur (det. Burrows). Streymoy: Argir (det. Burrows) (!). (!) O. S. I. *S. sonderi Kiitz. Eysturoy: head of Kaldbaksfj0rbur (det. Burrows). (!) S. Ulothrix consociata Wille On stones, intertidal, in sheltered localities. Found by H. J0nsson on Suburoy: Trongisvagur. S.I. U. flacca (Dillwyn) Thuret On rocks and epiphytic on the larger algae, intertidal, on exposed and sheltered shores. Common. O. S. I. THE FLORA 129 U. pseudoflacca Wille Epiphytic on various algae, intertidal, on exposed and sheltered shores. Common. S. I. Ulotrichales spp. See Codiolum gregarium and C. pusillum. Ulva lactuca L. On rocks and stones, and epiphytic on the larger algae, lower intertidal and shallow subtidal to 20 m, on sheltered and exposed shores. Common. Streymoy: Hoyvik (!), Torshavn (!). Eysturoy: Raktangi (det. Burrows) (!). (!) O. S. U. rigida (Agardh) Thuret In rock pools and shallow subtidal. Eysturoy: Skalafj0rbur (det. Burrows) (!), Raktangi (det. Burrows) (!).(!) O.S.I. Urospora mirabilis Aresch. On rocks and stones near high tide level on exposed coasts. Common and widespread. *U. penicilliformis (Roth) Aresch. On exposed intertidal rocks, Streymoy: Argir. (!) S. I. U. wormskioldii (Mert.) Rosenv. On rocks about high tide level on exposed to sheltered shores. Found by B0rgesen at various sites on Eysturoy and Streymoy. (!) I. 4. Conclusions This investigation covered only a limited period of the year, so doubtless missed many of the seasonally ephemeral species. Attention was concentrated on macroscopic species, and clearly there is considerable scope for a thorough investigation of, for instance, the microscopic brown algae, particularly the species described by B0rgesen from the Faroes. The crustose corallines, a particularly difficult group to study, require more consideration than we could give them, and the inner parts of the fjords and the two large southern islands, Sandoy and Suburoy, would almost certainly repay concentrated attention. Nevertheless, many interesting conclusions can be drawn from this study. Firstly, it is clear that B0rgesen's investigations were very thorough, that he missed or misinterpreted little, and that his analyses of the floristic units are sound. Secondly, the marine flora has changed little since his day, with few introductions of any significance. This is presumably due to the unfavourable current pattern illustrated in Fig. 2, which gives little help in bringing viable algal propagules from elsewhere, though it is perhaps strange that the flourishing Faroese shipping communications have not helped to introduce new species (except, perhaps, the Trailliella- phase of Bonnemaisonia hamifera). Thirdly, not only is the seaweed flora considerably poorer than that of the neighbouring Shetland Isles, but the cut-off is in many cases very abrupt. Several species which are abundant and widespread in the Shetland Isles have either not been recorded from the Faroes at all or are apparently rare and local. These include the following: Apoglossum, ruscifolium (Turner) J. Agardh Gelidium pusillum (Stackh.) Le Jolis Grifftthsia corallinoides (L.) Batters Halarachnion ligulatum (Woodw.) Kiitz. Hypoglossum woodwardii Kiitz. Laurencia pinnatifida (Huds.) Lamouroux Polyneura gmelinii (Lamouroux) Kylin Rhodophyllis divaricata (Stackh.) Papenf. Scinaia forcellata Biv. Desmarestia ligulata (Lightf.) Lamouroux 130 SEAWEEDS OF THE FAROES Fucus serratus L. Halidrys siliquosa (L.) Lyngbye Saccorhiza polyschides (Lightf.) Batters Alien species which have been firmly established in the Shetland Isles for several decades at least, yet so far are unrecorded for the Faroes, include Colpomenia peregrina Sauvageau and Codium fragile subsp. atlanticum (A. Cotton) P. C. Silva. Altogether nine Cyanophyta, 95 Rhodophyta, one Chrysophyta, 74 Phaeophyta and 44 Chlorophyta are recorded for the Faroes, making 223 species. This compares with 301 species reported from the neighbouring Shetland Isles (Irvine, 1980). A study of past records indicates that some species fluctuated considerably in their distribu- tion and populations: it may be that these species are indeed on the outer fringes of their range, and periodically suffer a catastrophic decline, recovering perhaps very slowly indeed, due to the lack of readily available sources of replenishing propagules. Laurencia pinnatifida, Desmarestia ligulata, Halidrys siliquosa, and perhaps Dilsea carnosa, may belong to this category. More long term studies of the marine algae of these isolated islands would be highly desirable. 5. Acknowledgements The descriptions of distribution features of the marine algae rely heavily on those of B0rgesen (1903) which, in general, could scarcely be improved. I am deeply indebted to Dr Elsie Burrows for the identification of many specimens of the Chlorophyceae. I am especially grateful to the Academia Faeroensis for the use of two laboratories and other facilities in Torshavn. The expedition itself would not have been possible without the financial assistance of the North Atlantic Treaty Organization (our main sponsors), the British Petroleum Company Limited, and the Carlsberg Foundation, as well as our own supporting institutions. I owe a personal debt of gratitude to the various other members of the expedition and associated personnel who contributed so much of the information incorporated in this paper and assisted in the collection of the material on which it is based. These are: Dr and Mrs P. E. Asen, Kristiansand Museum, Norway; Mr G. E. Asen (diver); Dr and Mrs W. F. Farnham, Portsmouth Polytechnic; Dr P. W. G. Gray, Portsmouth Polytechnic; Professor D. F. Kapraun, University of North Carolina; Dr K. Liming, Biologische Anstalt, Helgoland; Mr B. E. Picton, Ulster Museum, Belfast; Mr J. H. Price, British Museum (Natural History), London; Mr G. Ridley (photographer/diver); Dr and Mrs J. Rueness, Institut for Marinbiologi og Limnologi, Oslo; and Mr I. Tittley, British Museum (Natural History), London. I am also grateful to Mr G. Holt and Mr B. Rex for the opportunity to incorporate material from their unpublished manuscripts on the marine botany of the Faroes. 6. References B0rgesen, F. [C.E.] 1902. Marine algae. In [E. Warming (Ed.)], Botany of the Faroes based upon Danish investigations. Part II: 339-532. Copenhagen. - 1905. The algae-vegetation of the Faeroese coasts with remarks on the phyto-geography. In [E. Warming (Ed.)], Botany of the Faroes based upon Danish investigations. Part III: 683-834. Copenhagen and Christiania. Caram, B. & Jonsson, S. 1972. Nouvel inventaire des algues marines de 1'Islande. Acta hot. isl. 1: 5-31. Dixon, P. S. & Price, J. H. 1981 . The genus Callithamnion (Rhodophyta: Ceramiaceae) in the British Isles. Bull. Br. Mus. Nat. Hist. (Bot.) 9(2): 99-141. Holm, P. A. 1855. Skildringer af Naturen paa Faeroerne. Tidsskr. f. populaere Fremst. af Naturvidensk. Copenhagen. Holt, G. 1975. Marinbotaniske notaterfra Faer0yene 1975. [Unpublished report.] Irvine, D. E. G. 1974. The marine vegetation of the Shetland Isles. In R. Goodier, The natural environment of the Shetland Isles: 107-113. Edinburgh. - 1980. Marine algae. In R. J. Berry & J. L. Johnston, The natural history of Shetland: 267-272. London. Kain, J. M. 1976. New and interesting marine algae from the Shetland Isles. II. Hollow and solid stiped Laminaria (Simplices). Br. phycol. J. 11: 83-92. THE FLORA 131 Landt, J. 1810. Forsog til en Beskrivelse afFceroerne. Copenhagen. Lyngbye, H. C. 1819. Tentamen hydrophytologiae Danicae. , . . Hafnia. Parke, M. & Dixon, P. S. 1976. Check-list of British marine algae third revision. /. mar. biol. Ass. U.K. 56: 527-594. Rex, B. 1970. Rapport over algobservationer Faroarna sommaren 1970. [Unpublished report.] Rostrup, E. 1870. Faeroernes flora, . . . Bot. Tidsskr. 3: 5-112. Simmons, H. G. 1897. Zur Kenntnis der Meeres-algen Flora der Faroer. Hedwigia 36: 247-276. Seaweeds of the Faroes 2: Sheltered fjords and sounds Ian Tittley Department of Botany, British Museum (Natural History), London SW7 5BD William F. Farnham Marine Laboratory, Portsmouth Polytechnic, Hayling Island PO11 ODG Peter W. G. Gray Marine Laboratory, Portsmouth Polytechnic, Hayling Island PO11 ODG* Contents Synopsis 133 1 Introduction 133 2 Areas of investigation . 3 Materials and methods . 4 Sites investigated 134 135 136 5 Results 137 6 Discussion .... 145 7 Conclusions ..... 150 8 Acknowledgements 150 9 References . 150 Synopsis Some results of an international expedition to the Faroes during the summer of 1980 are included. The occurrence and distribution of benthic marine algae in sheltered fjords and sounds are described. Comparisons are made with the results of previous surveys undertaken by B0rgesen at the beginning of this century, and with the descriptions of vegetation presented recently for fjords in the Shetlands, Norway and Iceland. A reduced 'fjord effect' was seen in Faroese fjordic systems. The artificially divided Hvannasund did not show a 'fjord effect', but was an inlet of the sea. 1. Introduction The Faroes lie in the northern Atlantic Ocean between the Shetland Islands and Iceland. The island group comprises 17 inhabited and numerous small uninhabited islands and reefs. The main islands are separated by narrow sea-passages and the coastline is deeply indented with fjords. The present work was undertaken in July 1980 during a wider survey of the seaweeds of the Faroes (Irvine, 1982; Price & Farnham, 1982). The seaweed vegetation has been studied on only a few previous occasions. B0rgesen (1902, 1904, 1905) and B0rgesen & Jonsson (1905) undertook a comprehensive survey, which included both littoral and sublittoral habitats throughout the island group. More recent, but less detailed, studies which mention algae were "Present address: College of Agriculture, King Faisal University, PO Box 380, Saudi Arabia. Bull. Br. Mus. nat. Hist. (Bot.) 10 (3): 133-151 Issued 25 November 1982 134 SEAWEEDS OF THE FAROES Fig. 1 Maps of the Faroes (inset) and the Skalafj0rbur/Kaldbaksfj0ribur region (A on inset). Hvannasund is indicated B on inset. Numbers represent sites investigated (see text). Letters R = Raktangi; S = Storafles; E = Eystnes. undertaken by Rex (1970), Holt (1975), Johansen (1979), and Crothers (1981). B0rgesen's (1905) publication included ecological descriptions of the algal associations in sheltered waters. The main area selected for the present investigation was Skalafj0rbur, on the southern part of Eysturoy (Fig. 1). Other sites were visited for comparison in Kaldbaksfj0rbur (Streymoy), Hvannasund (Viboy) and Sundini, as well as the passage between Eysturoy and Streymoy (Fig. 1). 2. Areas of investigation Skalafj0rbur is a narrow, undivided sea inlet approximately 15 km in length and almost bisects the southern part of Eysturoy; this fjord opens to the Atlantic Ocean via Tangafj0rbur. The surrounding hills are drained by a number of small streams and rivulets, draining into the fjord. A larger river enters at the head-end. The foreshore surrounding Skalafj0rbur is narrow and formed of hard, basaltic rocks and boulders. SHELTERED FJORDS AND SOUNDS 135 Maximum depths of 72 m have been charted for the region of the fjord near Strendur and Glyvrar; elsewhere, in the middle regions, depths vary between 50 m and 60 m. At the head end and at the entrance to the fjord the maximum depths are less than 20 m. In many places the sea shore shelves abruptly into the shallow sublittoral zone and thereafter slopes more gradually. The slope of the sea bed varies little throughout the fjord. At most sites the sea bed consists of bedrock or boulders in the shallow sublittoral*; at greater depths there is an area of small boulders stones and shells which in the deepest waters merges into an extensive area of fine silt. The Admiralty Tide Tables record no appreciable tides for Skalaf j0rbur; during our periods of field work, however, a rise and fall of between 0-5-1 m was observed at the seaward end of the fjord. At the landward end, the tidal effect was negligible, although a fall of 0-4-0-5 m was noted on the harbour wall at Skalabotnur. Hydrological data for Skalafj0rbur are sparse, but B0rgesen (1905) gave a general account of the marine physical environment for the Faroes. This included measurements of surface salinities taken at the head-end near the mouth of the fjord, which showed no significant reductions; temporary, local reductions presumably occur following periods of heavy rain. The surface water temperature (July 1980) was 12C and this parameter may also vary during strong run-off. As B0rgesen (1905) indicated, the temperature of the open sea varies little throughout the year. Kaldbaksfj0rbur forms an indentation to the eastern side of Streymoy and opens to the Atlantic Ocean via Tangaf J0rbur; it lies almost opposite the entrance to Skalaf J0rbur. This fjord is smaller (approximately 7 km in length) than Skalaf J0rbur, and maximum depths of 60 m have been charted near its entrance; maximum depth decreases to 30 m near the head-end. The latter comprises an extensive foreshore of shingle and mud, traversed by a river and estuary. This foreshore extends for a considerable distance under shallow standing water of less than 0-5 m; it then shelves abruptly to the deeper sublittoral. Kaldbaksfj0rbur is otherwise fringed by a narrow foreshore of rock or boulders. Although the Admiralty Tide Tables again state there to be no appreciable tidal movement, a range of approximately 1 m was observed at the entrance of the fjord; by two-thirds of the way into Kaldbaksfj0rbur the range had decreased to a few centimetres, and at the head-end there was little tidal movement at all. Although rivers and streams enter, the waters were fully saline throughout. Presumably, local reductions in salinity at the head-end follow periods of rainfall. Hvannasund is a narrow channel separating the islands of Viboy and Borboy. The channel has recently been blocked at its narrowest point by a causeway constructed of large boulders. This man-made isthmus has artificially divided Hvannasund into two fjord-like arms; the northern arm was selected for investigation. The sound is surrounded by high hills and is fringed by a narrow, steeply sloping, and often almost vertical, rocky foreshore which continues into the sublittoral. Maximum depths of 80 m have been charted for the outer reaches of Hvannasund. At the inner end, the foreshore is more gently sloping and maximum depths are less than 15 m. The waters of the sound remain saline throughout. Sundini is the narrow sea passage (sound) which separates Streymoy from Eysturoy. 3. Materials and methods The intertidal and sublittoral vegetation was recorded by direct observations at eleven sites in Skalafj0rbur, four sites in Kaldbaksfj0rbur, two sites in Hvannasund and one area in Sundini. At intertidal sites species were listed, and the shore level at which they occurred, recorded. Subtidal bands of vegetation were identified by eye during scuba diving, samples of algae and substratum being taken from each band and brought back to the laboratory for checking. Vertical extents of sublittoral bands were recorded by depth gauge. * Shore zone terminology follows Lewis (1964). 136 SEAWEEDS OF THE FAROES 4. Sites investigated Skalafj0rfmr 1. Nes near Toftir Situated just out of the fjord on the eastern side of Eysturoy where conditions are fairly exposed. The intertidal comprised a narrow foreshore of rocks and boulders with occasional pools. A concrete quay was built over part of the foreshore. The foreshore of bedrock sloped into the sublittoral and was overlain at a depth of 3 m by a substrate of boulders and stones. At 10 m the substrate was of boulders, gravel, and horse-mussels (Modiolus modiolus L.); at 15 m there were fewer Modiolus and at 16 m (100 m offshore) the substrate was mainly gravel and mud. 2. Kumlavik Located on the western side of the fjord near its confluence with Tangafj0ribur. The intertidal comprised a small rocky headland which continued into the sublittoral to a distance of 80 m offshore (10 m depth). At greater depths the sea bed was covered by gravel and sand. 3. Saltnes Situated on the eastern shore of the fjord 2 km inland of the entrance. At this point the fjord was less than 1 km wide. A narrow rocky foreshore gave way in the shallow sublittoral to an extensive area of gravel and shells. At a distance of 50 m offshore (10 m depth) the sea bed was formed of mud and silt. 4. Strendur Situated on the western side of the narrows in the fjord, opposite site 3. The narrow rocky foreshore sloped into the sublittoral to a depth of 3 m. At greater depths the sea bed comprised gravel and sand deposits. 