5 Si c 4 ann eaerei if yisited Hote re att ratty Heees aa as Rares co =< es Satine Sore ate teh Aid Seiriest 7 sity icieteaat Gant an seed ; Wiehe Dbgat 4 ie ae oe S i nitat if ie ; bois WE i f LS OR Se vised. ¥ ‘pass ot 3 ¢ we th py ayy Sc PSE ees Poroetaeatit yt tN ih) rane ae \ Pee SHEA athe aa ] Redatssatt epep we (3 iy iete ss tee ileal eal ‘. Shal het r 5 a RNs es ait) x io 7 t;) ie fi ty a i 2 Neste inks a6 te e vist i Be ie ve hie, * aaa Risch Aas ie aks of arate es iy VAM cee rca wo arpa’ Re aie i ft ath Pnipat aaett — Sees me EE. Je oe Kes. Wactaidieteseia haa ie Ae ata * 3 Bi potatoe (A Wine iit eee 46 rat Ro i det g it} ay + *. ee pits 7 Siena ei Ape ey e rh eo ELS Sar one ot. A. ss . See = ore yack, sehese Kb} se ay "1 tee 7 eee ee, Sores > Seen cs * Ss Peer soce= see ee sia o5 See f tts A a i iene % ns Ae Ray as MiCeY ae rps eG ENO mites aed yt Aes, my bt) : ee AD iaciind ee a fOr fe a A Sonne ee ss > ral ee ‘ EM esa . oe hase es *! wheal: ¥ GCA iG aS S ag hone ed eet peer ¥ Sees eae re mh im Ae ae hayh soe een ui af ee ity war Re is x Rees eras ~~ a volty Mee eee Sp) : ray a Eh oe Se em @ WAS 4AGH N68. The {ible pose ohed comlinta page yt, 10 hore le2n 4 . rN te 11 4a ru Bulletin of the British Museum (Natural History) Taxonomic studies in the Labiatae tribe Pogostemoneae J. R. Press Botany series Voll10No1 30 September 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 SW7 5BD, 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 Ast Moe iS N f GENERAL ® \ OCT 1982 @ LIBRARY - Taxonomic studies in the Labiatae tribe Pogostemoneae > af » J.R. Press a Department of Botany, British Museum (Natural History), Cromwell Road, London SW75BD Contents LL PRED ARUP AEE faye NEE PE OPER ENTE N TE CETL ST TN OM RAR ASE BAD p TORN BARON SoS 2 MSN MAO pnd sits Vb vp ose aca Leadon aah Seep bcd eoaaneeoel Memes Met etacunn ees + ¥ ae MCRL CO VIO W oo o5 553 suger tons Aes ein tasind sd ysoaae a saaaey ean eclnGs ites obutbewsieelenciobac ts “ Taxonomic position of the Pogostemoneae..............5:..00.ccceccsescoesscasssoases 4 Previous taxonomic treatments of the Pogostemoneae ..................0..00eeeeees 5 Pe RR EEMION eh oon ey vey oes dacees EE eae cau Cet ex wh vcore eed audsa ean es ect whos 8 IRON EeCAE RIOT OF HIAUCIIAN cee SAt bd, foun evan hk ooanin ae aes rt asaeees a 8 MIMI COMMPUA OR cote ei oe eee Alc cars wu ca ap ytei cancoccdans aia eaten. 8 PACREATC RIEU OLCHALACIORS 1358504 ote oee cgeh thi sea raver ens aecideeee: 10 PTR ROT AIS 5 oc codecs te eee tet anc oee eee s dee Sue rereh aati aide cede Gonmasege 12 MM AAEMCUET VATIATION 560.05 cero vine penne secs aet coat canes uhien aNRGautel weave shoes 12 DAVOS eases cc. ons sunsneae hese EILat ec bee OT TMG eave secant eee me caneaeeee eta es PICLOIO IV UN soc coca est ea iaaninc gee ce oe Rega g oceans cvudswasv eine eee Te eee nee 12 j MEY gts « Raat oe re ACEP TET SET ree eer eye TERT emirate, test ie denbr Neer cae I. BIVVMOCMNIG cess corevn igs hy ton ke oe ce eacea rouse shea nemnea ke peg tones soeiGestestasa et oneal 12 iii tt) | Mean PEO ge Ey or ee AS EERE EE YET CRT eee EN Fae sgeapen mecha setae vox be | 6: =, SR eer RRR epee ORT Ar CO or ee ee i |S nol Cort (a Se prea ARON OE EE PRON. CS REAR ED Ee RUE SOAT CEE TT OE (aan meat ACRES eer ty PGMA TLUED 2 cee cos rae ct rot for eae ns canna tviey serene reiocs tan loka deuuenute ceemtiuee sae 17 GT SPR Ranen BPO SANE van Re eae Mmiy Uae Ninn CAs Seri eaMed a Aa TET T IS Sy Mer aA ree 19 Goo | aa eee ie RR teenies Aeknian) gaieryed Ten Rear ieee A nen Gn ey tne oe er a 19 SAI H BSCE O DEY ete phe PRC Ties CPN rey Rey ECA POP EP EP DR APY DEEP REU PUREE RC rt PLLA AIA 22 oe OSA ae TCO R Ran We Rie an om Aan :130; hildebrandtii RSIN 119. sinensis TFRU 131. fruticosa CFOR 120. Comanthosphace formosana KJAP 132. Keiskea japonica CSTE. M12), 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 GCOPP 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 Characters Type 1. Leaves : type f 2 rounded 0 homophyllus 3 truncate 1 heterophyllus 8. Leaves : apical shape 2 2. Leaves : petiole length 3 0 acute in millimetres 1 acuminate 3. Leaves : phyllotaxis 4 2 obtuse 0 opposite 9. Leaves : margin 2 1 verticillate 0 entire 4. Leaves : number per whorl 3 1 dentate 5. Leaves : comparative size of 2 2 doubly dentate members of a pair or whorl 10. Bracts : type 2 0 equal 0 not membranous 1 unequal 1 membranous 6. Leaves : shape 2 11. Bracts : shape 2 0 linear 0 linear 1 ovate 1 ovate 2 orbicular 2 at least as broad as long 3 trifid 12. Bracts : fusion 2 7. Leaves : basal shape 2 0 free 0 cuneate 1 attenuate 1 connate for at least part of their length TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 11 Characters Type Characters Type 13. Bracts : persistence 2 1 white 0 persistent 35. Corolla : colour Zz 1 deciduous 0 not yellow 14. Bracteoles 2 1 yellow 0 absent 36. Corolla : colour 2 1 linear 0 not purple 2 lanceolate 1 purple 15. Bracteoles : length in tenths of 3 37. Corolla : annulus 2 a millimetre 0 absent 16. Calyx : symmetry of teeth 2 1 complete ring of hairs 0 regular 2 interrupted ring of hairs 1 irregular 38. Corolla : invagination 2 17. Calyx : length of longest tooth 5) 0 invagination absent in hundredths of a millimetre 1 invagination present 18. Calyx : length of shortest tooth 3 39. Corolla : shape Zz in hundredths of a millimetre 0 corolla not gibbous above annulus 19. Calyx : annulus in throat p 1 corolla gibbous above annulus 0 absent 40. Stamens: exertion 2 1 present 0 exserted 20. Calyx : number of main veins 3 1 not exserted 21. Calyx : tube length at anthesis 3 41. Anthers : size D4 in hundredths of a millimetre 0 all anthers equal 22. Calyx : tube length in fruit 3 1 lower pair reduced in hundredths of a millimetre 42. Filaments : length 2 23. Calyx : tooth form in fruit 2 0 equal 0 not plumose 1 lower pair longer 1 plumose 2 upper pair longer 24. Calyx : tooth position in fruit Z 43. Anthers : locule number Ps 0 incurved 0 unilocular l erect 1 bilocular-fused 2 spreading 2 bilocular-free 25. Calyx : teeth hairs 2 44. Filaments : hairs 2 0 absent 0 glabrous 1 present 1 hairy towards base 26. Calyx : tube hairs 2 2 hairy towards middle 0 absent 45. Filaments : base 2 1 present 0 not bulbous 27. Cal: angles pe 1 bulbous 0 more or less terete 46. Style : type 2 1 strongly angled 0 gynobasic 28. Corolla : symmetry Z 1 terminal 0 lower lip with 1 lobe 47. Style : lobes 2 1 lower lip with 2 lobes 0 plain 29. Corolla : division 2 1 clavate 0 upper lip not emarginate 48. Style : basal shape 2 1 upper lip emarginate 0 not bulbous 30. Corolla : tube length 3 1 bulbous in hundredths of a millimetre 49. Disc: number of tumescent glands 2 31. Corolla : exsertion 2 0 none 0 exserted 1 one 1 not exserted 2 four 32. Corolla : length of upper lip 3 50. Nutlets : hairs 2 in hundredths of a millimetre 0 absent 33. Corolla : length of lower lip 3 1 present in hundredths of a millimetre 51. Nutlets : surface 2 34. Corolla: colour 2 0 not verrucose 0 not white 1 verrucose 12 J. R. PRESS Characters Type Characters Type 52. Nutlets : apex 2 1 sparsely hairy 0 not rostrate 2 densely hairy 1 rostrate 59. Indumentum : abaxial leaf surface 3 53. Nutlets : number at maturity 2 0 glabrous 0 one , 1 sparsely hairy 1 four 2 densely hairy 54. Hairs : septate, eglandular 2 60. Indumentum : stem 3 0 absent 0 glabrous 1 present 1 sparsely hairy 55. Hairs : septate, glandular 2 2 densely hairy 0 absent 61. Inflorescence : number of 3 1 present verticils per spike 56. Hairs : branched, stalked 2 62. Inflorescence : form of verticils Z 0 absent 0 not secund 1 present 1 sub-secund 57. Hairs : branched, sessile 2 2 secund 0 absent 63. Inflorescence : number of 3 1 present flowers per verticil 58. Indumentum: adaxialleafsurface 3 64. Flowers : length of pedicel 3 0 glabrous 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 Tetradenia). 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 cehe 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 Xx 1. (h) Colebrookea oppositifolia x 0.5. (i) Elsholtzia ciliata x 1. (j) Elsholtzia densa x 1. (k) Dysophylla stellata x 1. (I) 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 goudotii Briq.). 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. 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. Bulletin of the British Museum (Natural History) Botany series Vol 10 1982 British Museum (Natural History) London 1982 No 1 No 2 No 3 Dates of publication of the parts 30 September 1982 28 October 1982 25 November 1982 23 December 1982 ISSN 0068-2292 Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset No 1 No 2 No 3 No 4 Contents Botany Volume 10 Page Taxonomic studies in the Labiatae tribe Pogostemoneae 3,0 Tess : : ; : : : ; : : : ] The typification of Hudson’s algae: a taxonomic and nomenclatural reappraisal L. M. Irvine & P. S. Dixon : : : : : : ‘ 91 Seaweeds of the Faroes : : : : : : : Or The lichen genus Steinera A. M. Henssen & P. W. James : : : ; : 7 8227 TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE m4 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 Cyclostegia 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 gracilis, 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 Cyclostegia, 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 J. R. PRESS - i \ ) f = 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. Elsholtziu 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 i Sey ee LI ae a: SSS ISS < SST oS 2 SEES =e a LOD. — > > BOR mae: S LSS ae C7 a4 fees ORs ~o ace. 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 5S. (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 21 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 xX 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 flava (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 FL BRS 7 I Fig. 8 Stamen types in the Pogostemoneae. All stamens X 10. All anther details x 20. (a) Keiskea japonica. (b) Colebrookea oppositifolia. (c) Comanthosphace sublanceolata. (d) Elsholtzia ciliata. (e) Leucoscep- trum canum. (f) Dysophylla linearis. (g) Pogostemon parviflorus. (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. Number of nuclei Number of colpi Genus per cell per cell Elsholtzia Keiskea Tetradenia Eurysolen Comanthosphace Leucosceptrum Rostrinucula Pogostemon Dysophylla Colebrookea WWWW WWW OAD NNNN!I NI Www 26 J. R. PRESS \ y 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 Zi 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 Léveillé (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 14-16 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 14-16) 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 (Léveillé) 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 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, (8); Colebrookea, ©); Eurysolen, ©). TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 29 vector 1 aa A A AA ee . A TA é ? - 4 22 a (@) A A 4 Be $ On6 A °. © ee 4 4 = e cee oe = ee oO Brads © e , eS MOG ® Ou © 4 r) 7 ee 00°, ©) @ @ . One ®) A © eo a 4 A eo? A @ ‘ ‘ 7 ©© - A ¥" ) . A 4. ee P a, Pd ® 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 O P A & A A A @@e@® x aad a See A S A e@ A A © a e A A & Oo @® A @ © %e A A 008 08 0 4 4 @ 9 €020° rN © Cel 0 ee Se O fo) aaAAaA a @@o QO A ane nN a ¢ 0 €°o A AA OOO A O meee 2 eo) O A | A 22 ®© wi re (R) aan Bf & vector 4 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 Ss vector 1 a 65 AA Ve O A veh ry A A A a aad A Q) A a A A A 1262 x AA A A re A oe @) ee @) Sei A A 4 2 @ A A A A A AA A 127 Sain AA A @ A A | a a aa so o® 126 | | vector 2 Fig. 14 Two-dimensional plot of the Comanthosphace/Elsholtzia subset using principal co-ordinates analysis. Key: Dysophylla, O; Elsholtzia, A; Comanthosphace, @; Keiskea, A; Tetradenia, @); Leucoscep- trum, (); Eurysolen, ©). a2 J. R. PRESS vector 1 126 127 128 A A A A A A AAA A AA A A A ® aA 4 | 2 a “ a | a A A A 2 25 A A AA A vector 4 Fig. 15 Two-dimensional plot of the Comanthosphace/Elsholtzia subset using principal co-ordinates analysis. Key: Dysophylla, O; Elsholtzia, A; Comanthosphace, @; Keiskea, A; Tetradenia, ©; Leucoscep- trum, ©); Eurysolen, ©). TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE oo vector 1 128 2 65 127 A A A A A A . 4| A ad 4 A A @©4 @ e a r A 4a a 4 AA i | A A A A A A A A tl = = A A A ® A & A AA A A A A AA A 126 25 vector 4 Fig. 16 Two-dimensional plot of the Comanthosphace/Elsholtzia subset using principal co-ordinates analysis. Key: Dysophylla, O; Elsholtzia, A; Comanthosphace, @; Keiskea, A; Tetradenia, ©; Leucoscep- trum, ©); Eurysolen, ©). 34 J. R. PRESS VeCror 4 A @ aA A A 20 A AA A A x @ @ @ / O ee @ @ O e@ us, © eo @ e) e) 21 vector 2 Fig. 17 Two-dimensional plot of the Comanthosphace/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. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 35 vector 1 21 20 19 A ® Oo A oo A A A ®ee@ a e) C@ A A 8 AA a e ® A a 4 fe) AA A A A A A A vector 3 Fig. 18 Two-dimensional plot of the Comanthosphace/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 J. 36 J. R. PRESS vector 1 8 19 20 a e a A A A & a @ A A . e A A ox ) A e) e® @ A A e () ee A 8 a A AA A & A oe) A 21 vector 4 Fig. 19 Two-dimensional plot of the Comanthosphace/Elsholtzia 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 37 vector 1 | O oO 7) es ee ee @ es @ a e@ @ Oo @ a2 e oO | @ OO0A4O 61 6a e 00 CO OO ay 70 53-° Oo Oo e e refone! | eee ee i 69 e 59 O O 58 /O @ eee iat iat 3 6 2? $ 66 109 169 85 e e e ®& ry @ Po @ @ e je 82 e Ce vector 2 Fig. 20 Two-dimensional plot of the Dysophylla/Pogostemon subset using principal co-ordinates analysis. Key: Pogostemon, @; Dysophylla, O; Elsholtzia, A. J. R. PRESS 38 vector 1 e @ O oO @ @ e) 2 ® e % oe fe) & e So ee ee O = 68 e e @ © e) =o * ; * oO 62 (SH © ca ag O fone) 67 & x oO ", 85 e @ ae & 53 2 _? re) | @ ee @ ® % 43 e 59—O “oO p aes . o—69 110 109 é ee 55 —O sge6 a 86 & pi iS 58 ® 16 BS. . 66 82 Cd Q — Vg 70 71 vector 3 Fig. 21 Two-dimensional plot of the Dysophylla/Pogostemon subset using principal co-ordinates analysis. Key: Pogostemon, @; Dysophylla, O; Elsholtzia; A. vector 1 22 O O © oO TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 39 vector 4 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 Pogost yr Dysophylla YD LL, Elsholtzia Keiskea LA 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 canum Pogostemon Dysophylla Pogostemon Y Elsholtzia section Aphanochilus series Stenelasmeae WY Elsholtzia sections Elsholtzia and 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 Z 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 Te? 87-9 6 80-8 78:8 70-1 67:7 76:4 88-3 | 1 2 3 4 5 6 Key to groups: nee 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 85-8 2 67:9 90-8 a 70-3 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 W 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 Diss 76°5 79-3 9 72°8 73 76-4 70:8 72:2 78-3 73-0 68-5 86-7 1 2 5 4 5 6 7 8 9 Key to cote Be 1—Comanthosphace, Rostrinucula, and Leucosceptrum canum. 2—Keiskea. 3—Tetradenia. 4—Dysophylla section Verticillatae. 5—-species of Pogostemon and Dysophylla section Oppositifoliae. 6 hiholezia 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 88-5 2 69-5 90-8 3 71-3 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 heed 69-4 95-2 9 73-9 77:3 76:4 70-8 72:2 78:3 73-0 67-1 86-7 1 2 3 4 5 6 I) 8 9 Key to in 1—Comanthosphace 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. 9—Elsholtzia sections Cyclostegia 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. J. R. PRESS ot Se a i fa eS Bf fee eg O62 O18 O€B8 OSB O48 068 O16 O'F6 OSE 046 066 0°08 O'¢8 O'P8 098 088 006 026 OE 096 086 000! 45 TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE “SALLO 8€T JO} WeIsoIpusp ddeyUT] a[SUIS Sz “BIY 46 J. R. PRESS Leucosceptrum canum differs from Comanthosphace in two similar characters as C. nan- chuanensis. Bracteoles are present, but there are no annular hairs within the corolla, and the nutlets are glabrous. Also, it has a terminal style whereas all species of Comanthosphace, including C. nanchuanensis, have gynobasic styles. With the exception of the style character Leucosceptrum canum is scarcely less similar to Comanthosphace than C. nanchuanensis, as shown by the similarities in Tables 8 and 9; thus Comanthosphace might easily be expanded to include it. Table8 WGMS and BGMS values for Comanthosphace, Rostrinucula, and Leucosceptrum canum. 1 93-7 Z 81-6 94-8 3 84-3 Tee 100 1 Zz 3 Key to grou iS: i Caaaiios hace. 2—Rostrinucula. 3—Leucosceptrum canum. Table9 WGMS and BGMS values for Comanthosphace, Rostrinucula, and Comanthosphace nanchuanensis. 1 93.7 2 81-6 94-8 3 81-6 82:6 100 1 4 3 Key to grou Ss: 1 -Conaniss hace. 2—Rostrinucula. 3—Comanthosphace nanchuanensis. The gynobasic versus terminal style character is considered by most authors to be one of a number of heavily weighted characters distinguishing taxa at the tribal level. The analyses used here are all unweighted methods giving less consideration to characters of supposed high taxonomic value, such as those of the style, and thus raising the similarity between Leucoscep- trum canum and Comanthosphace. Many authors unquestionably accept the value of the gynobasic versus terminal style characters, especially in studies of the Labiatae and Verbe- naceae. I remain unsure of the absolute discrimination of these character states because the style type and nutlet attachment are not easily or clearly interpreted, at least for some species of the Pogostemoneae. The style character is linked with a nutlet character so that in the gynobasic condition the nutlets have a small basal attachment scar; in the terminal condition the nutlets have a large lateral attachment scar. A terminal style and lateral attachment scar are a characteristic of the Labiatae tribe Ajugeae, and on this basis Leucosceptrum canum is usually placed in that group. In most species this pairing of style and nutlet characters holds true, but in others the link appears to break down. For example in Elsholtzia densa (OTU 20), (Figs 9, 10, 28) the style is gynobasic but the nutlets have a relatively large, somewhat oblique scar; in E. flava (OTU 28) the style is apparently gynobasic and the nutlet scars small but clearly lateral (Figs 9, 10). In Leucosceptrum canum (Fig. 9) the style is not always obviously terminal, sometimes appearing to rise almost directly from the disc. This divergence from the defined condition is reinforced by the nutlet scars which are small and basal. In this respect they are very similar to the nutlets of species of Comanthosphace (Fig. 10). TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 47 The nutlet characters may have been wrongly assessed in Leucosceptrum canum, a claim which has already been made by Kitamura & Murata (1962). However, they failed to take into account the apparent disparity of style characters between the tribe Ajugeae and the tribe to which they transferred Leucosceptrum, the Satureieae. A more detailed study of gynobasic versus terminal styles in a wide range of species is required to assess the variation and importance of this character in the Labiatae. Should the style character prove to be less important than supposed, Comanthosphace and Leucosceptrum show sufficient overall morphological similarity to be considered congeneric. However, until the style character is reassessed, no conclusive statement can be made. Rostrinucula Rostrinucula dependens (OTU 118, Fig. 26) was originally described as Elsholtzia dependens by Rehder (1917) who stated that it seemed ‘not closely related to any other species of the genus’. My studies also suggest weak links with Elsholtzia. Instead, the phenetic affinities of Rostrinucu- la lie with Comanthosphace and Leucosceptrum canum, these taxa being constantly linked with it. Indeed, Rostrinucula sinensis (OTU 119) was originally described by Hemsley as Leuscep- trum sinense, only recently being recognized as a species of Rostrinucula by Wu (1965). Confirmation of Kudo’s (1929) separation of Rostrinucula, with bi-nucleate and tri-colpate pollen, from Elsholtzia, with tri-nucleate and hexa-colpate pollen, has already been pointed out by Wunderlich (1963). The pollen characters (Table 4) also confirm my placing of Rostrinucula with Comanthosphace and Leucosceptrum, which also have bi-nucleate and tri-colpate pollen. Rostrinucula bears a striking overall resemblance to Comanthosphace, particularly in the long flower spikes with their broad, membranous, deciduous bracts and in the whitish indumentum of unbranched and branched hairs which cover all parts of the plant. The calyces are of similar size and shape, as are the corollas, and species of both genera have unilocular anthers, hairy nutlets, and lack a tumescent gland on the disc. Since these are all characters which distinguish Comanthosphace and Leucosceptrum from other genera within the Pogostemoneae the phenetic affinities of Rostrinucula lie here. Rostrinucula, however, does not share all the characters of the Comanthosphace/Leucoscep- trum group. Some features found in Rostrinucula are unique within the Pogostemoneae while others, although occurring elsewhere in the tribe, are not found in Comanthosphace or Leucosceptrum. Rostrinucula dependens was named for its nutlets, each possessing a long curved beak at maturity (Figs 10, 26). Wu (1965) described the nutlets of R. sinensis as rostrate. In both species of Rostrinucula (Figs 7, 26) the annulus of hairs within the corolla is incomplete, unlike that of Comanthosphace. The hairs are borne on swollen disc-like excrescences at the base of the anther filaments and on the inner surface of a crescent-shaped invagination below the base of the lower lip. An identical arrangement is found in Eurysolen gracilis (OTU 1) (Figs 7, 29). Rostrinucula differs from Comanthosphace, Elsholtzia and Leucosceptrum by the entire upper lip of the corolla (Fig. 7). In this respect only Rostrinucula more closely resembles Dysophylla and Pogostemon. Rostrinucula is so closely linked with Comanthosphace in particular that the two might be considered congeneric. However, the species of Rostrinucula are less similar to Comanthos- phace than is Leucosceptrum canum (OTU 127) (see Table 8 and Fig. 25), and they are readily distinguished by. the nutlet and corolla characters given above. Elsholtzia The results of the analyses show that the species of Elsholtzia can be regarded as forming one group embracing all but one taxon, E. aquatica (OTU 22), or alternatively they can be regarded as two or three groups sharing high BGMS values. E. aquatica, which is now considered to be a species of Pogostemon (see pp. 64 & 66), is omitted from the following discussion. The clustering to maximize WGMS analysis (Tables 5-7) shows one or two groups. The principal co-ordinates analysis (Figs 14-19) shows one main group containing two sub-groups, whilst the single-linkage analysis (Fig. 25) shows three groups and several isolated species. Fig. 26 Rostrinucula dependens. (a) habit x 2. (b) calyx x 10. (c) corolla x 5. (d) dissected flower x 5. (e) stamen x 10. (f) annular hairs at the base of a stamen filament x 40. (g) nutlet, inner face, and pro- file x 5. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 49 When one group is formed it corresponds to the genus Elsholtzia as described by Bentham (1832-36) and other authors. When two groups are formed they correspond to section Cyclostegia with section Elsholtzia of Bentham, and section Apahanochilus series Stenelasmeae of Briquet. The section Aphanochilus series Platyelasmeae Brig. containing three species becomes divided up. Two species, E. densa (OTU 20) and E. manshurica (OTU 21), become included within the group comprised of sections Cyclostegia and Elsholtzia, and the third species, E. eriostachya (OTU 19), becomes included with the group comprised of section Aphanochilus series Stenelasmeae. When three groups are formed they correspond to sections Cyclostegia and Elsholtzia Benth., section Aphanochilus series Platyelasmeae Briq., and section Aphanochilus series Stenelasmeae Briq. Elsholtzia sections Cyclostegia and Elsholtzia The group corresponding to Bentham’s Elsholtzia sections Cyclostegia and Elsholtzia appears as a single close-knit group in every analysis. Even E. densa (OTU 20) and E. manshurica (OTU 21) are sometimes included within this group as in the clustering to maximize WGMS analysis (see Table 6, Appendix 4). Bentham (1829) described two genera, Aphanochilus and Cyclostegia, which he later reduced to sections of the genus Elsholtzia, distinguishing the sections on the basis of variation in bract and inflorescence characters. Section Aphanochilus has equal spikes with lanceolate or ovate bracts or secund spikes with lanceolate bracts. Section Cyclostegia has dense spikes and connate, imbricate, cyathiform, membranous, veined bracts with ciliate margins. Section Elsholtzia has broadly ovate spikes and broadly ovate, secund bracts. These sections were accepted by all subsequent authors with the exception of Kudo (1929), who reverted to Bentham’s original concept of Aphanochilus and Elsholtzia as separate genera. Kudo assigned to Elsholtzia two species of formerly unknown sectional affinities, E. heterophylla (OTU 16) and E. luteola (OTU 14) (Fig. 27). These were later assigned to section Cyclostegia by Wu & Huang (1974). However, the type species of section Cyclostegia, E. strobilifera (Benth.) Benth. (OTU 17), was not included in Kudo’s work and he did not make clear whether or not he considered Cyclostegia and Elsholtzia to be con-sectional. Several new species were assigned to both sections by Wu & Huang (1974), but the sectional definitions remained those of Bentham (1832-36). There seems to be no reason for recognizing two sections since in all of my analyses sections Cyclostegia and Elsholtzia form a single, close-knit group. Table 10 shows Bentham’s characters scored for all the species of sections Cyclostegia and Elsholtzia. All species have more or less dense inflorescences and imbricate, ciliate bracts. All species have veined bracts, although in E. concinna (OTU 8) and E. kachinensis (OTU 10), which have green, non-membranous bracts, the veins are less obvious than in other species which have membranous and often brown bracts. E. concinna and E. kachinensis (Fig. 4) are the only species with non-membranous bracts. Four combinations of the remaining three characters are represented: (1) Bracts free, non-cyathiform, inflorescence cylindrical: E. concinna (OTU 8), E. hunanensis (OTU 18), E. kachinensis (OTU 10). (2) Bracts free, non-cyathiform, inflorescence secund: E. argyi Léveillé (OTU 13), E. ciliata (OTU 9), ‘E. elegans’ (OTU 4), E. feddei Léveillé (OTU 3), E. nipponica Ohwi (OTU 6), E. oldhamii Hemsley (OTU 7), E. pseudocristata Léveillé & Vaniot (OTU 5), E. pygmaea (OTU 11), E. soulei Léveillé (OTU 12). (3) Bracts connate, non-cyathiform, inflorescence secund: E. /uteola (OTU 14). (4) Bracts connate, cyathiform, inflorescence cylindrical: E. bodinieri (OTU 15), E. hetero- phylla (OTU 16), E. strobilifera (OTU 17). Whichever combination of these three characters is used to divide the species into groups there is always some degree of overlap, and at least one species, E. concinna, E. hunanensis, E. kachinensis or E. luteola, will be an intermediate between the groups. Since four of Bentham’s characters, i.e. bracts dense, bracts imbricate, bracts veined, and bracts with ciliate margins have proved to be identical throughout both sections (Table 10) and the other characters, in any combination, always show intermediate species, the original gy a3 ~ >, £ <3 FLIES. SAA NE: Qa SS 4 Dy qed 57 See SS ——s SSE \ Y )) iS : Ss Fig. 27 Elsholtzia luteola. (a) habit x 1. (b) pair of fused bracts x 5. (c) pair of bracts opened out x 5. (d) corolla X 10. (e) dissected flower x 10. (f) stamen X 20. (g) nutlet, inner face, and profile x 10. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 51 distinction between the two sections is no longer valid. My data has produced no other evidence for a division of these species into two or more higher taxa. Therefore, I have placed all 14 species in a single group, Elsholtzia section Elsholtzia, which is characterized by orbicular, membranous bracts (except in E. concinna and E. kachinensis); the inflorescence may be cylindrical, strobilate or secund but always with imbricate bracts (see p. 69). Table 10 Bract characters in Elsholtzia sections Cyclostegia and Elsholtzia. Species Characters 4 5 6 7 — NO Ww oo E. concinna E. kachinensis E. hunanensis E. feddei section ‘E. elegans’ Elsholtzia E. pseudocristata E. nipponica E. oldhamii E. ciliata E. pygmaea E. soulei E. argyi E. luteola l l t++t+t+++¢¢¢i section E. bodinieri Cyclostegia — E. heterophylla E. strobilifera +++ ]+++4+4+4+4+4+4+4+4+44 +++ [+++ 4+4+4+4+4+4+4+4+4++4 +++ [+++ ++ 4+4+4+4+4+4+44 +++ [+++ 4+4+4+4+4+4+4+4+4+4+ +++ [+t++++¢eteetest l l +++ ]4+ 1 Key to the characters: i dense-t/lax—- 2 imbricate +/not overlapping —. 3 clearly veined +/obscurely veined —. 4 margins ciliate +/margins glabrous —. 5 membranous +/non-membranous —. 6 connate +/free —. 7 cyathiform +/not cyathiform —. 8 secund +/cylindrical —. (See also Figs 4 & 27.) Elsholtzia section Aphanochilus series Stenelasmeae The group corresponding to Briquet’s Elsholtzia section Aphanochilus series Stenelasmeae also appears to remain distinctive in each analysis, even though E. eriostachya (OTU 19) of series Platyelasmeae is sometimes included within it as, for example, in the clustering to maximize WGMS analysis (see Table 6, Appendix 4). Section Aphanochilus series Stenelasmeae is less well-defined than either section Aphanochilus series Platyelasmeae or section Elsholtzia sensu mihi. Some OTUs, e.g. E. alopecuroides Léveillé & Vaniot (OTU 32), E. communis (Collet & Hemsley) Diels (OTU 31), ‘E. glanduligera’ (OTU 34) and E. griffithii Hook. f. (OTU 33) snare high similarities with all the other OTUs in the group. Others, e.g. E. flava (OTU 28) and E. fruticosa (OTU 29), share high similarities with only a few other OTUs in the group. In the clustering to maximize WGMS analysis section Aphanochilus series Stenelasmeae seems to suffer more from the ‘rag-bag’ effect than any other group; loosely associated species such as Elsholtzia integrifolia (OTU 23) and Eurysolen gracilis (OTU 1) are included in the group together with the species of section Aphanochilus series Stenelasmeae simply because every species in the analysis must be placed in a group. The principal co-ordinates analysis (Figs 14-19) groups section Aphanochilus series Stenelasmeae within a large, open portion of the main species cluster. 52 J. R. PRESS Despite the associated species in the clustering to maximize WGMS analysis and the spreading cluster in the principal co-ordinates analysis, section Aphanochilus series Stenelas- meae is a consistently distinguishable group. Bentham (1829) and Kudo (1929) considered Aphanochilus to be a genus distinct from Elsholtzia, although Bentham qualified his view with the statement ‘perhaps my Aphanochilus might be united with it (Elsholtzia) as asecond section’ and later he (1832-36) did just that. Kudo (1929) recognized Aphanochilus by the verticillate, often long and lax flower spikes, the lanceolate or sublanceolate bracts, the divergent, ultimately confluent anther locules, the equal disc, and the shining nutlets. He distinguished Elsholtzia (in which he included some species of Bentham’s section Cyclostegia and Briquet’s section Aphanochilus series Platyelasmeae) from it by the many-flowered verticil in secund spikes, the broadly ovate, densely imbricate bracts, the emarginate upper lip of the corolla, the lower pair of stamens longer than the upper, the divergent anther-locules, and the disc with a swollen nectary. However, in all species of Aphanochilus and Elsholtzia sensu Kudo the upper lip of the corolla is emarginate, the lower pair of stamens is longer than the upper, the anthers are bilocular, the anther locules are partially fused, and the disc has a swollen gland or nectary. Species in both genera may have long, dense, secund flower spikes. Thus, the only credible distinguishing character for separating them is the lanceolate or subulate bracts versus the broadly ovate and densely imbricate bracts. Bentham (1829) similarly distinguished Aphanochilus, Elsholtzia, and a third genus Cycloste- gia by inflorescence, bract, corolla, and anther characters. Later Bentham (1832-36) reassessed these characters and commented that ‘The three sections of Elsholtzia differ, in many respects, from each other in habit; but on closer examination of their characters, these distinctions do not appear to be of sufficient importance to preserve the genera Aphanochilus and Cyclostegia, which I had originally established, but which I have now considered as mere sections of Elsholtzia.’ Bentham characterized sections Aphanochilus and Elsholtzia as follows: section Aphanochilus. Spikes equal, bracts lanceolate to ovate or secund with bracts lanceolate. section Elsholtzia. Spikes and bracts broadly ovate, secund. The characters match the unique characters used by Kudo (1929). Bentham’s reassessment corresponds with the results presented here although, as discussed above, section Cyclostegia cannot be differentiated from section Elsholtzia, and section Aphanochilus series Stenelasmeae Briquet, rather than section Aphanochilus sensu lato, forms a subgroup within Elsholtzia to correspond with Bentham’s ‘mere section’. Section Aphanochilus series Stenelasmeae was first described by Briquet (1897); the bracts were described as small, linear-lanceolate or stiffly pointed, and the nutlets shiny. Briquet created the series to differentiate the majority of the species in section Aphanochilus from E. densa and E. eriostachya which he placed in section Aphanochilus series Platyelasmeae, a series characterized by very short, broadly ovate or rounded bracts, and dark, dull nutlets. The group shown in my analyses corresponds almost exactly to Briquet’s concept of section Aphanochilus series Stenelasmeae (although it contains several species unknown to Briquet) and is comparable to the group formed by sections Cyclostegia and Elsholtzia. Since section Elsholtzia sensu mihi is regarded here as a section, section Aphanochilus series Stenelasmeae has been raised to the rank of section (see p. 69). The distinguishing features of section Aphanochilus sensu stricto given by Briquet are rather slight. Bentham’s description, although referring to section Aphanochilus sensu lato, provides a more suitable basis for characterizing section Aphanochilus sensu stricto. In order to accommodate some newer species of Elsholtzia Wu & Huang (1974) described eight new series within their section Aphanochilus subsection Stenelasmeae. However, these were confined to Chinese species and although the isolated positions of E. flava (OTU 28) and E. penduliflora (OTU 25) in separate series agree with the positions shown in my analyses, on the whole their divisions are not supported by my results. Only three of Wu’s & Huang’s series, Blandae, Communes, and Fruticosae, contain more than one species. Series Communes does emerge as a fairly clear group in the single-linkage dendrogram (Fig. 25), but the other two series are not evident in the analyses. In fact, series Fruticosae appears as a heterogenous group, five of the six species sharing their highest similarities with species in other series, e.g. E. winitiana TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 33 Craib. (OTU 36), in series Fruticosae, with E. blanda (Wallich ex Benth.) Benth. (OTU 38) in series Blandae (see Appendix 3). In other words characters given by the authors inadequately differentiate the series. As an example, series Capituligerae is described as: ‘frutices parvi; spicae sphaeroideae; corolla 4-fida, labio superiore integra’. The entire upper lip of the corolla is incorrectly observed; all species of Elsholtzia, including E. capituligera (OTU 42), have an emarginate upper lip. All species have a 4-fid corolla, not just those of this series. Similarly, series Communes is described as: ‘herbae erectae; spicae cylindriceae compactae; corolla 4-fida, labio superiore integro’. Again the description of the upper lip is incorrect and the 4-fid corolla is common to all species in the genus. Cylindrical, compact flower spikes are also found in many species in other series e.g. E. pilosa of series Pilosae, and E. winitiana of series Fruticosae. Elsholtzia section Aphanochilus series Platyelasmeae The most problematical group in the analyses corresponds to Briquet’s section Aphanochilus series Platyelasmeae of Elsholtzia which contains three species: E. densa (OTU 20, Fig. 28), E. eriostachya (OTU 19), and E. manshurica (OTU 21). It is only seen clearly in the single-linkage analysis (Fig. 25). The nearest neighbours list (Appendix 3) shows that E. manshurica shares a number of high similarities with species in section Elsholtzia sensu mihi. E. densa shares high similarities with ‘E. elegans’ (OTU 4) of section Elsholtzia sensu mihi and with ‘E. glanduligera’ (OTU 34) of section Aphanochilus sensu stricto. E. eriostachya does not share high similarities with any species from section Elsholtzia sensu mihi. In the clustering to maximize WGMS (Tables 5, 6, Appendix 4) and the principal co-ordinates analysis (Figs 19-21) series Platyelas- meae does not form a separate group. E. eriostachya is included in section Aphanochilus sensu stricto, while E. densa and E. manshurica are included in section Elsholtzia sensu mihi. If E. densa and E. manshurica are considered in isolation from E. eriostachya then on the basis of overall similarity they could acceptably be placed in section Elsholtzia sensu mihi. Similarly, if E. eriostachya is compared only with sections Elsholtzia sensu mihi and Aphanochilus sensu stricto then it would be placed in section Aphanochilus sensu stricto. This is exactly the treatment given in some earlier works. Bentham (1832-36) placed E. eriostachya (and later, in 1848, E. densa) in his section Aphanochilus since, although the bracts were broad, like those of his section Elsholtzia, the inflorescence was not secund. Kudo (1929) placed E. densa (as E. janthina Dunn) in Elsholtzia, which he regarded as a genus in the narrow sense, although he described the inflorescence of Elsholtzia pro majore as secund and that of E. densa (Fig. 28) cylindrical. However, E. densa and E. eriostachya are much more similar to each other than they are to any of the species of either section Elsholtzia sensu auct. or Aphanochilus sensu stricto. This is quite clear from the nearest neighbours list (Appendix 3) as well as the single-linkage analysis (Fig. 25) in which these three species form a distinct group. This view is similar to that of Briquet (1897) who placed E. densa and E. eriostachya in a separate group characterized by having short, broadly ovate or rounded bracts, and dark, matt nutlets. He considered section Aphanochilus series Platyelasmeae to merit only the rank of series, apparently on negative evidence; although series Platyelasmeae has broad bracts, these are neither secund as in section Elsholtzia nor connate as in section Cyclostegia. In fact, series Platyelasmeae seems to be intermediate between section Aphanochilus sensu stricto and section Elsholtzia sensu mihi. Table 11 gives a compari- son of the bract characters in each group. Species of section Aphanochilus sensu stricto generally have narrow, non-membranous, non-imbricate, and usually green bracts. Species of section Elsholtzia sensu mihi generally have orbicular, membranous, imbricate, and usually brown or bi-coloured bracts. Species of series Platyelasmeae have orbicular, non-membranous, non- imbricate, and usually bicoloured or brownish bracts. In addition E. densa, E. eriostachya, and E. manshurica resemble section Elsholtzia sensu mihi in lacking bracteoles, which are found throughout section Aphanochilus sensu stricto. Elsholtzia series Platyelasmeae has three distinctive characters: the fruiting calyx becomes greatly inflated (Fig. 6), the style lobes have clavate tips (Fig. 9), and the mature nutlets are distinctly verrucose (Fig. 10). E. densa, E. eriostachya, and E. manshurica are almost exactly similar to one another with respect to their qualitative character states. They differ significantly 54 J. R. PRESS Table 11 Bract characters in Elsholtzia. sect. Aphanochilus sensu sect. Elsholtzia sensu stricto ser. Platyelasmeae Briq. mihi not as broad as long at least as broad as long at least as broad as long non-membranous non-membranous membranous (except for E. concinna and E. kachinensis) non-imbricate non-imbricate imbricate usually green usually brown or bicoloured usually brown or bicoloured (except for E. concinna and E. kachinensis) (See also Figs 4, 27, & 28.) only in the variation of ten quantitative characters. These were scored as simple measurements and not converted into proportions. Consequently, there is the possibility of distorted separa- tion, particularly in the principal co-ordinates analyses, due to the size factors. This would explain the grouping of E. eriostachya with the generally small flowered species of section Aphanochilus sensu stricto, and the larger flowered E. densa and E. manshurica with section Elsholtzia sensu mihi. The calyx, style, and nutlet characters were utilized by Kitagawa (1935) who recognized Platyelasma as a genus distinct from Aphanochilus and Elsholtzia. Table 12 gives mean group similarities for section Elsholtzia sensu mihi, section Aphanochilus sensu stricto, and series Platyelasmeae. The similarities support Kitagawa’s recognition of Platyelasmaasa separate taxon at the same rank as Aphanochilus and Elsholtzia. However the high similarities shared by all three groups precludes recognition of Platyelasma at the generic level. Despite the distribution of E. densa, E. eriostachya, and E. manshurica in the principal co-ordinates and clustering to maximize WGMS analyses, the most suitable treatment for these species is to place them in a third section: Elsholtzia section Platyelasma. It is interesting to note here the monotypic genus Paulseniella described by Briquet (1908) from a plant collected in the Pamir. It has a campanulate calyx, becoming inflated in fruit, a barely exserted subequally five-lobed corolla, style lobes with globular swellings at the tips, and tuberculate-rugose nutlets. P. pamirensis was later recognized by Fedschenko (1908) as being based on a specimen of Elsholtzia densa. Although mistaking the identity of the plant, Briquet thought it sufficiently distinct to warrant its recognition as a new genus. Table 12 WGMS and BGMS values for the subgeneric divisions of Elsholtzia. qi 83-6 2 78-4 88-1 3 78-9 73-2 87-5 1 2 3 Key to groups: : 1—section Elsholtzia mihi. 2—section Aphanochilus sensu stricto. 3—-series Platyelasmeae. Relocated OTUs A number of specie of Elsholtzia appear to be somewhat divorced from the bulk of the genus, especially in the single-linkage analysis (Fig. 25). In almost every case the data were obtained from a single specimen and the resulting distortion appears to have affected the placing of some 56 J. R. PRESS species. However, most can be accurately placed by referring to the nearest neighbours list (Appendix 3). Elsholtzia beddomei and E. kachinensis Elsholtzia beddomei C. B. Clarke ex Hook.f. (OTU 24) and E. kachinensis (OTU 10) link at low similarities in the single-linkage analysis (Fig. 25). Although sharing few high similarities with other species of E/sholtzia there is no doubt as to where their phenetic affinities lie. Reference to the nearest neighbours list (Appendix 3) confirms that E. beddomei belongs to section Aphanochilus sensu stricto, and E. kachinensis to section Elsholtzia sensu mihi. Elsholtzia penduliflora Elsholtzia penduliflora (OTU 25) links at a low similarity in the single-linkage analysis (Fig. 25) and seems to be isolated from other species of Elsholtzia. It is rather surprising to find Pogostemon brevicorollus Y. Z. Sun (OTU 105) as the nearest neighbour to Elsholtzia penduliflora (Appendix 3). However, the data set for Pogostemon brevicorollus is incomplete and the similarity between it and Elsholtzia penduliflora may be considered as dubious. The similarity values shared by E. penduliflora with its remaining four nearest neighbours, E. elata Zoll. & Mor. (OTU 35), and E. blanda (OTU 38), Pogostemon travancoricus (OTU 92), and P. amarantoides (OTU 80) give no conclusive pointer to where the phenetic affinities of Elsholtzia penduliflora lie. Both the principal co-ordinates (Figs 14-17) and the clustering to maximize WGMS analysis (Appendix 4, Table 6) place it in Elsholtzia section Aphanochilus sensu stricto. Elsholtzia flava, E. fruticosa and E. hunanensis In the single-linkage dendrogram (Fig. 25), Elsholtzia flava (OTU 28), E. fruticosa (OTU 29) and E. hunanensis (OTU 18) are not referable to any section of Elsholtzia. As with the previous species the nearest neighbours list (Appendix 3) can be used to locate all three species. E. hunanensis belongs to section Elsholtzia sensu mihi, and E. fruticosa and E. flava to section Aphanochilus sensu stricto. However E. flava warrants further investigation. In this species the style is not always obviously gynobasic (see p. 22) and the nutlets have a small but laterally displaced attachment-scar (Fig. 10 and see p. 46). The bracts are unusually broad for section Aphanochilus sensu stricto and the very large, broadly-ovate leaves, and coarse habit help to identify this species. Should the style and nutlet-attachment characters prove to have greater value than they have been accorded here (see discussion of these characters in Leucosceptrum, pp. 46-47) the position of E. flava will need to be re-examined. Elsholtzia concinna Elsholtzia concinna (OTU 8) presents a more difficult situation and the evidence from the various analyses is conflicting. In the principal co-ordinates analysis (Figs 17-19) E. concinna might be included within either section Aphanochilus sensu stricto or section Elsholtzia sensu mihi. In the clustering to maximize WGMS (Appendix 4, Table 6) it is grouped with section Aphanochilus sensu stricto. In the single-linkage dendrogram (Fig. 25) it links on at the 86% level and is not referable to any section, although the nearest neighbours list (Appendix 3) shows it to be most similar to Dysophylla mairei (= Elsholtzia pilosa see pp. 57) which is a member of section Aphanochilus. The raw data were obtained from two rather poor specimens and should perhaps be regarded with some suspicion. Vautier (1959), when describing E. concinna (Fig. 4), stated that, although differing in bract and inflorescence characters, E. concinna and E. strobilifera (OTU 17) were similar in general appearance; E. concinna and E. ciliata (OTU 11) differed in leaf and inflorescence features. The clear implication was that E. concinna belonged to section Elsholtzia sensu mihi. E. concinna certainly shares the characters which define this section (Table 11) rather than those which define section Aphanochilus sensu stricto, although it in fact resembles E. ciliata more than E. strobilifera. In view of this, I have placed E. concinna in section Elsholtzia sensu mihi but the availability of more study material may necessitate reconsideration of its position. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE a Leucosceptrum plectranthoideum Leucosceptrum plectranthoideum (OTU 128) is placed in Elsholtzia section Apanochilus sensu stricto in all the analyses except the single-linkage dendrogram (Fig. 25), where it links with Elsholtzia, but is not referable to any particular section. Oringally described by Léveillé (1915-16) as Buddleja plectranthoidea it was transferred to Leucosceptrum by Marquand (1930). The type is a specimen of Elsholtzia fruticosa (OTU 29). E. fruticosa and Leucosceptrum plectranthoideum are not, as might be expected, inseparable, or at least very close, units in the analyses. Once again this seems due to inadequate sample size, only the type of Leucosceptrum plectranthoideum being available. This specimen did not fall completely within the variation range of Elsholtzia fruticosa as represented by the specimens used to generate the averaged data set (see p. 8). Dysophylla mairei Dysophylla mairei (OTU 65) was described by Léveillé (1912) as a species of Dysophylla. However all of the analyses clearly show it to belong to Elsholtzia and not to Dysophylla. Dunn (1915) gave Dysophylla mairei as a synonym of Elsholtzia pilosa and the type specimens compare very closely. I have accepted Dunn’s view. Elsholtzia integrifolia All previous authors who have considered E. integrifolia (OTU 23) place it in Elsholtzia section Aphanochilus, thus agreeing with Bentham’s original concept (1832-36). None of them appears to have seen the type specimen, a sheet only recently relocated by me at BM. As a supposed member of Elsholtzia section Aphanochilus, E. integrifolia was included in this study, but the analyses show it to be an isolated taxon, sharing an overall similarity of only 84-1% with its nearest neighbour, E. griffithii (OTU 33) (Appendix 3). Wu & Huang (1977) cite E. integrifolia as a synonym for Schizonepeta tenuifolia (L.) Briq. of the tribe Nepetae, apparently on evidence obtained from a photograph. When this work became available to me in 1979, after completion of the computor runs, I compared the type of E. integrifolia with available material and descriptions of Schizonepeta, although unfortunately not of Schizonepeta tenuifolia. Elsholtzia integrifolia does share a greater overall similarity to other species of Schizonepeta than to species of Elsholtzia, and therefore I exclude Elsholtzia integrifolia from the Pogostemoneae. ‘Elsholtzia japonica’ ‘Elsholtzia japonica’ (OTU 2) was thought to be a previously undescribed species represented by two specimens in the Léveillé herbarium at E. Both specimens were collected by d’Argy in the Kiangsu province of China, and were labelled ‘E. japonica’. In the clustering to maximize WGMS (Appendix 4, Tables 5, 6) and the principal co-ordinates analyses (Figs 14-17) ‘E. japonica’ clusters with Elsholtzia. In the single-linkage analysis (Fig. 25) it clusters with Pogostemon. However, ‘E. japonica’ lacks certain constant features possessed by species of Elsholtzia and Pogostemon. The corolla of Pogostemon has an entire upper lip, also hairy stamen-filaments and unilocular anthers. ‘E. japonica’ has an emarginate upper lip to the corolla, naked stamen-filaments, and bilocular anthers. Elsholtzia has bilocular-fused anthers and a tumescent lobe on the disc. ‘E. japonica’ has bilocular-free anthers and no lobe on the disc. It also has hairy nutlets while those of Elsholtzia and Pogostemon are invariably glabrous. These characters suggest that ‘E. japonica’ might be misplaced and a brief survey of genera outside the Pogostemoneae confirms that ‘E. japonica’ is more similar to species of Agastache Clayton ex Gronovius (tribe Nepetae) and, in particular, to A. rugosus Fish. & Mey, to which it might belong. I have therefore excluded it from the Pogostemoneae. The geographical distribution of Elsholtzia as accepted by other authors was rather striking. Most species occur in eastern Asia and Malesia with the weedy species, E. ciliata (OTU 9), extending over a wider area as far as central Europe (where it may be a relict of cultivation). A 58 J. R. PRESS single, very isolated species, E. aquatica (OTU 22), occurs in southern Malawi. However, the transfer of this species to Pogostemon rectifies the disjunction. Keiskea The five species of Keiskea form a discrete and homogeneous group in all the analyses. They are most similar to species of Elsholtzia and share slightly higher similarities with species of section Elsholtzia sensu mihi than with the other sections. This confirms the view of earlier authors (e.g. Bentham & Hooker, 1876; Ohwi, 1965) who placed Keiskea next to Elsholtzia. The two genera were said to differ mainly in features of the calyx, that of Keiskea being deeply divided and sometimes described as bilabiate, while that of Elsholtzia is shallowly toothed and not bilabiate. During this study, several other characters which distinguish the genera have been determined. As well as being deeply divided, the calyx in Keiskea (Fig. 6) has an annulus of hairs in the throat. No Elsholtzia possesses such an annulus. Similarly, Keiskea (Fig. 7) has a complete annulus of hairs within the corolla. Some species of Elsholtzia, e.g. E. rugulosa (OTU 26) (Fig. 7), also possess an annulus within the corolla but these annuli are always incomplete, with the ends of the open circle of hairs extending dorsally under the upper lip. In Elsholtzia the number of mature nutlets is always four. Fertilized flowers of all species of Keiskea have four very young nutlets but at maturity Keiskea japonica (OTU 132), the only species in which mature nutlets are known to me, has only one (Fig. 10), a character also noted by Ohwi (1965). However my sample was very small and the abortion of one, two or three nutlets is common in Labiatae. It may be that a larger sample will show the normal number of mature nutlets in Keiskea to be four. The confluence of the anther locules is a variable character. Keiskea has bilocular anthers in which the locules are free and separated by a short connective. Elsholtzia has bilocular anthers in which the locules are confluent, although still recognizably bilocular (Figs 8, 27). The single exception to this is E. hunanensis (OTU 18) which has anthers similar to those of species of Keiskea. Thus, although this appears to be a distinguishing feature between the two genera, the exceptional Elsholtzia species show it to be a linking character too. A clear phenetic similarity of Keiskea with Elsholtzia is established by a number of other characters. The shared characters of a (sometimes weakly) bilabiate corolla with an emarginate upper lip and spreading, three-lobed lower lip were recognized by earlier authors (Bentham & Hooker, 1876; Briquet, 1897; Ohwi, 1965). In addition both genera have a single tumescent gland on the disc (Fig. 9), a unique character within the tribe. In Keiskea the bracts are similar in shape and texture to those of anumber of Elsholtzia species, e.g. E. flava (OTU 28) (Fig. 4). The few-flowered verticils forming a loose, secund inflorescence in Keiskea are also typical of some species of Elsholtzia, particularly those of Elsholtzia section Elsholtzia sensu mihi e.g. E. ciliata (OTU 9). Only one species of Keiskea, K. macrobracteata, is omitted from this study for lack of available material. However, from the description all the comments made above equally apply. Its description suggests that it may be the most similar species to the taxa of Elsholtzia section Elsholtzia sensu mihi, particularly with respect to the bract characters. In the protologue Masamune (1940) considered that its conspicuous features distinguished it from other Keiskea species and divided the genus into two sections: Section I. Macrobracteatae. Bractea ovata-rotundata. Carex (sic) bi-labiatus supra barbatus. K. macrobracteata. Section II. Eukeiskea. Bracteata linearis. Carex (sic) late campanulatus extus subglaber. K. japonica K. sinensis. However the calyx characters used to distinguish the sections are inapplicable since all the species have calyces which are bilabiate and hairy above. K. elsholtzioides (OTU 133) (Fig. 4) has longer, broader bracts than other species and might be placed in section Macrobracteatae. The bracts of the other species are narrower but could hardly be described as linear. There seems to be little support for Masamune’s division, but the lack of specimens of K. macrobracteata prevents a complete assessment from being made. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 59 Tetradenia The three species of Tetradenia, like Keiskea, form an obvious group in all analyses. Again like Keiskea, Tetradenia is most similar to Elsholtzia. The clustering to maximize WGMS analysis (Table 7) suggests a greater similarity to Elsholtzia section Elsholtzia sensu mihi than to Elsholtzia section Aphanochilus sensu stricto, but this is not borne out by the principal co-ordinates analyses (Figs 14-16). In the genus protologue Bentham (1829) recognized Tetradenia by the four tumescent lobes on the disc and placed it ‘immediately before Elsholtzia’. In a later work (1832-36) he expanded this statement considerably: ‘it [Tetradenia] is intermediate between Elsholtzia and Col- ebrookea: it differs from the first by the calyx and by the more regular corolla; from Colebrookea by the calyx not plumose at the maturity of the fruit, by the style less deeply cleft; and from both by the remarkable glands of the ovarium’. Despite the link with Elsholtzia, Bentham (1848) and other authors (e.g. Bentham & Hooker, 1876) grouped Tetradenia with Colebrookea, Dysophylla, and Pogostemon; the group was characterized particularly by unilocular anthers or the anther locules confluent, the corolla tube rarely longer than the calyx, the lobes equal or the anterior lobes longer. Elsholtzia was placed in another group characterized by distant or sometimes confluent anther locules. The evidence of this study supports a link for Tetradenia with Elsholtzia rather than with Colebrookea, Dysophylla, and Pogostemon (although from the principal co-ordinates analyses, Figs 11-13, Colebrookea would appear to be more similar to Tetradenia than are either Dysophylla or Pogostemon). The proximity of Tetradenia and Elsholtzia in all of the analyses except the single-linkage analysis (Fig. 25) is clear evidence of the phenetic similarity between the genera. This is reflected by a number of characters. Perhaps the most obvious are the shape of the corolla, the shape and texture of the bracts, the bilocular (not unilocular) anthers, and the pollen types. Both Tetradenia and Elsholtzia have emarginate upper lobes to the corolla. The corolla in most species of Elsholtzia, as observed by Bentham, is more markedly bilabiate than that of species of Tetradenia, but some species of Elsholtzia, e.g. E. densa (OTU 20) and E. myosurus Dunn (OTU 39) have similar corollas (Figs 7, 28). Dysophylla and Pogostemon have entire upper lips to the corolla. The link between Tetradenia and Elsholtzia section Elsholtzia sensu mihi is emphasized by the similarity of the bracts (Fig. 4). In both the bracts are orbicular or somewhat broader than long, usually brown (or bi-coloured) and membranous. Elsholtzia section Platyelasma sensu mihi also has broad, brown or bi-coloured bracts, but they are not membranous. Similar bracts are found in Elsholtzia flava (OTU 28) (Fig. 4). Like Elsholtzia section Elsholtzia sensu mihi and Elsholtzia section Platyelasma sensu mihi, Tetradenia lacks bracteoles. This contrasts markedly with Dysophylla, Pogostemon, and Elsholtzia section Stenelasmeae sensu stricto which all possess bracteoles. The anthers of Tetradenia species are bilocular and confluent through fusion, not unilocular as given by Bentham (1848) and Bentham & Hooker (1876).. The pollen of Tetradenia is tri-nucleate and hexa-colpate, as is that of Elsholtzia and Keiskea (see Table 4). All other genera in the Pogostemoneae have bi-nucleate and tri-colpate pollen. Two other characters that help to link Tetradenia with Elsholtzia, while divorcing it from Colebrookea, Dysophylla, and Pogostemon, are the glabrous anther filaments (all species of Pogostemon and Dysophylla have hairy anther-filaments) and tumescent glands on the disc (absent in Colebrookea (Fig. 9), Dysophylla, and Pogostemon). The tumescent glands provide one of two unique features for Tetradenia by having four bright red glands encircling the disc. The young nutlets rise above the glands (Fig. 9), but as they mature the glands enlarge greatly, finally overtopping the nutlets. A second unique feature is a calyx character. The upper tooth of the calyx is very broad, much broader than, and overlapping, the teeth on either side of it, unknown in other Pogostemoneae (Fig. 6). The corolla in Tetradenia has a complete annulus of hairs within its throat (Fig. 7). Although some species of Elsholtzia, e.g. E. rugulosa (OTU 26), have annuli, they are never complete circles of hairs. Elsewhere, complete annuli are found only in Comanthosphace and Keiskea. 