5. Glyvrar Situated on the eastern shore approximately 5 km inland of the entrance to the fjord. The intertidal region consisted of concrete quaysides and slipways; elsewhere the narrow rocky foreshore comprised rock outcrops and boulders. At depths greater than 2 m the sea bed was formed of gravel and sand; beyond 12 m the sea bed was of mud and silt. 6. Lambarefoi Located on the eastern side of the fjord approximately 6 km inland of the entrance. A narrow rocky foreshore sloped into the sublittoral to 2 m (10 m offshore) at which depth it became covered by gravel deposits. At greater depths (5 m-10 m; 30 m-50 m offshore), the sea bed is formed of stones wedged in mud. 7. Gotuefoi Situated on the eastern side of the fjord almost opposite site 8. The narrow rocky foreshore sloped into the sublittoral and merged at 2 m depth into an extensive area of small boulders and shingle. Below 12 m (200 m offshore) the sea floor was of mud and occasional stones. 8. Skdli Situated almost midway along the fjord on the western side. The narrow rocky foreshore extended only to low water level. From low water level to depth of 7 m (50 m offshore) the sea bed was formed of stones wedged in mud. At greater depths the sea bed was almost entirely of mud deposits. 9. Anir Located on the eastern side of the fjord approximately 2 km seawards of the head-end. The narrow rocky foreshore sloped into the sublittoral, and at a depth of 1 m was covered by stones in mud. This substrate extended 40 m offshore where at 10 m depth it was replaced by silt. 10. Skdlabotnur Situated on the western side of the fjord, approximately 0-5 km from the head-on. The foreshore comprised fringing rocks and boulders; elsewhere the fjord formed a shallow 0-3-0-5 m deep lagoon which contained occasional rocks and boulders wedged in sand and gravel. A small harbour- wall constructed of large boulders was also investigated. The wall sloped steeply to a depth of 6 m; between 6-8 m the sea bed was formed of shingle deposits, and below 8 m mud formed the main substrate. 11. Fjafoard estuary The head-end of the fjord was formed of shingle deposits which were traversed by the River Fjarbara. Stones in the brackish reaches of the estuary provide a substrate for algal attachment. SHELTERED FJORDS AND SOUNDS 137 Kaldbaksfj0rftur 12. Hvitanes Located on the southern shore at the entrance to the fjord. The site comprised a small sheltered bay containing rocks and boulders, surrounded by sheer cliffs. The foreshore rock continued into the sublittoral. At a depth of 22 m (approximately 50 m offshore) the sea floor was bedrock with occasional boulders. 13. Sund Located on the southern shore almost mid-way along the fjord. The narrow, steeply sloping foreshore continued into the sublittoral to a depth of 7 m; at greater depths the sea bed was of small boulders. 14. Two-thirds of the way along the fjord The narrow, steeply sloping, foreshore rock continued into the sublittoral to a depth of 2 m. At greater depths (down to at least 17 m) the sea floor was formed of stones and boulders wedged in sand. 75. Kaldbaksbotnur The head of the fjord consisted of a shingle beach and sand flats, traversed by a river estuary. Occasional stones and boulders were partially immersed in an extensive area of shallow, 0-3-0-5 m deep, standing water. The sublittoral was not investigated. Hvannasund 16. Harbour by the causeway The wide foreshore rock sloped gently into the sublittoral; the harbour and causeway were constructed of large boulders and sloped more steeply. Adjacent to the harbour wall was a lagoon containing rocks and boulders immersed in the shallow 0-1 m deep standing water. The rocks of the harbour wall shelved to a depth of 2 m below low water level. At greater depths the sea floor was of gravel sand and mud. The centre of the sound reached a maximum depth of 8 m and the bed was of sand and mud. 17. Leiti Situated approximately midway along the eastern side of the sound and formed of a narrow rocky foreshore with a steep slope into the sublittoral; much of the foreshore was covered by dense growths of barnacles and there were occasional pools. The bedrock continued into the sublittoral to 10 m; thence downwards the sea bed was formed of gravel, shells and Modiolus. Extensive beds of Modiolus were a prominent feature at 14 m. Sundini 18. Nofoskdli; Langasandur Sundini is a narrow channel which separates the two large central islands of the northern Faroes. A road bridge crosses the sound at its narrowest point where it is 225 m wide. A narrow rocky foreshore is present on both sides of the sound and the rock continued to 4 m depth in the shallow sublittoral. At the bridge the sound reached a maximum depth of 7 m and the sea floor was formed of bedrock with a deposit of stones and boulders. A short distance to the north of the bridge at Langasandur a depth of 20 m was reached; the sea floor consisted of stones and shells held firm in mud deposits. 5. Results Eulittoral vegetation Band-forming species Table 1 indicates the principal band forming species of the eulittoral and the littoral fringe at 13 of the 17 sites investigated. Although Prasiola stipitata* was recorded from 5 sites it occurred as a recognizable band only at Kumlavik (site 2). Blidingia minima was found in both fjords and sounds and occurred throughout Skalaf J0rbur. At several sites B. minima formed a distinct band of vegetation near high tide level. Enteromorpha intestinalis was also present at most sites, but only formed a distinct band of vegetation on the narrow foreshore at Sund (Kaldbaksfj0rbur, site 13). Porphyra umbilicalis was found everywhere except at the head-ends of Skalaf j0rbur * Algal nomenclature follows Irvine (1982). 138 SEAWEEDS OF THE FAROES and Kaldbaksfj0rbur, where there was little firm substrate available for colonization. At six of the sites P. umbilicalis occurred as a distinct band. At site 14 (Kaldbaksfj0rbur) much of the narrow intertidal was colonized by P. umbilicalis. Table 1 Principal eulittoral band-forming species. Species/Site 1 2 4 5 6 10 11 12 13 14 15 16 17 Blidingia minima X X X X x X X x X Cladophora sericea X X X X Enteromorpha intestinalis X X X X X X X x X X Prasiola stipitata X X X X X Spongomorpha arcta X X X X Ulva lactuca X X X XXX X X X UlothrixlUrospora sp. X X X Alaria esculenta X X X X x X X Ascophyllum nodosum X X X x x X X Dictyosiphon foeniculaceus X X X x X Eudesme virescens X X X Fucus distichus anceps X F. distichus edentatus X X X X X x x X F. spiralis X X X X X X X X F. vesiculosus X X X X X X Himanthalia elongata X X X X X Laminaria digitata X X X X X X X X L. saccharina X x X x Pelvetia canaliculata X X Palmaria palmata X X X X X Porphyra umbilicalis X X X X X X x X X X X = Band-forming; x = present. In most fjords fucoids were an obvious component of the vegetation. The least common of these plants was Pelvetia canaliculata, detected at only two sites and present as a distinct band only at Hvannasund. Fucus spiralis was widespread in Skalaf J0rbur and Hvannasund but was not found in Kaldbaksfj0rbur. This species formed a distinct band on rocks just below high tide level. F. vesiculosus was less abundant than F. spiralis, and occurred as a distinct band at middle shore levels only at two sites in Skalaf J0rbur. Ascophyllum nodosum often grew together with F. vesiculosus and formed a distinct band at four sites; it was not detected in Kaldbaksfj0rbur. F. distichus subsp. anceps was only found on exposed rock faces at Hvitanes (site 12), at the entrance to Kaldbaksfj0rbur. F. distichus subsp. edentatus was widespread at lower shore levels in Skalafj0rbur, Hvannasund, and Sundini, where it formed a distinct band. In Kaldbaksfj0rbur a few plants grew on rocks and stones in shallow standing water at the head-end (site 15). At Sund (site 13), and on the harbour wall at Hvannasund (site 16), Palmaria palmata formed a band on rocks near low water level. This species was not detected in the inner regions of Skalafj0rbur and Kaldbaksf j0rbur. Himanthalia elongata was an obvious component of the vegetation on wave-washed lower shores at the entrances to the fjords; in Skalaf j0rbur the species was also found at a relatively sheltered locality, Glyvrar (site 4), approximately 4 km inland of the entrance to the fjord. SHELTERED FJORDS AND SOUNDS 139 Subflora (turf-forming) associations Table 2 lists the subflora or turf-forming species recorded during the survey. As Fucus was largely absent from sites in more exposed situations at the entrances to the fjords, the main algal vegetation comprised small turf-forming species which often grew on a dense cover of barnacles. Species such as Callithamnion sepositum, Ceramium shuttleworthianum, and Gigartina stellata were present as small tufts or as turf-like growths. A more detailed account of the algal vegetation of open sea conditions is presented in the following paper of this series (Price & Farnham, 1982). At less exposed sites a subflora of small species such as Ceramium diaphanum, Enteromorpha intestinalis, Gigartina stellata, and Viva lactuca grew beneath the cover of large brown algae. At Skalabotnur (site 10, near the head-end of Skalafj0rbur) the underflora solely comprised small plants of Ulva lactuca. In Kaldbaksfj0rbur, where fucoids were largely absent, the main vegetation of the eulittoral level comprised bands of Blidingia minima, Enteromorpha intestinalis, Porphyra umbilicalis, and Palmariapalmata. Lower littoral and sublittoral fringe vegetation The algae recorded in the lower eulittoral and sublittoral fringe levels are presented in Table 3. Species are listed separately for sites recorded in Skalafj0rbur and Kaldbaksfj0rbur. The vegetation at these shore levels was richer in species than that of the middle and upper eulittoral levels. Thirteen species were recorded only from sites in the outer areas of Skalaf j0rbur (sites 1-5); a further 13 species occurred throughout the fjord. Red algae such as Corallina officinalis, Ceramium rubrum, Dumontia contorta, Gigartina stellata, Polysiphonia brodiaei and Rho- domela confervoides, together with a few brown algae, such as Pilayella littoralis and Scyto- siphon lomentaria, were the most obvious components of the turf-forming vegetation at the more exposed entrance sites in Skalafj0rbur. At Hvitanes (site 12, Kaldbaksfj0rbur) these and other red algae such as Cryptopleura ramosa were recorded from lower shore pools. In the middle and inner regions of Skalaf j0rbur, and also at a few locally sheltered situations in the outer fjord, brown algae such as Chorda filum, Dictyosiphon foeniculaceus, Eudesme virescens, Myrionema strangulans (epiphytic on Ulva lactuca), Pilayella littoralis, Punctaria plantaginea and Scytosiphon lomentaria, together with bleached plants of Ceramium rubrum and Cystoclonium purpureum (red algae), were the most obvious components of the lower shore vegetation. A similar species assemblage was detected in the middle regions of Kaldbaksfj0rbur and in a sheltered lagoon near the head of Hvannasund. Our observations showed that very few algae were restricted to the inner regions of the fjords; Enteromorpha spp. were more common in these regions, although filiform and filamentous brown algae were the most obvious components of the vegetation; there were very few red algae. Near the head of Hvannasund, by contrast, red algae were the most obvious component of the lower eulittoral and shallow sublittoral vegetation. Himanthalia elongata H. elongata occurred at lower shore levels at sites 1,2, and 4 in the outer regions of Skalaf j0rbur. The species was particularly common where the shores were regularly washed by waves and swell. H. elongata was not found in the middle and inner regions of Skalafj0rbur. The species was also common on wave washed shores at Hvitanes (site 12) at the entrance to Kaldbaks- fj0rbur, but was not found at site 13 where conditions were more sheltered. H. elongata was not found at sites 16 and 17 in Hvannasund, although it was particularly common on the wave- washed foreshore at Vibareibi at the entrance to the sound. Alaria esculenta A. esculenta was an obvious component of the vegetation at the entrance to Skalaf j0rbur, where the species was widespread on rocks in both shallow sublittoral levels and deep pools. In contrast to Himanthalia elongata, Alaria esculenta was also found at sites located in the middle, sheltered, reaches of Skalafj0rbur. Its limit of distribution along Skalafj0rbur was found to be just to the west of S01dafj0rbur, 10 km into the fjord from site 1 at the entrance. At the former location J .g 1 ll.ll t-c "j3 ^ S j^^ "^ .^* ? 2J "^^* T3 'S <3 ^ Ci, 53 "Jjj u -a vj ? '1i sf 11 *- O ce "7^ ^ ^'S ^^ AJ ?* ^** fN sJ r\ U ~w ^3 ^ (3 c . ,^ ^ ^ fcj^ i* Vj N 1 * *^i * O o3 fe ^3^-2 v H,sc -SP^ t/3 Z ffl G c^fcUOUUQ^O^ s 1 1 " **c ~" ~-~ *"- CL 4s ^ " 03 ^r ^ ** GO * C S sllll'i ^> o u c^-S^^'S ^Sb^Q^^^ *^ -^i Cy ^ ^i O " "^ "5? ^O nJ OQ ^J ^^ ^^ x5 ^ S ^.ca2j _^ 1^ "< O "* ** ^** r^ f '2^^cs:J ||| g ||||l||| Site 10 Skalabotnur Fucoid cover Ulva lactuca Site 13 Sund No fucoid cover No subflora beneath cover of: Enteromorpha intesti Palmaria palmata Porphyra umbilicalis SHELTERED FJORDS AND SOUNDS 141 Table 3 Occurrence and distribution of lower littoral and sublittoral fringe species in A. Skalaf j0rbur and B. Kaldbaksfj0rt)ur. Table 3A OUTER SITES Alaria esculenta 1,2,4,5 Chaetomorpha melagonium 2 Chordariaflagelliformis 1,2,4 Cladophora sericea 2 Corallina officinalis 1,2,4,5 Ectocarpusfasciculatus 1 , 5 Elachista scutulata 2 Himanthalia elongata 2, 5 Palmar ia palmata 1, 5 Petalonia fascia 1, 2 Polysiphonia brodiaei 1, 2 Porphyra leucosticta 1 , 2 Spongomorpha aeruginosa 2, 4 S. arctal,2,5 Table 3B Kaldbaksfj0rfcur OUTER SITES Alaria esculenta Callithamnion sepositum Ceramium rubrum Corallina officinalis Cryptopleura ramosa Delesseria sanguinea Dermatolithon spp. Gigartina stellata Himanthalia elongata Plocamium cartilagineum Polysiphonia brodiaei THROUGHOUT FJORD INNER SITES MIDDLE FJORD SITES oCeramium rubrum 1, 2, 5, 7, 9 Enteromorpha Chaetomorpha capillaris 8 linza 10 >Chondrus crispus 8, 9, 10 Hildenbrandia oChordafilum 2, 7, 8, 9, 10 rubra 10 oCladophora rupestris 2, 5, 7, 8 Porphyra < > Cystoclonium purpureum 2,8,9 purpurea 9 < > Dictyosiphon foeniculaceus Ralfsia 1,2,4,5,8,10 clavatalQ oDumontia contorta 1, 3, 7, 8, 9, 10 Ulonema Enteromorpha intestinalis 7, 9 rhizophora 9 Erythrotrichia cornea 1 Eudesme virescens 4, 8 <> Gigartina stellata 1, 5, 8, 9 <>Laminaria digitata 2, 5, 8, 9, 10 < L. saccharina 2, 4, 5, 8 Membranoptera alata 4, 8 <>Myrionema strangulans 2 >Phymatolithon Ienormandii4, 10 oPilayella littoralis 1, 5, 7 Plocamium cartilagineum 8 <>Polysiphonia nigrescens 2, 9 P. urceolata Punctaria plantaginea 4 Rhizoclonium riparium 1 oRhodomela lycopodioides 2,4,9 <>Ulvalactuca 2, 5,9,10 THROUGHOUT FJORD INNER SITES MIDDLE FJORD SITES Chorda filum Chondrus Dictyosiphon foeniculaceus crispus Ectocarpus siliculosus Enteromorpha Eudesme virescens intestinalis Laminaria. saccharina Pilayella Myrionema strangulans littoralis Myriotrichia clavaeformis Palmaria palmata Scytosiphon lomentaria Viva lactuca < = occurs also at outer sites > = occurs also at inner sites 142 SEAWEEDS OF THE FAROES occasional small Alaria plants (less than 0-5 m in length) were detected among Laminaria digitata and the other small brown algae. In Kaldbaksfj0rbur Alaria esculenta grew abundantly at Hvitanes (site 12), an exposed site at the entrance to the fjord, and was also found at Sund (site 13), 5 km further in than Hvitanes. Alaria occurred throughout Hvannasund, even in the most sheltered areas by the causeway (site 16), where it grew among Laminaria digitata and L. saccharina. At site 17, mid-way along Hvannasund, the species formed a discontinuous band of vegetation on rocks near low water level. At several sheltered locations Alaria esculenta was found growing side by side with Ascophyllum nodosuml Sublittoral vegetation Occurrence and distribution of Laminaria spp. The horizontal and vertical distributions of Laminaria spp. at three sites in Skalafj0rbur and Kaldbaksfj0rbur are illustrated schematically in Figs 2, 3, and 4. The type of substrate present at each site, and recorded depths, are also shown. Laminaria digitata L. digitata occurred throughout Skalafj0rbur as a narrow band in the sublittoral fringe and shallow sublittoral (cf. Fig. 3). It was not detected, although sought, on the exposed sea shore at Nes (site 1). At Skali (site 7) the species was probably overlooked; at Skalabotnur (site 10) only small plants less than 1 m in length were detected. In Kaldbaksfj0rbur L. digitata grew in the shallow sublittoral at sites 11 and 12, but not at site 13. The species occurred throughout Hvannasund. Laminaria hyperborea At Nes (site 1, entrance to Skalaf j0rbur, Fig. 2) large L. hyperborea plants were the dominant vegetation in the sublittoral and grew attached to the rocks and boulders to a depth of 10 m. The largest plants measured 2 m in length (stipe approx. 