60 J. R. PRESS Eurysolen The taxonomic position of Eurysolen is an open question. Prain (1898, 1901) described the only species, E. gracilis (OTU 1) (Fig. 29), and tentatively placed it near Gomphostemma Benth. in his tribe Prasieae, citing only the one-celled anther character as supporting evidence for his choice. Briquet (see Prain, 1901) suggested that Eurysolen might be placed in the Prasieae or the Ajugoideae. However, the lack of mature fruits prevented the exact determination of its position. Later authors were similarly hampered by lack of fruiting material. Kudo (1929) placed Eurysolen in the Prasieae; Mukerjee (1940) placed Eurysolen in his tribe Ajugoideae. Wu (1959) placed Eurysolen in the Ajugoideae but noted an apparent relationship with Pogoste- mon. Unfortunately, he did not amplify this in any way. Chermsirivathana (1963) examined a number of fruiting specimens and described the mature fruits as dry and with a small basal attachment. On this evidence she placed it with Colebrookea, Dysophylla, Elsholtzia, and Pogostemon in her subfamily Stachyoideae. This was supported by Keng (1969, 1978) who considered that, in general habit, inflorescence, and flower structure Eurysolen was identical with another member of the Stachyoideae, Achyrospermum Blume. My study is restricted to the tribe Pogostemoneae and the exact position of Eurysolen cannot be ascertained without reference to the other tribes to which it might belong. However if Chermsirivathana’s and Keng’s reasons for rejecting the Ajugoideae and Prasieae are accepted, Eurysolen would seem well-placed with the other genera of the Pogostemoneae. My own examination of mature fruits of Eurysolen gracilis (Fig. 10) confirms Chermsirivathana’s observations; they are dry and not fleshy as in the Prasieae and the attachment scar is small and basal, not large and lateral as in the Ajugoideae. In general habit and inflorescence E. gracilis resembles some species of Pogostemon, e.g. P. wightii Benth. (OTU 100), and in floral structure resembles species of Elsholtzia and Rostrinucula (Fig. 7). The analyses give conflicting results for the affinities of Eurysolen gracilis within the Pogostemoneae. It is not very similar to any other OTU, although it shares its highest similarities with species of Elsholtzia section Aphanochilus sensu stricto. Appendix 3 shows that its nearest neighbour is Dysophylla mairei (OTU 65) (= Elsholtzia pilosa, see p. 000) with the very low similarity of 81.5%. The single-linkage analysis (Fig. 25) also shows Eurysolen gracilis to be an OTU of rather remote affinity with other members of the tribe. The principal co-ordinates (Figs 11-16) and clustering to maximize WGMS analyses (Tables 5-7) clearly and consistently show Eurysolen gracilis as a member of Elsholtzia section Aphanochilus sensu stricto, inseperable even in the principal co-ordinates analysis of the Comanthosphace/Elsholtzia subset (Figs. 14-16). The results should not perhaps be taken at face value. In the principal co-ordinates analyses the inclusion of Eurysolen gracilis in the Elsholtzia species cluster is almost certainly due to one of the flaws in this method. The axes used to produce the two-dimensional plots are those corresponding to the largest eigenvalues, i.e. those axes in the direction of maximum variance. Only the first four axes are normally used and this may be insufficient to split OTUs apart. The presence of Eurysolen gracilis in the Elsholtzia group in the clustering to maximize WGMS is probably due to the ‘rag-bag’ effect. Since every unit must be placed in a group and the pre-selected number of groups has already been formed at higher levels of similarity, Eurysolen gracilis has been placed with its nearest neighbour Dysophylla mairei (OTU 65). That the similarity of Eurysolen gracilis to Dysophylla mairei is very low (81-2%) does not affect the inclusion of these two OTUs in the same group; Eurysolen gracilis only appears to be a part of the Elsholtzia cluster. The single-linkage analysis (Fig. 25) suggests that this inclusion is unlikely and the nearest neighbours list confirms that they are separate taxa. The pollen data (Table 4) lends further support to the separation of Eurysolen and Elsholtzia. In all species of Elsholtzia examined the pollen is tri-nucleate and hexa-colpate. In Eurysolen gracilis the pollen is bi-nucleate and tri-colpate. Eurysolen gracilis does share some characters with other members of the tribe, most notably an invagination of the corolla and hairy bosses at the base of the stamen filaments as in species of Rostrinucula, and unilocular anthers as in Comanthosphace, Dysophylla, Pogostemon, and Rostrinucula. It is distinguished within the Pogostemoneae by a combination of three charac- Fig. 29 Eurysolen gracilis. (a) habit x 1. (b) calyx x 5. (c) corolla x 5. (d) dissected flower x 5. (e) stamen x 10. (f) annular hairs at the base of a stamen filament x 40. 62 J. R. PRESS ters: the absence of a tumescent gland on the disc, the emarginate upper lip of the corolla, and the corolla tube gibbous towards the base (Fig. 29). Pogostemon and Dysophylla Although usually treated as separate genera, Dysophylla and Pogostemon have been considered congeneric by several authors (Hasskarl, 1842; Miquel, 1856; Kuntze, 1891). Bentham (1870) retained the two as separate genera and commented that if they were to be regarded a single genus Pogostemon, Dysophylla would still be a distinct section. Keng’s (1978) arrangement in which species of Dysophylla are confined to Pogostemon section Eusteralis (Rafin.) Keng agrees with Bentham. El-Gazzar & Watson (1967) produced a modified treatment for the two genera. On the basis of a comparison of six characters: leaf shape, leaf arrangement and indumentum, petiole length, calyx inclusions, and presence of stem aerenchyma in 22 species of Pogostemon (approximately 37% of the genus) and 16 species of Dysophylla (approximately 47% of the genus) they transferred the four species of Dysophylla section Oppositifoliae to Pogostemon. Their emended genera were characterized as follows: Dysophylla. Leaves verticillate, 3-10 in a whorl, linear, sessile and usually glabrous. Corolla subequally quadrifid. Pogostemon. Leaves opposite, ovate or narrowly ovate, petiolate, usually more or less hairy or tomentose. Corolla usually subilabiate, upper lip trifid, lower entire. My results are in broad agreement with El-Gazzar’s & Watson’s view that species of Dysophylla section Verticillatae should be grouped together, while the species of section Oppositifoliae should be grouped with species of Pogostemon. However, all my analyses point to a greater degree of unity between these groups than is generally acceptable between two genera. Although El-Gazzar & Watson’s evidence and arguments are clear and persuasive, their division of the genera is based on a rather narrow sample. If the species of my wider sample are described in terms of the morphological features which characterize Dysophylla and Pogoste- mon sensu El-Gazzar & Watson, they form a transition series between the two extremes represented by the authors’ taxa. The stages in the series are as follows: 1. Leaves in whorls of three or more, sessile, linear, truncate at the base, glabrous, corolla with upper lip equalling or shorter than lower; this describes Dysophylla sensu stricto; Dysophylla stellata (OTU 53) is typical. 2. Leaves in whorls of four, sessile, linear, truncate at the base, sparsely hairy, corolla with upper lip equalling lower; e.g. Dysophylla linearis (OTU 54). 3. Leaves in whorls of three or more, sessile, linear, truncate at the base, hairy, corolla with upper lip equalling lower; e.g. Dysophylla cruciata Benth. in Wallich (OTU 55), D. gracilis Dalz. in Hook.f. (OTU 62), D. koehneana (OTU 61), and D. szemacensis Y. C. Wu & Hsuan (OTU 59). 4. Leaves in whorls of three or four, very shortly petiolate, linear, attenuate at the base, densely hairy, corolla with upper lip equalling or shorter than lower; e.g. Dysophylla falcata (OTU 58) and D. quadrifolia (OTU 56). Several specimens of D. quadrifolia in the herbarium at E have paired leaves instead of the more usual whorls of three. The isotype of Dysophylla falcata (the only specimen examined) appears to have leaves in whorls of three, changing to pairs of leaves at the two uppermost nodes. 5. Leaves in whorls of three, sessile, broadly ovate to orbicular, cuneate at the base, sparsely hairy, corolla with upper lip equalling lower; e.g. ‘Dysophylla trinervis’ (OTU 43) (Fig. 33). 6. Leaves in pairs, sessile, ovate, truncate at the base, densely hairy, corolla with upper lip equalling lower; e.g. Dysophylla andersonii Prain (OTU 66). 7. Leaves in pairs, petiolate, linear to narrowly ovate, cuneate at the base, sparsely hairy, corolla with upper lip equalling lower; e.g. Dysophylla salicifolia (OTU 69). 8. Leaves in pairs, shortly petiolate, ovate, cuneate at the base, glabrous, corolla with upper lip equalling lower; e.g. ‘Dysophylla glabrata’ (OTU 67) (Fig. 32). 9. Leaves in pairs, shortly petiolate, ovate, cuneate at the base, hairy (sometimes densely so), TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 63 corolla with upper lip equalling lower; e.g. Dysophylla auricularia (OTU 68), D. myosuroides (OTU 71), and D. rugosa (OTU 70). 10. Leaves in pairs, very shortly petiolate, ovate, cuneate at the base, sparsely hairy, corolla with upper lip longer than lower; e.g. Pogostemon strigosus (OTU 102). This species bears a striking resemblance to Dysophylla auricularia. 11. Leaves in pairs, shortly petiolate to subsessile, ovate, cuneate at the base, sparsely hairy to subglabrous, corolla with upper lip longer than lower; e.g. Pogostemon micangensis G. Taylor (OTU 85) and P. mutamba (OTU 86, Fig. 30). 12. Leaves in pairs, petiolate, linear, cuneate at the base, densely hairy, corolla with upper lip equalling lower; e.g. Pogostemon nilagiricus (OTU 116). 13. Leaves in pairs, petiolate, ovate to orbicular, cuneate, attenuate or rounded at the base, hairy, corolla with upper lip equalling or longer than lower; Pogostemon sensu Stricto, including e.g. Pogostemon hirsutus Benth. (OTU 109), P. menthoides (OTU 82), and P. rupestris Benth. (OTU 110). On this basis Dysophylla and Pogostemon should not be retained as separate genera and the consequences of this are that Hasskarl’s (1842) proposal to unite Dysophylla and Pogostemon must be accepted, and El-Gazzar’s & Watson’s (1967) proposal rejected. However, their thesis that Dysophylla section Oppositifoliae is much closer to Pogostemon sensu stricto than to Dysophylla sensu stricto is correct. Keng (1978) has written a classification which unites Dysophylla and Pogostemon incorporating the view of El-Gazzar & Watson. On the separation of Dysophylla and Pogostemon he stated that ‘the phyllotaxis is insufficient taxonomically for a (division at the) generic level’ and accepted Pogostemon in the widest sense, including Dysophylla. However, he made use of phyllotaxis to define two new sections. He accepted the joining of the opposite-leaved species of Dysophylla with Pogostemon sensu stricto to form section Pogostemon and placed the verticillate-leaved species of Dysophylla in section Euster- alis. Keng’s conclusions were based only on studies of 10 Malesian species but his eminently sensible arrangement fits the data from my wider sample too and is followed here (see pp. 71-74). With the exception of Dysophylla falcata (OTU 58) and D. quadrifolia (OTU 56) the species are easily divided between the sections. However to accommodate these exceptional species slight modification of Keng’s distinguishing characters of the sections is required: Pogostemon section Pogostemon. All leaves in opposite pairs, usually petiolate, the corolla with the upper lip equalling or longer than the lower. Pogostemon section Eusteralis. At least some of the leaves in whorls of three or more, usually sessile, the corolla with the upper lip equalling or shorter than the lower. Indirect support for this treatment comes from Cook (1978) who described from several unrelated families the occurrence of a complex of characters including erect, unbranched stems with simple, elongate leaves borne in whorls which he called the ‘Hippuris syndrome’. This syndrome is found only in aquatic or amphibious plants, appearing to be developed purely as a response to habitat. Among the plants named by Cook as exhibiting the ‘Hippuris syndrome’ are species of Pogostemon section Eusteralis. These species do indeed show all the required characters, including absence of hairs from most parts of the plants, stems which are creeping below, erect above, and small flowers with very short pedicels. The characters used by earlier authors (e.g. El-Gazzar & Watson, 1967) for the maintenance of Dysophylla as a genus distinct from Pogostemon are precisely those of the ‘Hippuris syndrome’. All species of Pogostemon section Eusteralis are plants of wet habitats, while many (though by no means all) species of Pogostemon section Pogostemon are plants of stony, dry grounds or forest margins. Some species of section Eusteralis, e.g. P. falcatus (C. Y. Wu) C. Y. Wu& H. W. Li and P. quadrifolius (Roxb.) Kuntze, inhabit the dryer parts of wet habitats or areas prone to drying out. It is noticeable that these species sometimes possess characters which are typical of species of section Pogostemon, e.g. toothed, hairy, petiolate leaves borne in opposite pairs. Similarly two species of section Pogostemon, P. micangensis (OTU 85) and P. mutamba (OTU 86) (Fig. 30) which grow along watercourses in southern tropical Africa resemble species of section Eusteralis in their general habit, branching and very shortly petiolate leaves. Both species are normally hairy but the type specimen of P. micangensis, which according to the field 64 J. R. PRESS notes was ‘an aquatic herb in the marshes of the river Micango’ is virtually glabrous. This evidence thus supports the union of Dysophylla and Pogostemon, since many of the characters formerly separating the genera are ecological adaptations in particular species. Subgeneric divisions Bentham (1832-36) divided Pogostemon sensu stricto into two groups § Paniculatae and § Racemosae on the basis of inflorescence structure. Briquet (1897) adopted Bentham’s divisions, expanded the characters on which they were based and further subdivided each group. His characters for these groups (Table 13) do not contrast; for example § Paniculatae has linear- subulate bracts while § Racemosae has stamens which are hairy all over. If the characters defining each group are scored for the species of the other group only one character remains valid for defining them, i.e. spikes paniculately branched versus spikes simple. This character varies between specimens and in my view the variation of the inflorescence is better expressed as verticils not secund versus verticils sub-secund (see character 62 in Table 3). The clustering to maximize WGMS analysis (Table 7, Appendix 4) indicates the presence of two groups (groups five and seven) which roughly correspond to Bentham’s and Briquet’s groups. However, there is sufficient overlap to prevent clear definition of these groups, nor do they appear in the other analyses, and I have not formally recognized them. Briquet’s characters for defining the subdivisions A and B in each group (Table 13) do contrast but are misapplied. His Glabriuscula and Barbata in § Racemosae are distinguished by the stamens being naked or hairy respectively. All species of Pogostemon have hairy stamens, although some species, e.g. P. travancoricus (OTU 92) (Fig. 8), bear hairs only on the lower portions of the filaments which are not clearly visible from examination of an undissected flower. However, even if Briquet’s character is substituted by the one ‘filaments hairy in the lower half versus filaments hairy in the upper half’ no useful division of the species can be made, since, for example P. rotundatus Benth. (OTU 101) with filaments hairy in the lower half would be placed in a separate group from P. mollis (OTU 91) with filaments hairy in the upper half, even though they share a very high overall similarity and are each other’s nearest neighbour (Appendix 3). Similarly the inflorescence character used by Briquet to divide A and B in § Paniculatae (Table 13) does not provide a clear or useful division of the species, and I have discarded his groups. Table 13 Names and diagnoses of Briquet’s (1897: 328-329) subdivisions of Pogostemon. §1. Racemosa Benth. Spicastra einfach, unterbrochen. Bracteen lineal-pfriemlich, kiirzer als der Kelch. A. Glabriuscula Briq. Stb. mit nacten oder fast kahlen Stf. B. Barbata Briq. Stb. mit deutlich dichtbartigen Stf. §2. Paniculata Benth. Spicastra rispig verzweigt. Stb. iiberall mit bartig behaarten Stf. A. Scheinwirtel meistens entfernt, in unterbrochenen Spicastris. B. Scheinwirtel meistens in dicken oder Spicastris (vergl. oben P. suave). Briquet also divided Dysophylla into two sections, section Rhabdocalicinae and section Goniocalicinae (Table 14). Here too, there is only one contrasting character. Members of both sections have verticillate, sessile and entire leaves, and may be annual, leaving only the character calyx-tube terete versus calyx-tube strongly five-angled. The opposite-leaved species of his section Rhabdocalicinae are those which form Bentham’s section Oppositifoliae and are removed to Pogostemon sensu stricto. As with Pogostemon sensu stricto the analyses show no evidence of groupings which match those of Briquet. Thus I have discarded these sections. Relocated OTUs Dysophylla mairei is discussed on p. 57. Elsholtzia aquatica All analyses group Elsholtzia aquatica (OTU 22) with Pogostemon sensu lato, and examination of the characters readily show that its inclusion in Elsholtzia is erroneous. It has linear, sessile, Fig. 30 Pogostemon mutamba. (a) habit x 1. (b) calyx x 10. (c) corolla x 5. (d) dissected flower x 5. (e) stamen X 10. (f) nutlet, inner face, and profile x 10. 66 J. R. PRESS verticillate leaves, an inflorescence of numerous small flowers in crowded whorls, a corolla with an entire upper lip which is shorter than the lower lip, stamens with unilocular anthers and hairy filaments, and no tumescent gland on the disc. None of these characters is shared by any species of Elsholtzia, but they all distinguish Pogostemon sensu lato and in particular Pogostemon section Eusteralis. Elsholtzia aquatica is thus transferred (see p. 73). Table 14 Names and diagnoses of Briquet’s (1897: 330-331) subdivisions of Dysophylla. Sect. I. Rhabdocalicinae Briq. Kelchrohre cylindrisch und stielrund oder sehr undeutlich, stumpf Seckig. B. gegen-oder quirlstandig, gezahnt oder ganzrandig. §1. Oppositifoliae Benth. B. gegenstandig A. Ausdauernde Arten, mit meistens holziger entwickelter Wurzel oder unteriridischen Stengel. B. Einjahrige Art, abstehend behaart, mit sitzenden oder kurz gestielten, eilanglichen, gesagten B, behaarten Spicastris und 3eckigen, zur Fruchtzeit nach innen gebogenen Kelchzahnen. §2. Verticillatae Benth. B. in Quirlen zu 3 oder 4 (selten bis 10). Sect. II. Goniocalicinae Briq. Kelchrohre Skantig, mit vorspringenden Kanten. 1jahrige Arten mit quirl- standigen, sitzenden, ganzrandigen B. Distribution of Pogostemon The genus Pogostemon has a disjunct distribution (Fig. 31); most species are confined to the Indo-Chinese and Malesian regions and Japan with P. stellatus (Lour.) Kuntze extending as far as northern Australia, but three species, P. aquaticus (C. H. Wright) Press (OTU 22), P. micangensis (OTU 85), and P. mutamba (OTU 86) (Fig. 30) are confined to southern tropical Africa. There is no evidence for any species of Pogostemon occurring in the intervening areas and the nearest members of the tribe are the three species of Tetradenia which are found in Madagascar. The analyses do not separate the African species from the Asian and Australian species in any way and there is no morphological evidence to help explain the distribution. Colebrookea Colebrookea is the most isolated of all the genera within the Pogostemoneae and has no strong affinities with any of the other taxa. The nearest neighbours of C. oppositifolia (OTU 138) and C. ternifolia (OTU 137) occur in Pogostemon and Elsholtzia section Aphanochilus sensu stricto (Appendix 3). Previous authors have neither discussed nor commented on the affinities and relationships of Colebrookea beyond placing it next to Pogostemon (see Bentham, 1832-36; Bentham & Hooker, 1876; Hooker, 1885; Briquet, 1897; Kudo, 1929; Wu, 1977) with which it shares the characters of unilocular anthers and an equal disc. These authors have given undue emphasis to the rather tenuous relationship between it and Pogostemon. While my analyses confirm this, they also show that Colebrookea is not significantly more similar to Pogostemon than it is to Elsholtzia. Grouping Colebrookea with Pogostemon has also tended to obscure the recognition of Colebrookea as a distinct genus which is so apparent in my results. Bentham (1832-36) believed Tetradenia to be an intermediate genus between Colebrookea and Elsholtzia, but I have found no supporting evidence for this. Colebrookea is easily recognized by the distinctive features of the fruiting calyx, particularly the calyx teeth becoming greatly elongated and plumose (Fig. 6). The single, hairy nutlet (Fig. 10) does not fall from the calyx when ripe, but remains firmly attached to the disc, fruit and calyx acting as a diaspore. The plumose calyx teeth give the infructescence a characteristic fluffy appearance. Colebrookea may also be recognized by a combination of characters which are found in other genera within the Pogostemoneae. Like Pogostemon sensu lato it has an equal disc and unilocular anthers, but the filaments are naked (Fig. 8) and the upper lip of the corolla (Fig. 7) is emarginate. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 67 Se J U T£IDYO BP Fig. 31 Distribution of Pogostemon sensu lato. 7. Taxonomic conspectus Key to genera and sections 1a. Anthers bilocular; disc with one or more tumescent lODES................:eceeceeeee ee eee ee ee eee ee een ee ees Ps 1b. Anthers unilocular; disc without tumescent IObES ................00ccceceeeeeeceeecererreeeeeeeseseecenenseees 4 2a. Disc with four large, bright-red, tumescent lobes; upper tooth of calyx broad and overlapping the lateral teeth................-...esseseeeeeeeeee ees VI. Tetradenia (p. 70) 2b. Disc with one tumescent lobe; upper tooth of calyx not overlapping Hive lateral tC ety os comet rer eee cea Onegin amg ein ak ones Reeuaodniath race tes tdomens ean Ties =] 3a. Calyx with a well-defined annulus of white hairs at the mouth of the tube: mature nutlets.OME .ccsicceteossorccs con cxenssa reeves npisenaectsreessorgcsasreene V. Keiskea (p. 70) 3b. Calyx without an annulus; mature nutlets usually four .............----sssseeeeeeeeees IV. Elsholtzia (p. 69) ia. Bracteoles present; bracts longer than broad.................- IVb. section Aphanochilus (p. 69) ib. Bracteoles absent; bracts at least as broad as LON ...............ecee sees ee ee eee ee ee ee eee ee ne nese ees il iia. Fruiting calyx inflated, much broader than at anthesis; nutlets verrucose; style tips clavate ............::ecseeeeereeeees IVc. section Platyelasma (p. 70) iib. Fruiting calyx sometimes enlarged but never inflated, not much broader than at anthesis; nutlets not verrucose; Style tips SUbUIALE ce. ccrs dacaactsstiewasonietnnn shor seenentnenecssenen IVa. section Elsholtzia (p. 69) 4a. Bracts at least as broad as long, membranous, caducous; the whole plant covered in a dense indumentum of branched, stalked hairs ..................eeeeee sees ee eens eee eee es a 4b. Bracts longer than broad, not membranous, persistent; the indumentum variable but never of dense, branched, stalked hairs...............::seeeeeeee esse eee ee ee ee nese ee sess ee ness a! 68 J. R. PRESS 5a. Mature nutlets with a pronounced rostrate tip; corolla with an entire upper lip, hairy disc-like excrescences at the base of the stamen filaments and a hairy crescent-shaped invagination at the base of the Fea Uy oa ne iE ee RHE Oa WAR AEC rN SR ayer CRU eR AR III. Rostrinucula (p. 68) 5b. Mature nutlets without rostrate tips; corolla with an emarginate upper lip and lacking excrescences at the bases of the stamen filaments and an invagination at the base of the lower lip ......o.-dcncssnacssssevadcecares sein esdwsw anes 6 6a. Stamen filaments shortly hairy towards the base; corolla Without an annwlus Of NAS. cyeg cess: eas sens. 20s Ponwcoceas neomnteancaeeepuaonsaetcs II. Leucosceptrum (p. 68) 6b. Stamen filament completely glabrous; corolla with an SUMUNIS ORM AINS hace occ l ot havas cpus eoeesrece saan suss oun ore teng ote seaiees I. Comanthosphace (p. 68) 7a. Stamen filaments glabrous; upper lip of corolla emarginate.................ccccececeeceeeeeeee sees eeeenees 8 7b. Stamen filaments with long, often purplish hairs; MNpeclipor corolla entire oes sgrce coves os Lites drain wacde ere ociouaeetecasnaets VIII. Pogostemon (p. 71) iiia. All leaves opposite, usually petiolate; upper lip of corolla equalling or longer than the lower lip......................05. VIIIa. section Pogostemon (p. 71) iiib. At least some leaves in whorls of three or more, usually sessile; upper lip of corolla equalling or shorter than tHe lO WET UN seo ere ches 5 Vcc tans Adee haan ct eae mes VIIIb. section Eusteralis (p. 73) 8a. Calyx teeth plumose in fruit; corolla tube straight ...................cseceeeeeee eens IX. Colebrookea (p. 74) 8b. Calyx teeth not plumose on fruit; corolla tube gibbous towards 1 07] Ok Cl nee Aart terete en Rian USUAL A PETAR SR EN Sr eh AMER Ee eR MPN mR AOE VII. Eurysolen (p. 70) Conspectus In the following conspectus diagnoses are given for supra-specific taxa only. Species are listed alphabetically within each genus and section. This study is concerned primarily with supra- specific groups and does not extend to an assessment of individual species. However, some species share very high similarity values and may well prove to be conspecific. In the conspectus species which are indented are possibly conspecific with the species which immediately precedes them. I. Comanthosphace S. Moore in J. Bot., Lond. 15:293 (1877). Leaves ovate, petiolate; indumentum composed of branched and unbranched eglandular hairs; bracts at least as broad as long, membranous, caducous; bracteoles narrow, caducous; calyx sub-equally five-toothed; corolla bilabiate, upper lip emarginate, the throat with a complete (interrupted in C. nanchuanensis) annulus of hairs; stamen filaments glabrous, anthers unilocu- lar; disc equal; mature nutlets four, sparsely hairy (glabrous in C. nanchuanensis). Pollen grains bi-nucleate/tri-colpate. C. barbinervis (Miq.) S. Moore C. formosana Ohwi C. nanchuanensis C. Y. Wu & Li C. ningpoensis (Hemsley) Hand.