1 m; blade approx. 1 m) and grew at depths between 6-8 m. Below 10 m occasional L. hyperborea were detected on the few stones and boulders lying on the gravel sea floor. No L. hyperborea was found below a depth of 14 m, where the sea floor was entirely of small stones and gravel. Dense stands of L. hyperborea were also observed on bedrock and boulders at Raktangi headland, and at Storafles, which lie seawards of the sites investigated in Skalafj0rbur. Unlike L. digitata, L. hyperborea was not found in the inner reaches of Skalaf j0rbur between G0tueibi (site 8) and the Fjarbara estuary (site 11); it occurred at all other sites except Skali (site 7). At Kumlavik and Saltnes (sites 2 and 3) in the outer fjord, L. hyperborea occurred sporadically in the shallow sublittoral to a depth of 4 m. At these depths and below only one or two small plants were sampled from one square metre. At sites 4, 5, and 6 in Skalaf j0rbur (Fig. 3) L. hyperborea was present as a narrow band of vegetation in the shallow sublittoral below the band of L. digitata and was not found at depths greater than 3 m. At Lambareibi (site 6) plants measured approximately 1 m long but at Glyvrar (site 4) occasional plants with short stipes and wide blades were detected. At its inner limit of distribution in Skalafj0rbur, site 8, occasional L. hyperborea grew among L. faeroensis* in the shallow sublittoral at depths between 1-3 m. L. hyperborea was present at all sublittoral sites investigated in Kaldbaksfj0rbur, Hvanna- sund, and Sundini. At Hvitanes (site 12) the bedrock supported a dense stand of L. hyperborea at depths between 1-9 m. The maximum growth of L. hyperborea at this site occurred at a depth of 4 m where there were many large plants with stipes 1-5 m long; L. hyperborea was not found at depths below 13 m. At Sund (site 13) the species formed a band of vegetation between 4 m and 10 m below low water level; between 10 m and 15 m L. hyperborea grew together with L. saccharina, while plants collected at 15 m had short stipes and very broad laminae. This form of L. hyperborea was also detected at site 14, a more sheltered locality in the inner region of Kaldbaksfj0rbur, where it grew among L. faeroensis and L. saccharina on a substrate of small *For comments on the current taxonomic status of L. faeroensis B0rg. see Kain (1976). For convenience we have used this name. SHELTERED FJORDS AND SOUNDS 143 2 Gigartina stellata & Himanthalia elongata Alaria esculenta Laminaria hyperborea 10 _-]5 Boulders Rock L. hyperborea & L. saccharins L. saccharina Stones, shells Sand ,100 Fig. 2 Site 1: schematic representation of vegetation and transect profile; numbers on vertical axis indicate depth in metres, and on the horizontal axis indicate distance offshore. Laminaria digitata & L. hyperborea Fig. 3 Site 6: schematic representation of vegetation and transect profile; details as for Fig. 2. -2 Porphyra umbilicalis & Ulva lactuca Chorda filum, Laminaria saccharina & small brown algae L. hyperborea, L faeroensis L. hyperborea Cruoria 14 Rock Stones & boulders Fig. 4 Site 14: schematic representation of vegetation and transect profile; details as for Fig. 2. 144 SEAWEEDS OF THE FAROES boulders and stones (Fig. 4). A dense band of L. hyperborea occurred in the shallow sublittoral (2-4 m depth) at site 17 in Hvannasund. At greater depths the species grew together with L. saccharina; at 10-14 m the sea floor was covered with extensive Modiolus beds and these were widely colonized by L. hyperborea. At the head of Hvannasund (site 16) the form of L. hyperborea with broad fronds colonized stones and shells at depths between 4-8 m. Typical forms of the plant grew on the rocks and boulders of the harbour and causeway in the shallow sublittoral (2 m depth). Laminaria saccharina The typical form of L. saccharina was found at most sites in Skalafj0rbur, but was noticeably absent from sites 9 and 10 near the head of the fjord. L. saccharina grew among L. digitata and L. hyperborea in the shallow sublittoral regions (between 0-2 m). At sites 4 and 5 the species formed a narrow band of vegetation at slightly greater depths (3-4 m) between an upper band of L. hyperborea and a lower, extensive, area of L. faeroensis. At Nes (site 1, Fig. 2) L. saccharina and L. hyperborea formed a mixed community of plants inhabiting the boulder and gravel sea floor at depths between 10-14 m. L. saccharina grew on small stones, or even unattached, at depths between 14-16 m (approximately 100 m offshore) ; it did not grow on the muddy sea floor below 16 m. In Kaldbaksfj0rbur the species grew together with L. digitata and L. hyperborea in the shallow sublittoral (0-2 m). At Hvitanes (site 12) L. saccharina was present at greater depths (10 m and below) in open stands of L. hyperborea; mixed stands of the two laminarians were also seen at site 13 at depths between 2-8 m. At greater depths L. saccharina developed wider blades and longer stipes, and was not easily distinguishable from L. faeroensis. The typical form of L. saccharina was only found in shallow waters at the sheltered site 14 in Kaldbaksfj0rbur. It was found among L. digitata and L. hyperborea in both shallow and deep sublittoral regions in Hvannasund (sites 16 and 17). At site 16 (head end) plants with long stipes and broad laminae formed a mixed community with the broad form of L. hyperborea. Laminaria faeroensis The L. faeroensis form of L. saccharina occurred widely in the sheltered regions of fjords. In Skalafj0rbur this species was absent from the outer sites (1, 2, and 3), but was present everywhere else on unstable substrates in the deeper sublittoral (Fig. 3). At Glyvrar (site 5), for example, L. faeroensis not only colonized small stones on the muddy sea floor, but in some places appeared to be unattached. L. faeroensis was the dominant species between 4-12 m (approximately 100 m offshore). Plants were often large in size, and one specimen plant measured 2-7 m in length (stipe 1-2 m; lamina 1-5 m long, 0-6 mwide). At G0tueibi (site 8) beds of L. faeroensis spread to a distance of 200 m offshore. At sites 13 and 14 in Kaldbaksfj0rbur, it was difficult to distinguish L. saccharina from L. faeroensis; most L. saccharina had wide laminae and long, but solid, stipes. L. faeroensis was not detected in Hvannasund, but was widespread in Sundini. Subflora vegetation Three associations of algae were recognized in the subflora vegetation in Skalafjorbur, Kaldbaksfj0rbur, and Hvannasund. The constituent species of the subflora vegetation are listed in Table 2 and the sites at which they were recorded is given. An association, mainly of red algae such as Audouinella purpurea, Callophyllis (Euthora) cristata, Chaetomorpha melagonium, Epilithon membranaceum, Membranoptera alata, Pal- maria palmata, Phycodrys rubens, Polysiphonia urceolata, Ptilota plumosa, P. serrata and the Aglaozonia stage of Cutleria multifida was detected on the stipes of Laminaria hyperborea and on adjacent rocks and boulders. At most sites the number of species present decreased with increasing depth. This association occurred in the outer regions of fjords where L. hyperborea was more abundant; an impoverished form of the association, containing many fewer red algae, was detected at Lambareibi (site 6) in the middle regions of Skalafj0rbur. A distinct association of plants was recorded on or among Laminaria saccharina on unstable substrates in the outer fjord or on/among L. faeroensis in the middle and inner regions. SHELTERED FJORDS AND SOUNDS 145 Ceramium rubrum, Chorda filum, Desmarestia aculeata, D. viridis, and Viva lactuca colonized stones and boulders within the stand of L. saccharina, while small brown algae such as Ectocarpus fasciculatus and Litosiphon filiformis were common epiphytes on the laminae of L. saccharina. The crustose species Cruoria pellita, Lithothamnium glaciate, and Pseudolithoderma exten- sum were detected as a distinct association on rocks and boulders in deeper waters at the entrance to Skalafjorbur, and occurred sporadically in the middle and inner regions, where solid substrates were available (Fig. 4). Several species were found only at the inner end of Skalafj0rour (sites 9 and 10); these included Asperococcus turneri, Erythrotrichia carnea, Polysiphonia elongata, Sphacelaria plu- mosa, and the Trailliella phase of Bonnemaisonia hamifera. Several of these algae were growing on plastic bristle-like debris strewn over the sea floor. Antithamnion floccosum and Sphacelaria rigidula were also found on this substrate. . At most sites in Skalafj0rbur and Kaldbaksfj0rbur, subflora was sparse beneath the canopy of Laminaria faeroensis. In Hvannasund, by contrast, a subflora rich in both numbers of species and individual luxuriances was detected in the deeper sublittoral zone. Several species occurred here which were not recorded in Skalafj0rbur and Kaldbaksfj0rbur; these included Callocolax neglectus parasitic on Callophyllis laciniata, Fimbrifolium dichotomum, Lomentaria clavellosa, Phyl- lophora truncata, and Polyides rotundus. Unusual growth forms of several red algae were detected at the head end of Hvannasund (site 16). A crisp, more branched, form of Odonthalia dentata was common in the deep sublittoral, together with a narrow and more branched form of Callophyllis cristata and the lingulata form of Phycodrys rubens. Site 18. Sundini The Streymoy shore of Sundini near the road bridge was mainly of sand and gravel deposits uncolonized by algae; boulders standing in fast-flowing water supported large growths of Porphyra umbilicalis. Near high tide level the gravel beach was colonized by maritime flowering plants, and Vaucheria coronata was found on damp soil surrounding the plants. The narrow foreshore on the Eysturoy side of the sound carried a dense cover of fucoids; filiform and filamentous brown and green algae were detected in lower shore pools. Alaria esculenta and Laminaria hyperborea formed a distinct but narrow band of vegetation on bedrock in the shallow sublittoral. L. faeroensis grew attached to stones and boulders at depths between 2-5 m. Although Desmarestia aculeata grew among the L. faeroensis, almost no other subflora was found associated with the cover of the latter. Stones and boulders at depths below the L. faeroensis level were colonized by crustose algae such as Lithothamnium glaciale and Pseudolithoderma extensum. At Langasandur (2 km north of the road bridge) most of the narrow intertidal was covered by dense growths of Ascophyllum nodosum. A narrow band of Laminaria digitata was present in the shallow sublittoral, and the broad form of L. hyperborea grew on bedrock at depths between 1-4 m; a few epiphytes colonized the stipes of L. hyperborea. At greater depths (4-10 m) L. faeroensis was detected growing on a deposit of stones and boulders covering the sea floor; Desmarestia aculeata was occasionally present. Phycodrys rubens, Polysiphonia elongata, and crustose species formed an open community of plants and stones and shells between 10-18 m. No vegetation was detected below a depth of 20 m. 6. Discussion Comparison with B0rgesen's observations Our observations on the algae of the littoral fringe differ little from those made by B0rgesen (1905), who described an upper eulittoral 'formation of Chlorophyceae' which was composed of three bands of vegetation. The predominant species present in these bands were Prasiola stipitata, Blidingia minima and Enteromorpha intestinalis . B0rgesen also described a 'Fucaceae formation', which comprised five bands of algae; these 146 SEAWEEDS OF THE FAROES were Pelvetia canaliculata, Fucus spiralis, F. vesiculosus, Ascophyllum nodosum and F. distichus subsp. edentatus. We detected these bands of algae and also noticed that these fucoids were absent from exposed sites at the entrances to the fjords; there, F. distichus subsp. anceps was the dominant fucoid. Fucoids were recorded throughout Skalafj0rbur, and were particularly abundant at sites where a firm substrate prevailed; in Kaldbaksfj0rbur these algae were conspicuously absent from the inner regions where littoral rocks were colonized by dense growths of Enteromorpha intestinalis, Palmariapalmata, and Porphyra umbilicalis. The natural foreshore in this area has recently been covered by a steeply sloping boulder foreshore which supports a new road. Although A. nodosum was widespread in the sheltered parts of Skala- fj0rbur, it was also found in sheltered locations at very exposed sites. At both Hvitanes (site 12) and Kumlavik (site 2) the species grew together with Alaria esculenta. A subflora or turf-forming vegetation was detected at most sites; in exposed situations Callithamnion sepositum, C. hookeri, Ceramium shuttleworthianum, Corallina officinalis, and Gigartina stellata were particularly abundant; C. sepositum and C. shuttleworthianum did not occur in sheltered areas and the other species were less common. In the sheltered regions of Skalafj0rbur and Kaldbaksfjorbur species of red algae were a less obvious component of the vegetation of the eulittoral level, and at Skalabotnur (site 10) there were no red algae beneath the cover of Fucus spp. B0rgesen (1905) described a thick turf of Corallina officinalis from lower shore levels at exposed sites; he commented that this vegetation is replaced in sheltered areas by prolific growths of filamentous and filiform brown algae which he named a 'Stictyosiphon association'. Our observations revealed that this type of vegetation was widespread in the inner regions of Skalafj0rbur, Kaldbaksfj0rbur and in a sheltered lagoon near the head end of Hvannasund. Both B0rgesen's and our surveys recorded a few red algae (Ceramium rubrum, Cystoclonium purpureum, Dumontia contorta, and Rhodomela lycopodioides among the vegetation of filiform brown algae. We detected Devaleraea (Halosaccioh) ramentacea at the head end of Hvannasund where it grew in shallow standing water in pure stands; B0rgesen found the species in similar conditions near Klaksvik. Although we noted more green algae, particularly Enteromorpha spp., in the shallow sublittoral regions at the head ends of Skalafj0rbur and Kaldbaksfj0rbur, we did not locate the large growths of Monostroma fuscum noted by B0rgesen as characterizing a 'Monostroma- Enteromorpha Association'. B0rgesen indicated that this association occurred in shallow water at the heads of fjords with considerable inflow of freshwater, and considered it to be perennial. We too detected the loose-lying mats of Ceramium rubrum, Chorda filum, Enteromorpha spp., Rhizoclonium riparium, and Scytosiphon lomentaria in very shallow water at the head end of Skalafj0rbur, and additionally found Capsosiphon fulvescens, growing in shallow, brackish water in the estuaries at the heads of Skalafj0rbur and Kaldbaksfj0rbur. B0rgesen (1902) remarked that Alaria esculenta was widespread in the Faroes and, that while it showed a preference for open sea shores, it may also occur in the more sheltered interior of a fjord; our observations of some Alaria plants growing side by side with Ascophyllum confirms this and supports Russell's (1978) comments that 'abundance measures are a more reliable indicator of environmental conditions than presence data . . .'