-Mazz. C. stellipila (Miq.) S. Moore C. japonica (Miq.) S. Moore C. sublanceolata (Miq.) S. Moore | II. Leucosceptrum Smith, Exot. bot. 2:113, t.116 (1805). Leaves ovate, petiolate; indumentum composed of branched and unbranched eglandular hairs; bracts at least as broad as long, membranous, caducous; bracteoles narrow, caducous; calyx equally five-toothed; corolla bilabiate, upper lip very short, emarginate; stamen filaments shortly hairy towards the base, anthers unilocular; disc equal; mature nutlets four, glabrous. Pollen grains bi-nucleate/tri-colpate. L. canum Smith III. Rostrinucula Kudo in Mem. Fac. Sci. Agric. Taihoku imp. Univ. 2(2):304 (1929). Leaves ovate, petiolate; indumentum composed of branched and unbranched eglandular hairs; TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 69 bracts at least as broad as long, membranous, caducous; bracteoles narrow, caducous; calyx sub-equally five-toothed; corolla bilabiate, upper lip entire, the throat with an interrupted annulus of hairs borne on disc-like excrescences at the base of the stamen filaments and on a crescent-shaped invagination at the base of the lower lip; stamen filaments naked, anthers unilocular; disc equal; mature nutlets four, sparsely hairy, the apices extended to form a strongly-hooked beak. Pollen grains bi-nucleate/tri-colpate. R. dependens (Rehder in Sargent) Kudo R. sinensis (Hemsley) C. Y. Wu IV. Elsholtzia Willd. in Bot. Mag. 4:3 (1790). Leaves narrowly ovate to orbicular, petiolate; indumentum composed of unbranched eglandu- lar hairs or a mixture of unbranched glandular and eglandular hairs, sometimes with branched stalked hairs; verticils sometimes secund with one-many flowers; bracts linear to broader than long, sometimes membranous, persistent; bracteoles, when present, narrower than the bracts, persistent; calyx with five equal teeth or the upper three shorter than the lower two; corolla bilabiate, upper lip emarginate, the throat with or without an interruted annulus of hairs; stamen filaments glabrous, anthers bilocular, the locules confluent through partial fusion at the apex or free (E. hunanensis); disc with a single tumescent lobe; mature nutlets usually four, rarely one, glabrous. Pollen grains tri-nucleate/hexa-colpate. IVa. Section Elsholtzia Cyclostegia Benth. in Bot. Reg. 15: sub t. 1282 (1829). Elsholtzia section Cyclostegia (Benth.)Benth., Lab. gen. sp.:163 (1833). Spikes usually secund, sometimes cylindrical; bracts at least as broad as long, usually membra- nous, veined, imbricate, the members of a pair sometimes fused to form a cyathium; bracteoles absent; fruiting calyx sometimes accresant but never inflated; style lobes subulate; nutlets rugulose or nearly smooth. E. bodinieri Vaniot E. ciliata (Thunb.) Hylander E. concinna Vaut. E. feddei Léveillé ‘E. elegans Franch.’—an apparently unpublished name applied by Merrill to material in E. E. heterophylla Diels E. hunanensis Hand.-Mazz. E. kachinensis Prain E. luteola Diels E. oldhamii Hemsley E. argyi Léveillé E. nipponica Ohwi E. pseudocristata Léveillé & Vaniot E. soulei Léveillé E. pygmaea W. Smith E. strobilifera (Benth. in Wallich) Benth. IVb. Section Aphanochilus (Benth.) Benth., Lab. gen. sp.:161 (1833). Aphanochilus Benth. in Bot. Reg. 15: sub t. 1282 (1829). Spikes usually cylindrical, rarely secund; bracts linear to ovate, not membranous or imbricate, always free; bracteoles similar to bracts but usually narrower; fruiting calyx sometimes accresant but never inflated; style lobes subulate; nutlets rugulose. E. alopecuroides Léveillé & Vaniot E. beddomei C. B. Clarke ex Hook. f. E. blanda (Benth.) Benth. E. capituligera (Dunn) C. Y. Wu E. communis (Collett & Hemsley) Diels E. elata Zoll. & Mor. E. flava (Benth. in Wallich) Benth. 70 J. R. PRESS E. fruticosa (D. Don) Rehder Leucosceptrum plectranthoideum (Léveillé) Marquand ‘E. glanduligera’—an unpublished name applied by C. B. Clarke to material in K which might represent a new species. More material is required for assessment. E. griffithii Hook. f. E. myosurus Dunn E. ochroleuca Dunn E. penduliflora W. Smith E. pilosa (Benth. in Wallich) Benth. Dysophylla mairei Léveillé E. pubescens Benth. E. rugulosa Hemsley E. stachyodea (Link) Raiz. & Saxena E. stauntonii Benth. E. winitiana Craib. IVc. Section Platyelasma (Briq.) Press, stat. nov. Elsholtzia section Aphanochilus series Platyelasmeae Briq. in Engl. & Prantl, Natiirl. Pflanzenfam. 4 (3a):327 (1897). Platyelasma Kitagawa in Rep. Scient. Exped. Manchoukuo 1933, 4 (2):26 (1935). Spikes cylindrical; bracts at least as long as broad, not membranous or imbricate, free; bracteoles absent; fruiting calyx inflated in fruit; style lobes with clavate tips; nutlets verrucose. E. densa Benth. E. manshurica (Kitagawa) Kitagawa E. eriostachya (Benth. in Wallich) Benth. V. Keiskea Mig. in Annls Mus. bot. Lug.-Bat. 2:105 (1865). Leaves ovate, petiolate; indumentum composed of unbranched eglandular hairs only; verticils secund, with two flowers; bracts ovate, not membranous, persistent; bracteoles absent; calyx five-toothed, the upper three teeth shorter than the lower two, the throat with an annulus of white hairs; corolla bilabiate, upper lip emarginate, the throat with a complete annulus of hairs; stamen filaments glabrous, anthers bilocular, the locules free; disc with a single tumescent lobe; mature nutlets one (?), glabrous and with a somewhat reticulate pattern of ridges. Pollen grains tri-nucleate/hexa-colpate. K. elsholtzioides Merrill K. glandulosa C. Y. Wu K. japonica Miq. K. sinenensis Diels K. szechuanensis C. Y. Wu VI. Tetradenia Benth. in Bot. Reg. 15:sub t. 1300 (1829). Leaves ovate, petiolate; indumentum composed entirely of unbranched eglandular hairs or mixed with branched stalked hairs; bracts broader than long, somewhat membranous, persis- tent; bracteoles absent; calyx five-toothed, the upper tooth much broader than, and overlap- ping, the two lateral teeth; corolla subilabiate, upper lip deeply emarginate, the throat with a complete annulus of hairs; stamen filaments glabrous, anthers bilocular, the locules confluent through partial fusion at the apex; disc with four bright-red, tumescent lobes equally spaced around the edge; mature nutlets four, glabrous. Pollen grains tri-nucleate/hexa-colpate. T. fruticosa Benth. T. hildebrandtii Briq. T. goudotii Briq. VII. Eurysolen Prain in Scient. Mem. med. Offrs Army India 11:43 (1898). Leaves ovate, petiolate; indumentum composed of unbranched eglandular hairs only; bracts ovate, persistent; bracteoles similar but narrower, persistent; calyx subequally five-toothed; corolla bilabiate, upper lip emarginate, tube gibbous, the throat with an interrupted annulus of TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE eal hairs borne on disc-like excrescences at the base of the stamen filaments and on a papilla-like invagination on the ventral surface of the tube just below the gibbous curve; stamen filaments glabrous, anthers unilocular; disc equal; mature nutlets four, glabrous. Pollen grains bi- nucleate/tri-colpate. E. gracilis Prain VIII. Pogostemon Desf. in Mem. Mus. Hist. nat. Paris 2:154 €1815). Leaves linear to orbicular, sessile or petiolate; indumentum composed of unbranched eglandu- lar hairs; unbranched eglandular and glandular hairs or branched sessile hairs; verticils sometimes subsecund; bracts linear to ovate, not membranous, persistent; bracteoles similar but narrower, persistent; calyx sub-equally five-toothed or with the three upper teeth shorter than the two lower; corolla bilabiate or subilabiate, upper lip entire; stamen filaments hairy in the middle or towards the base, the hairs long and often purplish, anthers unilocular; disc equal; mature nutlets four, rarely one, glabrous. Pollen grains bi-nucleate/tri-colpate. VIIla. Section Pogostemon Dysophylla section Oppositifoliae Benth., Lab. gen. sp.:157 (1833). Leaves in opposite pairs, usually petiolate; corolla with the upper lip equalling or longer than the lower lip. P. amarantoides Benth. in DC. P. andersonii (Prain) Press, comb. nov. Dysophylla andersoni Prain in J. Asiat. Soc. Beng. 59:298 (1891). . atropurpureus Benth. in DC. . auricularius (L.) Hassk. . benghalensis (Burm. f.) Kuntze P. parviflorus (Benth. in Wallich) Benth. . brachystachyus Benth. in DC. . brevicorollus Y. Z. Sun . cablin (Blanco) Benth. in DC. . championii Prain . dielsianus Dunn . elsholtzioides Benth. in DC. P. fraternus Miq. P. formosanus Oliver in Hook.f. P. gardneri Hook.f. P. glaber Benth. in Wallich P. glabratus Chermsirivathana ex Press, sp. nov. Hen a ~ a - Jaen - Bi = i = Sa a | (Fig. 32) A P. auricularius (L.) Hassk. caulibus et foliis glabris, nodis subito contractis, differt. Herbae glabrae vel interdum remotis pilis eglandulatis. Caules subquadrangulati sulcis non profundis, nodis subito contractis. Folia opposita, 4-6 X 2-3 cm, ovata acuta cuneata, serrata vel dentata, petioli 2-5 mm. Inflorescentia terminalis ad 13 cm longa, densa, verticillastris numerosis multifloribus. Bracteae 2-3 mm longae, ovatae; bracteolae ad 2 mm longae, lanceolatae. Calyx campanulatus, tubo 1 mm longo, dentibus quinque 0-5 mm longis, triangularibus, sub fructu incurvatis. Corolla + bilabiata purpurea, tubo 1-5 mm longo exserto, labio superiori 0-5 mm longo integro, lobis lateralibus labio superiori similibus, labio inferiori 0-5 mm longo integro. Stamina quator filamentis exsertis, pari inferiori longiori, prope medianum pilis longis barbatis; antherae uniloculares. Discus non lobatus. Nuculae maturae quatuor. Type: Siam, Hin Dat, Kanburi, 14 July 1926, Put 132 (BM !-holotype). Other material: Siam, Ta Kanun, Kanburi, ‘flowers pale purple, cultivated by Karens’, 20 January 1926, Kerr 10274 (BM !) LOcaL NAMES: Niam [Kerr]; Pak hom hang nu [Put]. Known only from the Kanburi area of Thailand. P. griffithii Prain P. heyneanus Benth. in Wallich P. hirsutus Benth. P. hispidus Prain P. litigiosus Doan in Humbert b Fig. 32 Pogostemon glabratus Chermsirivathana ex Press [ = ‘Dysophylla glabrata’, see p. 19]. (a) habit 1. (b) calyx x 20. (c) corolla x 20. (d) dissected flower X 20. (e) stamen x 40. (f) nutlet, inner face, and profile x 20. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 73 . macgregorii W. Smith . menthoides Blume . micangensis G. Taylor . mollis Benth. . mutamba (Hiern) G. Taylor nelsonii Doan nigrescens Dunn . nilagiricus Gamble paludosus Benth. in DC. . paniculatus (Willd.) Benth. in Wallich phillipensis S. Moore pubescens Benth. in DC. purpurascens Dalz. in Hook f. . reflexus Benth. in DC. . reticulatus Merrill . rotundatus Benth. in Wallich . myosuroides (Benth. in Wallich) Kuntze P. rugosus (Hook.f.) El-Gazzar & Watson . rupestris Benth. . Salicifolius (Dalz. ex Hook.f.) Kuntze P. speciosus Benth. in Wallich P. strigosus Benth. in DC. P. travancoricus Beddome P. tuberculosus Benth. in Wallich P. velatus Benth. in DC. P. williamsii Elmer P. villosus Benth. P. wattii C. B. Clarke P. battakianus Ridley P. wightii Benth. VIIIb. Section Eusteralis (Rafin.) Keng in Fl. Males. 8(1):352 (1978). Eusteralis Rafin., Fl. Tellur. 2(4):95 (1837). Dysophylla section Verticillatae Benth., Lab. gen. sp.:158 (1833). At least some leaves in whorls of three or more, usually sessile; the corolla with the upper lip equalling or shorter than the lower lip. P. aquaticus (C. H. Wright in Dyer) Press, comb. nov. Elsholtzia aquatica C. H. Wright in Dyer, Fl. Trop. Afr. 5:451 (1900). P. crassicaulis (Benth. in Wallich) Press, comb. nov. Dysophylla crassicaulis Benth. in Wallich, Pl. As. Rar. 1:30 (1830). P. cruciatus (Benth. in Wallich) Kuntze P. deccanensis (Panigr.) Press, comb. nov. Eusteralis deccanensis Panigr. in Phytologia 32(6):475 (1978). Dysophylla tomentosa Dalz. in Hooker’s J. Bot. 2:337 (1850), non Pogostemon tomentosa Hassk. (1844). P. falcatus (C. Y. Wu) C. Y. Wu & Li P. faurei (Léveillé) Press, comb. nov. Dysophylla faurei Léveillé in Reprium Spec. nov. Regni veg. 9:248 (1911). P. erectum Kuntze Dysophylla gracilis Dalz. in Hooker’s J. Bot. 2:337 (1850), non Pogostemon gracilis Hassk. (1843). P. griffithii (Hook.f.) Press, comb. nov. Dysophylla griffithii Hook.f., Fl. Brit. India 4:641 (1885). P. helferi (Hook.f.) Press, comb. nov. Dysophylla helferi Hook.f., Fl. Brit. India 4:640 (1885). US RUSS VV VV UV VU DUD 74 J. R. PRESS P. koehneanus (Muschler) Press, comb. nov. Dysophylla koehneana Muschler in Reprium nov. Spec. Regni veg. 4:269 (1907). P. linearis (Benth. in DC.) Kuntze P. lythroides (Diels) Press, comb. nov. Dysophylla lythroides Diels in Notizbl. bot. Gart. Mus. Berl. 9:1031 (1926). P. peguanus (Prain) Press, comb. nov. Dysophylla peguana Prain in J. Asiat. Soc. Beng. 59:298 (1891). P. pentagonus (C. B. Clarke ex Hook.f.) Kuntze P. pumilus (Graham) Press, comb. nov. Mentha pumila Graham in Edinb. New phil. J. 4:393 (1828) [non Host (1831)]. Dysophylla pumila (Graham) Benth. in Wallich, Pl. As. Rar. 1:30 (1830). P. quadrifolius (Benth. in Wallich) Kuntze P. sampsonii (Hance) Press, comb. nov. Dysophylla sampsoni Hance in Annls Sci. nat. V , 5:284 (1866). P. stellatus (Lour.) Kuntze P. stocksii (Hook.f.) Press, comb. nov. Dysophylla stocksii Hook.f., Fl. Brit. India 4:642 (1885). P. szemacensis (C. Y. Wu & Hsuan) Press, comb. nov. Dysophylla szemacensis C. Y. Wu & Hsuan in Acta phytotax. sin. 10:238 (1967). P. trinervis Chermsirivathana ex Press, sp. nov. Fig. 33. Inter species section Eusteralis (Rafin.) Keng foliis latis triplinervibus distinguilibus. Herbae hirtae parcae pilis glanduliferis vel eglandulatis. Caules ad 30 cm longi, bases versus prostratae supra erectae ad nodos radicantes. Folia sessilia in verticilli e tres dispositi, 6-13 x 4-7 mm, ovata vel rotundata, acuta vel obtusa, cuneata, serrata vel dentata, triplinervia nervo medio et pari laterali impresso subtus elevata. Inflorescentiae terminales, ad 5 cm longae, bases interruptes, verticillastris 5-10 (—15), flores 6-20 in omnis verticillaster. Bracteae 2-3 mm longae, lineares; bracteolae ad 2 mm longae, lineares vel lineares-subulatae. Calyx campanulatus, tubo 1-1-5 mm longo, dentibus quinque 0-5 mm longo, triangularibus sub fructu erectis vel leviter patentibus. Corolla + bilabiata purpurea, tubo 0-75 mm longo exserto integro, lobis lateralibus labio superiori similibus, labio inferiori 0-25 mm longo integro. Stamina quatuor filamentiis exsertis, pari superiori longiori, prope medianum pilis longis barbatis; antherae uniloculares. Discus non lobatus. Nuculae maturae quatuor. Type: Siam, Hui Taleng, Korat, 24 December 1928, Put 2223 (BM !-holotype). Other material: Siam, Korat, Pak Tong Chai, ‘flowers purple; among short grass on open ground’, c. 200 m, 25 December 1923, Kerr 8115 (BM). Known only from the Korat area of Thailand. P. tsiangii (Y. Z. Sun) Press, comb. nov. Dysophylla tsiangii Y. Z. Sun in Acta phytotax. sin. 11:50 (1966). P. yatabeanus (Makino) Press, comb. nov. Dysophylla yatabeanus Makino in Bot. Mag., Tokyo 1:55 (1898). IX. Colebrookea Smith, Exot. bot. 2:111, t.115 (1805). Leaves ovate, petiolate; indumentum composed of unbranched eglandular hairs only; bracts linear, not membranous, persistent; bracteoles similar but smaller, persistent; calyx with five equal, long, slender, plumose teeth; corolla bilabiate, upper lip emarginate; stamen filaments glabrous, anthers unilocular; disc equal; mature nutlets one, hairy. Pollen grains bi-nucleate/ tri-colpate. C. oppositifolia Smith C. ternifolia Roxb. Species excludendae Elsholtzia integrifolia Benth. = Schizonepeta tenuifolia (Benth.) Briq. ‘Elsholtzia japonica’ sensu d’ Argy, non Miq. = Agastache sp. (? rugosus Fisch. & Mey.). inner face, and ,’ see p. 14]. (a) habit x 1. (b) calyx x 20. (c) corolla x 20. (d) dissected flower x 20. (e) stamen x 40. (f) nutlet, Fig. 33 Pogostemon trinervis Chermsirivathana ex Press [ = ‘Dysophylla trinervia profile x 20. 76 J. R. PRESS Appendix 1. Specimens used for scoring and producing averaged data For the names to which the acronyms refer see Table 2. EGRA 1: Putin Herb. Kerr 4404 (BM), Robinson & Kloss 130 (BM), s.n. 1914 (BM). EJAP 2: d’Argy s.n. Kiangsu (E). EFED 3: Forrest 22386 (E), Soulie 227 (E—holotype). EELE 4: Bau Hwa Shan 1490 (E). EPSE 5: Taquet 1223 (K—paratype), 1224 (K—paratype). ENIP 6: Ikeo 9567 (TI—isotype), Kurata s.n. 1964 (BM), Mayebara 4125 (TI). EOLD 7: Oldham s.n. 1864 (K—holotype), Tagawa 4162 (TI), Yamazaki 3059 (TI). ECON 8: Ludlow & Sherriff 9057 (BM), Polunin 1020 (BM), 1612 (BM). ECIL 9: Togasi 787 (BM), Wilson 5717 (BM), Yu 10728 (BM). EKAC 10: Forrest 6729 (E), Kingdom-Ward 20365 (BM), Put in Herb. Kerr 3478 (BM). EPYG 11: Forrest 17128 (E—holotype). ESOU 12: Maire 321 (E), 599 (E), Soulie 226 (E—holotype). EARG 13: Cavalerie 7993 (E), Léveillé s.n. (E). ELUT 14: Forrest 11145 (E), 15184 (E), Yu 13952 (BM). EBOD 15: Bodinier 2 (E—holotype), Forrest 2997 (BM), 7371 (BM). EHET 16: Forrest 934 (E—holotype), Maire 1203 (E), [Collector unknown] 096035 (TAI). ESTR 17: Ludlow, Sherriff & Taylor 7139 (BM), Nicolson 2652 (BM), Stainton, Sykes & Williams 8526 (BM). EHUN 18: Handel-Mazzetti 2702 (E—isotype). EERI 19: Dhwoj 0180 (BM), Ludlow, Sherriff & Hicks 16682 (BM), Stainton, Sykes & Williams 8102 (BM). EDEN 20: Ludlow, Sherriff & Elliot 14130 (BM), Stainton, Sykes & Williams 2169 (BM), 8102 (BM). EMAN 21: Nakai, Honda & Kitamura s.n. 1933 (TI—holotype), Togashi 828 (TI), 1578 (TI). EAQU 22: Johnson 15 (K—holotype), Fanshawe 8503 (K), 8512 (K). EINT 23: Staunton s.n. (BM—holotype). EBED 24: Beddome 147 (BM—holotype), MacGregor 69 (E), 608 (E). EPEN 25: Forrest 11686 (E—holotype), Yu 17600 (E). ERUG 26: Forrest 6527 (BM), Maire 633 (E), Yu 16748 (E). ESTA 27: Jackson s.n. 1929 (BM), Licent 2996 (BM), Staunton s.n. (BM—holotype). EFLA 28: Forrest 6283 (BM), Stainton 5090 (BM), Wallich 1553 (BM—isotype). EFRU 29: McLaren’s native collectors 326d (BM), Polunin, Sykes & Williams 3126 (BM), Young s.n. 1880 (BM). EPUB 30: Horsefield 337 (BM). ECOM 31: Maire 561 (BM), Yu 14498 (BM), 14712 (BM). EALO 32: Cavalerie 1426 (E—holotype). EGRI 33: Haines s.n. 1914 (E), Lace 4391 (E). EGLA 34: Clarke 37393 (K). EELA 35: Koorders 37621B (K), Herb. Kuntze 5613 (K), Ridley s.n. 1915 (K). EWIN 36: Kerr 1607 (BM), s.n. (BM), Put in Herb. Kerr 4412 (BM). ESTC 37: Buchanan s.n. 1802 (BM—holotype of Elsholtzia leptostachya), Clarke 23765 (BM), Polunin, Sykes & Williams 5827 (BM). EBLA 38: Ludlow, Sherriff & Taylor 7075 (BM), Norkett 8631 (BM), Stainton, Sykes & Williams 8323 (BM). EMYO 39: Forrest 7220 (E—holotype), Wilson 3533a (K). EOCH 40: Maire s.n. (E—holotype of Elsholtzia lampradena). EPIL 41: Forrest 28965 (BM), Maire 1204 (BM), Stainton, Sykes & Williams 4452 (BM). ECAP 42: Forrest 1680 (BM), 20697 (BM), 22956 (BM). DTRI 43: Kerr 8115 (BM), Putin Herb. Kerr 2223 (BM). DHEL 44: Helfer 194 (BM), 3968 (K—holotype). DTOM 45: Buchanan s.n. (BM), Hohenacker 371 (BM), Young s.n. 1879 (BM). DPEN 46: Carlke 20438 (BM—isotype), Garrett in Herb. Kerr 59 (BM), Kerr 1465 (BM). DSTO 47: Stocks s.n. in Herb. Hooker (K—holotype). DSAM 48: Sampson in Herb. Hance 10946 (BM—holotype). DPEG 49: Kurz 2401 (K), 2405 (K), Putin Herb. Kerr 1971 (BM). DGRI 50: Chattaya 5224 (K), Gamble 13748 (K), Griffith 3968 (K). DYAT 51: Science College imp. Univ. Japan s.n. 1883 (K—isotype), Tanaka 7338 (TAI). DCRA 52: Clarke 23691 (BM), Griffiths 1024 (BM), Wallich 1545 (BM—isotype). DSTE 53: Clarke 8073 (BM), Loureiro s.n. (BM—holotype), Simpson 9189 (BM). DLIN 54: Clarke 45731 (BM), Kingdom-Ward 14260 (BM), Ludlow, Sherriff & Hicks 21019 (BM). DCRU 55: Beddome s.n. (BM), Clarke 18059 (BM), Polunin, Sykes & Williams 5875 (BM). DQUA 56: Roxburgh s.n. (BM—isotype), Rugel s.n. (BM), Wallich 1538 (BM). DTSI 57: Tsiang 9449 (KUN—isotype). DFAL 58: Wang 79441 (KUN— isotype). DSZE 59: Tsiang 12713 (KUN—isotype). DFAU 60: Faurie 760 (E—holotype). DKOE 61: Hosseus 704 (BM—isotype). DGRA 62: Dalzell s.n. (K—holotype), Stocks & Law s.n. Malabar & Concan (BM). DLYT 63: Fan & Li 563 (BM). DPUM 64: Wallich 1546 (K). DMAI 65: Maire s.n. 1911 (E—holotype). DAND 66: Anderson s.n. 1867 (K—holotype). DGLA 67: Kerr 10274 (BM), Putin Herb. Kerr 132 (BM). DAUR 68: Clarke 26464 (BM), Forbes 89 (BM), Stainton, Sykes & Williams 6481 (BM). DSAL 69: Dailzell s.n. (K—holotype), Lain s.n. in Herb. Hooker (K), Young s.n. 1882 (BM). DRUG 70: Beddome s.n. (BM), Herb. Rottler s.n. 1828 (K), Wallich 1547 (K-W—holotype). DMYO 71: Fricker 4724 (K), Koenig s.n. (BM), Wallich 1547 (K-W—holotype). PBEN 72: Roxburgh s.n. India (BM), Stainton 13 (BM), 5220 (BM). PPAR 73: Andrews 348 (BM), Beddome s.n. (BM), Metz 1393 (BM). PPAN 74: Stocks & Laws.n. Malabar & Concan (BM), Thomson s.n. Mont. Nilghiri & Kurg. (BM), Wallich 1561 (BM). PTUB 75: Clarke 26368 (BM), Ludlow, Sherriff & Taylor 6759 (BM), Sarail 23 (BM). PGLA 76: Flatt 161 (BM), Hooker s.n. Sikkim (BM), Nicolson 2941 (BM). PHEY 77: MaCrae 737 (BM), Robinson & Kloss 88 (BM), Thomson s.n. Maisor (BM). PCAB 78: Clemens s.n. 1924 (BM), Horsefield s.n. Java (BM), Ramos 22432 (BM). PELS 79: Griffiths 1018 (BM), Kingdom-Ward 14234 (BM), Ludlow, Sherriff & Taylor 7211 (BM). PAMA 80: Hooker s.n. Sikkim (BM), Murata 06306529 (BM), Stainton, Sykes & Williams 9276 (BM). PFOR 81: Kao 4883 (TAI), Kudo & Mori 2401 (TAI), Simada 5419B (TAI). PMEN 82: J. & M. S. Clemens 32575 (BM), 40267 (BM), Pételot 5110 (BM). PFRA 83: Clarke 44065 (BM), TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 77 Horsefield s.n. Java (BM), Kerr 9658 (BM). PBRA 84: Clarke 40303 (BM), Griffiths 222 (BM), Hooker & Thomson s.n. Khasia (BM). PMIC 85: Gossweiler 2545 (BM—isotype), 9668 (BM), Raynal 12161 (BM). PMUT 86: Gossweiler 12134 (BM), Welwitsch 5496 (BM), 5990 (BM—holotype). PNIG 87: Henry 9082 (K), 11174 (K—holotype), 12563 (K). PPHI 88: Ramos 33320 (BM), Ramos & Edano 45012 (BM), Whitehead s.n. 1896 (BM—holotype). PREF 89: Herb. Hooker 80 (K), Thwaites 154 (BM), Walker s.n. Ceylon (K—lectotype). PRET 90: Ahern’s collector Forestry Bureau no. 3395 (BM—holotype), Edano 48842 (BM). PMOL 91: Clarke 10673 (BM), Gardner s.n. in Herb.Miers (BM), Vine 216 (BM). PTRA 92: Beddome 109 (BM—holotype). PATR 93: Beddome s.n. Anamallays (BM), s.n. W. slopes of Nilgherries (BM), Herb. Wight 2127 (K—holotype). PSPE 94: Metz 1225 (BM), Schmidt s.n. Nilghiries (BM), Vine 215 (BM). PVEL 95: Cuming 1097 (BM—isotype), MaGregor 11339 (BM), Mendoza 40923 (BM). PWIL 96: Elmer 22225 (BM—isotype). PPUR 97: Herb. Dalzell s.n. (K—holotype), Stocks s.n. Concan (BM), Vasnoli s.n. 1881 (BM). PPAL 98: Gamble 17853 (BM). PVIL 99: Masters s.n. in Herb. Hooker (K), Roxburgh s.n. India (BM), Wallich s.n. 1831 (K). PWIG 100: Gamble 18389 (BM), Schmidt 74 (BM), Stocks & Law s.n. Malabar, Concan (BM). PROT 101: Lawson s.n. 1884 (BM), Wallich 1535 (K- W—holotype). PSTR 102: Clarke 5501 (BM), 15617 (BM), Kingdom-Ward 18763 (BM). PHIS 103: Herb. Hooker s.n. (K), Jenkins 346 in Herb. Hooker (K—paratype), Kerr 6631 (BM). PPUB 104: Kerr 2384 (BM), 3113 (BM), Pételot 5297 (BM). PBRE 105: Chen 3206 (KUN—isotype). PNEL 106: Cook’s 3rd voyage s.n. (BM). PBAT 107: Ridley s.n. 1921 (K—holotype). PWAT 108: Clarke 41719 (K—holotype). PHIR 109: Beddome s.n. (BM), Simpson 9049 (BM), Thwaites 283 (BM). PRUP 110: Cramer 4277 (K), MaCrae 396 (K—holotype), Thwaites 343 (BM). PMAC 111: Hansen & Smitinand 12661 (K), Iwatsuki, Fukuoka & Chintayungkun 9659 (K). PGAR 112: Gardner 1847 (K—holotype), Herb. Wight s.n. (K), V. Row 3229 (K). PCHA 113: Champion 339 (K—holotype), Shiu Ying Hu 12421 (K). PDIE 114: Forrest 875 (K—isotype). PGRI 115: Griffith 3962 (K—holotype). PNIL 116: Bourne 5094 (K), Herb. Hooker s.n. (K). PLIT 117: Evrard 1834 (K), Poilane 24241 (K), s.n. (K). RDEP 118: Cavalerie s.n. in Herb. Léveillé (E), Wilson 3534 (E—holotype), 4313 (BM). RSIN 119: Bodinier 2709 (E—holotype of Leucosceptrum bodinieri), Henry 7765 (E—holotype). CFOR 120: Huang 7040 (TAI), Huang & Hsieh 7297 (TAI), Kao 8611 (TAI). CSTE 121: Murata 19058 (TI), 19171 (TI), 36037 (TI). CBAR 122: Humsawa s.n. 1947 (TI), Kanai 6012 (TI), Yamozaki 6 (TI). CSUB 123: Bisset 102285 (BM), Hara s.n. 1953 (TI), Togasi & Matsuoka 180 (BM). CJAP 124: Maximovicz s.n. 1862 (BM), in Herb. Hance 13214 (BM), Tschonoskis.n. 1864 (BM). CNIN 125: Law 1012 (TAI), Liou 1378 (E), Handel-Mazzetti 2602 (E). LCAN 127: Bowes Lyon 37 (BM), Ludlow, Sherriff & Taylor 6475 (BM), Stainton 5140 (BM). LPLE 128: Maire s.n. (E—holotype). TGOU 129: Goudot s.n. (G), Hildebrandt 3471 (G—holotype). THIL 130: Hildebrandt 3971 (G—holotype). TFRU 131: Forsythe-Major s.n. 1895 (G), Lyle 278 (K—holotype). KJAP 132: Bot. gard. Tokyo s.n. 1880 (TI), Herb. Terasaki s.n. 1906 (K), Yano s.n. 1890 (TI). KELS 133: Jiangsu group 2667 (KUN). KGLA 134: Y. Ling 728 (KUN—isotype). KSIN 135: Chen Quan 999 (KUN). KSZE 136: T. P. Tsung 39354 (KUN—isotype). CTER 137: Roxburgh s.n. Mysore (BM), s.n. Mysore Bot. gard. Calc. (BM—isotype?), s.n. Hort. Calc. (BM). COPP 138: Clarke 3464B (BM), Hooker s.n. Sikkim (BM), Thomson s.n. Mont. Nilghiri & Kurg. (BM). J. R. PRESS 78 O2 BL O SL O€ O02 O G2 Of OF O 02 O€ OL O S2 S 02 0 Of OL 2b°0 2: OL Ob O SL G6 8 OG OL 9. O42 OL O2 O SL OL 2 002 6 9 00 OL OL O 02 OL OL O SL OL 9 202 GS 4 202 OL OL 2 SL Gob, “2-Sb € OL OSL OL OL 2 02 OL g O2L OL 9 2 02 OL 9 2 StL S OL 2 SL OL 8 2 SL OL 9 2 S&L OL 9L O @L O€ OF O G2 oS 2 co oOo © °o owe o-oo o 9° ° °o °o °o °o ° ° Oo +6 © o o oO o °o o o o o °o °o ° °o °o {o) °o N o sed LtLoood SALLO 6€I 40J B}ep papo| *7 xIpuaddy L OOL 002 OOL O 00S OOL O 00h 002 O 00€ OOL O 002 OOL O 062 0S 0 002 OOL L OSL OS O OSL G2 L OSL OS 1 OOL O0L O 002 0S 0 006 OOL O 008 OSL O 056 OOL O 004 0S O 062 OOL O O0¢ OOL O 009 OOL O O04 0S O O0£ OOL O O0¢ OOL O O04 OOL O 00¢ OOL O O0¢ OSL O 00S OOL O OOh OSL O O0€ OOL O OOH o Oo ° °o ° °o °o °o fo) fe) Oo L L L °o Oo oOo jo) °o °o ce) fe) 00S 002 00h O2L 002 OOL 00 002 OSL OOL 0S2 OSL 002 OOL OOh 0S 00S OOL 0S2 OOL +» OOL 002 OSL 00S OSL +» OGL OOL OOL 002 OOL OO€ OOL + 002 OSL OOL 002 OSL 002 OSL +» OOL OS2 OOL +» OOL OOh OSL OOh OOL OOE OO¢ OLtLL O OSE 00E OL OL SL OL OL OL ° °o °o o o ° OOL G2 0S OOL OSL OSL 002 002 SZ L L 1o] x\ - °o o °o o °O OF O72. OF aa °o °o °o °o o °o o o ao 9° 0 02 O SL O OL 0 02 O SL Qeviaa 2evisd geonegt SeNada STACKS ICA S2LNId eendva LONVWa OzNaCda 6L uaa QUNNHG Laisa OLLTHE SLaoga 4LLNTa CLDava 2Lnosa LLDAdH OLOVNa 6T10a QNOOT 2a oa Od INE 79 TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE WN WW MN WN OL OL OL 02 02 SL SL Ge h2 Ww ie) 0 fe) 0) O ce) 10} 09 02 St 02 $f 02 On c oS: 6 °o ° o oOo °o o os oO o ° oo. 6. 6. -o .6 eo (Ox ° So oo oo OSL OOL OOL 0S OOL OOL OOL 0 O 002 OOL Oo :O CO: tO c o Oy Om, Oo 002 OSL OO OSL OL OL 0S OS O 0S OS O 0S 0S 0S OS O 0S 0S 002 OS OOLL GZ OOL L OSL L OS OOLL 02 02 °o o oO ° O77 OF aS °o °o ° o-" :O °o On cS jo) o ° fo} °o °o Co Na. N — Nn ma mM A OM nN DAN AN DN AM ° fo} o ° ° mM Dre © Ft a a ao = 9Svnea Ssnuod +SNI'TC €GaLsa 2Svaod LSLVAC oSTuDa 6hDada SHWvSa 24OLSa OyNada SHWOLd 4+/THHO ¢yI ala 2Hd VOR Ly Tada OnHOOT 6£0AWT gcv ida L¢EOISa QENI MT Sov Taa EVID oT ao 2¢OTWT LEWOOe o¢andat 6endag J. R. PRESS 80 02 02 82 02 02 02 eL eL 02 ce) L L L L fe) fe) O GL o Oo oo: So ° oS oS © Oo oO o oOo oo Oo °o ° °o ° ° ° ° °o ° — 002 0S2 OSL OSL OSL fe} O Oo Oo" °o © 0 002 002 O O0€ 002 O OSL OOL 0 062 0S2 O 002 002 O 00% 00h O OO£€ ODE 0 O0€ 0S2 O OOF O0¢ O.« 062 O O00€ 00¢ O O0£ O00¢ O 00€ 0S2 O OOL 0S O OOL OOL O S2t 0S © G2L OOL O OOL OOL O OSL OOL O 002 OOL © OOL 0S Os OOL O OOL OOL O OOL OOL O OOL SZ O« OOL O« OOL O.« OOL NARS ti Ne NT EN UN CU OLIN Wy oN OS 4 OL OL OL LOS OOLL L OOL O02 L L OOL OSL L O OOL OOL O O OOL OOL O © OOL OOL O O 002 002 O O OSL OSL O O OSL OSL O O OOL OOL O 0 OGL OSL O O OOL COOL O O OOL OOL O 00€ OF O 00€ OF O 00S OS O 00S OS O 00S OS O 00S OS O © OOL O02 L 00S OS O 006 0S O 00S OS O 00S OS O 00S OS O 00S OS O 00S OS O O OOL OOL O OL GL GL 02 °o °o o °o ° o so NY oN \a ak kay L OL fe} fe) fe) “gvadd €Qvadd ZQNAW Lgdoad ogynvd 6éSTad gcavod L2KaHd gdvTdd Gdanid 4dNVdd ¢davdd ZdNadd LZOAW ozonad 69TvSa gounva 29VTOd 99aNva S9aVWG *9WNdd ¢OLKTC Z9veOd L9qONa ognvaa 6SazSa QSTvia dSISid "JUO0D Z xipuaddy 81 TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE — - — — OOL 002 OSL OSL OSL OSL OSL 0 0S2 0 0S2 O 00¢ 0 0S2 0 006 O OOF So oo 00g 00+ O0¢ 002 OSe S OO£ S O0S2 OL OS2 S 0) OSL OOL GeL OOL O2L OSL OSL L OL 0 09 0 02 O S¢ O 02 0 0S O O02 0 02 00 O 2LLYWdd 0 0 O LLLOWNd O OL O OLLdndd 0 OL O 60L4THd 0 02 0 gSOLLVMd 0020 d0OLivad O OF O 9OLTHNd 0 020 SOLauad 0020 40Landd O Oh O OLSIHd 0S O 20Lu4LSd 0 € OO LOLLOUd 0 02 O OOLDIMd 0S O 661IAd 0020 = g6TY¥dd 0020 d64ndd 0020 96TIMd 00h O G6THAd 0090 *#6adSd OOLO 6aLvd 00S 0 26VvuId OOLO Lé6TOWd 0020 O6Laad 0020 6 9amud 0020 = ggIHdd O+ O ZQDINd 02 0 99LNWd OOLO SgdIWd J. R. PRESS 82 O02 02 Of O OF 2 42 002 2 ce. Git 2. -2'0n Cre ORE 22GeL ce ht g O0¢2 8 OO 2 9 00 2 42 0 O€ 2 QL O Sh 2 8 O02 91. 