. We were unable to locate A. pylaii which B0rgesen (1905) recorded from sheltered waters in Skalafj0rbur. B0rgesen (1905) described the 'Halidrys Association' from the shallow infralittoral region at Glyvrar in Skalafj0rbur; apparently this was the only site from which the species was known in the Faroes. H. siliquosa grew among Laminaria spp. and supported a number of brown algal epiphytes such as Punctaria latifolia. We failed to find Halidrys at Glyvrar despite an intensive search. Laminaria spp. are the most obvious component of the vegetation of the sublittoral region in the fjords and sounds. B0rgesen (1905) described a 'Laminariaceae Association' which he noted as being widespread in the Faroes. The occurrence and distribution of Laminaria follows the patterns described by B0rgesen (1905) and other workers (Kain, 1960; Norton & Milburn, 1972). Laminaria hyperborea was the dominant species on firm substrates where more exposed conditions prevailed. In the sheltered parts of Skalafj0rbur and Kaldbaksf j0rbur it occurred as a SHELTERED FJORDS AND SOUNDS 147 narrow band in the shallow sublittoral but was absent from the innermost regions; a broad form of the species which resembled the cucullata form of L. digitata was found in deeper waters in the sheltered parts of fjords. The Laminaria faeroensis form of L. saccharina occurred abundantly in Skalafj0rbur and Kaldbaksf j0rbur. Our observations parallel those of B0rgesen who recorded Laminaria with large laminae and long, hollow stipes from the deeper parts of fjords. Although the species was common in the sheltered parts of Sundini, it also occurred near the narrows where there was a strong current. The normal form of Laminaria saccharina grew in situations where there was more water movement. At most of the sites investigated the species grew in the shallow sublittoral levels among L. digitata and L. hyperborea; it was also widespread on unstable substrates in deeper waters at the entrance to Skalafj0rbur. We confirmed B0rgesen's observations that in certain situations it was difficult to distinguish between L. saccharina and L. faeroensis. We detected Desmarestia aculeata and D. viridis growing among L. faeroensis on unstable substrates in deeper waters; the epiphyte Porphyropsis coccinea was also found. B0rgesen described a 'Desmarestia Association' which appeared to comprise almost pure stands of Desmarestia spp. from similar habitats. We failed to find eel-grass Zoster a marina in sheltered habitats. B0rgesen commented that the plant was rare on the Faroes and cited only one locality (Vaag fjord) for this community. The leaves of Zoster a bore a number of small algal epiphytes. A well developed subflora was detected at the entrances to Skalaf j0rbur and Kaldbaksf j0rbur and throughout Hvannasund. The subflora consisted mainly of red algae growing either as epiphytes on the stipes and holdfasts of Laminaria hyperborea or on firm substrate between L. hyperborea plants. This subflora vegetation differs little from the vegetation described by B0rgesen as a 'Laminaria hyperborea Association'. This assemblage of species was not found in the inner regions of fjords because either L. hyperborea was absent or suitable substrate was not available. Almost no subflora was detected beneath the cover of L. faeroensis, which appears to form a blanket covering over large areas of the sublittoral in Skalaf j0rbur and Kaldbaksf j0rbur. In contrast, a well developed subflora was found throughout Hvannasund; this vegetation was recorded both among Laminaria spp. and from levels below the lower limits of Laminaria. Species were detected which were not listed by B0rgesen under his 'Laminaria hyperborea Association' but included by him in a 'sublittoral Floridae formation', which usually occurred in open sea conditions rather than in sheltered waters; the formation is characterized by such species as Callophyllis cristata, C. laciniata, Delesseria sanguinea, Fimbrifolium (Rhodophyllis) dichotomum, and Phycodrys rubens. The causeway which divides Hvannasund was constructed in the middle 1970s. The main effect of this was the elimination of water current through the narrows between Borboy and Viboy and the creation of still water conditions; vegetation nevertheless retains many of its open water characteristics (such as the abundance of red algae and the presence of species such asAlaria esculenta) although sheltered conditions are suggested by the occurrence there of the broad form of Laminaria hyperborea. It was hoped that the investigation of the narrows between Streymoy and Borboy would reveal a similar type of vegetation. At Nordskali, near the road bridge, Alaria esculenta and Laminaria faeroensis were found but the impoverished subflora did not resemble the vegetation in Hvannasund. B0rgesen described a 'Lithoderma Association' from deep waters in open sea situations; this association comprised a few small filamentous and frondose algae together with a number of crustose species. We detected a similar community inhabiting stone and shell substrates at the entrance to Skalafj0rbur but did not find the community in the inner regions of Skalafj0rbur. A community of crustose algae dominated by Cruoria sp. was present in deep waters in Kaldbaksf - j0rbur. Both surveys recorded Phycodrys rubens as the most frequent and sometimes the only macroalga in deeper waters; the species occurred in the sheltered middle regions of fjords, but was not found in the innermost regions. Plants resembling the lingulata form of P. rubens were recorded in deep waters where there was little current. Several species which were not recorded by B0rgesen (1902; 1905) were found in deeper 148 SEAWEEDS OF THE FAROES waters at the head end of Skalafj0rbur; Asperococcus turneri was found on stones and shells lying on the muddy sea floor; the species has not been found elsewhere in the Faroes but is common in the Shetlands. The tetrasporophyte stage of Bonnemaisonia hamifera (Trailliella) was detected growing on plastic bristle-like debris also on the muddy sea floor. Members of the Bonnemaisoniaceae have in recent times spread throughout much of northern Europe. Irvine et al. (1975) first recorded B. hamifera (as Trailliella) and Asparagopsis armata (as Falkenbergia) from the Shetlands, and Printz (1952) similarly noted Bonnemaisonia hamifera and B. aspa- ragoides from Norway. Neither Asparagopsis armata nor Bonnemaisonia asparagoides game- tophytes were found on the Faroes despite intensive searching. A few other uncommon species (Antithamnion floccosum, Polysiphonia elongata, and Sphacelaria rigidula) were also recorded from this habitat and are also known from the Shetlands and Norway. Comparison with other fjords The algal vegetation of fjord systems in Iceland, Norway, Shetland, and Scotland has only recently attracted scientific interest. Tittley etal. (1977) described how Sullom Voe (a fjord-like formation on the nearby Shetlands) differed from fjord and loch formations elsewhere in Norway and Scotland. Sullom Voe is a long inlet of the sea with an almost constant salinity throughout, unlike the Hardangerfjord in Norway in which freshwater drainage from the surrounding hinterland has a direct effect on the salinity regime of the fjord. Similar types of reduced salinity regimes were described by Munda (1972, 19780, 19786) for fjords on Iceland. River estuaries drain in the head ends of Skalafj0rbur and Kaldbaksfj0rbur, and, according to B0rgesen (1905), the freshwater inflow can bring about a local depression in salinity. Our observations indicated that in July 1980 there was no appreciable reduction in salinity at the heads of Skalafj0rbur and Kaldbaksf j0rbur. Hvannasund resembled Sullom Voe in that it was a narrow inlet of the sea without any significant inflow of freshwater at the head. A comparison of the sublittoral vegetation of Sullom Voe with that in Kaldbaksfj0rbur and Skalafj0rbur revealed a number of differences. Laminaria hyperborea was present as dense forests on bedrock at the entrances to Sullom Voe, Skalafj0rbur, and Kaldbaksfj0rbur, but in Sullom Voe the species was also found at the head, where it formed a narrow but distinct band in the shallow sublittoral. In both Skalafj0rbur and Kaldbaksf j0rbur, L, hyperborea did not grow at the landward ends. The broad ('cucullate') form of L. hyperborea, found in the inner regions of Kaldbaksfj0rbur and Hvannasund, was not detected in Sullom Voe. Laminaria faeroensis showed a very restricted distribution in Sullom Voe and was recorded only in the most sheltered regions. In Skalafj0rbur and Kaldbaksfj0rbur, L. faeroensis occurred throughout save only for the entrances. Sheltered, deeper waters in Sullom Voe were populated by the normal form of L. saccharina, whereas in Skalafj0rbur and Kaldbaksfj0rbur the normal form of the species was only found in the shallow infralittoral levels. Although a mainly red algal subflora was detected in both Sullom Voe and Skalafj0rbur, the vegetation was much less extensive in Skalafj0rbur and was absent from the inner regions. In both systems, a mainly brown algal subflora was associated with stands of L. saccharina/ L. faeroensis. The turf-forming vegetation in the shallow sublittoral was also similar in these systems. Species such as Phycodrys rubens, Polysiphonia elongata and Phyllophora crispa were the principal components of the vegetation in deep water throughout Sullom Voe. Phycodrys rubens was the most frequently observed species in the deeper parts of Skalafj0rbur, but it did not occur in the inner regions of the fjord. Polysiphonia elongata, by contrast, was found only once at the inner end of Skalafj0rbur and Phyllophora crispa was not found at all in Skala- fj0rbur. 'Trailliella' grew on mud and silt in the deeper parts of Sullom Voe, and was found in a similar habitat in Skalafj0rbur. Although some fucoids grew permanently immersed in shallow standing water at the heads of Skalafj0rbur and Kaldbaksf j0rbur, others formed a distinct and dense zone of vegetation where intertidal substrate was available; a well developed fucoid vegetation was present on rocks at the head end of Sullom Voe, but was absent in the shallow sublittoral levels. The vegetation in Hvannasund was similar to that in the inner regions of Sullom Voe in several respects: (i) there was a distinct zonation of eulittoral species on the harbour and causeway walls; (ii) a narrow but SHELTERED FJORDS AND SOUNDS 149 distinct band of Laminaria hyperborea was present in the shallow sublittoral zone; (iii) a well-developed subflora with many red algae was present in deeper waters. In Norway and Iceland fjords are much longer, more sheltered, and receive a greater inflow of freshwater. Consequently different patterns of algal occurrence and distribution have been recorded. Jorde & Klavestad (1963) described a 'fjord effect' which involved '. . . an impover- ishment of the vegetation and an abrupt rise in the lower limit of continuous vegetation of larger brown and red algae on passing from the outer fjord areas inwards . . .'. The occurrence and distribution of algae in Skalafj0rbur and Kaldbaksfj0rbur are in some respects similar to the patterns of distribution in Hardangerfjord, Norway (Jorde & Klavestad, 1963) and in Dyrafjor- dur, Iceland (Munda, 1972, 19780, 1978b). Brackish water species Capsosiphon fulvescens, Percursaria percursa, and Rhizoclonium riparium were recorded from saline meadows in Hardangerf jord and from oligohaline habitats in the inner part of Dyrafjordur; C. fulvescens and R. riparium occurred in the estuaries at the heads of both Skalafj0rbur and Kaldbaks- fj0rbur. On the other hand, Fucus ceranoides occurred in river estuaries in fjords in Norway and Iceland, but has not been found on the Faroes. An association of filamentous green and brown algae (Chaetomorpha capillaris, Dictyosiphon spp. , Enteromorpha spp. , Stictyosiphon spp. , and ectocarpoids) was widespread in the inner brackish region of Hardangerf jord and Dyrafjorbur; a similar association was restricted to the inner ends of Skalafj0rbur and Kaldbaksfj0rbur. F. spiralis and F. vesiculosus occurred throughout the Norwegian, Icelandic and Faroese fjord systems. In Norway, F. serratus was the dominant fucoid at lower shore levels, whereas on the Faroes F. distichus subsp. edentatus occupied these levels. F. serratus was found growing in the shallow sublittoral in the inner reaches of the Hardangerf jord, although B0rgesen (1902) did not find it and dismissed previous records. The limit of distribution of Laminaria saccharina in Dyrafjorbur, Iceland, is salinity depen- dent and coincides with the 28%o isohaline (Munda, 19786). The species occurs throughout the middle and outer parts of Hardangerf jord, these remaining more or less saline throughout the year; its limit of distribution coincides with a summer salinity reduction to 15%o (Jorde & Klavestad, 1963). L. digitata shows a similar salinity dependent distribution in both fjords. On the Faroes, both L. saccharina (incl. L. faeroensis} and L. digitata were found at the head of Skalafj0rbur, where presumably the salinity remains high for most of the year. In the Norwegian, Icelandic and Faroese fjordic systems, a subflora of predominantly filamentous brown algae is associated with Laminaria saccharina/ L. faeroensis, and this community is particularly common in the sheltered inner reaches. The occurrence of Laminaria hyperborea in the Dyrafjorbur, Hardangerfjord, and Skala- fj0rbur is largely restricted to the outer exposed regions where it can form dense stands. In more sheltered conditions, L. hyperborea occurs only as a narrow band in the shallow sublittoral levels; in Hardangerfjord, Dyrafjorbur, and Skalafj0rbur, the inner limit of distribution of L. hyperborea is in a region where the salinity only very occasionally falls below that of sea water. There, and further into the fjords, L. hyperborea is replaced by L. saccharina. The broad form of L. hyperborea was widespread in the inner parts of Kalbaksfj0rdur but has not been recorded from the Norwegian and Icelandic fjords. The subflora associated with L. hyperborea is also restricted to the outer regions of fjords in Norway, Iceland, and the Faroes; an impoverished subflora was, however, recorded from the middle and inner regions of Skalafj0rbur. A few species of red algae, such as Ceramium rubrum and Polysiphonia elongata recorded from the inner part of Skalaf j0rbur, were also found in the inner regions of Hardangerfjord. A dense turf of Corallina officinalis was present in the lower littoral and sublittoral fringe levels at the entrances to fjords in Norway, Iceland, and the Faroes. In Norway and on the Faroes Polysiphonia brodiaei was also commonly present. In all fjordic systems this vegetation did not occur in sheltered waters, and was replaced by filamentous brown algae associated with Dumontia contorta, Cystoclonium purpureum, andRhodomelaspp. 150 SEAWEEDS OF THE FAROES 7. Conclusions The occurrence and distribution of marine algae are similar in Skalafj0rbur and in Kaldbaks- fj0rbur; the vegetation in Hvannasund and Sundini differs considerably from these locations. In Hvannasund and Sundini species which are characteristic of both exposed and sheltered conditions were detected, although in Sundini the subflora of the eulittoral and sublittoral levels was rather impoverished. The luxuriant subflora, particularly of red algae, found in the inner parts of Hvannasund indicated that there was little or no 'fjord effect', and that Hvannasund was an inlet of the sea. The vegetation in Sullom Voe, Shetlands, similarly showed little or no 'fjord effect'. A reduced 'fjord effect' was detected in Skalaf j0rbur and Kaldbaksf j0rbur, and was indicated by the absence of Laminaria hyperborea and of red algae from the inner regions of these fjords. The reduced 'fjord effect' was probably brought about by the rivers which drained into the heads of these fjords. The resulting estuarine conditions were only local and in July 1980 were restricted to an area within a few hundred metres radius of the river mouth. In Dyrafjorbur on Iceland by contrast the estuarine area was more extensive and the 'fjord effect' noticeable over a greater distance along the fjord. In the much larger Hardangerfjord system, which drains an extensive hinterland, the flow of freshwater into the fjord has a more pronounced effect on the occurrence and distribution of algae. Despite the less extensive 'fjord effect' in Skalaf j0rbur and Kaldbaksfj0rbur than in Hardangerfjord and Dyrafjorbur, the occurrence and distribution of marine algae in these systems is more closely similar to the latter fjords than to inlets of the sea such as Hvannasund and Sullom Voe. 8. Acknowledgements We wish to thank our colleagues Dr D. E. G. Irvine (expedition leader), J. H. Price, Dr K. Liming, and Mrs P. Farnham for their considerable help with field work. Thanks are also due to the Academia Faroensis for providing laboratory facilities. The expedition was funded by N. A.T.O. , the British Museum (Natural History), and the Carlsberg Foundation. 