0021 g@ OCHL g OSH tL g@ OO 1L g@ OO¢2L 4t O G2 2 cL O OW 2 02 0 OF L 9L OGL UL 02 O OF 2 Helbeel .c AU SG aL O2-l OL-L oOo. oc Oo © °o fo} — -_ GZ G2 O 0S OF O 002 002 O 002 OOL O 002 002 O 002 002 O 0S2 OSL O OSL S2L O OSL OOL O 00¢ 002 O OSL OOL O 00g 0S O 002 OOL O 0G2 OOL O 002 OOL O 002 OOL O 002 OOL O CoOL GZ O 004 002 O 0G2 002 O 002 OOL O OOL COOL L 002 002 O OSL 002 O OOL 002 O 002 002 O 002 L 0S OOL OL OOH eL fe) O O 0 0 O O oo¢ OSL OSL OOL OOL fe) O O oO O L O L O 40 oO « O 02 02 OL OL OL 02 oO O gE Lddoo LE LUaLO 9 LAZSy SELNISH HELVION €CLSTaM 2ELdV LM LELnedL O€ LTIHL 62LNODL Q2LATdT L2LNVOT 92LNVNO G2LNINO heLdvlo €2Lanso 22Luvao L2LaLSO O2LYO49 6LLNISH GLidada 2LLLITd OLLUTINd GLLIaDd HLLaIdd SLLWHOd ‘Ju09 7 xIpueddy Appendix 3. List showing the five nearest neighbours for each OTU with the similarities expressed as percentages OTU 1 2 3 4 5 oTu 4% oTU % otuU 4% OTU ¥% oTU 4 1 65 81.5 34 81.1 3279.8 38 79.6 102 79.5 2 102 «87.1 83 85.6 109 85.6 88 85.3 33 84..9 3 i 9755 12 94.0 7 91.8 5 91.6 6 91.3 3 97-3 12 92.0 7 91.4 5 -9ilse 6 90.9 P 6 97.4 7 97-3 12 96.9 11 96.3 13 96.0 6 ? 98.5 5 97.4 12: “97 s2 IS, 96.5 11 95.4 ? 6 98.5 & 9725 13 97.0 12 96.7 Or 29556 8 65 88.5 35 88.3 38 §=685.9 Sie bb y5) Oroos5 9 7 956 DF 90eD 6 94.8 13, 94.3 Ue bs 10 16 85.7 12 84.8 15 83.9 6 83.8 7? 82.6 11 5 96.3 6 95-4 (fr) Vee 39567) 15. (9537 12 6 97.2 5 96.9 6 96.7 13, 94.7 3 94.0 13 7 97.20 6 96.5 5 96.0 12 94.7 9 94.3 14 6 90.7 5 90.7 7 90.6 12 90.0 17 89.6 15 16 96.7 47 88.1 7 84.2 5 84.0 10 83.9 16 15 96.7 127 90.1 6 86.5 7 86.0 10. 85.7 alg 5 92.4 6 92.1 1291.8 2.9128 13 91.0 18 12? 87.3 B Ohse 7 86.5 6 86.2 3 84.9 19 20 90.7 34 83.9 20 83.4 39 83.3 65 82.6 20 19 90.7 21 88.4 34 85.6 ie R05 62 3 84.0 21 20 88.4 6 86.3 5 86.2 12 85.4 33 85.2 22 52, 90:7 51 88.6 60 88.6 63 88.4 64 88.4 23 33 84.1 38 = 82.3 30-—«2..1 31 81,5 2508 8055 24 34 86.7 26 «86.1 35. 85.8 32 «85.2 3 84.7 25 105 85.7 35 84.7 38 (B44 92 83.8 80 83.6 26 35 (91.3 42 92.0 30 «= 89.8 34 88.4 36 = 87.9 27 31 89.9 32 88.2 33 88.1 28 87.5 30 «86.4 28 30 =87..9 lf EE SS 26 «686.4 14° 85.0 31. 85.0 29 40 = 87.8 42 86.4 36 «= 85.8 38 85.6 30 684.6 30 32-922 36 =: 922 39 92.0 33-9142 31 91.6 p! Se Obese 33-9303 34 92.6 30 91.6 4191.2 32 a1 95.2 33 94.5 34 92.8 30 92.2 36 9067 33 32 94.5 Se Bee hSs: 30 «91.8 39-905 36 «= 89.8 34 32 (92.8 31 92.6 35 92.4 30 91.6 38 90.1 35 36 ©=©95.0 38 = 94 8 34 92.4 26 «91.3 65 90.5 84 Appendix 3 cont. OTU 4 J. R. PRESS 3 oTU = % 30-9242 36 =—90.5 4O 93.4 34 88.9 35 90-3 33 = 88.6 128. 8757 85 90.4 Slee 9illed 64 92.9 4g 92.4 57 89.9 4g - 92.9 63 9502 60 92.8 63 96.7 63 95-4 55° 92.6 52 90.8 66 93.9 59 92.0 63 90.7 55 94.1 62 93.2 4g 94,1 4G O5a7 D9 95ee 60 95.7 5296.8 37 89.8 78 90.5 70 83.7 71 =89.2 58 92.5 69 92.5 90.7 89.8 92.8 88.5 90.1 88.4 87.5 90.2 90.5 91.8 92.3 89.6 92.8 9520 91.7 96.4 95-0 92.5 90-5 93.2 91.1 90.6 93.0 92.7 93.8 93.0 93.0 . 95.4 95-4 89.2 90.3 83.5 89.0 92.4 91.5 OTU 101 102 103 104 105 TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE 105 103 114 116 dite TS 110 105 103 2 OTU 58 112 112 72 79 104 113 112 110 102 101 107 3 OTU 66 113 113 (e 115 108 108 113 115 114 114 70 81 89 102 115 101 89 96 96 87 90 96 100 88 88 73 103 72 98 87 69 tHE 77 108 4 OTU 56 rid, 76 ie, is 115 75 108 He 113 78 109 88 86 82 70 86 95 83 83 85 AOS 92 83 89 101 72 108 72 105 85 113 108 113 68 85 86 Appendix 3 cont. OTU 1 otU % 106 107 90.9 107 108 97.6 108 107 9746 109 Ate. 9654 110 102 93.8 111 102. 93.1 112 at 209560 113 108 95.5 114 75 93.4 445 99 9561 116 86 992.4 117 83 86.3 118 119 94.8 119 118 94.8 120 125 9305 121 124 9747 122 123 96.1 123 124 99.2 124 Mos 2966 125 122 93.7 126 4123 »85.1 127 124 85.7 128 42 87.7 129 130 «94.1 130 13 9757 131 130 9747 132 135 94.0 133 136 96.0 134 551.9565 135 132 94.0 136 133 96.0 137 138 = 9502 138 137 OTU 112 113 113 110 108 100 108 115 “ 113 91 82 126 125 122 123 124 121 121 120 120 123 29 131 129 129 133 135 136 139 135 105 105 J. R. PRESS 3 oTtU 4% 113, 88.9 112 93.3 112 9542 69 91.6 81 91.2 2s ene 113 94.9 112 94.9 ie SESS 739 9eee 70 90.6 109 86.0 125 83.4 120 82.7 124 89.4 122 95.7 121 9567 122 96.1 122 95.9 725, “95-4 118 83.6 122 85.0 24 84.1 101 79.6 24 7929 13. 81.0 136 88.7 134 93.3 135-8767 136 9247 134 89.5 80 79.4 71 hte? oTU Gl Lh 77 174 69 108 74 77 te, te 102 102 124 126 123 125 125 125 125 124 119 121 Appendix 4. Distribution of OTUs in the six and nine group schemes for clustering to maximize WGMS analysis Six group scheme Nine group scheme Group 1. Group 1. 66° 70: 712.82: 845 °85: 118.:719::1202121-:122 :1232 86: 87: 91: 98:101 :102: 124)-125'3126:3127. 103 :105 :109 :110 :111 :116: #51 158. Group 2. Group 2. OSs 722 134. 14 “15.7716: 192 13321342135 2136. Ts 161d Oe Ort 9k 99 :103 :106°:107 :108 :112: TS.21 14:1 15: Group 3. Group 3. yd Ria he, ee SS Em « 12613021312 134 :<135 7136; 48 AO O02 OL 3240.53 Group 4. D4 3D 200s les 8 too. 2182419 120 121 51225123: 60: 61: 62: 63: 64: 65: 124 :125 :126 :127 69 Group 5. Group 5 $3:2-88 7 892. 90>-92 =°93: TOS TAs. 82:5- 83: 84 BO): Group 6. Group 6 $2 45 44 AS: 46: 47: Te 2 8 oe 19 eco es ga toe | ee | Bei Pee oe 22 20.) 21a een Le OU =”: SE es Berka pee Acai. eae bf Ss 25203 -95 2 O4 So O08 GU GP? 02.- 63;: 642 G7: 37 36 200 440241 a2 69 65 :128 Group 7 io caste oS l One Oe ea oes 1G so ea A ays 1G et 16S ses 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*), Genéve (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 Léveillé’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 trgnslations 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. Jn G. Bentham & J. D. Hooker, Genera Plantarum 2 (2): 1160-1223. London. Blume, C. L. 1826. Bijdragen tot de Flora van Nederlandsch Indié 3. Batavia. Briquet, J. 1897. Labiatae. In A. Engler & K. Prantl, Die Natiirlichen Pflanzenfamilien 4 (3a): 183-374. Leipzig. 1908. Lieutenant Olufsen’s second Pamir-expedition. Plants collected in Asia-media and Persia by Ove Paulsen. VII. Labiatae. Bot. Tidsskr. 28 (1): 233-248. Chermsirivathana, C. 1963. Labiatae of Thailand. M.Sc. thesis. Aberdeen. Chopra, R. N., Nayar, S. L. & Chopra, I. C. 1956. Glossary of Indian medicinal plants. New Delhi. Cook, C. D. K. 1978. The Hippuris syndrome. Jn H. E. Street (Ed.), Essays in plant taxonomy: 161-176. London, New York, San Francisco. Desfontaines, R. L. 1815. Description d’un nouveau genre de Labiée. Mem. Mus. Hist. nat. Paris 2: 154-156. Diels, L. 1924. Miscellanea sinensea 1. Notizbl. bot. Gard. Mus. Berl. 9: 197-199. Don, D. 1825. Prodromus florae nepalensis. London. Dunn, S. T. 1915. A key to the Labiatae of China. Notes R. bot. Gdn Edinb. 6: 12-190. El-Gazzar, A. & Watson, L. 1967. Consequences of an escape from floral minutae and floristics in certain Labiatae. Taxon 16: 186-189. —— —— 1968. Labiatae: taxonomy and susceptability to Puccinia menthae Pers. New Phytol. 67: 739-743. 1970. A taxonomic study of the Labiatae and related genera. New Phytol. 69: 451-486. Endlicher, S. 1838. Genera plantarum. Vindobonae. Erdtman, G. 1945. Pollen morphology and plant taxonomy. IV. Labiatae, Verbenaceae, and Avicenniaceae. Svensk. bot. Tidskr. 39: 279-285. —— 1952. Pollen morphology and plant taxonomy. Angiosperms. Waltham, Mass. Fedtschenko, B. A. 1908. [Book review] J. Briquet. Lieutenant Olufsen’s second Pamir-expedition. Plants collected in Asia-media and Persia by Ove Paulsen. VII. Russk. bot. Zh. 1908 (1-2): 65-66. Gower, J. C. 1967. Multivariate analysis and multidimensional geometry. Statistician 17: 1-16. —— 1971. A general coefficient of similarity and some of its properties. Biometrics 27: 857-871. Hasskarl, J. K. 1842. Plantarum genera et species novae aut reformatae javanenses. Flora, Jena 25, Beib. 2 (1-2): 1-32. Hermann, P. 1717. Musaeum Zeylanicum 3. Lugduni Batavorum. [Not seen. ] Hooker, J. D. 1885. Flora of British India 4. London. —— 1896. Comanthosphace japonica. Curtis’s bot. Mag. 122: t.7463. Keng, H. 1969. Flora Malesianae precursores 48. A revision of Malesian Labiatae. Gdns Bull., Singapore 24: 13-180. —— 1978. Labiatae. In C. G. G. J. van Steenis (Ed.), Flora Malesiana ser. 1, 8 (3): 301-394. Keys, J. D. 1976. Chinese herbs. Rutland, Vermont. TAXONOMIC STUDIES IN THE LABIATAE TRIBE POGOSTEMONEAE Kitagawa, M. 1935. Contributio ad cognitionem florae Manshuricae. Plantae novae Jeholensis II. Rep. scient. Exped. Manchoukuo 1933, section 4 (2): 14-41. —— 1939. Lineamenta florae Manshuricae. Rep. Inst. scient. Manchoukuo 3 (App. 1): 1-477. Kitomura & Murata, G. 1962. The union of Leucosceptrum and Comanthosphace. Acta phytotax. geobot., Kyoto 20: 165-171. Kudo, Y. 1929. Labiatarum sino-japonicarum prodromus. Mem. Fac. Sci. Agric. Taihoku imp. Univ. 2 (2): 37-332. Kuntze, O. 1891. Revisio generum plantarum 2. Leipzig. Lamarck, J. B. A. P. M. 1789. Encyclopédie méthodique 3. Paris, Liege. Léveillé, H. 1912. Nouvelles Labiées chinoises. Bull. Géogr. bot. 22: 236. — 1915-16. Catalogue des plantes du Yunnan. Le Mans. Linnaeus, C. 1747. Flora Zeylanica. Holmiae. —— 1753. Species plantarum 2. Holmiae. —— 1764. Genera plantarum 6th ed. Holmiae. — 1767. Mantissa plantarum 1. Holmiae. Loureiro, J. 1790. Flora Cochinensis 2. Ulyssipone. Marquand, C. V. B. 1930. Revision of the old world species of Buddleja. Kew Bull. 1950: 177-208. Masamune, G. 1940. Miscellaneous notes on the flora of the eastern Asia 18. Trans. nat. Hist. Soc. Formosa 30: 337-343. Merrill, E. D. 1937. Miscellanea sinensia. Sunyatsenia 3: 246-262. Miquel, F. A. W. 1859. Flora Indiae Batavae 2. Amsterdam. —— 1865. Prolusio florae Iaponicae. Annis Mus. bot. Lug.-Bat. 2: 69-212. Moore, S. le M. 1877. Alabastra diversa. J. Bot., Lond. 15: 289-298. Mukerjee, S. K. 1940. A revision of the Labiatae of the Indian empire. Rec. bot. 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. nJ.B. A. P.M. Lamarck, Encyclopédie méthodique Botanique suppl. 5: 663. Paris. Prain, D. 1898. On three new genera of plants from the Kachin hills. Scient. Mem. med. Offrs Army India 11: 41-44. — 1901. Eurysolen gracilis. In G. King, J. F. Duthie & D. Prain, A second century of new and rare Indian plants. Ann. R. bot. 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. Plants of the coast of Coromandel 3. London. —— 1832. Flora Indica 3. Serampore. Rumphius, G. E. 1750. Herbarium Amboinense 6. Amsterdam. Smith, J. E. 1805. Exotic botany 2. London. Sneath, P. H. A. & Sokal, R. R. 1973. Numerical taxonomy. San Francisco, London. Taylor, G. 1931. Pogostemon mutamba. In A. W. Exell, R. D’O. Good & G. Taylor, Mr John Gossweiler’s plants from Angola and Portuguese Congo. Dicotyledones, Gamopetalae. J. Bot., Lond. 69 (suppl. 1): 166. Vautier, S. 1959. Resultats des expéditions scientifiques Genévoises au Népal en 1952 et 1954 (partie botanique). 14 Labiatae. Candollea 17: 41-52. Willdenow, C. L. 1790. Novum vegetabile genus Elsholtzia nominatum. Bot. Mag. 4 (11): 1-6. Wu, C. Y. 1959. Revisio Labiatarum sinensium. Acta phytotax. sin. 8: 1-66. 1965. Rostrinucula sinense. InC. Y. Wu, H. W. Li, S. J. Hsuan & Y. C. Huang, Materiae ad floram Labiatarum sinensium (2). Acta phytotax. sin. 10: 215-242. —— & Huang, Y. C. 1974. Materiae ad floram Labiatarum sinensium 4. Acta phytotax. 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. Acta phytotax. 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). J. Indian bot. Soc. 42A: 321-330. Yunnan Xian Weishenju (Ed.) 1973. Chinese herbal medicines of Yunnan: supplement. Kumming. oe ageslty 5 re Oe ae oe oa a Pe ee ee —— FO Ne ee woe wee 2S eS SS 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). 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 The typification of Hudson’s algae: a taxonomic and nomenclatural reapprais Linda M. Irvine & Peter S. Dixon Botany series Vol10No2 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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(Bot.) © Trustees of the British Museum (Natural History), 1982 The Botany Series is edited in the Museum’s Department ga age er Keeper of Botany: Mr J. F. M. Cannon ME (en [4 ~ Editor of Bulletin: Mr P. W. James Vi a oe ULICA TG Alg es, HEN Assistant Editor: Mr J. R. Laundon ft i ae 7 O ACT 489 ij blow gy ZOww'l i: L ie i if 4 —o Gamuinde ) , { 4 \ p / / YEN y hy, * ll vp Zz As fee ; : “i ts es ae ; - a ISSN 0068-2292 “Botany-seriés- Vol 10 No 2 pp 91-105 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 SW75BD Peter S. Dixon Department of Ecology and Evolutionary Biology, University of California, Irvine, California 92717, U.S.A. Contents Synopsis . : : ; : : : ; ; : : ; ; ; : 91 1. Introduction : ; : ’ : : : : : ‘ ; : OZ 2. Hudson’s species concept : ; 2 : ‘ : : ; : : 92 The algae of Flora anglica: first edition : : ; ; : ; 92 The algae of Flora anglica: second edition , : ; : : ; ; 5 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 ; 2 ; : ; : ; 98 III. Lectotype (provisional): a Hudson specimen : : ; i : ; : 99 a. Conferva elongata . : 2 : : : : : ; , ; 99 b. Fucus plicatus . 5 : ‘ : ; : : 100 IV. Lectotype: a Hudson description ‘ ; : , ; , : ; 102 a. Conferva fucoides : : ‘ : : : : : 102 b. Conferva nigra . ‘ ; : ‘ : ; : : : : 102 V. Species inquirenda : : : ; : ; ; ; : , : 103 a. Conferva fulva . 5 . : ; ° ; : ¢ : ; 103 5. Acknowledgements. : : : : , : ; ; ; : 104 6. References . : : : : ; 5 | ; ; : ; : 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. nat. 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, 1977a), 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, 1959b, 1960, 1962, 1963, 1967; Dixon & Irvine, 1977a, 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 resumé of his life and of the history of his personal herbarium has been previously presented (Dixon, 1959a). 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 ALGAE: 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 (1959a) 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 Buddle, Petiver and Plukenet. These are all now in the Sloane Herbarium (BM-SL), of which Dandy (1958) has given a detailed account, anid were probably consulted by Hudson when he was employed as a resident sublibrarian at the British Museum between 1757 and 1758. Buddle 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, Ulva 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 Stearn (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 et al., 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 Ia). 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 Ila), Fucus pinnatifidus (Example IIb), and F. plumosus (Example IIc). 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 IIIa) and Fucus plicatus (Example IIIb), 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 a/., 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 Caloplaca flavorubescens (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: pl. 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.) Kitz. ] The original description of Conferva imbricata first appears in the second edition of Flora anglica (Hudson, 1778:603), printed as followed: 44. CONFERVA filamentis geniculatis ramosissimis, imbricata. ramis acutis, ramulis verticillatis imbricatis dicho- tomis. Muscus marinus hirsutus, flagellis longioribus, rarius divisis, ruber. Hist. Ox. IIT. 650. s. 15. t. 9. f. 7. Anglis, imbricated Conferva. Habitat in rupibus saxis et fucis 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: pl. 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 frondibus planis ramosis, ramis dentato- pinnatifi- pinnatifidis marginibus callosis. dus. Fucus dealensis pedicularis rubre folio. Mus. pet. 405. R. Syn. 48. Alga cervi cornu divisura. B. hist. II. 797. Anglis, jagged Fucus. Habitat in saxis et rupibus marinis. Found about Deal by Mr. Dandridge. R. Syn. in littore Devoniz fre- quens. This description was clearly based upon four elements: the previous treatment by Petiver (1695—1703:39); the previous treatment by Dillenius in Ray (1724:48); the previous treatment by Bauhin (1651:797); 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 J.B. IIT. 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 rubre 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 Buddle’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. ic sae teak 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. ExamMPLEc. 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. duplicato pinnatis coloratis. Fucoides purpureum eleganter plumosum. R. Syn. SOc te 2: fed: Anglis, Feathered Fucus. Habitat in littoribus marinis passim. This description was based on two elements: 1. an illustration given by Dillenius in Ray (1724: pl. 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 Buddle 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) aspecimen from the Lambert herbarium, now in BM-K; (b) aspecimen from the Forster herbarium, at BM; (c) aspecimen 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 Buddle 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 Buddle 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. FUCUS fronde cartilaginea compressa ramosa, ra- plumosus. mis duplicato-pinnatis, fructificationibus peduncu- latis globosis radiatis. Fl. angl. 473. Gmel. fuc. 152. Fucus frondibus cartilagineis lanceolatis bipinnatis plumosis, caule filiformi compresso ramoso. Mant. 134. Syst. nat. 718. Fl. dan. 350. Fucoides purpureum eleganter plumosum. R. syn. so ae ere Anglis, plumous Fucus. Habitat in rupibus et saxis 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: pl. 152 and Oeder, 1767: pl. 350) are of the alga to which the binomial Prtilota plumosais 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. musc. 35. Gf. 38: Anglis, Pointed Conferva. Habitat in littoribus marinis frequens. 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: pl. 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 Buddle Hort. Sicc. 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 Buddle 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 Buddle 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 et pedalia, crassitie fili emporetici tenuioris, articulata, levia, fusco-purpurea, basi ra- mosa; Rami dichotomi, longissimi, setaceti; 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. 11 1778 p. 599. . . A. R. A. Taylor 1.1X.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. ExaMPLeb. 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. f. 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: pl. 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. Alm. 160. T. 184. f.2. Alga exi- gua dichotomos arenacei coloris Syn. //. 4. 10. Fucus ceranoides ramosus tenuissime divisus Dood. Syn. II. App. 329. Palmaris est, corneus tenax, albus, per siccitatem rigidus, ubique ejusdem fere crassitudinis, que filum parvum superat. In littore Essexiano, Sussexiano & alibi. Both Dillenius in Ray (1724) and Hudson (1762) refer to an illustration by Plukenet (1696: pl. 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 IIc) and Fucus pinnatifidus (Example IIb), 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, 1977a). 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. CONFERVA filamentis geniculatis ramosissimis ra- 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 filamentis geniculatis ramosissimis fucoides. ramulis multifidis, inferioribus fasciculatis fructi- feris. Anglis, fucus Conferva. Habitat in rupibus, saxis et fucis marinis passim. [perennial]. I-XII. Desc. Fila pedalia, geniculata, levia atro-rubescentia, ramosissima, Rami alterni; ramuli multifidi, subdicho- tomi, inferiores fasciculati, fructiferi. Fructifica- tiones terminales, radiate, parve. 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: pl. 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 filamentis equalibus ramosis, 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 equalibus ramosis lon- nigra. gissimis, ramis alternis multifidis brevissimus. FI. angl. 481. Anglis, black Conferva. Habitat in littore Eboracensi passim. [annual]. V—X. Desc. Fila qguinque pollicaria, equalia, 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: pl. 70) had described C. atro-rubescens but stated later (Dillwyn 1802-09 fasc. 15: pl. 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 rubescens 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 et fucis 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: pl. 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. J. Bot., Lond. 40 (suppl. 2): 1-107. Bauhin, J. 1651. Historia plantarum universalis. 3: 1-866+12. Ebroduni. Clokie, H. N. 1964. An account of the herbaria of the Department of Botany in the University of Oxford. viiit+280 pp. 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. xvit+576 pp. Oxonii. Dillwyn, L. W. 1802-09. British Confervae. 16 fasc., Introduction, Synopsis. 109+A-G pls, 94 pp. London. Dixon, P. S. 1959a. Notes on two important algal herbaria. Br. phycol. Bull. 1 (7): 35-42. 1959b. 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 of Fucus cartilagineus L. and F. corneus Huds. Blumea 15: 55-62. — & Irvine, L. M. 1977a. Seaweeds of the British Isles. 1. Rhodophyta (1). Introduction, Nemaliales, Gigartinales. xi+252 pp. London. — & —— 1977b. 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: 404-427. 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. xxxviiit+690 pp. London. — 1798. Flora anglica. 3rd ed. [4]+iv+xxxii+688 pp. London. Juel, H. O. 1936. Joachim Burser’s Hortus Siccus mit erklarungen heraus gegeben. Symb. bot. 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 8: 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’]. cones 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. J. 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. [xiti]+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. Jn 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. xlvit+400 pp. London. — 1808-19. Fuci, sive plantarum Fucorum generi. . . 4 vol. London. Vines, S. H & Druce, G. C. 1914. An aceount of the Morisonian herbarium: \xviii+350 pp. Oxford. oe, ie eee) © rat) Pte Ot UT ee Byis a ; ie te are : ef 2 He hate ane \ lat British Museum (Natural History) Seaweeds of the British Isles Volume I 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 0 565 00781 5 1977 Approx. £12. Titles to be published in Volume 10 Taxonomic studies in the Labiatae tribe Pogostemoneae. By J. 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 Vol10No3 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 SW7 5BD, 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 3 pp 107-225 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 25 November 1982 Seaweeds of the Faroes Contents 1: The flora. ByD.E.G.Irvine_. 2: Sheltered fjords and sounds. By I. Tittley, W. F. Farnham and P. Wats: Gray 3: Openshores. By J. H. Price and W. F. Farnham . ; ; ; ; 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 . : . ; ; : A , ? : , . 109 2. Methodology . : , : ; ‘ : ‘ ; : ; 110 3. Species list : 5 i F : : é : , ; aa | Cyanophyta ‘ : ‘ : : ‘ : : : : Pi by Rhodophyta. . : ‘ ; : : : ; : i ly be Chrysophyta. : : : ‘ F : ; ; : . 120 Phaeophyta : , ; : ‘ : : , ; - . 120 Chlorophyta. : ‘ ; . ‘ j g : : . 126 4. Conclusions . : : : : : : : : A : 129 5. Acknowledgements . : , : : ; : : F : . 130 G;. References: 4 : ‘ : ; ; : : as : : rot) 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 Bgrgesen 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 (Fergerne; Faeroes; Fgroyar) 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, Térshavn, 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 KALSOY VIDOY STREYMOY Cp ye BORDOY cs 7 EYSTUROY VAGAR Térshavn = SANDOY h Q | Q fay 0 10 20 25Km SUDUROY 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. Bérgesen and various colleagues and published in a series of papers from 1895 to 1905. Bérgesen 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. Jonsson 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 Bérgesen’s account of the marine algae (1902). Since the papers of Bgrgesen, 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 Bgrgesen, had not found it, this record has been discarded by B¢rgesen. 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 Pit 0 250 500Km Ce ane FN Fig.2 Map showing position of the Faroes in relation to main ocean currents and neighbouring land masses (after Ryder in Bgrgesen 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 Vidareidi in the northern island of Vidoy, followed by a week in Térshavn. Most of the major islands, apart from Suduroy, were examined both intertidally and subtidally (using inflatables and scuba equipment), and particular attention was given to Skalafjgréur, 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. Recorded from the Orkneys. Recorded from the Shetland Isles (Irvine, 1980). Recorded from Iceland (Caram & Jénsson, 1972). ] Name used by Bérgesen (1902). Note that Bgrgesen’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, (');0: S_E E. deusta (Menegh.) Drouet & Daily [Hyella caespitosa, H. endophytica| In shells of molluscs, subtidally to 40 m. Found on Streymoy by B@grgesen. O. S. I. Microcoleus lyngbyaceus (Kiitz.) 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 Bgrgesen on Kalsoy: Husar, and on Streymoy: Tinganes. I. Oscillatoria lutea Agardh ex Gomont [Lyngbya lutea] Intertidally on somewhat exposed shores. Recorded by B¢grgesen from Vidoy: Vidvik, and Streymoy: Torshavn. O. Schizothrix calcicola (Agardh) Gomont Listed by Holt from Torshavn (as Phormidium fragile). O. S. 1. S. tenerrima (Gomont) Drouet [Microcoleus tenerrimus| Found once by H. Jgnsson at Hvannasund. Spirulina subsalsa Oersted ex Gomont In rock pools on moderately exposed shores. Recorded by Bérgesen from near Térshavn 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. (YOuseh Antithamnion boreale (Gobi) Kjellman [A. plumula var. boreale] Subtidally to 50 m, mainly on sheltered shores. O. I. A. floccosum (Miill.) Kleen Intertidal and shallow subtidal, mainly on sheltered shores. Occasional, widely distributed. (9'O7S.1, THE FLORA I ee A. plumula (Ellis) Thuret Subtidal on rocks, shells, and epiphytic, exposed and sheltered shores. Widely but sparingly distributed. (!) O. S. (1?). Audouinella alariae (JOnsson) Woelk. [Chantransia Alariae] Shallow subtidal on Alaria fronds, especially on exposed shores. Widely distributed and generally abundant, although Bérgesen 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 B¢rgesen as epiphytic on Ceramium shuttleworthianum on an exposed coast, near high-water mark. Streymoy: Velbastadur. 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 Bérgesen, 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 Bgrgesen, 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 Borgesen. 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) Kitz. [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. ) Kitz. 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, pl. 37) from Torshavn and Eidi, but these records dismissed by Bérgesen 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 Bgrgesen as referring to a form of C. rubrum]. C. shuttleworthianum (Kitz.) 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 Bgrgesen 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 Bérgesen (1905: 773). (!) O. S. Clathromorphum circumscriptum (Stromf.) Foslie Found once by Bgrgesen on Streymoy: in Sundini between Hosvik and Hvalvik. Later placed by Foslie in Phymatolithon compactum (Bergesen 1905: 773) but probably belongs to Clathromorphum. 1. 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. ():0-S- 1 THE FLORA 15 Corallina officinalis 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 Bgérgesen 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. Suduroy: Lobra. I. D. hapalidioides (P. Crouan & H. Crouan) Foslie Noted by Ostenfeld on a Patella shell in Suduroy: 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. (0. S25 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. Bordoy: Haraldssund (!) and near Klaksvik. Vidoy: Hvannasund (!). Streymoy: Vestmanna. It has also been recorded by Jgnsson 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 Bérgesen (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: Skalafjgréur, 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 of Laminaria 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 sanguinea 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 1d] Furcellaria lumbricalis (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.1. Goniotrichum alsidii (Zanard.) M. Howe Recorded by Holt from Térshavn 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 Bgrgesen as a single small specimen (Subduroy: Trongisvagsfjgrour) and during the present expedition by P. A. Asenasa single small tuft. (!) O. S. Harveyella mirabilis (Reinsch) Reinke Parasitic on Rhodomela confervoides. Apparently rare. Kunoy (Jonsson). Suduroy: Trongis- vagur (Bérgesen). 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). Suduroy: Tvgroyri and @ravik (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 (Strémf.) Adey [Lithothamnion laeve] In deep water, on sheltered and exposed coasts. Found by Bgrgesen on Bordoy: Haraldssund; Streymoy: Argir; Suduroy: Trongisvagsfjgrdur and Lopra. (!) S. I. Lithophyllum incrustans P. Philippi Intertidal, on Phymatolithon polymorphum. Recorded by Jonsson on Vidoy: Vidareidi, and by Bgrgesen on Suduroy: Hvalba. O. S. Lithothamnion glaciale Kjellman 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 /ingulata, 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 Bordoy: near Klaksvik, but Bgrgesen 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. 1. *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. 1. 