9. References B0rgesen, F. [C. E.] 1902. Marine algae. In [E. Warming (Ed.)], Botany of the Fceroes based upon Danish investigations. Part II: 339-532. Copenhagen. 1904. Om Algevegetationen ved Fcer0ernes Kysterenplantegeografi.sk unders0gelse. pp. [6] + 122+ [4]. K0benhavn and Kristiania. - 1905. The algae-vegetation of the Faeroese coasts with remarks on the phyto-geography. In [E. Warming (Ed.)], Botany of the Fceroes based upon Danish investigations. Part III: 683-834. Copenhagen and Christiania. & Jonsson, H. 1905. The distribution of the marine algae of the Arctic Sea and of the northernmost part of the Atlantic. In [E. Warming (Ed.)], Botany of the Fceroes based upon Danish investigations. Part III: Appendix I-XXVIII. Copenhagen and Christiania. Crothers, J. H. 1981. Shell-shape variation in Faroese dog-whelks (Nucella lapillus (L.)). Biol. J. Linn. Soc. 15: 327-337. Holt, G. 1975. Marinbotaniske notaterfra Faer0yene 1975. [Unpublished report.] Irvine, D. E. G. 1982. Seaweeds of the Faroes. 1: The flora. Bull. Br. Mus. not. Hist. (Bot.) 10 (3): 109-131. , Guiry, M. D., Tittley, I., & Russell, G. 1975. New and interesting marine algae from the Shetland Isles. Br. phycol. J. 10: 57-71. -, Tittley, L, Price, J. H., & Farnham, W. F. 1982. Seaweeds of the Faroes. [Abstract.] Br. phycol. J. 17: 234. Johansen, J. 1979. 30f0roysk tarasl0g. Torshavn. Jorde, I. & Klavestad, N. 1963. The natural history of the Hardangerfjord 4 the benthonic algal vegetation. Sarsia9: 1-99. Kain, J. M. 1960. Direct observations on some Manx sublittoral algae. J. mar. biol. Ass. U. K. 39: 609-630. 1976. New and interesting marine algae from the Shetland Isles. II. Hollow and solid stiped Laminaria (Simplices). Br. phycol. J. 11: 1-11. SHELTERED FJORDS AND SOUNDS 151 Lewis, J. 1964. The ecology of rocky shores. London. Munda, I. 1972. General features of the benthic algal zonation around the Icelandic coast. Acta nat. islandica 21: 1-36. 1978a. Salinity dependent distribution of benthic algae in estuarine areas of Icelandic fjords. Botanica mar. 21: 451-468. 1978ft. Survey of the benthic algal vegetation of the Dyrafjordur, northwest Iceland. Nova Hedwigia 29: 281-403. Norton, T. A. & Milburn, J. A. 1972. Direct observations of the sublittoral marine algae of Argyll, Scotland. Hydrobiologia 40: 55-68. Price, J. H. & Farnham, W. F. 1982. Seaweeds of the Faroes. 3: Open shores. Bull. Br. Mus. nat. Hist. (Bot.) 10 (3): 153-225. Printz, H. 1952. On some rare or recently immigrated marine algae on the Norwegian coast. Nytt. Mag. Bot. 1: 135-151. Rex, B. 1970. Rapport over algobservationer Farodrna sommaren 1970. [Unpublished report.] Russell, G. 1978. Seaweeds as environmental indicators. Porcupine Newsl. 1: 109-110. Tittley, L, Irvine, D. E. G., & Jephsen, N. A. 1977. The infralittoral marine algae of Sullom Voe, Shetland. Trans, hot. Soc. Edinb. 42: 397-419. Seaweeds of the Faroes 3: Open shores James H. Price Department of Botany, British Museum (Natural History), Cromwell Road, London SW7 5BD William F. Farnham Marine Laboratory, Portsmouth Polytechnic, Hayling Island PO11 ODG Contents Synopsis ............. 153 1. Introduction ............ 153 2. Methods 155 3. Sites, descriptions, and profiles . ........ 155 4. Results: comparative species distributions ....... 170 i. Subtidal: major organisms ......... 171 Alaria 171 Laminaria hyperborea and other Laminaria spp. ..... 173 Populations at depths greater than laminarian limits .... 183 ii. Subtidal: epiphytes and underflora ....... 183 Laminaria hyperborea epiphytes and underflora . . . . .211 Epiphytes/underflora to other major subtidal organisms. . . .211 iii. Subtidal: other specialized situations ....... 212 Detrital conditions ......... 212 Epizoic growths and animal associations ...... 212 Pools and standing waters ........ 216 iv. Intertidal ........... 217 5. Discussion and conclusions ......... 218 i. General 218 ii. The subtidal 219 iii. The intertidal ........... 224 6. Acknowledgements . . . . ' . . . . . . 224 7. References ............ 224 Synopsis Detailed studies on the intertidal and, more especially, the subtidal of sheltered and exposed open shores are presented. Only comparative changes, or detailed differences for particular groups or taxa, are given for the intertidal, since B0rgesen's 80-year-old data were so accurately observed and well expressed. The earlier information lacked details derived from direct observation of the subtidal flora, now available for the first time. Vertical and horizontal distribution patterns of the major subtidal canopy organisms at the 44 open-shore sites examined are discussed and profiles of particularly detailed transect studies presented. Data on subordinate organisms (epiphytes; epizoic and epilithic underflora) and on flora at depths below the laminarian limits are considered, after presentation in tabular summarized form. Specialized situations (detrital, scour-tolerant species; consistent epizoic or other animal associations; the effects on distribution of lower littoral pools or standing waters) are examined. Amplification or revision of previous information by present data, and apparent comparative changes in the last 80 years, are discussed for the subtidal flora and its distribution. Particular rarities, unusual distributions, and new records are briefly considered. 1. Introduction There is no easy and absolute distinction between fully marine macroflora distribution patterns in sheltered waters and those in the open, more wave-exposed situations. This appears true Bull. Br. Mus. nat. Hist. (Bot.) 10 (3): 153-225 Issued 25 November 1982 154 SEAWEEDS OF THE FAROES whatever the geographical location in temperate and cooler waters. Transitions from one set of most usual patterns to the other are gradual, or (at most distinct) step-wise. Aside from that, most shores present complex mosaics of exposed and sheltered local areas that, as with the overall impressions of shores, are clear enough in themselves until one attempts detailed analysis of the individual biotic or environmental facets, when blurring of hitherto clear-cut patterns is immediately experienced. 'Exposure' (as a concept applied to areas experiencing strong water-movement) is not limited to the effects of wave-action; swell, up-carry, scour, tide-race and so on are often as important as the direct effects of true wave-impact, are more widespread as phenomena, and can often produce similar results. This is particularly true for the much-sculpted northern and southern Atlantic oceanic islands, where sheer geographical/ physiographical position of an open shore area is, as with its topography, no necessarily firm guide to its exposure/shelter characteristics. It will therefore be appreciated that the open shores of the heavily-glaciated, volcanic Faroe Islands are by no means necessarily all of extreme exposure to strong water-movement effects, and that it is not easy to draw clear-cut limits in coverage between the floras of sheltered fjords/sounds (Tittley etal., 1982) and those of the open shores. Essentially, the present paper is concerned with non-fjordic shores, although with such a complex configuration as that of the Faroese shores, even the concept of 'fjord' is not always clear. Outer parts of fjords or sounds, especially where (as in Raktangi, at the confluence of Tangafj0rbur and Skalafj0rbur, Streymoy) a wide entry has permitted the direct effects of essentially open-shore wave characteristics to impact a rock promontory within a fjord, show few distinctions from open shore conditions in either intertidal or the first 30 m of the subtidal. Some data from such shores are briefly included for comparison within the present coverage, although overlap between the second paper in this series and the present text has been kept to a minimum. A general background to the work pattern has already been presented by Irvine et al. (1982), Irvine (1982), and Tittley et al. (1982). Data on certain taxa of importance on open shores have also been incorporated in papers of more specific coverage (e.g. on Callithamnion; Dixon & Price, 1981). General background presented here therefore concerns characteristics of open shore biota not already adequately covered elsewhere. Whatever the style of marine vegeta- tional study in the Faroes, continual reference to the very effective work by B0rgesen (1895-1902, as to visits; 1902-1905, as to publication dates) is unavoidable. Although certain changes seem to have taken place in the 80-odd years since then, the possibility of such changes being one of the major considerations behind the present work scheme, it is impressive that he was able to make such detailed and accurate descriptive accounts, as will emerge later of considerable validity still today. The dredging work then carried out gave good insight into the nature of the subtidal open shore flora and its distribution, but (from absence of direct observation) that still represents the least detailed part of his account. His descriptions of the open shore (sheltered or wave-washed) intertidals are of such widespread application and sufficiently comprehensive as to make detailed repetition unnecessary, save where there are changes or differences in detail. Points already well made in his accounts of those intertidals are therefore not further elaborated here, the present text concentrating on subtidal detail from direct observation and on critical variation from the earlier intertidal data. The configuration of the generally rugged coastlines of the Faroes and the mosaic nature of the habitat distribution provided thereby are well summarized in B0rgesen's analysis of the overall algal vegetation (1905: 705-706), reference to which is recommended for full appreciation of certain background characteristics. A comment on terminology is appropriate. We have generally accepted the limits of vertical divisions of the shore suggested by B0rgesen (1905) for the Faroes. For the purposes of the present paper, the most important shore levels requiring clear definition are those concerned with the lower intertidal, subtidal fringe, and subtidal proper. In presenting definition, we also clarify our terminology. We accept that generally speaking the lower limit of Himanthalia is an appropriate lower limit for what B0rgesen (1905: 708-710; 733-735) referred to as his Littoral Region; we use the term intertidal to cover the same vertical amplitude, and include within that general term all those areas, including the splash or spray upper levels (often far inland) still OPEN SHORES 155 within the sphere of marine influence such that some algae also found lower on shore in direct tidal contact will grow there (e.g. Audouinella purpurea) . Subtidal is therefore applied to all levels, uncovered or not in usual tidal sequences at springs periods, below the lower limit of Himanthalia. Where the latter is not present in shelter, we take the upper limit of whatever laminarians are present at band-forming density as the limit between intertidal and subtidal. Across these limits usually lies the subtidal fringe, created by tidal movement, swell, and/or any form of variable water run that gives a variable regime of immersion/emersion to any organisms present there. The width and height on shore are locally variable, as are the detailed presences of organisms, but Alaria and some outliers of Himanthalia (or even most of the latter, where present in less wave-exposed locations) are usually involved. We have therefore employed the term subtidal fringe in a somewhat general sense, without definition as to rigid upper and lower limits, to indicate rather an area around the usual calm low water level of medium tides and subject to a fairly similar variation in ambient conditions. Configuration of this fringe area varies considerably between shores and is much affected by the presence of incut channels, downwash surfaces and upcarry grooves. The relationships of lower shore pools and standing waters to such a fringe area are debated later. Minor differences of concept between this treatment and B0rgesen's (1905) approach are easily appreciated by reference to his earlier (1905: 708-711) comments. 2. Methods All observational data, quantitative and qualitative, were directly derived by shore examination or scuba. Profiles related to low water level were constructed from diving and intertidal notes, including depths recorded from wrist gauges or by direct measurement. Vegetational densities have been derived from counts of larger flora and from subjective estimates of smaller species. Quadrat reading was not possible, except in the case of specialist studies on particular taxa. Material of doubtful determination in the field was subsequently confirmed or revised in the laboratory. 3. Sites, descriptions, and profiles Logistics and accessibility resulted in slight limitation on the establishment of study sites, but not such that the results obtained here show major lacunae or bias. Observations have been derived from a wide variety of open shores and open or more exposed fjordic/sound situations on the islands of Borboy, Eysturoy, Fugloy, Koltur, Kunoy, Mykines, Nolsoy, Sandoy, Streymoy, Suburoy, Svinoy, Vagar, and Viboy. The particular locations concerned are indicated on the island group map (Fig. 1). All visits were made during July and August, 1980. In the following descriptive list, sites productive of supporting data only are in italic; more fully documented, especially profiled, locations are in bold face. Figure numbers in the lists refer to detailed profiles presented. The site order does not reflect sequence of working since, for clarity, sites have been re-ordered on a geographical basis. Except in the case of the detailed Hoyvik transect area, intertidal profiles are not presented. As indicated above, we have accepted B0rgesen's (1905) utilization of Himanthalia lower limits to mark the lower limit of his 'Littoral' on open tidal coasts of adequate exposure to water movement. Elsewhere, the same limits can be represented by the upper limits of continuous dense laminarian growth (usually of Alaria), and these are mostly the highest level organisms of which the vertical limits appear on the profiles. Pools, creating upcarry of subtidal flora, are a separate matter in this context and we have adopted the same strategy as did B0rgesen (1905), in dealing with that habitat in a different section where comments on additions or differences are required. Fig. 1 Faroe Islands: open shore and comparative stations. Scale approx. 