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 Bgrgesen 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 Bérgesen 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. (!) O25). 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. Polyides rotundus (Huds.) Grev. On stones subtidally to 20 m, on sheltered coasts. Locally abundant, widely distributed. (!) OS. F. 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 B¢grgesen to be creeping filaments of P. urceolata]. P. nigra (Huds.) Batters [P. atrorubescens]| A few specimens collected by Ostenfeld from 6-8 m subtidally. Suduroy: Trongisvagsfjgréur. 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. MIO. S21. 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, Suduroy: Trongisvagsfjgrdur (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 Bérgesen (1903) found by Lyngbye to be abundant on Suduroy: Hvalbiar- fjgrdur. 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. umbilicalis (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 coccinea] 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. (!) One. 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. (!) OFe bd ed Od x «> De x x va * * > OO be x > x > be > al ms x Me Me x x mx KK > > be wx KK mK x MK bt x x > * > x wm a al al ele 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 Skalafjgrdur and Hvannasund but was not found in Kaldbaksfjgrdéur. 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 Skalafjgréur. Ascophyllum nodosum often grew together with F. vesiculosus and formed a distinct band at four sites; it was not detected in Kaldbaksfjgrdur. F. distichus subsp. anceps was only found on exposed rock faces at Hvitanes (site 12), at the entrance to Kaldbaksfjgréur. F. distichus subsp. edentatus was widespread at lower shore levels in Skdlafjgrdéur, Hvannasund, and Sundini, where it formed a distinct band. In Kaldbaksfjgréur 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 Skalafjgréur and Kaldbaksfjgréur. Himanthalia elongata was an obvious component of the vegetation on wave-washed lower shores at the entrances to the fjords; in Skdlafjgrdur 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 Ulva lactuca grew beneath the cover of large brown algae. At Skalabotnur (site 10, near the head-end of Skalafjgréur) the underflora solely comprised small plants of Ulva lactuca. In Kaldbaksfjgréur, where fucoids were largely absent, the main vegetation of the eulittoral level comprised bands of Blidingia minima, Enteromorpha intestinalis, Porphyra umbilicalis, and Palmaria palmata. 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 Skalafjgréur and Kaldbaksfjgrdur. 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 Skalafjgrdur (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 Skalafjgréur. At Hvitanes (site 12, Kaldbaksfjgrdéur) these and other red algae such as Cryptopleura ramosa were recorded from lower shore pools. In the middle and inner regions of Skalafjgrdur, 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 Kaldbaksfjgrdéur 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 Skalafjgrour. 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 Skalafjgrdur. The species was also common on wave washed shores at Hvitanes (site 12) at the entrance to Kaldbaks- fjgrdur, 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 Vidarei6i at the entrance to the sound. Alaria esculenta ; Sa A. esculenta was an obvious component of the vegetation at the entrance to Skalafjgrdur, 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 Skalafjgr6ur. Its limit of distribution along Skalafjgrdur was found to be just to the west of S¢ldafjgrdur, 10 km into the fjord from site 1 at the entrance. At the former location DIDIJAIS DUIADSIDH DJ40]UOD DYUOUNG siujsadna vaoydopy]) siupjpjidvs vydiowojavy) winiqnd WNIUD4ay I9A09 ploon] IQIONIDD 8 I$ WNUDIYJAOMALIINYS WNIWD1ID sopeuleg I9A09 ploony ON INPUdIS ¢ a11S DINIID] DAI] DID] A1S DUNJADS1H sypUuysajul Dydiowosajuy pj4ojuod pyuounqg unaindind wniuojI0jsk) s1ajsadna "Dd vaoidas vAéoydopy]D winignd UnIWUDdsaD I9A09 ploon{ IRIAATD pf US wnuviyjaomapyInys WnNIWddaD sopoeuleg I9AO09 PION} ON MST LT Ag sypoyiquin vakydiog pudiunu vIsuIpIg :JO 19A09 asUdp B YJBOUDq BIOYQns ON I9A09 ploonj ON vl SUS DID]JaIS DUIJADSIH) pj40JU0) DYUOUNG siajsadna vioydopy]D winiqna wuniuDsay I9A09 ploon.j punseuueAy [] NS DINIIV] VAIN sypuysaqul pydiouuajuyq DjAOJUOD DYUOUNG winiqna uuniuDpsaZD I9A09 ploon.j INPUIIIS ¢ 11S wunuviyjaomapynys uniuDpsay Meyooy *D winjisodas uo1umDyjyyjvD sopeuleg sdaouv snyoysip snonq SOUBAH ZT OS sypoyiquin vakydiog ppiujod viuvUjvg sypuysayul vydiowosajuq :JO 19A09 asuop B YJeOUNQ BIOYQns ON I9A09 Pploony ON PUNS ET 2S DINIJIV] VAIL) I9A09 prloon.y INUIOQLIPYS (IT HS DINIIV] VAI] pjoav pydiowosuods Npuvusoua] UoYyINOIDUAYd pipupd DiUuDUlDg DIDIJAIS DUNADSID sypuysaqul vy diowosajuyq pj40JUOD DYUOUNG sypuisyffo puljv10D pandas *) siajsadna vaoydopu]D I9A09 ploony yravpun’y 7 aS Npuvusoua] UOYIIOJOUAY DIDIJAIS DUIJADSIDH syouidyffo puyjv40D padidas vAoydopy]y uinuDdIYJAOMaLIINYS WNIWUDAID sopoeuleg IQAO9 ploony 9]}3T SON T OHS ‘so1oods eroyqns [ediouug Zaqey SHELTERED FJORDS AND SOUNDS 141 Table3 Occurrence and distribution of lower littoral and sublittoral fringe species in A. Skdlafj@rour and B. Kaldbaksfjgrour. Table 3A OUTER SITES Alaria esculenta 1,2,4,5 Chaetomorpha melagonium 2 Chordaria flagelliformis 1,2, 4 Cladophora sericea 2 Corallina officinalis 1,2, 4,5 Ectocarpus fasciculatus 1,5 Elachista scutulata 2 Himanthalia elongata 2, 5 Palmaria palmata 1,5 Petalonia fascia 1, 2 Polysiphonia brodiaei 1,2 Porphyra leucosticta 1,2 Spongomorpha aeruginosa 2, 4 S.arcial 2.5 Table 3B Kaldbaksfjgrour 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 MIDDLE FJORD SITES «Ceramium rubrum 1, 2,5,7,9 Chaetomorpha capillaris 8 »Chondrus crispus 8, 9, 10 «©Chorda filum 2,7, 8,9, 10 «Cladophora rupestris 2,5, 7,8 «Cystoclonium purpureum 2, 8, 9 « Dictyosiphon foeniculaceus £25 4,94,0,10 <«»>Dumontia contorta 1,3,7, 8,9, 10 Enteromorpha intestinalis 7, 9 Erythrotrichia carnea 7 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 lenormandii 4, 10 «Pilayella littoralis 1,5, 7 Plocamium cartilagineum 8 « Polysiphonia nigrescens 2,9 P. urceolata Punctaria plantaginea 4 Rhizoclonium riparium 7 «Rhodomela lycopodioides 2, 4,9 «Ulva lactuca 2, 5,9, 10 THROUGHOUT FJORD MIDDLE FJORD SITES Chorda filum Dictyosiphon foeniculaceus Ectocarpus siliculosus Eudesme virescens Laminaria saccharina Myrionema strangulans Myriotrichia clavaeformis Palmaria palmata Scytosiphon lomentaria Ulva lactuca INNER SITES Enteromorpha linza 10 Hildenbrandia rubra 10 Porphyra purpurea 9 Ralfsia clavata 10 Ulonema rhizophora 9 INNER SITES Chondrus crispus Enteromorpha intestinalis Pilayella littoralis <= 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 Kaldbaksfjgrdur 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 nodosum! Sublittoral vegetation Occurrence and distribution of Laminaria spp. The horizontal and vertical distributions of Laminaria spp. at three sites in Skalafjgrdur and Kaldbaksfjgrour 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 Skdlafjgréur 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 Kaldbaksfjgrdur 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 Skalafjgrdéur, 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 St6rafles, which lie seawards of the sites investigated in Skalafjgr6ur. Unlike L. digitata, L. hyperborea was not found in the inner reaches of Skalafjgrdur between G¢gtueidi (site 8) and the Fjardaré 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 Skalafjgrdéur (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 Lambareidi (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 Skalafjgréur, 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 Kaldbaksfjgréur, 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 Kaldbaksfjgréur, where it grew among L. faeroensis and L. saccharina on a substrate of small *For comments on the current taxonomic status of L. faeroensis Borg. see Kain (1976). For convenience we have used this name. SHELTERED FJORDS AND SOUNDS 143 Gigartina stellata & Himanthalia elongata Alaria esculenta Laminaria hyperborea L.hyperborea & L. saccharina -10 y L. saccharina e4 Ed A im ee @!, GZ A A L16 Boulders Stones, shells Chiottads = 26 SP —<—— tyke Vale Sate BAVA A rtlentape eet. Puat Rock 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 YE 2 L. faeroensis r9 Rock Stones, gravel Mud cer res 10 50 l Fig. 3 Site 6: schematic representation of vegetation and transect profile; details as for Fig. 2. Porphyra umbilicalis & Ulva lactuca Cruoria si : NE ess, Secs H14 “as oe a ¥ Oy | ‘ f Sok eel i, Cos nee Rock Stones & boulders Sand & stones 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 (24 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 Skalafjgrdur, 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 Kaldbaksfjgréur 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 Kaldbaksfjgr6ur. 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 Skalafjgrour 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 min length (stipe 1-2 m; lamina 1-5 m long, 0-6 m wide). At Ggtueidi (site 8) beds of L. faeroensis spread to a distance of 200 m offshore. At sites 13 and 14 in Kaldbaksfjgrdéur, 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 Skalafjordur, Kaldbaksfjgrdéur, 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 Lambareidi (site 6) in the middle regions of Skdlafjgroéur. 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 Ulva 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 glaciale, and Pseudolithoderma exten- sum were detected as a distinct association on rocks and boulders in deeper waters at the entrance to Skalafjordur, 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 Sk4lafjgrdur (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 Skalafjgrdur and Kaldbaksfjgréur, 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 Skalafjgréur and Kaldbaksfjgréur; 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 Bgrgesen’s observations Our observations on the algae of the littoral fringe differ little from those made by B¢grgesen (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. Borgesen 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 Skalafjgrdur, and were particularly abundant at sites where a firm substrate prevailed; in Kaldbaksfjgrdur these algae were conspicuously absent from the inner regions where littoral rocks were colonized by dense growths of Enteromorpha intestinalis, Palmaria palmata, 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 Skdla- fjgrdur, 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 Skalafjgréur and Kaldbaksfjordur 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. Bgrgesen (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 Skalafjgrdur, Kaldbaksfjgréur and in a sheltered lagoon near the head end of Hvannasund. Both Bégrgesen’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 (Halosaccion) ramentacea at the head end of Hvannasund where it grew in shallow standing water in pure stands; Bgrgesen 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 Skdlafjgrdur and Kaldbaksfjgréur, we did not locate the large growths of Monostroma fuscum noted by Bé¢rgesen as characterizing a ‘Monostroma- Enteromorpha Association’. Bérgesen 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 Skalafjordur, and additionally found Capsosiphon fulvescens, growing in shallow, brackish water in the estuaries at the heads of Skalafjgréur and Kaldbaksfjgrdur. Borgesen (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 B¢grgesen (1905) recorded from sheltered waters in Skdlafjgréur. Bgrgesen (1905) described the ‘Halidrys Association’ from the shallow infralittoral region at Glyvrar in Skalafjgrdur; 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. B¢rgesen (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 Bgérgesen (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 Skalafjgrdur and Kaldbaksfjgr6ur 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 Skalafjgrdur and Kaldbaksfjgréur. Our observations parallel those of Bérgesen 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 Skdlafjgréur. We confirmed Bégrgesen’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. Bérgesen described a ‘Desmarestia Association’ which appeared to comprise almost pure stands of Desmarestia spp. from similar habitats. We failed to find eel-grass Zostera marina in sheltered habitats. Bérgesen commented that the plant was rare on the Faroes and cited only one locality (Vaag fjord) for this community. The leaves of Zostera bore a number of small algal epiphytes. A well developed subflora was detected at the entrances to Skalafjgréur and Kaldbaksfjgréur 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 Bgrgesen 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 Skalafjgrdur and Kaldbaksfjgrdur. 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 Bérgesen 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 Bordoy and Vidoy 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 as Alaria 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 Borbdoy 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. Bégrgesen 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 Skalafjgrdur but did not find the community in the inner regions of Skalafjgrdur. A sc oe al of crustose algae dominated by Cruoria sp. was present in deep waters in Kaldbaksf- jorour. 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 Bgrgesen (1902; 1905) were found in deeper 148 SEAWEEDS OF THE FAROES waters at the head end of Skalafj@rdur; 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 et al. (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, 1978a, 1978b) for fjords on Iceland. River estuaries drain in the head ends of Skalafjgréur and Kaldbaksfjgrdur, and, according to Beérgesen (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 Skalafjgréur and Kaldbaksfjgréur. 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 Kaldbaksfjgréur and Skalafjgrdur revealed a number of differences. Laminaria hyperborea was present as dense forests on bedrock at the entrances to Sullom Voe, Skalafjgréur, and Kaldbaksfjgréur, 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 Skalafjgrdur and Kaldbaksfjgrdéur, L. hyperborea did not grow at the landward ends. The broad (‘cucullate’) form of L. hyperborea, found in the inner regions of Kaldbaksfjgréur 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 Skaélafjgrdur and Kaldbaksfjgr6ur, 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 Skalafjgr6ur and Kaldbaksfjgréur 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 SkaAlafjgrdur, the vegetation was much less extensive in Skalafjgrdur 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 Skalafjgréur, 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 Skalafjgréur and Phyllophora crispa was not found at all in Sk4la- fjgrdur. ‘Trailliella’ grew on mud and silt in the deeper parts of Sullom Voe, and was found in a similar habitat in Skalafjgréur. Although some fucoids grew permanently immersed in shallow standing water at the heads of Skalafjgré6ur and Kaldbaksfjgr6ur, 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; (ili) 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 Skdlafjgréur and Kaldbaksfjgrdur are in some respects similar to the patterns of distribution in Hardangerfjord, Norway (Jorde & Klavestad, 1963) and in Dyrafjor- dur, Iceland (Munda, 1972, 1978a, 1978b). Brackish water species Capsosiphon fulvescens, Percursaria percursa, and Rhizoclonium riparium were recorded from saline meadows in Hardangerfjord and from oligohaline habitats in the inner part of Dyrafjérdur; C. fulvescens and R. riparium occurred in the estuaries at the heads of both Skalafjgréur and Kaldbaks- fjgrour. 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 Hardangerfjord and Dyrafjérdur; a similar association was restricted to the inner ends of Skdlafjgréur and Kaldbaksfjgroéur. 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 Hardangerfjord, although Bérgesen (1902) did not find it and dismissed previous records. The limit of distribution of Laminaria saccharina in Dyrafj6rdur, Iceland, is salinity depen- dent and coincides with the 28% isohaline (Munda, 1978b). The species occurs throughout the middle and outer parts of Hardangerfjord, 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 Skalafjgréur, 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 Dyrafjordur, Hardangerfjord, and Skala- fjgrdur 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, Dyrafj6rdur, and Skalafjgréur, 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 Kalbaksfjgrdur 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 Skdlafjgrdur. A few species of red algae, such as Ceramium rubrum and Polysiphonia elongata recorded from the inner part of Skdlafjgrdur, 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, and Rhodomela spp. 150 SEAWEEDS OF THE FAROES 7. Conclusions The occurrence and distribution of marine algae are similar in Skdlafjgréur and in Kaldbaks- fjgréur; 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 Skalafjgréur and Kaldbaksfjgréur, 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 Dyrafjér6ur 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 Skdlafjgrdur and Kaldbaksfjgrdur than in Hardangerfjord and Dyrafjérdur, 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. Lining, 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 Borgesen, F. [C. E.] 1902. Marine algae. Jn [E. Warming (Ed.)], Botany of the Ferdes based upon Danish investigations. Part II: 339-532. Copenhagen. — 1904. Om Algevegetationen ved Fergernes Kyster en plantegeografisk undersggelse. pp. [6]+122+[4]. Kgbenhavn and Kristiania. — 1905. The alge-vegetation of the Feréese coasts with remarks on the phyto-geography. Jn [E. Warming (Ed.)], Botany of the Ferées based upon Danish investigations. Part III: 683-834. Copenhagen and Christiania. — & Jonsson, H. 1905. The distribution of the marine algz of the Arctic Sea and of the northernmost part of the Atlantic. Jn [E. Warming (Ed.)], Botany of the Ferées based upon Danish investigations. Part III: Appendix I-X XVIII. 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 notater fra Faergyene 1975. [Unpublished report. ] Irvine, D. E. G. 1982. Seaweeds of the Faroes. 1: The flora. Bull. Br. Mus. nat. 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, I., Price, J. H., & Farnham, W. F. 1982. Seaweeds of the Faroes. [Abstract.] Br. phycol. PATS 234. Johansen, J. 1979. 30 foroysk taraslog. Torshavn. Jorde, I. & Klavestad, N. 1963. The natural history of the Hardangerfjord 4—the benthonic algal vegetation. Sarsia 9: 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. 1978b. Survey of the benthic algal vegetation of the Dyrafjérdur, 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 ver algobservationer Férodrna sommaren 1970. [Unpublished report. ] Russell, G. 1978. Seaweeds as environmental indicators. Porcupine Newsl. 1: 109-110. Tittley, I., Irvine, D. E. G., & Jephsen, N. A. 1977. The infralittoral marine algae of Sullom Voe, Shetland. Trans. bot. Soc. Edinb. 42: 397-419. Pk 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 0DG Contents Synopsis . ‘ ; ; ; ‘ , : : ‘ : ; eoetS3 1. Introduction . : : ; ‘ ‘ ‘ ‘ ‘ : ; kos 2. Methods ; : 5 ; ; ; : : pee bhp, 3. Sites, descriptions, and profiles . ; ; ‘ : : : 7 Pre ek, 4. Results: comparative species distributions . : ‘ : ; ; 170 i. Subtidal: majororganisms . : : : : ; ; ; Peavy! Alaria . ; : ‘ : : , tse Laminariah yperborea and other Laminaria: spp. : é : Samet 6: Populations at depths greater than laminarian limits : é : - 183 . Subtidal: epiphytes and underflora : . ; : é 7 133 Laminaria hyperborea epiphytes and underflora ; ‘ ; : nes ul Epiphytes/underflora to other major subtidal organisms . : : ae | iii. Subtidal: other specializedsituations 3 ; , ‘ ; cae AV. Detrital conditions : : : ; ‘ : ke Epizoic growths and animal associations ; : , ‘ ; ele Pools and standing waters ; , : : : ‘ : gy 216 iv. Intertidal : ‘ ‘ : : ; : é : apa A a oe and conclusions ; : : : : , . 2 2/218 . General . : : ; : : : ; ‘ : ‘ a 218 i Thesubtidal : ; : : P : : : : e219 iii. Theintertidal . : : ' : : : i : : 2224 6. Acknowledgements , ; , aT ee ; F ; : « 224 7. References . : : ; : A : : , : 3 2 224 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 Bgrgesen’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 et al., 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 Tangafjgrdur and Skalafjgréur, 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 Bégrgesen (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 Bgrgesen’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 Bgrgesen (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 Bérgesen (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 Bgrgesen’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 Bordoy, Eysturoy, Fugloy, Koltur, Kunoy, Mykines, Ndlsoy, Sandoy, Streymoy, Suduroy, Svinoy, Vagar, and Vidoy. 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 B¢grgesen’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 Borgesen (1905), in dealing with that habitat in a different section where comments on additions or differences are required. ALD ‘, 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 = Suduroy Ko = Koltur Ka = Kalsoy LD = Litla Dimun No = Nélsoy Ku = Kunoy SD = Stéra Dimun Str = Streymoy Bo = Bordoy Sk = Sktivoy Va = Vagar Vi = Vidoy Sa = Sandoy My = Mykines Sv = Svinoy He = Hestur Ey = Eysturoy 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. Depths 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 var. demersum Ho Halicystis ovalis Ae Alaria esculenta Hr AHildenbrandia rubra An Ascophyllum nodosum Le Lomentaria clavellosa Ap Ahnfeltia plicata Ld Laminaria digitata Ba ___ Barnacles, in general Lg __Lithothamnion glaciale Bb Brongniartella byssoides Lh Laminaria hyperborea Be —_ Balanus crenulatus Li __Lithothamnia, in general Bp Bryopsis plumosa Ll Leptophytum laeve Bs _ Brittle stars, in general Lo Lomentaria orcadensis Ca = Callophyllis cristata Ls Laminaria saccharina Cc = Clathrina coriacea Ma Membranoptera alata Cd = Callithamnion decompositum Mm Modiolus modiolus Ce Ceramium shuttleworthianum Od Odonthalia dentata Cl = Callophyllis laciniata Pa Palmaria palmata Cm Chaetomorpha melagonium Pe Plocamium cartilagineum Cn Callocolax neglectum Pe Pseudolithoderma extensum Co Corallina officinalis Phe Phyllophora crispa Cr Ceramium rubrum Php Phymatolithon polymorphum Cs = Callithamnion sepositum Pht Phyllophora traillii Da Desmarestia aculeata Phtr Phyllophora truncata De __ Dilsea carnosa Pl = Polysiphonia lanosa De Dermocarpa prasina Pm Porphyra miniata (= Entophysalis conferta) Po Porphyra umbilicalis Dl. = Desmarestia ligulata Pp __ Pterosiphonia parasitica Dm_ Derbesia marina Pr Phycodrys rubens Ds __ Delesseria sanguinea Ps _ Ptilota serrata Dv _ _Desmarestia viridis Pt Ptilotaspp. Ec __ Ectocarpoids, in general Ptp Ptilota plumosa Ee = Echinus esculentus Pu _—_— Polysiphonia urceolata Ei = Enteromorpha intestinalis Pv Patella vulgata Fd = Fimbrifolium dichotomum Sa Spongomorpha arcta Fl = Furcellaria lumbricalis Se — Serpulids, in general Gs __ Gigartina stellata Sp __ Sponges, in general Ha _ Halichondria Ul = Ulvalactuca He Himanthalia elongata Ve -Verrucaria spp. General symbols l = epilithic Pp = epiphytic used in connection with taxa abbreviations or on distribution bars. Z = epizoic } in( ), after1, p. or z (see above) = substrate types or ‘host’ organisms involved. S =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). een ees = organism scattered, but consistent to occasional, over the depths concerned. Rata enasler eee he = organism present, but detected as very sparse to rare. 158 SEAWEEDS OF THE FAROES Site list 1. Akraberg, south Suouroy. 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 Suduroy. Intertidal. Gullies and faces of stacks just off-shore. Generally exposed positions. 3. Trollkonufingur, Skarvanes, Sandoy. Intertidal. Asen and Rueness. Tide-pools in rocky but not excessively steep shore. Pools showing strong upcarry of usually subtidal flora. 4. Husavik, 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. Gdbdidrangur, Mykines. Subtidal. Off rocky steep small islets west of Mykines, to 20 m depth. 6B. Tindagjégv, Mykines. Intertidal. Sloping rock with pools, north-west side of island. 7. Borgagjégv, Mykines. Subtidal. Vertical to slightly overhanging slab rock-face, with uneven large boulders on bottom below. All depths to 17 m. 8. Skerhélmur, Sorvagsfjgrour, Vagar. Subtidal. Seaward face of exposed steep rocky islet. Collections from between 6/7 and 15 m depths. 1 35 o Sheltered 2: . 3 5 positions «300 = 1 Outside and away LW. I from harbour walls lo Depths(m) ij <—— 22m ” 2 Horizontal distance Depths (m) Symbols Fig. 2 Sandur, Sandoy (site 5). Sketch map ins Bl showing main observations area and locations eet Sp(encrusting) of profiled stretches. Profiles represented are: j Le 1. Outside and away from harbour walls. 2. Harbour wall end: concrete-faced blocks. f) bo 3. Configuration on boulders and bedrock below wall 9 vm end base. See p. 157 for lettered symbols. pica oy ars bel concrete-faced blocks OPEN SHORES 159 9. Stack in Dragasund, opposite Tindhdélmur. Subtidal. Vertical face and easing slope/boulders below, in 5-9 m depths. South side exposed, north-west side less directly so. 10. Oyrargjégv, Vestmannasund, Vagar. 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. Skrioutangi, north-east (seaward) shore of Nélsoy. 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. Kirkjubgur, 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 Kirkjubgur cove. 14. Velbastadur, 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. c £9 35 Fy °F 3 7 Ti = =) ia 5 Orodacn i ‘ I ae 3. Configuration below rs H wall end base Up lea & i) Ss ees eet oa is i g iz 1b a a i 9- 152 ‘wa ee Le eg OTe pL LAL H 10- i : £ 2 H £ 1 a Ike H Y aa TTL 11% i aoe OE TAAL RERUN deaD 8 MD o>. tea hee ee limit of detailed 12- + ! Pe BP eat ES I poate keat nae observations pon el Popipiytes vunsiertiocs $ pel ig saradty flora patches cf “ty, = re) 5 io Horizontal distance (m) Fig. 2 (cont.) Caption, see p. 158. =a 5a 2h 4g ce ha= ay _- = a i eae ee ee © Tem ae Oe ne ee a o mil NssthA 8828 8829 328 i] i r TT le eae | 5- 15 | 1 s 1 10- = 1 ae at 1 | ot oe ae i! re er. Jog Sines ee 1 | | | \ \ { r-Al 1)! ae 1 | 1 ol ; | | 1 15- ! a eee ae ary | ! Or ess \ ! | Votes aiticos i] I | | | | ! 1 \ a | I | ! | -I a io 1 | bog : ! AD) Balti “ | Seen Bis jae ese os l fy he | £ a | an ae : pay <= | 1 I re | | J S| ! I @ 25- 1! ~ t+jijaa A f | lle ! 1 THe ae, ou | I | aes 4 pce avd : 30- 4 Observation yi > ! | limit - 11 1 | | 1 | | | Cc, | | s, iy 35> 9, ‘ 1 Y MA f ! : I tay, ery Xey aly Yak tth 40- «<—Observation limit—12_______ 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 (Nélsoy). 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 saccharina). 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 Tangafjgrour, 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. Stérafles. Subtidal. Isolated rock in the middle of Tangafjgrour 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 (Skdlafjgrour) 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. A/aria 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 Noroskali, 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. Sakshovn, south side, 200 m seaward of Grétdalsgjégv, 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 Fig.4 Hoyvik transects (site 19) 1. 1T profile $ is ” Oo > 8 ot 2-3 Peek ata aa et ae | ac ( ==4 (jS0105 9n1} Ou) Table, ——_parejyenss] ears! F----------4 fF *--ysaedg4 i P= UIpIOn Alb UST 92 ree = aac (w) syydag gae below line even Cp absent NO al strong E—©& W tidal stream rough; boulder-strewn 25- 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. Cony 2 So a 2 £2 d 1 8% © 5 a — nH 986 2 88 o oo r) = © OG 6. hr c 228 : tee Sok eye o & ” = —_ ih SaSadéo 6f€6as5da 6 Fig.6. To west of and below Kollur Peak (site 30) ® ‘hi oo | Orrigeet 9 110 | wn no | a | | | | | \ | | cer ! | aa 1 | ty | | ees | hal tna \ ae | | | | | hoa | ee | l a 1 il 1 | 19 el \ RP het ! V of \ ca | [pe lest | ie pa | | | | ae | | et \ ks | ! | limit of tn y i ole, a IER [elds Oe observations large the ular ~* 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 c open shore profile west of Scan’ a cleft length: 175m —-4------}-------------------4-+--5---------------4 ! cleft centre dips 1m B south wall WES 2 ie oe ane only Dm on ] Ay shallower diagrammatic: not scaled _Co | Lo in Sp iccark “8 wii Li Li (bracket s ungus") 3 77) a W: > overnangs = T C0 d -10 with sparse red upCarry from Lh depths Fig. 7 Gjégv (site 31). Plan of the Cleft, indicating the positions of the profiled sections A—A,; B—B;; C—C, (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. Oyndarfjgrour, 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, Kalsoyarfjgrour. 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 167 Cc WL ) Open shore profile, Lh west of cleft -4 -8 Od and Phc Pa underflora | Pl2 © S | = ! luxuriant mixed a : red algae: = abundant Ds; Pr|_|. ~~ Phc ;Cl Gome Cn); small Pc; small la single Co isolated; small slope with red algae continues C, Fig. 7 (cont.) Caption see p. 166. 