1 : 300000. For details of stations, see the numbered site list in the text. Numbers underlined represent profiled sites. Abbreviations of island names on the map represent the following: Su = Sufouroy LD = Litla Dimun SD = Stora Dimun Sk = Skuvoy Sa = Sandoy He = Hestur Ko = Koltur No = Nolsoy Str = Streymoy Va = Vagar My = Mykines Ey = Eysturoy Ka = Kalsoy Ku = Kunoy Bo = Borboy Vi = Vfeoy Sv = Svinoy Fu = Fugloy Figs 2-10 General Notes. Certain symbols and abbreviations are standard throughout the profiles covered by these figure numbers. All profiles carry bars representing the depth distributions for the site of all detected, or the most important, marine biota. Depth's are always in metres, although the scales employed vary with quantity and style of data to be presented. Horizontal distances, where given, are also in metres. In some cases, standard symbols are augmented by pictorial symbols for further clarification; since the quantity and form of these vary between figures, they are detailed on the profile(s) concerned. For further development of these comments and the information background on which they depend, see the appropriate parts of the main text. Standard symbols or abbreviations are as follows: Taxa Ad Antithamnion plumula vat . demersum Ae Alaria esculenta An Ascophyllum nodosum Ap Ahnfeltia plicata Ba Barnacles, in general Bb Brongniartella byssoides Be Balanus crenulatus Bp Bryopsis plumosa Bs Brittle stars, in general Ca Callophyllis cristata Cc Clathrina coriacea Cd Callithamnion decompositum Ce Ceramium shuttleworthianum Cl Callophyllis laciniata Cm Chaetomorpha melagonium Cn Callocolax neglectum Co Corallina officinalis Cr Ceramium rubrum Cs Callithamnion sepositum Da Desmarestia aculeata DC Dilsea carnosa De Dermocarpa prasina (= Entophysalis confertd) Dl Desmarestia ligulata Dm Derbesia marina Ds Delesseria sanguinea Dv Desmarestia viridis EC Ectocarpoids, in general Ee Echinus esculentus Ei Enteromorpha intestinalis Fd Fimbrifolium dichotomum Fl Furcellaria lumbricalis Gs Gigartina stellata Ha Halichondria He Himanthalia elongata Ho Halicystis ovalis Hr Hildenbrandia rubra Lc Lomentaria davellosa Ld Laminaria digitata Lg Lithothamnion glaciale Lh Laminaria hyperborea Li Lithothamnia, in general LI Leptophytum laeve Lo Lomentaria orcadensis Ls Laminaria saccharina Ma Membranoptera alata Mm Modiolus modiolus Od Odonthalia dentata Pa Palmaria palmata PC Plocamium cartilagineum Pe Pseudolithoderma extensum Phc Phyllophora crispa Php Phymatolithon polymorphum Pht Phyllophora traillii Phtr Phyllophora truncata PI Polysiphonia lanosa Pm Porphyra miniata Po Porphyra umbilicalis Pp Pterosiphonia parasitica Pr Phycodrys rubens Ps Ptilota serrata Pt Ptilota spp. Ftp Ptilota plumosa Pu Polysiphonia urceolata Pv Patella vulgata Sa Spongomorpha arcta Se Serpulids, in general Sp Sponges, in general Ul Ulva lactuca Ve Verrucaria spp. General symbols I = epilithic ^ p = epiphytic > used in connection with taxa abbreviations or on distribution bars. z = epizoic J in ( ), after 1, p. or z (see above) = substrate types or 'host' organisms involved. 5 = sand, in patches or stretches. S G = blackish sandy gravel. Symbols in distribution bar types = organism present in band-forming, 'forest', or otherwise dense growths (therefore common, abundant, or characterising). = organism scattered, but consistent to occasional, over the depths concerned. = organism present, but detected as very sparse to rare. 158 SEAWEEDS OF THE FAROES Site list 1. Akraberg, south Suburoy. Subtidal. Dive off steep rocky head-and-cove type exposed shore beneath light-house. Clear water in 15 m depth. General algal collections on zoological dive. 2. Beinisvort, south Suburoy. Intertidal. Gullies and faces of stacks just off-shore. Generally exposed positions. 3. Tr0llkonufingur, Skarvanes, Sandoy. Intertidal. Asen and Rueness. Tide-pools in rocky but not excessively steep shore. Pools showing strong upcarry of usually subtidal flora. 4. Husuvik. Sandoy. Intertidal and subtidal. Concrete-faced boulder harbour wall and steps. Broken boulders seaward of base of harbour wall (in 6 m) gave way to firm bedrock. At 10 m depth, gullies with sand and small stones penetrated the gently sloping bedrock. Latter completely covered by bare sand below 10 m. 5. Sandur, Sandoy. Subtidal and intertidal. Harbour wall of cemented vertical blocks faced with cement; base of wall at 7 m depth. Adjacent open sheltered rocks at and around low water level. Bouldered bottom below and to seaward of harbour wall in 7-12 m depths, with scattered and broken sandy patches. See Fig. 2. Also in Holt (1975). 6A. Gtfbidrangur, Mykines. Subtidal. Off rocky steep small islets west of Mykines, to 20 m depth. 6B. Tindagjogv, Mykines. Intertidal. Sloping rock with pools, north-west side of island. 7. Borgagjogv, Mykines. Subtidal. Vertical to slightly overhanging slab rock-face, with uneven large boulders on bottom below. All depths to 17 m. 8. Skerholmur, S0rvdgsfj0t > bur, Vdgar. Subtidal. Seaward face of exposed steep rocky islet. Collections from between 6/7 and 15 m depths. l Outside and away from harbour walls Symbols Fig. 2 Sandur, Sandoy (site 5). Sketch map showing main observations area and locations of profiled stretches. Profiles represented are: 1. Outside and away from harbour walls. 2. Harbour wall end: concrete-faced blocks. 3. Configuration on boulders and bedrock below wall end base. See p. 157 for lettered symbols. f Ba Sp (encrusting) Lc Lo Dm 2 +1 T- - ZlAe w Lh 2. Harbour wall end: concrete-faced blocks OPEN SHORES 159 9. Stack in Dragasund, opposite Tindholmur. Subtidal. Vertical face and easing slope/boulders below, in 5-9 m depths. South side exposed, north-west side less directly so. 10. Oyrargjogv, Vestmannasund, Vdgar. Subtidal. Subtidal fringe and subtidal proper to 5 m depth. Strong water-flow, but less exposure than fully-open shores. 11. Koltur, mid-north coast. Subtidal. Barnacle-dominated exposed steep shore continuing subtidally initially as vertical cliff in shallower depths, with kelp forest to 12 m depth; easing slope and thinning kelp to 20 m depth. Bare rocks with a few crusts thence to 30 m depth. Details observations to 20 m, then general scan. See Fig. 3. 12. Skrfbutangi, north-east (seaward) shore of Nolsoy. Subtidal. Detailed data to 32 m depth, more general below that to 40 m depth. Uneven but fairly steep exposed rock lacking laminarians other than fringing Alaria (1-2 m) and Laminaria hyperborea. Scattered plants of latter on easing slope between 10 and 20 m depths. True forest deeper than elsewhere, from 10 m to 20-22 m according to location. Firm rock and boulders to about 30 m, then scattered rocks amongst gravel substrata, with overlying or partly embedded small boulders down to and beyond 40 m; algae still then present. See Fig. 3. 13. Kirkjubmir, point near Arnanes, Streymoy. Intertidal, with observations on superficial subtidal. Exposed steep (but not high) cliffed headland of individually steep and fallen blocks, with gullies and upcarry niches at intervals. The steep cliffs generally continue descent as steeply to and into the subtidal; occasional very local areas form shallow intertidal shelves or shallowly but consistently subtidal shelves at fringe of cliffs. Downwash areas with easing slopes penetrate the boulders and narrow platforms. Headland grades to south into stones and boulders of the sheltered Kirkjub0ur cove. 14. V dbastahur, harbour wall and adjacent subtidal, Streymoy. Subtidal. Detailed observations to 10 m depth, then more general scan beyond. Firm rock and boulders, penetrated throughout (in 2 m and below) by gullies where slope eases. 3. Configuration below wall end base epilithics on stones in sandy patches Horizontal distance (m) Fig. 2 (cont.) Caption, see p. 158. WL 20- 25- 30- 35- 40- ^_Qbservstian_jjmit_-_i2. Fig. 3 Semi-diagrammatic composite showing structure of steep exposed subtidals and distribution of biota present. Based on data from sites 11 (Koltur) and 12 (Nolsoy). No distinctions between sites, or absence of the organism from one site bars plain; critical differences between site patterns bars appropriately numbered. See p. 157 for symbols. OPEN SHORES 161 15. Argir, south of Torshavn, Streymoy. Intertidal. Semi-exposed moderately-sloping firm rock slab and bedrock shore, with occasional small boulder overlay. Penetrated by runnels and upcarry/downwash areas. Dense Alaria fringe; locally sheltered niches with good Fucus spiralis. Laminarians in adjacent small harbour. 16A. Hospital Rocks, Torshavn, Streymoy. Intertidal. Minor low headland of smooth rock slabbing with downwash channels, swell channels, pools, and a flat inwash area on north side of outer point. Occupies north side of the inner depths of the haven, just south of the sewage pipe and mole below hospital. 16B. Tinganes, Torshavn, Streymoy. Intertidal, with shallow subtidal observations. Gently sloping firm rock and artificial facings (at intervals), on central headland in Torshavn harbour; usually pollution pattern associated with functional port. Presumably the same as the station in Rex (1970) and close in position to Holt's (1975) station Tinganeset. 17. Shore below Old Fort, north Torshavn, Streymoy. Intertidal. Rocky shore with sheltered to semi- exposed rock tongues, contained coves, and pools, just north of mole enclosing ferry and unloading harbour areas. Shore slopes rather gentle. Approximately the position of Holt's (1975) station Skansen. 18. Torshavn, heads and coves to north, between Old Fort Head (station 17) and coves/heads transect area, Hoyvik (station 19), all Streymoy. Intertidal, with shallow subtidal observations. Semi-exposed, mainly gently-sloping, bedrock headlands, with contained gullies, pools, lagoons and channels. Only local detrital and broken loose rock areas. Flora a mosaic, but some accepted major exposure indicators locally present. Coves and inlets between heads rather more sheltered, but carrying exposure indicators (e.g. Himanthalia) alongside shelter indicators (e.g. Ascophyllum, massive Laminaria saccharind) . 19. Hoyvik, coves and headlands, Streymoy. Subtidal and intertidal. Detailed transects on and seaward from semi-exposed firm rock minor headland. Comparative observations intertidally and subtidally in adjacent more sheltered coves to north and south. See Fig. 4. Close to one of the stations in Rex (1970). 20. Hoyvik, opposite Byrgistangi islet, Streymoy. Intertidal, with shallow subtidal observations. Pools and shallow subtidal along south side of firm rocky headland, semi-sheltered, north of station 19. Flora an equivocal mosaic of exposure and shelter characteristics. Pools and shaded undercut verticals showing considerable subtidal upcarry. 21. Eystnes, Eysturoy. Subtidal. Dive off steep firm rocky shore at the mouth of the Tangafj0rbur, east of and opposite Hvitanes. Firm rock substrate slope eases as depth increases. Strong Laminaria hyperborea forest in first 10 m, opening out to smaller and sparser plants below. Algal growth continuing on rock below dive limit (24 m) at which depth red underflora was not well-developed. 22. Storafles. Subtidal. Isolated rock in the middle of Tangafj0rfour mouth. Dive to 20 m over steep but shallowing rock slope bearing Laminaria hyperborea forest in upper half of penetrated depths. 23. Raktangi and Abbin, Eysturoy. Intertidal with shallow subtidal observations. Peninsula tip and rocks lateral on east (Skalaf j0rbur) side of headland. Largely exposed or semi-exposed flora in wash channels, on rock slabbing, on undercut sheer rock, on fallen boulders and in retained pools. Considerable subtidal upcarry in shaded standing water. Shallow subtidal wave-washed edges with mosaic exposed flora. Sheltered-water flora on some locally protected mobile substrata. Alaria and Ascophyllum noted growing side-by-side. Modiolus near peninsula tip. 24. Kvivik, Streymoy. Subtidal. Sloping rock surface penetrated by shallow subtidal channels in bedrock, leading to upcarry of the dominant Laminaria hyperborea forest; latter continuing down to 8/9 m depth, where slope levelled off. Quite extensive sand-covered intrusive gullies in depths of 4 to 8 m ; deeper gullies tended to be bare of algae. Bedrock scored by niches and locally with boulders in declivities and gullies. Modiolus clumps at bedrock/boulder/gully boundaries and in the bedrock niches. Dive terminated at 9 m. 25. Sundini, under middle span of bridge, near Norfokali, Eysturoy-Streymoy. Subtidal. Comparative station deep into sound between the two islands. Included because of characteristics sheltered as to wave-action but aping exposure through very fast tidal current effects. Total available depths 5 to 7 m, according to location. Solid bedrock, with small overlying boulders. Luxuriant Alaria mixed patchily with Laminaria hyperborea. Locally equal admixture of Laminaria faeroensis, with thick stipes and long blades. Generally impoverished underflora and epiphytes. Echinus and asteroids in fair abundance. 26. Saksh0vn, south side, 200 m seaward of Grotdalsgjogv, Streymoy. Intertidal and shallow subtidal. Narrow and smoothly steep shaded intertidal descending in about 2 m or locally less beneath the sand/silt substrate forming the lateral base of the haven. Deep cut and high overhangs from above, with continuous off-fall of strong freshwater flow. Restricted and specialized sheltered-water flora. See also Holt (1975) and Rex (1970), both of whom examined parts of the same haven. 162 SEAWEEDS OF THE FAROES Comparative r 1. Headland IT profile 2. Subtidal transect, 4Om long (s haded areas -general observations) Fig.4 Hoyvik transects (site 19) 1. IT profile CD Q- CO , I lower IT grooves widen bands by upcarry^ and slope easing (A O x | CO 0> I O w 0) X E a J a> I\^ vi O o O LW Fig. 4 Caption, see p. 163. 27. #', harbour wall (south) side, within the artificial mole, Eysturoy. Subtidal. Mole wall of large broken boulders, grading out in 9-10 m depths to blackish shell-sand having only diatom cover. Sheltered-water flora. 28. Eibi, open (north) side, beyond the inland lake; strong wave-action; Eysturoy. Subtidal. Laminaria hyperborea forest in 5 to 7 m; oldish plants on bedrock, with density varying mosaically. Sandy patches overlying bedrock in places; large boulders overlying bedrock elsewhere in places. Water movement above 5-7 m too strong for significant observations. OPEN SHORES 2.ST transect as 5m, but with Ma and Pc(p); Ca (I; few) 163 _f WL_. flat area (less than 0.25m depth variation 14*i30m)omitted. V gentle slope 30 ^ 36m rw^^ /as 5/6m,except > (jess Cp. more Pel). omitted Horizontal distancefm) 14 30 Fig. 4 Hoyvik (site 19). Transect and observational area sketch map showing positions of profiles and limits of Cove 1 and Cove 2 general observational and comparative areas. Profiles cover (1) intertidal patterns, and (2) subtidal patterns. See p. 157 for symbols. 29. Near Stakkur, north-west coast of Streymoy. Subtidal and intertidal. Very steep to vertical cliff descending to 23 m depth and then levelling off to rough boulder-strewn bottom, latter with no evidence of even crustose algae. No true 'forest', but Laminaria hyperborea plants present from 8 to 15 m depths. Very strong west-to-east running tidal stream during dive; if consistent, may have influence on state and distribution of flora. See Fig. 5. 30. Below and west of Kollur peak, north-west Eysturoy. Subtidal. Similar steeply-sloping rocks to those of station 29, but showing more irregularity than there and the overlying boulders larger and more irregular. Laminaria hyperborea forest over depth range 10-16 m, with plants still present at depths well beyond that. See Fig. 6. 31. Gjogv, north-east Eysturoy. Subtidal and intertidal. Deep (175 m long) cleft into the solid bedrock of the exposed northern shore; shows gradation of effects of configuration, depth, substrate type and water-movement as inter-relatedly determinating species presence, density and distribution along and throughout the cleft. Comparative transect also examined in the subtidal of the vertical to steep rock face outside the cleft to the north. South side with similar but lesser steep face and with additional off-cliff shallowly-sloping seaward slabbing and retained pools, all fairly well exposed to strong wave-wash and intertidally largely animal-dominated. Grades into bouldered cove on south. See Fig. 7. 