35. Anir and adjacent narrows, Haraldssund, Bordoy-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 throughfiow 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 (Bordoy) 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), Vidoy. 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 2S o ® x oy eet ~ fae Sos 3 E © oa SS 5 e¢€ OCA tGtAG6E BO Sa C4c86 ' “s ey r 4-+ ----------- / ag r.) 2 le g S74 7 TT ! F3 ie | no) oO ow \ Le ci v : ni AS \87 aG| & | # & =| / so | 3 ! 1 = —| \ wl y : fod ne | o| 8 . 3 2 oO a c ae Ts é E | ma a 2 ut: l ae T 7 tke 8 = rs 1 1216 & - 2 | 7 3 a p= | § 3 oO —_ 5 S ba i 3le a) — 1 : St=] ‘on 1; & — | _| : ia &) = ig ee : _ — ——7, 1 3 26 2 B Ze ese 25 ” ! o o a Lah = so rar x! Zo a T — <=: 3 Ss! fA =a S5 & © re) a2 am en) — es ae a ! in fie Be BRS a i ! sea alakahss | 1 o: "aaa | ' ae i Lait 2On tote ice fa} 3 MR 2 Sects ake Observations 21 Fig. 8 Off Kuvingafjall, Kalsoyarfjgrour, 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. Vidareidi, Vidoy. 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. Vidareidi, Vidoy; 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 Depths (m) n ae Es as) 33 oO-= & S56 A £2 @ of oe os = oO Is ag 12 : a — fs I< § 9 Wk ei 8 f WN ee Sees i ee nS a a ee ® = es. 35 2a = 8 eae : aie Fas f . +3 Bs i a ee ote To /} ‘ a 8i5 wena se ofr: | fo ' ! 2 oc) iis sa = ian Sri 3 i ' ' re ee : Yjiig = ee eee. < 4 et flea res eee — ar ®©%5 ive 4 o : ' £ ieeeay = ae ° = ! rs af 2 {8 | i ig gan! Dc He 05 SS! i pare asmall to ee 2g ad + ' 2 m He | ‘ 4 = I! i} k = ae a eerrre a2) + Py ar Ss Pf aa Ss = 0} ~ th vib Z = : Pr oS ae eee ee re a. eee UI stones on silt PaGe boulders on silt Cr or gravel Fig. 9 Central Cleft, Vidareidi (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. Laminaria 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. Hamarsgjégv, north of Eidsvik, Viboy (= Vibareibi, east side). Intertidal and subtidal. Intertidal chiefly of concrete steps of Eidsvik 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. Marknagjégv cave, east Vivoy. 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. Svinoyareidi, 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 Svinoyareioi. 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). Depths(m) 26 28 170 SEAWEEDS OF THE FAROES 43. Promontory in Scarosvik, Fugloy. Intertidal only. Collection by boat from steep rock surface; animal-dominated exposed rocky shore throughout. 44. Svgoutangi, 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 ey’amined at Nes (Tangafjgréur) and Hvitanes (Kaldbaks- fjgréur entrance, described by Borges, 1905: 694, as carrying a‘. . . perfect “open sea” algal vegetation . . .’) would have been appro riate for comparative purposes but for two reasons: (i) both are situated in areas close to ope%;shore sites and with existing, more appropriate, comparative sites of similar characteristics; \@) both are described in some detail by Tittley et al. (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 coo ~ Zz 2 2 > &$ $ Exposed rock standing offshore; 2 =e partial transect of deeper subtidal 9 £ 8 a E i 3 8 3 2 ROBY < S56 KA BKSSK LASSE B Ned ee { as fen eee 1 a 4 es selaa | 1 238 sg age | 1 ee E i | | B85 3 ar e| ts cia Tay - i] 2 : — Ne, LIMITS OF DETAIL WL: p /p(Lh) on horizontals. Fig. 10 Svg@dutangi, 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 17k subtidal, although the extent and frequency of occurrence may be much greater in the one milieu than the other. Species that are cqgmparatively 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 Bérgesen’s observations. Bérgesen (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. apclaiione 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 ¢.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 Tablel Alaria on the Faroes. SEAWEEDS OF THE FAROES a. Epilithic Station 20 2D 24 25 21 29 30 31 Substrate wall broken boulders below/away from wall wall boulders away from wall steep bedrock steep rocks and irregular large boulders firm rock rocks and large boulders rocks rocky firm areas rocky firm areas stony/bouldered patches rocks and firm boulders rocks/boulders rocks rocks large broken boulders very steep firm rock forming cliff-face steep rocks and irregular large boulders steep rocks of very steep to vertical cleft wall Depths (m) 0-1 (1-)6-7 +0-5 to +0-6 0-4 1-2 around ELWS 0-2 around 0 0-1 +0-2 to 0-2 2°5 5-6 around ELWS around ELWS 0-2 0-1(-2) 2-3 (0—)5-9 (0—)5-10 0-1 Notes 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 Tablel cont. Station Substrate Depths (m) Notes 34 firm steep to vertical 0-5-2 sharp demarcation Laminaria rock 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. boulders on gully bottom 8-10 some small clumps in the deeps. 35 small rocks in dark sand subtidal fringe somewhat patchy; fertile. to2m 36 moderately steep rocks 0-2 patchy, but locally dense band; fertile. oF steepish firm rock subtidal fringe band-forming; fertile. (not boulders) to 1 m; to +1 in upcarry areas 38 steep firm rocks of cleft +0-5 to 0-5 fertile on south (scoured) side. sides 42 rocks 1-2 band-forming. 44 very large boulders over 15 a few sporelings only. bedrock, or on other boulders embedded in sand b. Epiphytic Station Host Position Depths(m) Notes 11 Laminaria hyperborea _ stipes 1-2 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/Jaria growths from the fringe at c.0 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). 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Harbour conditions elsewhere, both in and outside fjords, tended to be dominated in otherwise L. hyperborea depths by either L. saccharina in various combinations with large algae of other groups, or mixtures in varying proportions of L. saccharina and L. faeroensis* (as, for example, in the artificial harbour on the north shore of Kollafjgréur). Elsewhere in Kollafjgréur (Jgkilsgjdgv, on the steep western shore of Tangafjgréur opposite Raktangi—station 23— and at the extreme outer entrance to Kollafjgrdéur), L. saccharina and L. faeroensis were sufficiently vigorous and dense to form replacement forest, for the more usual dominant L. hyperborea, between 1 m and 5-6 m depths. Very large L. saccharina/L. faeroensis within these depths bore laminae up to 16 ft (c.5 m) long. Within fjords but outside harbour conditions, at least some L. hyperborea was usually present, often at its ‘cape’ form (f. cucullata); this form occurred in 2-3 m depths at Jokilsgjégv. The circumstances in which the open shore presence of L. hyperborea did not result in the formation of forest were detected at Station 29 (near Stakkur, north-west Streymoy) and presumably reflected too steep an angle on the cliff face in very turbulent conditions, followed by too mobile a substratum where the angle eased to a boulder-strewn bottom at 23 m. Elsewhere, particularly at station 12 (N6lsoy), distribution depths of forest density of Laminaria hyperborea varied quite markedly from the usual situation. Development of forest density commonly occurred on firm but not excessively steep natural substrata between (0—)4-16(—22) m depths. At station 12, forest growth was definitely deeper than anywhere else examined around Faeroese shores, beginning at about 10 m and descending to 22 m and below before thinning to become scattered but still large plants. Small plants with long stipes and attenuate blades were still present attached to scattered rocks amongst the gravel substrate down to 30 m depths. This tallies well with the 15 fathoms maximum depths for the species noted by Bérgesen (1905: 758-759), although he elsewhere (1902: 462) commented that the ‘. . . proper habitat is from a depth of a few to as much as 10 fathoms, sometimes even to almost twice that depth.’ Because it is a relatively larger plant with erect stipe and usually grows in depths greater than such species as L. digitata, L. saccharina or Alaria, Laminaria hyperborea generally shows less overt and identifiable effects of upcarry by local configuration in its vertical distribution. Nevertheless, it is occasionally possible to identify such effects, as at station 24 (Kvivik) where definite overlap with the Alaria band (occupying depths of 0 to 2 m) was promoted by the upcarry of L. hyperborea plants in shallow subtidal channels in the bedrock. The mosaic situation that occurs with the major laminarian bands in some localized areas, even in the absence of substrate variation, is exemplified by data recorded at station 34 (Kalsoyarfjgrdur, off the west coast of Kunoy). The lower intertidal and shallow subtidal there is generally, down to at least 8 m, a very steep to vertical rock-face; below that, the slope eases and the substrate changes to boulders over detritus, penetrated by sandy channels clear of boulders. Sandy channels persist amongst the bedrock that succeeds the boulders at 16 m. The vertical rock that terminates in about 8 m is cleft in places rather deeply; the inner ends of clefts show erosion caves with basal detrital build-up that submerges the lower 1 metre of the rear and side walls. Laminaria digitata formed a fairly consistent one-metre band between 0-5 m above and 0-5 m below the low-water level; it lacked epiphytes and had only straggly Plocamium as underflora. Below —0-5 m and sharply demarcated (but not spaced) from Laminaria digitata was a band of Alaria, occupying —0-5 m to 1(—-2) m depths. Small outlying clumps of Alaria also occurred on boulders on gully bottoms in 8-10 m depths. The lower limit of the Alaria upper band was complex; in places (and this was most common) was a graded but continuous transition from Alaria to fully-developed, but straggly and sparse, Laminaria hyperborea over the 2-3 m depths band. In other places, young Laminaria hyperborea plants occurred between Alaria and Laminaria hyperborea, mixing into the upper fringe of the latter. The straggly and sparse L. hyperborea quickly (by 3-4 m depths) became more abundant, larger and denser, in a continuous sequence to about 8 m, the bottom of the almost vertical wall. By then, true forest *A name used for convenience only; see Irvine (1982). OPEN SHORES Table 3 Other laminarians on the Faroes 179 Laminaria saccharina a. Epilithic Station 4 19 20 24 25 va) 31 i (Anir) 42 Substrate broken boulders (1-10 m) grading to bedrock with stones in sandy gullies (8-10 m) firm boulders sheltered positions on rock firm rock and boulders firm rock and boulders on stony/bouldered patches firm rock extensive sand-covered gullies in bedrock small boulders on solid substrate large broken boulders boulders stabilized in sand small boulders in dark sand; concrete artificial substrata shallowly-sloping firm rock, giving way to steeper face b. Epiphytic Station 36 Host Laminaria hyperborea Laminaria digitata a. Epilithic Station 9 Substrate steep to vertical firm rock Depths (m) 1-10 4-12 shallow subtidal shallow subtidal shallow subtidal to 2 m 5-6 all deep pools and shallow subtidal 4-6 1-7 1-9 c.5-6 shallow subtidal, to 34m 1-2 Position on stipes Depths (m) 1-5 Notes mixed with Alaria, 1-6 m; plants on stones in sandy gullies at c.10 m often buried to 7-10 cm up stipe. mixed with Desmarestia aculeata and massive Chaetomorpha melagonium inside shelter of harbour back from exposed headland. only in harbour formed by artificial moles; not on open natural coast. masses of large plants, mixed with L. digitata, in sheltered ‘lagoon’. in cove | alongside headland; not on fringing bedrock; equal intermix of large fertile Alaria and L. saccharina. in semi-exposed positions on head. some on larger fragments and on lateral bedrock. Laminaria faeroensis; as locally scattered, patchy mixture in equal proportions with Alaria and Laminaria hyperborea; with thick stipes and long blades. codominant with Desmarestia aculeata below Alaria. common and locally dense; inner end of main cleft. large vigorous plants; iridescent. in sheltered positions; few plants, mixed with Alaria. Depths (m) s Notes above L. hyperborea forest. 180 Table 3 cont. SEAWEEDS OF THE FAROES Station 13 18 19 (head and cove 2) 20 23 34 S00) (Anir) 36 37 38 42 Substrate firm rock of irregular but steepish slope firm rock and large boulders steep firm rock firm rock on headland firm rock and boulders firm steep to vertical rock some boulders on detritus; also on artificial concrete/ metal/wood quay moderately steep firm rock steepish firm rock on steep upper firm rock sides of cleft shallowly-sloping firm rock, giving way to steeper face b. Epiphytic Depths (m) shallow subtidal shallow subtidal to 2 m; lower intertidal pools ; lower intertidal/shallow subtidal, low water +15 cm, as band of laminarians all deep pools and shallow subtidal 1-2 +0-5 to 0-5 subtidal fringe, to2 m lower intertidal pools; shallow subtidal to about 2 m 1-2 mixed with Alaria in 0-0-5 m on north (scoured) side of cleft; forming narrow band over 0-33 to 1 m, below Alaria (+0-33 to 0-33) on south (non-scoured) side 2-3; also some intertidal pools Station Host Position Lambareidi, Skalafjgrour. Laminaria faeroensis on stipes Chorda filum a. Epilithic Station Substrate Depths (m) 20 firm rock and large subtidal fringe, to2 m boulders Notes narrow fringe around most of head; more luxuriant into bay to south. below or mixed with Alaria in exposure; with Laminaria saccharina in shelter. few plants admixed in Alaria band. in semi-exposed positions. good plants, below Alaria. young plants lower, forming interband between Alaria and L. hyperborea; no epiphytes noted. strong fringe, especially on quay. densely present at about 2 m; above more sporadic, amongst Alaria and in pools. exposed shore; firm band below Alaria, o¢casionally above it where Alaria patchy; grades to L. hyperborea. see depths comments. fringing band, patchy, below Alaria. Depths (m) shallow subtidal (within 2 m) Notes in more shaded and wave-sheltered parts; few plants only. OPEN SHORES 181 Table 3 cont. Station Substrate Depths (m) Notes Ps) larger fragments in shallow subtidal, in — mobile substrate channel deepening to lm Hvannasund __ boulders on detritus in deeper parts of sheltered by artificial (causeway/ lagoon only, c.1—2 m closure by causeway and lagoon) protection from harbour moles. Qualifications 1. Omitted stations lack appropriate data. 2. For details of Alaria distribution, see Table 1; for Laminaria hyperborea, see Table 2. 3. Presence of Laminaria faeroensis, Alaria, and Laminaria hyperborea far into Sundini (25) derives from fast flow in sheltered sound. , 4. Depths are below low water unless sign + appears. 5. All Chorda filum records from stations sheltered from wave-action. density being reached, the plants were large and dense; the forest persisted over boulders and in bedrock channels to the limit (16 m) of the dive, by which depth L. hyperborea plants were very large and strong, with unbreakable stipes, and the forest was very dense. Although the Laminaria hyperborea forest was by no means devoid of underflora (see station profile and the appropriate later section), stipe epiphytes were the more obvious subordinate flora in the upper 4-5 m of the Laminaria hyperborea depths; richer and more dense underflora developed in thinning areas and with increasing depths, although the boulders over detritus in 8-16 m depths showed very few of the usually very obvious crusts (e.g. of Cruoria, Pseudolithoderma, Lithothamnia), probably due to canopy outshading (see Discussion and conclusions for detailed comment). Where slightly more sheltered locations produced conditions that would also support growths, at least locally, of Laminaria saccharina, as at Anir (Haraldssund, station 35), fairly strong mosaic intermixing of all four of the more common ‘open shore’ laminarians occurred. Anir has an intermittently strong throughflow of water just off-shore, but this hardly directly affects the shore-wise conditions in the subtidal of the boathouse-and-quays area examined. There, and immediately offshore, juxtapositions of Laminaria hyperborea (‘forest’ to 2 m in the shallow subtidal, on small boulders buried in dark sand); Alaria (the usual fringe on same substrate and in same depths as L. hyperborea, but locally patchy only); Laminaria digitata (strong fringing growths at and just below extreme low water mark of spring tides on the artificial substrata of the quay, concrete, metal and wood; some plants on boulders in the detritus); Laminaria saccharina (principally on small boulders in the dark sand in shallow subtidal depths to 3-4 m; some specimens present on concrete artificial substrata at the water level), are impressive if irregular. Similar mosaic juxtaposing of all four common species was noted at stations 13; 18; 19; 23 (the latter with Chorda filum substituted for L. saccharina); and 42. The numbers of instances where three of the four species were juxtaposed, the missing fourth usually being either L. saccharina or L. digitata, are legion (see Tables 1, 2, 3). Open shore populations of Laminaria saccharina are in our experience rarely present in considerable depths, although Johansen (1979) gave limits on the Faroes as in 10-15 m. This shallow growth was at least in part due to the fact that along open shores the appropriate sheltered water conditions most frequently occur in protected channels, coves and inlets, where available depths are shallow. Reference to Table 3 will show that only at two stations (4 and 5), both with a high degree of extra protection from adjacent harbour walls in already wave- sheltered conditions, was L. saccharina detected at 10 m depth or greater. Similarly at Eidi (station 27), where the species was detected down to 9 m, harbour conditions were again 182 SEAWEEDS OF THE FAROES involved. Maximum depths otherwise noted were, according to local configuration, 6-7 m, and most populations grew within the first 5m. Depth limits of the more exposure-tolerant Laminaria digitata were even more rigorous; the great majority of observed populations grew within the narrow fringe covered by the first 2 m depth. Only a few plants in the growths at stations 9 and 42 attained the 5 m and 3 m depths respectively detected as maxima. Jéhansen (1979), however, recorded 10-15 m depths for Faroese L. digitata. Chorda filum was noted in very shallow fringe conditions only, but in too few instances for assessment on a general basis, particularly in view of Johansen’s (1979) statement of 1-10 m as the depth range. All Laminaria hyperborea populations, of forest density or not, showed with increasing depths an increase in density and luxuriance up to a maximum that occurs most commonly between 4 and 12 m, and a subsequent decrease in such parameters that occurs over a greater depth range than did the initial increase, usually 8 m or so against 2 m or so (Table 2). This negatively skewed density curve is more or less paralleled, although with the individual size of plants somewhat more positively skewed, by curves of plant size and epiphyte density. This latter is also more positively skewed, but less so than is plant size. Variation in plant size between average in the densest depth range and average in the sparsest depth range can be quite impressive—absolute measures between largest and smallest even more so. Even at station 29 (near Stakkur), where plants were relatively poorly-developed and forest density was not formed at any depths, plants at the upper limits of L. hyperborea growth (8 m depth) were up to 60 cm total length, whilst at the lower limits (15 m), total lengths were less than 15 cm. In populations more representative of full potential development, similar proportional growth and characteristics are revealed, but plant and density dimensions are very much greater. Size reduction from the optimal depths range at a particular location is usually manifest as a reduction in blade size and a thinning and shortening of the stipe, especially where the plants are growing in more shallow depths. Similar changes will very often be apparent between optimal growths present at favourable locations and those at adjacent less favourable ones as, for example, at station 19 (Hoyvik transects), where plants off to seaward of the head were much larger and more robust than the small etiolated ones (with smaller blades and thinner stipes) representing optimum growths in the adjacent cove to the south. Reduction in stipe length is often not manifest with increasing depth below the lower fringe of forest density growth. Indeed, plants below the forest will often have even longer stipes than in optimal conditions; in Faroese populations observed (e.g. at station 12) such plants always possessed markedly thinner stipes than did those from the forest depths. In forest depths, plants often had long thick stipes and relatively small or attenuate blades (for example at station 24), although this is not always so since blade size increase over adjacent less favourable greater or lesser depths may be in the same proportion as stipe augmentation (as at stations 30 and 34). Station 34 (west Kunoy) perhaps best exemplifies the idealized sequence of changing size and proportion of Laminaria hyperborea growth with depth (see profile, Fig. 8). Locally variable in shallow depths according to shoreline and substrate configuration, plants most usually were straggly and sparse below Alaria at about 2 m depth. At around 3 m depth (although in places not until 4 or 5 m depth), there ensued considerable rapid increase in abundance, size and density over the depths (3—)4-8 m, the latter usually being the optimum for all characteristics. Not much change thenceforth to 16 m was noted, plants at this latter depth being very large and strong, with virtually unbreakable stipes. Substrate changes over forest depths from about 5 to 16 m, from very steep or vertical rock (to 8 m), followed by rock channels or gullies amongst large boulders on detritus, giving way in turn at about 12 m or so to smaller boulders more closely arranged, with bedrock tongues and sandy channels coming in at 16 m, may well have contributed to slight thinning of forest density at about the 16 m depth, the limit of the detailed dive. More casual observations below that indicated more marked thinning with increasing depth. The effects of artificial available substrata on this usual growth and distribution pattern were noted at a few locations; harbour walls, moles and quays were the most important constructions. The most common overall effect was an increase in homogeneity of growth size and density over the available depth range of artificial substrate. A narrower higher Alaria band giving way much more quickly to dense L. hyperborea growths than occurred on adjacent boulders at similar OPEN SHORES 183 depths away from the harbour wall was also usual, as at station 5 (Sandur, Sandoy; see profile, Fig. 2). Concomitant changes in the pattern of under- and epiphytic flora and fauna are clearly shown by the profile and are treated in detail later; the steep to vertical nature of the substrate and the wash-patterns created probably are also implicated in such changes, since some similar characteristics are manifest on the very steep sides of the outer cleft at Gjdgv (station 31; Fig. 7). Populations at depths greater than laminarian limits Considerable amounts of previous descriptive literature on eastern Atlantic subtidal depth- distribution of organisms indicate algal populations often characterized below laminarian limits by variably-constituted assemblages of principally filamentous red algae. No such assemblages were detected in any of the subtidal stations examined in detail on the Faroes. Certain red filamentous forms are luxuriant and widespread in lesser depths and in other conditions, but these play little or no part in characterizing the deeper populations. Pterosiphonia parasitica, Ceramium rubrum, Audouinella purpurea, Polysiphonia urceolata, Ptilota serrata, and P. plumosa, as examples, are all of importance in subtidal situations, but it will be clear from the profiles and other data presented that they do not occur as assemblages of the kind concerned. Distributional characteristics of all detected ecologically significant individual species (other than major laminarians and fucoids) are summarized in Table 4. Although Polysiphonia urceolata was occasionally (as at stations 12, 22, 35 and 44) detected as substantial tufts on rocks and horizontal surfaces of boulders in detritus or on bedrock, algal populations below the lower limits of significant laminarian growth were commonly characterized by crustose growths, of which the major constituents were Cruoria pellita, Pseudolithoderma extensum, and the Lithothamnia (especially Lithothamnion glaciale and Leptophytum laeve). Locally dense clumps of certain other erect forms occurred on rocks, stones, or boulders at considerable depths. Such clumps never obscured the characterizing contribution by the crusts mentioned. Most widespread erect forms were the species Dilsea carnosa, Delesseria sanguinea (also recorded in Johansen, 1979, as between 10 and 40 m depths), Callophyllis lacinata, Membranoptera alata, Ptilota serrata, Phycodrys rubens, and Plocamium cartilagineum/Callophyllis cristata (station 12), Odonthalia dentata (station 44), and Phyllophora crispa (station 35) were less often noted in depths greater than 20 m, although Odonthalia dentata has been recorded (in J6hansen, 1979) at between 10 and 25 m depths. Not all erect growths that occurred amongst the deeper populations of marine algae were Rhodo- phyta. Desmarestia aculeata (stations 11, 21, 22, 31, 44) and, less often, Desmarestia viridis (stations 22, 44) were frequently noted, but only infrequently fully below the limits of tlie Laminaria hyperborea populations at the same localities. Similarly rarely appearing below the thinning lower areas of L. hyperborea growth were Ulva lactuca (station 44), Fimbrifolium dichotomum (station 12), and Corallina officinalis (stations 29, 31). Rare species such as Brongniartella byssoides (station 44) occasionally occurred in open spaces nearing the L. hyperborea limits. It should be clearly noted that none of the species referred to above demonstrated restriction to these greater depths. All occurred in much shallower subtidal situations; as epiphytes on Laminaria hyperborea and/or other larger species; in shaded pools within the lower intertidal levels of the shore; or in combinations of these other circumstances. Detailed treatments of these other occurrences are presented in other sections and the habitat tolerance limits of individual species are summarized in Table 4. ii. Subtidal: epiphytes and underflora There is everywhere considerable similarity in major constituent species of the underflora and epiphytic flora associated with the Laminaria hyperborea forest and its upper/lower fringes (Price & Tittley, 1978; Hiscock & Mitchell, 1980). This is also true for a comparison of either flora, (a) with erect and crustose species growing below the laminarian limits; (b) with species associated with Alaria/L. saccharina/L. digitata levels; (c) with that of shaded lower-shore pools manifesting subtidal upcarry. 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snyoysip > ~ Se > S BS eS S: ES Ss ee: 3° thalline hyphae below subhymenium in the process of supplementing short cells to the 3 fe) fe) oO 3 3 gS AMY Rosca ‘CfoF oOoIIIAT Ss \ Hoesce SA 10 Steinera radiata subsp. radiata (holotype, BM), part showing compound proper margin composed of excipulum proprium (e) and adjacent fused thal- line margin (tm) formative layer. B Median part with lens-shaped subhymenium. Scale OF, “AOZTOe © Fig. section of mature apothecium, camera lucida drawing of the , (tm) thalline margin. apothecium illustrated in Fig. 10, (e) excipulum proprium Fig. 11 Steinera radiata subsp. radiata, 240 A. M. HENSSEN AND P. W. JAMES SS CR OS pS aa Fig. 12 Thallus anatomy of Steinera species. A S. radiata subsp. radiata, B S. radiata subsp. aucklandica, phycobiont a member of the Rivulariaceae, note pseudoparenchymatous upper cortex. C S. polymor- pha, phycobiont Nostoc, pseudoparenchymatous cortex less well developed. Scale = 20 wm. strands of hyphae in a broad algal zone in the upper part of the thallus (Figs 17B, 19C, 21A-B). The filaments of the phycobiont of S. radiata are more evenly distributed throughout the thallus. They are discrete, often convoluted and sometimes folded back (Fig. 23A—B). In this species haustoria are rather frequent, often associated with moribund or dead algal cells. 