32. Elduvik, Funningsfj0rf)ur, Eysturoy. Intertidal with some subtidal observations. Subsidiary small cove on open south coast of the wide mouth area of this fjord; studies on bedrock and large boulder strewn, fairly steep, shores of both east and west sides. Pools on both sides of cove, the west (Litlanes) showing larger and richer low water level pools and channels, protected by the immediate off-lying large rocks taking main wave impact. Subtidal of east side examined to 4 m depth within the cove; substrate a continuation of the large boulder strewn aspect of the intertidal. T I <& (0 0) W } CO O D. OO CO O UU T r _. o - Q. O Q Q_ _l "O > _i O Q o 3 O Q (TO O 5- 10- w +1 a. O 20- WL. NO algae below line even Cp absent W tidal stream 25- rough; boulder-strewn Fig. 5 Near Stakkur, Streymoy (site 29). Very steep subtidal cliff-face showing certain unusual character- istics of distribution. Detailed comments in appropriate parts of main text. See p. 157 for symbols. IU _Z Q 2 Fig. 6. To west of and below Kollur Peak (site 30) limit of _, observations large irreqular boulders Fig. 6 Below and west of Kollur Peak, north-west Eysturoy (site 30). For physical distinctions from site 29 (see Fig. 5), see comments in the site list descriptions. Sponge-associations, a facet of the biota here and elsewhere, are discussed in a special section of the text. See p. 157 for symbols. 166 SEAWEEDS OF THE FAROES -SLIPWAY^ Ul open shore profile west of cleft with sparse red upcarry from Lh depths -15 Fig. 7 Gjogv (site 31). Plan of the Cleft, indicating the positions of the profiled sections A A!; B B t ; C Q (the latter outside the Cleft, to the west). Special characteristics of biota distribution are discussed throughout appropriate sections of the main text. See p. 157 for symbols. 33. Oyndarfjoi^bur, Eysturoy. Intertidal and subtidal. North side of this short open fjord; intertidal of the two minor flat headlands and broken/bouldered flats at south end of the village. Rich intertidal flora, showing principally only moderately sheltered aspect with local mosaics of rather greater exposure, especially in damp shaded continuous wash areas. Shallow subtidal between the two minor heads showed standard Laminaria hyperborea 'forest', with penetrating sandy gullies in 2-3 m depths. Some sand-buried species, but the majority on bedrock under the forest, as epiphytes, or on sand-embedded boulders/stones. 34. Off Kuvingafjall, west coast of Kunoy, Kalsoyarfj 01^0111-. Subtidal. Very steep to vertical, ledged, rock face down to about 8-10 m; penetrated at lower depths by gullies with boulders in the bases, and by clefts with sandy bottoms. Slope eases markedly at about 8-10 m, where substrate changes to bedrock with many sandy channels and gullies, all overlain by boulders. Dive limit 16 m. Laminaria hyperborea forest present over 3-16 m depths and beyond. See Fig. 8. OPEN SHORES c 167 WL Lh Open shore profile, west of cleft luxuriant mixed red algae: abundant Ds ; Pr Phc;ClomeCn); small PC; small "a slope with red algae continues Fig. 7 (cont.) Caption see p. 166. -4 -8 -12 -16 -20 -24 -28 35. Anir and adjacent narrows, Haraldssund, Borfoy-Kunoy. Intertidal and subtidal (shallow only). Front of boathouses, north of Klaksvik. Concrete and old metal intertidal and subtidal surfaces, with small boulders buried in dark sand. Downflow of freshwater across a few intertidal strips. Potentially strong throughflow of tidal water in the sound, but shore appears largely to be a sheltered one; Alaria, Laminaria hyperborea and Ascophyllum nodosum locally juxtaposed. Deeper subtidal at point in centre of narrows just north of Strond (Borboy) and Haraldssund (Kunoy). No real forest of Laminaria hyperborea; only scattered small plants at 10-20 m depths. Substrate largely small rocks, boulders, and stones embedded in dirty loose layer of sand and gravel; occasional bedrock outcrops. Modiolus throughout. Observations between 10 and 30 m depths. 36. Hvannasund (Leiti), Vfooy. Intertidal and subtidal. Narrow intertidal rocky fringe of irregularly- surfaced slabbing, varying from smooth domes to much-dissected and stream-incised stretches carrying pools and wash-channels. Shallow subtidal with similar configuration, grading into gravel with embedded shells and stones; Modiolus abundant throughout. Laminaria hyperborea present over range (3-)5-15 m; epiphytes and red underflora dense on stipes and stones. 168 SEAWEEDS OF THE FAROES w en _ o _ t. .c 0. Q O Q u_ O Q- 8 CO E E O 0- _QlT_tllQL9__L observations Fig. 8 Off Kuvingafjall, Kalsoyarfj0rbur, west Kunoy (site 34). Subtidal profile of the ledged and varyingly sloped rock face and overlying substrates, described in the site list. Substrate symbols represent as follows: A = Li-covered, almost vertical, rock penetrated by vertical clefts with sandy bottoms and inner erosion caves. B = Boulders on detritus, with sand channels between, to 16m depth. C = Bedrock with sand channels. See p. 157 for other symbols. 37. Vfbarefbi, Vfooy. Intertidal and shallow subtidal. Firm rocky promontories of moderate slope, west side of island, near church. Draining surfaces, strong upcarry channels, and pools present. Occasional deep clefts in shallow subtidal (see station 38) and upstanding protective off-shore or bottom shore rocky knolls at lower shore levels. Freshwater downflow over some parts of intertidal. Usual major brown subtidal flora (see station 38). 38. Vfbarefbi, Vfooy; central cleft, west side. Subtidal and intertidal. Deep incut cleft in firm bedrock of centre of the promontory area (see station 37). Strong swell and wave-run creating greater scour on north side than on south. Firm steep rocky sides terminating in 7 m depth in a gravel and silt base bearing OPEN SHORES 169 Fig. 9 Central Cleft, Vibarefoi (site 38). Comparative distribution around base and both sides of the differentially scoured and wave-run affected cleft. Position of the detailed transection indicated on the semi-diagrammatic sketch plan of the whole cleft. See p. 157 for other symbols. boulders and stones. Laminar la hyperborea forest on south side; richer in cover, species and individual luxuriance than north. Narrow intertidal with patchy coverage on steep areas. Overall slightly calmer than surrounding wave-beaten coast, despite the swell and wave-run. See Fig. 9. 39. Hamarsgjogv, north ofEtbsvik, Vfooy (= Vtbarefoi, east side). Intertidal and subtidal. Intertidal chiefly of concrete steps of Eibsvik slipway, bearing rich flora; subtidal on steep rock in 5 to 7 m depth, bearing Laminaria hyperborea forest. Configuration uneven. Collections from rock and of epiphytes. 40. Marknagjogv cave, east Vtooy. Intertidal and shallow subtidal. Boat collection from walls and accessible parts of roof of large, deep, shaded cave (to 4 m above waterline); strongly swell-affected, causing continual wetting to high level and roof spray. Animal-dominated lower bands, due to light reduction and strong wave-run. 41. Havnartangi, Svinoy. Subtidal. Substrate of large boulders, rocks, and very steep to vertical surfaces of bedrock. Large boulders in 5 to 9 m depths bearing Laminaria hyperborea forest. Flora showing little variation over 1 to 9 m depths, with very sparse red flora on verticals. 42. Svmoyarefbi, Svinoy. Intertidal and shallow subtidal. Flat rock-slabbing shores, stepping and shelving to low water level and beyond, north and south of landing stage, west coast to north side of Svinoyareibi. Many dissected and draining surfaces; deep to shallow and wide pools, some in shade between upstanding knolls of protective rock at lower levels of the shore. Few deep incut caves with dripping freshwater, behind main shore line. Overall a semi-exposed shore, but with wavewash or swell most of the time. Subtidal shows usual shallow-water dominance by large forms on the mainly firm substrata. Upcarry of subtidal flora into large, deep and shaded, pools. A shore stretch close to this site was examined by Holt (1975). 170 SEAWEEDS OF THE FAROES 43. Promontory in Scafosvik, Fugloy. Intertidal only. Collection by boat from steep rock surface; animal-dominated exposed rocky shore throughout. 44. Sv^butangi, north Fugloy. Subtidal. Exposed rock standing offshore. Initially steep, fairly soon (in 12 to 15 m) shelving and easing, firm bedrock surface overlain in 15 to 29 m by large boulders of house-size. Latter lie on other large embedded boulders in blackish sandy gravel where that intrudes as patches and tongues over bedrock. Extensive barnacle (B. crenulatus) cover, reaching 100% of surface on horizontals of boulders. Laminaria hyperborea forest shows reduction in density, plant size, and epiphyte numbers with depths between 15 and 29 m. See Fig. 10. Note that the south-eastern bay (Hattarvik) on Fugloy was examined by Holt (1975). The inclusion of certain other sites e; amined at Nes (Tangaf J0rbur) and Hvitanes (Kaldbaks- fj0rbur entrance, described by B0rgesai, 1905: 694, as carrying a '. . . perfect "open sea" algal vegetation . . .') would have been appropriate for comparative purposes but for two reasons: (i) both are situated in areas close to ope vshore sites and with existing, more appropriate, comparative sites of similar characteristics; ( : both are described in some detail by Tittley etal. (1982). 4. Results: comparative species distributions The visually more impactful (level-, band-, or zone-characterizing) organisms in shore and subtidal populations are so because of unusual size, morphology, density of growth, or consistent association with certain recognizable combinations of environmental conditions. Consistency of association with environmental conditions is always comparative, not absolute: epiphytic species generally, if less abundantly, also occur epilithically; underflora species beneath the canopy are rarely consistently so, usually also appearing away from the canopy dominants, or epiphytically on the canopy, or both; species characteristically associated with particular subtidal or intertidal levels below and above Chart Datum commonly, if variably densely, occur also elsewhere. Thus, very few species are entirely intertidal, very few entirely Exposed rock standing offshore; partial transect of deeper subtidal 25 2$ Fig. 10 Sv0butangi, north Fugloy (site 44). Detailed profile and distributions of subtidal deeper levels only, with unusual characteristics indicated on profile and/or in site description. See p. 157 for symbols. OPEN SHORES 171 subtidal, although the extent and frequency of occurrence may be much greater in the one milieu than the other. Species that are comparatively consistent tend to be utilized as the indicators of their preferred situations and it is primarily such species, rather than interesting rarities, that receive comment here. The interesting rarities and their distributions are summarized alongside more frequent species in Table 4, and comments appear as appropriate throughout later parts of this text. i. Subtidal: major organisms Alaria This, generally the uppermost major organism specific to the subtidal, shows considerable consistency in appearance, depths colonized, and conditions tolerated, in the Faroes. It is generally less confined to situations of strong water movement than in the British Isles, although the richer, denser growths more frequently occur in those conditions. A summary of its detected occurrence patterns at our sites is presented in Table 1 . It will be noted that detected depths in non-turbulent situations require some revision of comments made on the basis of B0rgesen's observations. B0rgesen (1902: 450) indicated the presence of A. esculenta '. . . gregariously in enormous masses from about low-water mark to several fathoms below it . . .', but elsewhere (1905: 754) qualified this by stating that the association preferred steep or vertical rocks in most exposed places, there developing '. . . a dense covering, often many feet thick, at varying but hardly ever very great depths. Yet the Alaria may be found at a depth of several fathoms, as in sounds where there is a rapid current . . . [where] Alaria is often richly represented in the Laminaria hyperborea-association.' It will be clear from profiles presented in the figures and from the summary in Table 1 that this is not wholly supported here. Sound areas with fast currents (e.g. at station 25) certainly carry Alaria on bedrock and sufficiently stable boulders to substantial depths. But these circumstances are neither the only conditions in which Alaria occurs at such depths, nor indeed the greatest depths at which Alaria occurs. An additional qualification rests in the nature of the depth penetration. The requirement for strong water movement does not seem to be a restricting factor in the occurrence of Alaria at depths beyond the subtidal fringing band, nor does the penetration of greater depths always occur as a continuous pattern of penetration stretching down from the lower limit of the subtidal fringing populations to the greatest depths achieved. Station 4 (Husavik, Sandoy), for example, is a comparatively calm area close to a harbour wall where the only induced movement of water at depths results from the effect of the wall itself; that movement is not great. Quite vigorous mixed growths of Alaria and Laminaria saccharina occurred there to depths of 7 m, with the most dense growths being in the 6-7 m band, clearly separated from the other dense fruiting band on the wall and adjacent boulders at 0-1 m depths. Between the two major growth depths were only irregular straggly plants of Alaria. Another example of discontinuous depth distribution in by no means strongly exposed situations is provided by station 19, the Hoyvik area of detailed transects. There, the peripheral subtidal fringe Alaria growths on bedrock gave way to quite dense L. hyperborea growths at about 2-5 m depth; Alaria re-established itself at greater depths only as a clearly delimited mixed and vigorously fertile population with Laminaria digitata on stony and bouldered patches in 5-6 m depths. Even the growths at 2-5 m depth were only a few scattered large but healthy plants amongst Laminaria hyperborea on a ledge of favourable configuration; the true subtidal fringe growths terminated without noticeable downcarry well above that, at a few centimetres depth. Populations in depths beyond the first 2 m often represent only sporelings, as at stations 27 (sporelings over 2-3 m depths) and 44 (a few sporelings at 15 m, the deepest at which we detected this species). Sites with rather stronger water movement had either discontinuous depth distribution of Alaria (e.g. station 34), with the main band on steep rock down to c.2 m depth and smaller (but still vigorous and mature) clumps in greater depths (often in the shelter of gullies) of some 8-10 m, or a continuous depth distribution with very different vigour at different depths. Stations 29 and 30, both steep rock areas with very strong water movement, 172 SEAWEEDS OF THE FAROES Table 1 Alaria on the Faroes. a. Epilithic Station Substrate Depths (m) Notes 12 13 14 15 18 19 20 23 24 25 27 29 30 31 wall 0-1 broken boulders below/away (l-)6-7 from wall wall +0-5 to +0-6 boulders away from wall 0-4 steep bedrock 1-2 steep rocks and irregular around ELWS large boulders firm rock 0-2 rocks and large boulders around rocks 0-1 rocky firm areas +0-2 to 0-2 rocky firm areas 2-5 stony/bouldered patches 5-6 rocks and firm boulders around ELWS rocks/boulders around ELWS rocks 0-2 rocks 1-5 large broken boulders 0-1 (-2) 2-3 very steep firm rock forming (0-)5-9 cliff-face steep rocks and irregular (0-)5-10 large boulders steep rocks of very steep 0-1 to vertical cleft wall fertile. mixed with Laminaria saccharina. dense band 10 cm deep. band-forming. abundant dense fertile band along subtidal fringe of exposed head; alongside Ascophyllum passing into bay. Audouinella alariae on blade, dense fringe along whole slab shore, heads and coves: dense narrow fringe in wave-/water-wash; mixed with Laminaria digitata. head profile and comparative cove 2; admixed with Laminaria digitata and Himanthalia. head profile; few scattered large healthy plants amongst Laminaria hyperborea on ledge, comparative cove 1 ; intermix of well-developed fertile plants with similar Laminaria digitata. band peripheral to pools and shallow subtidal on semi-exposed head; fertile, masses forming dense band, fertile sometimes alongside Ascophyllum; occasional large rounded plants amongst populations would appear to represent ' Alaria pylaii'. some overlap with Laminaria hyperborea below, patchy, but luxuriant and dense locally, amongst patches of Laminaria hyperborea and L. faeroensis; fertile, band-forming, sporelings only. 1 m overlap with Laminaria hyperborea (8-15 m); densest in 5-9 m, where plants also larger; all plants distally tattered, tattered throughout range; most dense in lower half, band-forming. OPEN SHORES 173 Table 1 cont. Station Substrate 34 firm steep to vertical Depths (m) 0-5-2 Notes sharp demarcation Laminaria 35 36 37 38 42 44 rock boulders on gully bottom small rocks in dark sand moderately steep rocks steepish firm rock (not boulders) steep firm rocks of cleft sides rocks very large boulders over bedrock, or on other boulders embedded in sand b. Epiphytic Station Host 8-10 subtidal fringe to 2 m 0-2 subtidal fringe tol m;to+l in upcarry areas +0-5 to 0-5 1-2 15 11 Laminaria hyperborea Position stipes Depths (m) 1-2 digitata (to 0-5 m) to Alaria (0-5 m downward): Alaria to Laminaria hyperborea junction either direct or via young plants of L. digitata forming second digitata band. some small clumps in the deeps. somewhat patchy; fertile. patchy, but locally dense band; fertile, band-forming; fertile. fertile on south (scoured) side. band-forming. a few sporelings only. Notes on few stipes only. Qualifications 1. Omitted stations lacked recorded data of sufficient precision. 2. All entries relate to Alaria esculenta unless notes specify 'A. pylaii'. 3. Notes concern outstanding characteristics; they do not summarize all available data. 4. All depths are below low water unless + sign appears. revealed continuous A laria growths from the fringe at c.O m and a little below down to 9 or 10 m. In both cases, the more vigorous and dense growths were in the levels below 5 m depth, plants being tattered even there, probably as a direct result of the strength of water movement. The effect on distribution where water movement induced by presence or introduction of vertical surfaces results in upcarry of fringe conditions is dramatically demonstrated by Fig. 2, where adjacent natural boulder slopes and harbour (mole) wall are profiled for station 5 (Sandur, Sandoy). Unlike station 4 (see above), water movement at this wall around low water positions remains quite strong and, combined with the verticality of the surface presented for colonization, has resulted in both considerable upcarry of the narrowed but vigorous Alaria band and introduction of sponge-associated algal populations (see later). Laminaria hyperborea and other Laminaria spp. The Laminaria hyperborea forest was a general, but by no means completely consistent, facet of the subtidal flora in both exposed and sheltered open-shore situations. The species itself was represented at least patchily at almost all subtidal sites examined, although there were some exceptions (Table 2). It was not, for example, detected within the sheltered conditions of the 0) 0) 1 a i 3 | | o o % ^!=! t| , 00 _c 'S CO i underflora depth. ata incomp v E <-> *2 Q. e IS os "1 . 8 8 < Q, o> S > T3 "3 a *- O -a h fe *- C [I c ^ Q Q) Z ^ C3 Q-) OJ C ;> C (D pH C 5 2' - 1 -1^ 3 -arg "g : S 8-8 5 g-8 2 Z i i 1 Q U-, i-l UH o |fi C 8 ON 10 c 0) " " C 00 "g CO 1 CO CO 8 nusual data nusual data plants below c o) co co co co t^ T3 T3 T3 ~c^ -^ ^3 0) 3 3 S l; 1 1 1 I l-i js i) '^ - "iL i "Q t5 i) ^ TO io - o i .a o^ If fSa 3*s 3 -? 2 c 3 3 u oo -3 3 -3 C " c C O "7 3 e 5 ;S CO H o o o * s Z Z JS ^) C*^ CN (N e E 's" 1 1 1 > "S ' 1 JD E "^3 o Q JC> - "O *^ ? 0) o ^ oo o 1 TD 3 E > d, 1 42 2 ^^ "o S" C CO o X3 73 ^^ in co cfl rn tn T3 CO O- ^H J* ^ 00 c cO -C c/) 8,8 C o) o O Q Q ^ >> "T* T3 08 'S Q^ cs r^ " o ^2 M - 00 E 00 ;r.S fli i^ 7^ ^ linaria hyperborea on the Substrate harbour wall (boulders faced with concrete) (1-6 m) broken boulders below (6-10 m) grading to bedrock (c.8-10 m) concreted vertical wall; boulders below, or al (N 4 15 c E S on boulders) firm rock of vertical fao large boulders on eas slope below vertical 3 *" o) *" ^ >^ C 5 & Tig S-S S E ^22 o ^ i) ti o> 3 43 ^ T3 ?1 SiTJoo 3 " > '- i ^ >CcO ^S^ 2^^2^) ^ 2 l! 2 s 8g| - |.2 g | -giS^ E^^ 1) 1- D O - > ys > S cc c 3 ^ o n s c ^ 'B. .2 2 W rt H CO V5 ^ m r^ ^H ~i <-i Of C " T) 04 Absent from sto and sandy pat Forest thick abo opening to spj with depths. T3 C/3 &JQ C (/3 >-> t> <4-i C r3 "c ra 03 03 1 O a E IT) C3 "o. v _ .5 se ' .s -o >,^ ^ "> o c ' ,*i OO Q & ^OX3 t? o -o CO w 11. !)_'-' S c -o Z E V5 (JU o . 3 " ^ 2 o M "c 3 ' - aE O T ^ CO O 2 . >^ o * tS : characteris 5 eo -'i s 1 J o rt "O T *3 t^ 1 *? *-" ^ * C3 ^ ^ ^j!^ f^ JS S >;o>-o ^ o. 22 c ^ 4> t (Ut.3 w "33 3 S o.Xi c - W 3 Q g^^CJD CU t-y -^^ C/5 Z t/5 Z Q - S > CO ^ c C >v ZS o S o o O *y 1 CO *- M-i l ^o JS C^ '^ Substrate "35 1 wg-c^ S _j= OT3 ^-v.ti ^Dto^J^OIUx-vOgfe 55^^ 3 gpgf ^.2 Sv^-2 2 1 1 Sf^7 8 (so^ u^iS-Sr^ ot "- 3 52, T3 O (U O x> x> <-> U g CJ J3 C - ^ GOlC .S ^ $ 7 o g- 2 % ^ -"S J^ 8^ 5 ^ ^ o D ^-Q -35 -2 *s _ 5 S ^^TJT5^C0^2^-C: ot =5 S g o 8.|- 8 S C^ gTSo^-^O^o""^ 1 o g 8 u a o o "^ o UlS fill T3 ^ ^ 2-a O.Q Or- U on \- t3 T3 g ari w l wt lu ckin m lo ro ncl lilll o-o o OD "z. ' T3 ^ U .."N U uo =3 1) e > =s o cui:co ^cc OW>C-Oc E'en t '?" a >,^ ^" f> la ~ n i -^ la >^ u 13 u- *^ i 1 o at 1 1 ^^ C c "e j2 C II O C3 locally few plant in few locations. rare, but locally luxuriant on individual lamin Overall depth range characteristics as epiphytic; data sparse. ELWS (IT); 0- c.lO(ST). Locally ELWS BO 1 ^ <2 -* ^2 f i 5 ^2 mid to low IT pools and wet; shallow ST in sheltered harbours. i H J3 C/5 t3 as epiphytic. 5 J2 O rll f 8 in E f*! OX) -* Slgll fe"2 rl 2 1 % 1 U w o C 04 o o _^ (U Q> OJ (U C < C ^ *- t c C Ca O J- C, J El C 3 .__. . tvj u^.w y./*-^ E E tS .9- E .9- E J2J2 TD tS -S to -2 c tS T3 CS ^ O O. _2 ei 1 3 2 /tic subordina all 111 Laminaria hyperborea C 2 V; N O O -C -3> | ! a'jill 5 ^ < 2 -C | S ^ ^ -g 2 S3 !;: S w -5 H c 1 (N 1 f 1 - (S o" .'o N 1 'S. IS C/5 u u ^ 12 > 1H Ug-f'R UJ-o 2JS Uc^ ^ (/5 'a BO is X) 2 2 u SS> "O l_ rt o - c ^ "o 2 S -j3 o O 'S ^^ H 8-S 22C ro (N ^ 03 E IW 3 "~ -^ (/) r* S -~i 2 05 H i C/5 'Aglaozonia parvula' Antithamnioi floccosum 'S .a ^ |i|? < < < < < T < 3 s S 5| j-l /l^perococc^ turneri Audouinella alariae i blades LWSto LWSto probably more distal shallow shallow ST. widespread than ST. recorded; Palmaria host on Laminaria hyperborea stipes. stipes 0-5. continuous, some on walls of adding (1) to ruined Kirkjub0ur (2). cathedral; in caves; turf usually luxuriant where present. f young only mid IT mid IT pools/ more widespread than n to3(ST). wet, to 3m. recorded; terminal hairs occasional (16B); dense branching occasional (15). lower IT/ as epiphytic. well-developed plates ST fringe, present; probably in more widespread. exposure. top shore as epiphytic. probably more to widespread than n lower IT; recorded; 1 -layered shallow at base (15). ST. tips mid to as epiphytic. massive sporangial lower IT; growths on E. shallow compressa (16B); more um ST. than one undetermined ?fl species involved in these records. \ 1 3 a s * >< o. s: 5 Palmaria palma Cladophora sericea Laminaria hyperborea Phycodrys rube Ceramium rubr Callithamnion sepositum Pilayella littoralis Ectocarpus fasciculatus esp. Ceramiaceae Lomentaria orcadensis Porphyra umbilicalis Ceramium rubri various other sp Plocamium cartilagineum Ceramium shuttleworthic Cladophora seri and rupestris Callithamnion sepositum Enteromorpha compressa Antithamnion floccosum O .*S to - J/Tra u jfl 4= a u> u | S o 3 ,0 '5 c rt CO ~ ^ ^C "*7* T3 > O -o OJ .C ~ *- . g- < 'B. B- 03 o V |J 'S -3 .a .h 43 . o. -a . as epilithic; IT mostly in shad r we lit st o ps; onti fc.l iii o a > ^ CO 3 _i 5 .S .s ^Z? '* 2 H 2 E I S II -2 T3 ^5 43 'IE! Ifi . -a^'u u ^^ n * s "* .a S u w s "8 1 1 1 1 1 sj tl 1 1 1 s | ^2 oq 03 CX a OQ ;2 "^s I -.a 1-22+ . often best mostly with developed in Callocolax, although d middle depths latter patchy; often (5-15 m). more epilithic than epiphytic, when present together; epilithic plants in depths (as e.g. at 29) sometimes small and resetted. lower IT continuous over detected only in and shallow narrow Torshavn area and ST. amplitude. adjacent fjords as- new records; not abundant but possibl; more widespread; epilithic plants with and cf . ST:5-7. continuous ST4-8m; through depth cosmopolitan to the mid to range (upper island group; very low IT; mid IT to ST common as both pools 16m); always epiphyte and and wet. in wet places. underflora. ST: 7-16, in sandy conditions. mid IT continuous over probably more abunc pools and narrow and widespread than wet. amplitude. even recorded; vast swards at some sites (e.g. 32). % ^ 4> tS >. CO ffl CO 1111 1 ts>b.s Ji laminae laminae laminae stipes and laminae . laminae laminae laminae w =1 S '3 3 J i-t 1 saccharina . faeroensis ucus spiralis vesiculosus aminaria hyperborea orallina officinalis ucus vesiculoi scophyllum ther fucoids olysiphonia lanosa urcellaria lumbricalis orallina "3 S u 1 -j -j -^ "- it. ~J u (*, **, o a. tt, O. *- \ CO C/3 ^jocC.-DQJug gw o w' 3u- > S- _: rt w -Sas-Sb3^2''^? c-in>^-i;_es2^i-> -^^ Soo2uo^^-s2 o (C JD M oo-O o O > ^~- u-. J2 O eo i/T D H *- L- C W 6-'E S o 5 111111 2 E 2 * 111 1 1 'i ~" ti ; d epilithic/epiphy status doubtful; .s: C" oo O _c W ^ ^ CO a g -33 o a. %j M ce 1111 ^ 3"? 2 -%~ - S -5 S w E B4.SJI * 3 O o C 5 3 SO g o Ci. .., O u a^ 8-d c w >-, -0 open ro omm mos flora tme sce ; rec 4-6 plants; iride sites 20, 23; E-sl O "u C/3 - 2 -5 "3 C ^ ^ a-g u^2 a "0 o U _C C O T3 "tS 3 O. c 8- U CO TD -a 3 j= C C X) e > ca co .:; C8 r S fe 2 a 2 c I sifeiii e , c o < ^ o .2 a, =3 '55 o ? in O ^ t *j v g" s M s i . 3 fi 4= _) -S "s rarely detected: probably more in C o W Q. Cfl o eo o tca5 t: C - "I fei i -S a M oo JJ S e "2 a.S = C m ^ rt r^ * U O OB QQ f'^'22 5 r^ b 5 o So W5 C -^1 s O tn 3 > O -g O J= C H t^ t^3S g.S^g.8 B S '6 = S-= '5 3 e r ,S CQ *3 c 73 -S E.S C C C C C 1 1 1 1 I % = 2 ll s -a -c .& 2 S C -Q S t 5 ex. if q _ 3" . 5 -ti ^ i fl> 3-2 % S -c fc- O *'*-'' u O ^^ - g i- 0..5 T3 I TD O 5 ff^-|;s,^lss ) 7 o>oJ3 G^^o^J2cu.SCSEJ2iA OT >. .a 3 P a h o -o 5 c g >; M .ts -o .S = J3 < fc U ' t 8 3 8 . 2 2 S .S * e *2 S s .a I 1^1 1 11 1 s | 1 .2 i o 2 -2 o - ' o i/i I T3 C " O f= O 8111 liilli - 5 a, f oo I-H "2 < =3^ ;r ^^ s* . c a 1 -S .s TS^ 4^ l 1-itnCGcQ.H C~^^ r*^ c i! 2 > Q x = to > ex u 2 - Q j= c^ .5 on -" 2 o c so 3 |J5 rr /*! UH T^ debble whether first animal or on often few to S o cs ij I A 3 C w JB -a E g C S . 1J g .2 3 o u 0. O.^ T3 TJ O 3? 1> 2 V T3 3 IT < IT i *- ti E i) P - r" D " I els o c 3 "O *- XS -C 3 CS w ? js o. O, U > s^-g sfei^ gt - eo r^ lb-52 nil I '8'r I UKafJliSsS UOOT3Mto t O S < <2 S 3 i ? ' S b ^ o y -P a 11 rallina fficinalis ei a .2.2 co ha "3 O llllll 5 EX t ^ ^ K l^s>^ II ^ 6 tS a o 11 1* -J ll > ^S 3 Si .p s? 2 = e03T3i3 2 a c o i_ . O tS fll ^~* r~ : g-s o c I tly found equ pla 19 CO Q U - sporadic, mo drift, as liste by B0rgesen; oderately fr Rex; single takkur in os d epilithicopen, not usually under-flora; probably more widespread than recorded (cf . B0rgesen); abundant and luxuriant locally: a ^_ -~: f^ O 00 15 -H la *"> ^i t/) *J in river estuary at fjord head; single location with plants requiring further confirmation; cold and murky waters. predominantly in fjords, enclaves and sounds; few plants only at some locations; usually in turbid waters (e.g. harbours) fertile (unilocular sporangiz sometimes. data inadequa for assessment C w ' .- s H S 1/5 c ca s "3 ^"2 38-8S to a. g ~c, u^-a ^8 b lni II 11*11 oiilfesl d) o 1C JS jD X) OX! o *s o c o . II ^ S E Jo 6 CO u 1 jfi^i?t.J* ^ a ^ oo^- C w c o S C "O . , 4> ' c - O G,"O ^ Z'a. c .t; a ^ ^ s ^^ ty . ^ jl O "G OO C "^ S*O ool *t"SinJJ(uSk- s^oosgcg-rs^ .S v -i/3-*nl .S 1 1 s a js - S g||S a o ' M C 1) t/i <*- O. S sssstj&se-sacjs *0^ In ^ JS s"- 3 e > >^ & ce *) y5 X) >?^i c o E3 " 2 ^ > ^ "O " "ffi -2 o ^ '3 3 S2 ^ - cQ C J2 ** O s c/i TI t" H 3 CJ W Ic 3 ^ o 2 O .^ ^ ~- "^ *u ^^ ^0 >^ >^ pfc Q "^ +-* ~^ Is 1 in ca TO l-( a "o u eu (l j-e.2'o. ~ s 1 ~ s i . g. a 5 S 2 &H T3 T3 O. -Si 3 C S S gt . ^Si,^ E S-=^2 > vi-i in O o c c ^ " C U O ^ 9 a. > o TO a j L. faeroensis i ^S? in PN u ^; o . P JD 1 H u 1 " C - c a ^ C3 f-H C W U IM B o-5 E 2 <- o "3 uT i> 1 1 a.5 8-S 8 lllii I* > in u a u c u = .-"c ca 2 "^ . r, QO O CA m 1 1 f[-; cQ a c c -5 u C O O u. 3 . o o CS in CJ 42 M { O .tj -2 ^ g -5 o 5 2 to C > 2 3 >>4= S . - "c .& e fc -a _r o a. o , ^2 g s I 1 S 1 1 s 5 15, llittlsl tsl S-lt's.sl.aB 8 < 8l ^^ g-E S ^^= " 1 ll^lllll S..E 8 o S-^i 5 S: '% &K U $ ; 1 ! || l-glal 2 S^ g fee 1 -al a=s^g.^ 1 gi |lllll | la iSit^J | U u- 8 * = ^ C ^5 JJ^o.E&.ES not clear whether calcareous surface of Lithothamnia really 1 0.1 8.S2 S 1 |l 8 11 ^ HQtS co CO CO cO a 3 T3 'CL,~O c TD TJ O .E data for clear assessment; masses at 15 m (36). ST fringe in ll 1 S2 w ^io^ 1&fe|||l CO C O C CO *- i/5 43 4) iJ C O 2 I^ 1/3 CO '/i - ^- ^4 t CO see epiiithic range column; locally continuous; very abundant, luxuriant and widespread. = 4= JQ o a O. &o ii i E 1? C 00 c o o E U T3 c '7, Tio g 13 ca E c s H" E 4> 4> CO O 1 ff ?*^ Si " u w O I- O oo u T3 C <_, &. O V] o JS c C 1 Q -J ^ 1 -3 -j Q^ 1 ^ i-J 2 u c oo 1 u c ~ 1) CO O I/) U- CJ bljr. Hll 2 S.ts ^ 63 oH o &2 a .ti 15 i/i T3 s If i 2^-8 a E >3 c a e pr ria -2 o vj .- E 2 c co o & u s ~ o o ' S -fi ~co c* 3 -s <> c -n 4) .S E 2 2 S ~Q s O -S r- U r- JT oo5 2 3 5 2 w u o C i-T o , a ^ ^ Q> it-s'M ts*l T3 U eg .2 *^ "*^ CO - i 73 CO <- ' ^"S g-3 I' i"S a 1 1 1 I 1 1 - S * I . ^ " 5 c "o 8 2 w g o D E O" Sft T3 CO CO t- .S ~ 0/5 w 2 T3 CO - C ,O ti u |^6 | I 2 g ! | If II 1 5 *o o QI ^-^ fc 0.2 "cO O 2 S "2 ? El a _x -S Mj5 s u Q If I O J 1"S E t H r HH - - - o .. 1 v- T3 El C "> - 111 1C to JO .0 E2 .2 S ^ S ?^ o 2 S 22 a: cr 3 o c S o often abundant and/< luxuriant locally, but not frequent; usually as shaded underflora or aj a. 'a. lll sA 3 j : " S J-'S - S : ^2 ^ +0 apparently relatively intolerant of scour; prolific locally on vertical slopes with sponges; sometimes ; L. hyperborea- underflora. specimens in exposui or below laminarians in depth are smaller; sometimes codomina L. hyperborea epiphyte in c. 15-20 m; epiphyte underflora to L. digitata in shallow ST. identified by E. M. Burrows. o T3 o 55 C ii ( W , 4S ta . . C/3 E 3 CO ._ u . 'e3 i O ^ L^ C O [__, o-^ O Q> > C ^"* i/i -! % O O ^ <3 ! - 2 ^ '3 ^ . 3 ^' ^c/5 u to t! u ti .2.0 discontin || ||| continual locally mi or stepwi distributi oT ^ u S g '-5 |3 llsl^si .B-'C j a S > S 8-^loJ & 1 r, | &$ c =3 ^ d, HUES locally continuol ||sfci4fl^fi i f 1 K 1 l-sr O C % < C i (S) ^ 2 8^.SS^ i cr u CO -O 1.1 T3 "cO 13 13 : j C oH S c c (U o J= E-T-S "cO C ^ O ~* J <3 .a ^ ^ to c a V Q C -13 K r\ "^^ *t* ?* *** ! n *"^* Si, "*"* ^ "d o "^s C r\ "5^ R """* "^ ^ . ^ 1SP Q ^ cs J*"| ^ ^ ! a 6 ^^1 1 a. ;; J"^ 351-1 1 c^ (N -If voi 2 1 | C =3 PH "cO '"* "cO CO c/5 l-i 1> C fe G 1> C U c i* j o a! M 1 o a 1 == (U C V C (U CO O 95 h o - si 8 S 8 2 8 *-s t. o D u% if T3 | 8 = u 2 a u ^ c y -s . _ Cy TO uo <* J< ( o u u T3 ^ 5 ^ c if ta J 111 1 - cc J2 c -( _ ** M el . a .^ " -2 '^ s O Q a -5 is -2 CS ^J ^ s s z -s; c- .|--| 2 -2 11 11 -J c"| *> ~ S-S ^ U o o jl p very rarely on Dumontia; young whol 1 sacs not infrequent, e; n. recorded from sheltered conditions in fjords. perhaps more than one taxon, since 'hosts' differ. inuous o _>, estricte< oo e c tu S 2S. S | | co T >> a* .-3 JS U U CO a- c o o 13 8 CO T3 g-sH-l 31 CO CO a CO . -2 o c CO CO oo o 59 fc ai CO (U -O |s T3 00 | ex J CJ O -C , ^ ^J 18 tu j? ex 4J H .1 H S> 5& " -C H ex c 2^ S ts .S & o o H 'K y? E 11 S E? E 5j u ^ "1 e en Q S O B-2 5 . cT "c '" E ''co 8 2 - 111? S?,sl, CO 0/5 X) ? a J * a "a. fill 1 as host, most probably. 1 11s |i!|II o 'ex S e A 'ex u u eo.3 ^ u continuous over ranges; details, see range columns. 2 Q ^ as^ .0 . 8,-S.S^ |L g j J2 7T (/j litib S .1 2 3 E j 'ex a T CO (U XI O "O M C/J "O / * xT 1 <-" ^ ^ " - _2 3 >>" *~ > J3 e o .y ex n E -S 'ex o 10 *" C 2 Life's U C i- S.S2 > oo'-a o 8.1-02 8.2 ||S ^ <*- C O O O CO O CO -C v: -C s. w -C s "a a S e a v. O" 3 G 3 1 3 o c? S I -283 a c 3 ^*. ^ a a I Is. 2 -^ til |i| a. a. Scytosiphon lomentaria minaria yperbo .2 o 8. o E - 8, 8 a 1^. O xi H V . g o -5 - r c = ^3 ' c -"0 -5 c n. " O ^ OS ^ o ac I 1 occasionally as dense and continuous c "S % '> o | wl-ii H ji 3 I" '-3 .y 'C .0 o o O O *""> :.*43 2 43 OO c .H o O.-B S3 .9 2 g C ** > ^ 2"T73 C rz D ^ o LH ^ ^ "cL'cL S "cL o % E a^Q _ a -<=~ S 1 S !| 1 J a C^o 1 ^ n-,.2 C t/1 -" lS 3131 j! |i|lf Biifililii'll si! Ilitlril i3^yu5 8 I! S T3 -g S ^ 'C 1 J 4 f i S a 8 8^^ 8 s a "O * 3 : B / a E E ~ C "3 c'^-S , ^^ (N D (-; u^E E: l > ^ fe 1 i+^2 go 12 H 1 43 ^ o.^ "S ^O.Oct?C B.C C?^^ O O > **J ^-J O e*"^ a O a w llis'l 5 ex, to oo "s 1*3 i 1 "J "-J So 1