5. Fructifications Apothecia Apothecia are abundant in Steinera polymorpha, S. radiata, lessso in S. glaucella, and rare in the sorediate species, S. sorediata. In S. polymorpha the apothecia are aspicilioid, sunken in the thallus, and have no, or a barely discernible, thalline margin (Fig. 18B—C). In S. glaucella the apothecia are also deeply set in the thallus; in this species a coarsely scabrid, more or less THE LICHEN GENUS STEINERA 241 elevated thalline margin is developed (Fig. 16C). The apothecia of S. sorediata and S. radiata are emergent. In the former the thalline margin is sorediate and eroded (Fig. 20B); in S. radiata it is smooth and variable in colour (Figs 11, 22, 24). The amyloid reaction of the ascus is not uniform in the genus. A ring structure is seen in the thickened apex of the ascus of Steinera glaucella and an amyloid cap in the three other species. This variation is similar to that which occurs in species of Coccocarpia (Keuck, 1977). The spores of Steinera radiata are simple (Fig. 13D-E) as in Coccocarpia, Peltularia, and Spilonema, the other genera of the Coccocarpiaceae. In Steinera glaucella the spores are three-septate, ellipsoid with rounded ends (Fig. 13A), whereas those in S. sorediata and S. polymorpha are elongate, fusiform, multiseptate, and + acicular at one or both ends (Fig. 13B-C). In Steinera glaucella and S. sorediata the paraphyses are richly branched for much of their length with occasional anastomoses. In the former the apical cells are elongate and not enlarged, whereas in S. sorediata the upper parts of the paraphyses are somewhat enlarged and sub- moniliform (Fig. 13A—B). The paraphyses in S. polymorpha are simple below, becoming bi- or tri-furcate near the apices; the apical cells in this species are not noticeably swollen (Fig. 13C). In S. radiata the paraphyses are simple for much of their length and only branched near the tips. In subsp. radiata the apices of the paraphyses have a candelabra-like appearance but are not submoniliform, whereas in subsp. aucklandica the paraphyses are mostly simple and the terminal row of 3-4 cells more or less submoniliform (Fig. 13D-E). In all species the epithecium is brown; the pigmentation is external to the apices of the paraphyses. jee ne a Fig. 13 Asci, spores and paraphyses in Steinera species. A S. glaucella, B S. sorediata, C S. polymorpha, D S. radiata subsp. radiata, E S. radiata subsp. aucklandica. Scale = 20 wm. 242 A. M. HENSSEN AND P. W. JAMES Pycnidia Pycnidia have been observed in two species, Steinera polymorpha and S. radiata. In both species the pycnidia are relatively large, the wall is convoluted, and the conidiophores are short-celled. However, in spite of a superficial resemblance they belong to two different pycnidial types; in S. radiata the short conidiophores are aggregated and line the wall, leaving a relatively large, central cavity (Figs 14A, 15A-B) closely resembling the Umbilicaria type (Vobis, 1980: 23, Vobis & Hawksworth, 1981: 252). In S. polymorpha the long, richly branched-anastomosing conidiophores form a filigree arrangement which entirely fills the cavity (Figs. 14B, 15C, 19D), corresponding to the Xanthoria type (Vobis, 1980: 27; Vobis & Hawksworth, 1981: 254); the pycnidia of Peltularia gyrophoroides also belong to the same type but a central cavity is present (Fig. 15D). The conidia are shorter in Steinera polymorpha where they are mainly produced laterally by the conidiogenous cells (Fig. 14B); in S. radiata.they are mainly developed terminally (Fig. 14A). Fig. 14 Type of conidiophores in Steinera species. A S. radiata subsp. aucklandica, short aggregated conidiophores lining the pycnidial wall, conidiogenous cells producing conidia mainly terminally, B S. polymorpha, filigree network of long conidiophores, conidiogenous cells producing conidia mainly laterally. 6. Key to the species 1 + Thallus copiously sorediate, rarely fertile. Apothecia with a sorediate margin. Spores elongate-fusiform, 5—7 septate, 45-57 5-5-7 um. New Zealand ............... iii. S. sorediata (p. 249) — Thallus lacking soredia, usually fertile. Apothecia without a sorediate margin .................... 2 2(1) Spores (5—) 7-septate, elongate-fusiform, 65-75 X5-5-5 wm. Thallus thick, polymorphic, in part radiating, loosely attached to mosses, stones and soil. New Zealand .. ii. S. polymorpha (p. 246) — Spores simple to 3-septate, ellipsoid or subglobose, not exceeding 20 wm in length. Thallus closely affixed to stones and rock, margin always radiate ................ccceeceeeeeeeeeeeeneeeeneeeene a 3(2) Central areoles convex, surface more or less scabrid. Asocarps more or less innate, remaining concave, disc pale brown to brown. Spores (1—) 3-septate, ellipsoid. 19-216-7 um. Rereaelen stand cee nese ok ot acs tna sncesees wyateeaas vetomwoeen aosee cease i. S. glaucella (p. 243) — Central areoles plane, surface smooth. Asocarps emergent to sessile, disc dark brown. Spores simple and with a plasma bridge, less than 40 wminlength ................ cc cce cee eee ee eee eee eee eee es 4 4(3) Spores subglobose or shortly ellipsoid, 8-10-56-5—7-5 wm. Paraphyses branched at the, apices, not submoniliform. New Zealand ....................0.eeeeees iv. S. radiata subsp. radiata (p. 252) — Spores ellipsoid, 12—-14x3—4-5 wm. Paraphyses simple or bifurcate at the apices, upper part submoniliform. Auckland Islands ....................aseseceesees iv. S. radiata subsp. aucklandica (p. 253) 243 THE LICHEN GENUS STEINERA Wy Fig. 15 Types of pycnidia in Steinera and Peltularia. A-B S. radiata subsp. aucklandica (holotype, BM); A Irregularly shaped pycnidium within algal zone, B Upper part in higher magnification showing aggregated conidiophores. C S. polymorpha, part of pycnidium filled with branched conidiophores. D P. gyrophoroides, pycnidium of same type as in S. polymorpha. Scale A, C, D = 50 wm; B = 20 wm. 7. The species i. Steinera glaucella (Tuck.) Dodge in B.A.N.Z. Antarct. Research Exped. 1929-31, Reports B, 7: 66 (1948). Figs. 4A—C, SA-E, 6A-B, 16A-C, 17A-D. Pannaria glaucella Tuck. in Bull. Torrey bot. Club 6: 57 (1875). Type: Kerguelen Island, 1875, Kidder (U.S. Transit Expedition) (FH—holotype). Amphidium molybdoplacum |‘molybdopheum’| Nyl. ex Crombie in J. Bot. Lond., 13: 333 (1875); [‘molybdoplacum’] in J. Linn. Soc. (Bot.) 15: 181 (1876). Steinera molybdoplaca (Nyl. ex Crombie) 244 A. M. HENSSEN AND P. W. JAMES Zahlbr. in Deutsche Stidpolar-Expedition 1901-1903 8: 43 (1906). Type: Kerguelen Island, Swain’s Bay, December 1874, A. E. Eaton (Venus Transit Expedition) (BM—lectotype; BM, H— isolectotypes). Thallus irregular or forming + complete rosettes, to 6 cm diam., closely appressed to the substrate and not easily detached without damage, uniformly pale grey or becoming darker grey towards the centre, sometimes with an olive tinge, margins shortly radiating, + lobate. Marginal lobes flabellate, broadening to 2 mm towards their convex ends and there occasionally notched, separated by deep intersecting fissures with dentate edges; centre of thallus with coarse, angular, predominantly convex, complete or incomplete, areoles derived from secondary rimose crack- ing between the main fissures; entire surface of the thallus roughened, sometimes coarsely scabrid, matt. Ascocarps rather scarce, scattered or sometimes 2 (-—4) contiguous, developed on the uppermost part of the older, inner areoles, 3-8 mm diam., deeply innate at first, eventually more or less level with the surface of the thallus, urceolate; disc dull brown to deep red—brown, smooth or slightly roughened, thalline margin not or slightly elevated, markedly scabrose, sometimes appearing spuriously sorediate. # , ar amma eae.) ‘ ee Fig. 16 A—D Habit photographs of Steinera glaucella (lectotype of S. molybdoplaca, BM). Scale = 2 mm. THE LICHEN GENUS STEINERA 245 Fig. 17 Anatomy in Steinera glaucella (lectotype of S$. molybdoplaca, BM). A—C Sections of thallus; A L.s. of lobe tip, B T.s. of central area, C L.s. in higher magnification. D Ascus with thickened apex (arrow) and spores. Scale A, C, D = 50 um; B = 20 wm. Thallus c. 1000 wm thick, without a distinct upper cortex but older parts of the thallus with a thin, amorphous, semi-translucent, uneven layer overlying a + algal-free layer, 5-40 um thick, of mostly vertically aligned, thin-walled, oval, rounded or slightly cuboidal cells, + uniform in size, 7-10 um. Phycobiont a species of Nostoc, cells 6-8 zm, mainly concentrated in a broad zone c. 60 wm thick, occurring in vertically orientated fascicles between the fan-shaped ascending hyphae, algal cells 15S—18 x5—7 wm. Below the phycobiont zone the hyphae are closely compacted, either vertically aligned as in the central areoles, or + horizontally radiating towards the apices of the marginal lobes, generally interspersed with simple chains of algal cells. Lower 246 A. M. HENSSEN AND P. W. JAMES part of thallus brown—black, composed of periclinal hyphae with blackened walls. Simple or rarely branched rhizoidal hyphae anchor the thallus to the substrate. Thecium 150-180 wm, epithecium more or less pale brown. Asci 120-140 13-16 um, elongate-clavate or sometimes almost cylindrical, apices up to 10 wm thick, with an apical ring structure. Spores mainly biseriately arranged (4-7) —8 per ascus, (1—) 3-septate, ovoid, straight or slightly curved (15—) 19-21 x6~7 (-9) um, thin-walled. Paraphyses distinct, in a gelatinous matrix, richly branched with scattered interconnecting anastomoses, apical cells shortened and somewhat swollen. Excipulum-like structure (formed by formative layer) slightly pigmented, c. 50 wm thick, cupular, composed of small, deeply staining (in lactophenol-cotton blue), nubili- ated, cuboid cells, 5-7 xX 5-6 um; at the outer edge, extending upwards to form the inner part of the overarching thalline margin which contains clusters of algal cells. Subhymenium lens- shaped, up to 200 um thick, of short-celled, vertical hyphae permeated by thicker ascogenous hyphal filaments. Pycnidia not observed. Specimens seen: Kerguelen: Kidder (holotype of Pannaria glaucella); Swain’s Bay, Eaton (type collections of Amphidium molybdoplaca). Observations: Steinera glaucella is distinguished by the rosette-shaped, closely appressed thallus with convex central areoles and more or less radiating marginal lobes (Fig. 16A—B), the urceolate ascocarps with only slightly elevated, scabrid thalline margins (Fig. 16C), and the short, three-septate spores (Figs 13A, 17D). Pannaria glaucella Tuck. was published one month earlier (October 1875) than Amphiloma molybdoplaca Nyl. ex Crombie (November 1875), a fact to which Tuckermann drew attention in a subsequent paper (1877). The holotype on which Tuckermann’s description of Pannaria glaucella is based is a small specimen, barely 1 cm in diameter, with immature apothecia. Although this specimen has been badly damaged by mites and abrasion, the few lobes which remain intact have the characteristic roughness to scabrid surface and internal anatomy, including ‘Collogonidia 0-004-9 mm in diameter, in chains often 4-10’, found in the abundant type material of Steinera molybdoplaca (Nyl. ex Crombie) Zahlbr. The two gatherings are therefore considered to represent the same species, the name S. glaucella taking priority as indicated by Dodge (1948). Even if Steinera glaucella and S. molybdoplaca had been found to be different entities the specific epithet molybdoplaca was rendered nomenclaturally superfluous when published (Art. 63). This is because the earlier name Pannaria glaucella, which should have formed the basionym, is cited in the synonymy of Zahlbruckner’s new combination Steinera molybdoplaca, which was based on Amphidium molybdophaeum which was published a month later. With regard to the difference in spelling, Crombie (1876, loc. cit.) remarks: “This was erroneously b I De 4 printed as molybdopheum in “Journ. Bot.’’’. ii. Steinera polymorpha P. James & Henssen, sp. nov. Figs 7A—D, 12C, 13C, 14B, 15C, 18A—C, 19A-D. Thallus supra solum, muscos lapidesque, plus minusve placodioideus, laxe affixus, ad 5 cm diam., polymorphus, pallide cremeus vel caesiellus, marginibus plus minusve lobatis, crustaceus areolatusque in centro fissus, sorediis nullis praesentibus. Alga ad genus Nostoc pertinens. Apothecia crebra, 0-4-0-9 mm diam., 3-4 saepe confluentia, innata, urceolata, marginibus inconspicuis; discus obscure rufescenti-fuscus, concavus semper. Hymenium usque ad 150 ym; asci 100-110 16-25 um, apice incrassato et amyloideo. Sporae (5—) 7-septatae, elongato-fusiformes, (55—) 65—75 x (4-5—) 5-5-5 (-6:5) um. Pycnidia numerosa, in sectione ad 0-08 mm lata; conidia praecipue terminaliter formata, bacilliformia, 4-5x1-5 wm, con- idiophoris longis, articulatis et anastamosantibus. Typus: New Zealand: South Island, Fiordland, Dusky Sound. In iugo super Cascade Cove, 1000-1200 m, supra solum et lapides muscis vestitos in pratis subalpinis, 10 February 1967, D. J. Galloway (BM— holotypus; CHR—isotypus). THE LICHEN GENUS STEINERA 247 Thallus + placoid, spreading, to 5 cm diam., polymorphic, overgrowing rocks, bryophytes and soil, rather loosely attached to the substrate, margin lobate; marginal lobes radiating, + distinct up to 2 mm wide, contiguous, rarely overlapping, apices rounded, sometimes notched, flabel- late. Upper surface often coarsely scabrose, pale cream to pale blue-grey, older parts conspi- cuously radially fissured, areolate with numerous, anastomosing, coarse ridges and fissures. Apothecia 0-4-0-9 mm diam. when mature, at first urceolate, deeply immersed in the thallus, gradually emergent, thalline margin at first inflexed, later not apparent or only slightly elevated above the disc. Apothecia discrete at first, later becoming 2-4-confluent, sometimes the combined discs dissected by fine to rather coarse fissures; discs dark red—brown, slightly roughened, often shining. Pycnidia numerous, seen as small, rather irregular, dark brown spots on the surface of both young and old areas of the thallus, not easily distinguishable from apothecial primordia without microscopic examination. Thallus c. 1000 «wm thick, upper algal-free zone very irregular in thickness, 70-150 um thick, composed of thin-walled, isodiametric or slightly elongate, closely compacted cells in the upper part, forming a + pseudoparenchymetous layer, of enlarged cells, 7-11-57—-13-5 um, overlaid by a semi-transparent, uneven surface layer of necrotic cells. Phycobiont a species of Nostoc, mainly concentrated in a zone c. 500 um thick, individual cells, 4-5-6 um. Filaments crowded into elongate, vertically orientated fascicles between the fan-like ascending hyphae, hyphal cells 15-23 4-5—-5-5 um. Hyphae at thallus base elongate, closely contiguous and + horizontally Fig. 18 Habit photographs of Steinera polymorpha (holotype, BM). Scale = 2 mm. 248 A.M. HENSSEN AND P. W. JAMES . oth ; D re (Seay : Fig. 19 Anatomy in Steinera polymorpha (holotype, BM). A part of hymenium. B Asci and paraphyses. C L.s. of thallus. D Thallus section with large pycnidium. Scale A, C, D = 50 wm; B = 20 um. % aligned ascending towards the centre of the thallus or horizontally spreading into the marginal parts of the lobes by scattered single algal filaments. Lower surface + brown-black developing dark-walled, rhizoidal, simple hyphae which penetrate the substratum. Thecium c. 150 high, epithecium pale brown, semi-opaque. Asci 100-110 16-25 um, clavate or elongate-clavate, widening towards the apex, apices of ripe asci thickened to up to 5 wm, an amyloid cap but no ring structure present. Spores elongate-fusiform, with one or both ends more or less tapered, (55—) 65-75 x (4-5-) 5-5-5 (-6-5) um, (5—) 7-septate, spore cells equal in length. Paraphyses conspicuous in a gelatinous matrix, simple below, often furcately branched towards the apices, cells of lower part c. 1-7 wm wide, to 2-5 wm towards the apices, terminal cells sometimes collapsed and necrotic. Exciple-like structure of the formative layer pigmented, cupuliform, c. 50 wm thick, composed of small, closely compacted cells with deeply staining contents (in lactophenol-cotton blue), incorporated at the overarching inner edge of the thalline margin in young apothecia; thalline margin not distinct in older ascocarps. Subhymenium up to THE LICHEN GENUS STEINERA 249 150 um high, of closely compacted, short-celled hyphae which include more or less vertically aligned ascogenous hyphae in the upper part. Pycnidia of Xanthoria type (see p. 242), in section large, globose to ellipsoid, up to 600 wm broad and 800 wm wide; conidiophores repeatedly branched to form a filigree-like network inside the pycnidium (Figs. 15C, 19D); conidiogenous cells mainly producing conidia laterally; conidia 4-5 X1-5 um, ellipsoid to slightly rod-shaped (Fig. 14B). Specimen seen: New Zealand: South Island, Fiordland, Dusky Sound, Cascade Cove, D. J. Galloway (type collection). Observations: As indicated by the choice of specific epithet, Steinera polymorpha has a very variable morphology (Fig. 18A—C). Although the centre of the thallus may be considerably contorted, convoluted, and ridged, the marginal lobes and effigurate shape are characteristic for the genus. The entire thallus is deeply and widely fissured. The new species can be easily distinguished from other taxa in the genus by the aggregated aspicilioid apothecia and the relatively long spores. S$. polymorpha is closely related to S. sorediata, which has similar but shorter spores tapering at one or both ends; further differences are the abundantly sorediate thallus, and the formation of a distinct thalline margin around the apothecia. The new species was collected from Chionochloa-dominated subalpine grassland on an exposed west-facing ridge above Cascade Cove. The climate in the region is severe with a high rainfall (up to 90 cm per annum) and often long periods of mist. It occurs on poorly drained, podsolized soil associated with the moss Andreaea rupestris and the lichens Siphula complanata, S. fragilis, and Placopsis spp.; in nearby grassland Cladia inflata and Menegazzia inflata are more or less frequent. iii. Steinera sorediata P. James & Henssen, sp. nov. Figs. 8A—B, 13B, 20A—D, 21A-D. Thallus supra rupes, per solum et bryophyta extendens, usque ad 6 cm diam., irregulatim placodioideus, plus minusve laxe affixus, substramineus vel pallide lilacino griseus, marginibus plus minusve lobatis, crustaceus areolatusque in centro fissus, sorediatus. Soralia numerosa, saepe confluentia, dilute caesia vel lilacina. Alga ad genus Nostoc pertinens. Apothecia rara, 0-4-0-7 mm diam., innata, plus minusve emergescentia, margine paulo elevato, saepe fere ubique sorediato; discus rufo-badius vel brunneus, paulo concavus vel planus. Hymenium ad 220 wm; asci 160-17012-14 wm, apice incrassato et amyloideo. Sporae (3—) 5—7-septatae, aciculari-fusiformes, (37—) 45— (-75) x (S—) 5-5-7 (-8) wm. Pycnidia non visa. Typus: New Zealand: South Island, Otago, Dunedin, Mount Cargill. Prope verticem ad rupes plus minusve nudas, cum Menegazzia circumsorediata, 9 January 1963, P. W. James NZ 2099/2 (BM— holotypus; BM, CHR, MB, US—isotypi). Thallus predominantly placoid, spreading, to 6cm diam., overgrowing rock, soil, and bryophytes, rather loosely attached to the substrate, pale stramineous, pale mauve-grey, or pale grey, margin conspicuously lobate. Lobes radiating, apices and margins slightly raised, con- tiguous except near their ends, + flabellate towards the tips, 1-5—2-5 mm wide, often overlap- ping, radially fissured, surface becoming conspicuously and irregularly radially undulate and corrugate, in places folded, often with radial and some transverse anastomosing fissures and radiating ridges. Surface markedly scabrose-roughened, especially towards the tips of the lobes. Central part of the thallus subcrustose, + continuous with deep, partially interconnected fissures. Soralia laminal, numerous, originating especially from a breakdown of ridges, rounded or more frequently oval or elongate, becoming irregular and more or less confluent, in some specimens soredia + covering the entire upper surface of the thallus. Soralia efflorescent, pale blue-grey or lilac, coarsely granular, occasionally with a few finger-like, coralloid isidia which become partly dissolved in soredia. Apothecia rather rare, widely scattered, 0-4-0-7 mm diam. , at first innate, gradually more or less emergent with a slightly elevated thalline margin when mature; disc bright red-brown to 250 A. M. HENSSEN AND P. W. JAMES & is ’ ~ ett ee € Fig. 20 A-—D Habit photographs of Steinera sorediata. Scale = 2mm. brown, smooth or slightly roughened, slightly concave or plane, margin rather thin, 0-5 mm wide, in part smooth, mostly or entirely granular sorediate. Thallus c. 400 wm thick, with an upper, algal-free zone, c. 50 wm high, composed of + anticlinally organized, + isodiametric cells, 8-12 x5—9 um, thin-walled and colourless, becom- ing collapsed and moribund at the surface to form a very thin, hyaline surface layer. Phycobiont a species of Nostoc, violet—blue, cells 3-5-7 um, filaments concentrated in a zone 200-250 wm high in elongate, + vertically orientated clusters between anticlinal strands of hyphae, hyphal cells thin-walled, 14-20x4-5-5 um. Lowest thallus zone c. 150 um, mainly composed of horizontally aligned, rather compacted, thick-walled hyphae. Scattered, single chains of Nostoc THE LICHEN GENUS STEINERA 251 Fig. 21 Anatomy in Steinera sorediata (holotype, BM). A L.s. of lobe tip. B L.s. of older part. C Development of soredia. D Squash preparation of asci and paraphyses, note the multi-septate spores. Scale A, B, D = 50 wm; C = 20 um. also occur between these hyphae.The hyphae in proximity with the substrate have dark brown walls and tend to elongate into rhizoidal extensions into the substrate. Soredia up to 60 wm, often between 30-50 wm diam., composed of extruded, irregular or rounded clusters of algal cells surrounded by a compact envelope of hyphae with thin-walled, cuboidal or rounded cells. Thecium c. 200 um high, epithecium more or less red—brown, colour contained in the gelatinous matrix surrounding the tips of the paraphyses. Asci 160-170 12-14 um, clavate, broadening toward the apex, apex up to 5 um thick. Spores 8 per ascus, bunched, (37—) 45-57 (-75) x (5—) 5-5-7 (-8) um, (3-) 5—7-septate, acicular—fusiform with one or both ends tapered. Paraphyses distinct in a gelatinous matrix, simple at their base, becoming furcately branched and sparingly anastomosing above, cells rather short, sub-moniliform, enlarged to c. 5 yw at apex. Annular exciple-like structure of the formative layer, poorly developed, c. 45 ym thick, composed of very small, deeply staining periclinally arranged hyphae with cuboidal cells, 3—4 wm in size. Subhymenium up to 300 wm thick, of unorientated anticlinally arranged hyphae with short cells and vertically aligned ascogenous hyphae in the upper part adjacent to the thecium. Thalline margin similar in structure to the sorediate part of the thallus, the cortex dissolved in characteristic soredial clusters. Pycnidia not observed. 252, A. M. HENSSEN AND P. W. JAMES Specimens seen: New Zealand: North Island, Mount Egmont, Tahurangi Bluff, J. K. Bartlett, 14 December 1977 (BM, CHR). South Island, Otago, Dunedin, Mount Cargill, fertile, P. W. James NZ 2099/2 (type collection); Otago, Dunedin, on mossy rocks near the summit of Flagstaff Hill, 580 m, sterile, December 1958, J. Murray 3696 (BM, OTA); Otago, Dunedin, Mount Cargill, near summit in sheltered, rather damp, mossy clefts, 640 m, sterile, January 1959, J. Murray 3791 (BM, OTA); Otago, Dunedin, Mount Cargill, with Placopsis parellina, 670 m, sterile, January 1959, J. Murray 3793 (BM, OTA). Observations: Steinera sorediata is distinguished from all other species of the genus by the presence of soralia. These are abundantly produced in all specimens seen and often cover extensive areas of older parts of the thalli (Fig. 20B—C). In outward appearance S. sorediata might superficially be mistaken for a species of Physconia, but the presence of Nostoc as the phycobiont in the former distinguishes it immediately. The habit of Steinera sorediata is also reminiscent of the cyanophi- lic species Vestergrenopsis isidiata, although this species is smaller in size and has isidia. The presence of multiseptate, + acicular spores suggests that Steinera sorediata may be closely related to S. polymorpha. Even so, there are some differences in their ontogeny and anatomy which suggest that they cannot be considered as a species pair. Variation in the species is confined to the size, distribution, and organization of the marginal lobes and the degree of soredial development. Steinera sorediata is a saxicolous species on rock outcrops in subalpine grassland with Hebe elliptica, Dracophyllum longifolium, Phormium tenax, and Aciphylla spp. Associated lichens are Placopsis parellina, P. perrugosa, P. cribellans, P. gelida, Parmelia signifera, and Menegaz- zia circumsorediata. iv. Steinera radiata P. James & Henssen, sp. nov. subsp. radiata Figs 10A-B, 11, 12A, 13D, 22, 23A, F. Thallus saxicolus, placodioideus, perfecte rosulatus, ad 4 cm diam., pallide griseus vel caesius, arcte appressus, marginibus regulatim lobatis, radiatus in centro crustaceus, regulatim areolato-fissus, sorediis nullis praesentibus. Alga verosimiliter ad familia Rivulariacearum pertinens. Apothecia crebra, 0-5- 0-9 mm diam., innata, gradatim plus minusve sessilia; discus planus vel convexus, obscure vel laete rufo-badius, margine thallino coloribus variis, pallide griseo vel rufo-badio. Hymenium ad 60 wm; asci 50-55 10-15 um, apice incrassato et amyloideo. Sporae curte ellipsoideae vel plus minusve globosae, eseptatae, ponticulo e plasmate mediano, (7-5) 8-10-5x(5-5) 6-5-7:5 wm. Paraphyses cellula apicali ad 7 wm aucta. Typus: New Zealand: South Island, Fiordland Botanical District, Mount Barber. Supra saxa in amne, in asperis subalpinis super Deep Cove, 3 March 1927, G. Einar & Greta Du Rietz 2063: 1 (BM— holotypus; CHR, MB, UPS (Du Rietz)—isotypi). Thallus to 4cm diam., rosette-shaped, very closely appressed, not detachable from the substrate without damage, placoid, radiate, markedly effigurate, the centre becoming areolate—crustose, pale grey to pale blue-grey. Marginal lobes regular, contiguous for their entire length but frequently separated by deep, coarse, radiating fissures, lobe ends slightly flabellate, 0-25—0-5 mm wide, 34 mm in length; central areoles derived from rimose cracking, 0-5-0-9 mm diam., irregular with angular margins, fissuring particularly distinct when dry. Surface overall plane or slightly uneven, matt, sometimes slightly scabrose on the marginal lobes. Apothecia frequent, mostly restricted to the central, crustose area of the thallus, 0-5—0-9 mm diam., at first innate, hemiangiocarpic, slowly becoming emergent when mature, eventually appearing superficial and adnate on the thallus surface, rounded or slightly elliptical, discrete, rarely confluent; disc dull to bright red—brown, convex from the first, remaining so when mature or becoming plane, surface finely roughened. Young apothecia covered by rupturing thalline tissue, proper margin gradually exposed so that in old apothecia it is well developed to + 1 mm thick. THE LICHEN GENUS STEINERA 253 Fig. 22 Habit photograph of Steinera radiata subsp. radiata (holotype, BM). Scale = 2 mm. Thallus c. 450 um thick, upper zone pseudoparenchymatous, 60-90 um thick, cell walls thin, well defined cells 18-25 x 15-20 um, collapsed at the thallus surface to form a thin necrotic layer. Hyphae in algal zone thin-walled, cells 10-15 x6-8 um, algal filaments inspersed between the hyphal strands. Phycobiont a member of the Rivulariaceae, concentrated in a zone of c. 230 wm high, filaments 8-12 um thick, discrete, contorted and folded, + vertically aligned or radiating fan-wise near the lobe ends, intercalary and basal heterocysts present, ends of the filaments not distinctly tapered. Basal zone of thallus c. 140 wm thick, including scattered filaments of the alga, hyphae closely compacted, vertically to horizontally aligned according to their location at the centre or the margin of the thallus respectively, cells 10-164 um, walls thickened, a few rhizoidal hyphae with darkened walls penetrating into the substrate. Thecium c. 60 um tall, epithecium red-brown appearing semi-opaque in thin sections. Asci 50-55 10-15 um, elongate-clavate, broadest towards the apex, apex c. 6 wm thick with an amyloid cap. Spores 8 per ascus, simple, with a very thin and more or less median plasma-bridge but without a true transverse septum, shortly ellipsoid to subglobose, (7-5—) 8-10-5x(5-5—) 6-5-7-5 um. Paraphyses rather sparse, more or less conglutinated in a gelatinous matrix, simple below, irregularly branched above, cells often irregular, apical cells to 7 wm, rounded. Margin compound, annular exciple well developed in old apothecia, to 0-1 mm broad, cells of radiating hyphae to 6-8 mm long. Subhymenium to 150 um high, of short-celled hyphae interspersed by vertically aligned ascogenous hyphae in the upper part. Specimen seen: New Zealand: South Island, Fiordland, Mount Barber, above Deep Cove, G. Einar & Greta Du Rietz 2063:1 (type collection). subsp. aucklandica P. James & Henssen, subsp. nov. Figs 9A-G, 12B, 13E, 14A, 15A-B, 23B-E, 24. A subsp. radiata ascis angustioribus, 55-608-12 um, sporis longioribus ellipsoideis, (9-5—) 12-14 (15) x 3-4-5 (—5) wm, cellulis apicalibus paraphysium maioribus, usque ad 9 wm differt. Pycnidia in sectione 0-2mmlata; conidia praecipueinlateribusformata, bacilliformia, 5 x 24, conidiophorisbrevibus, articulatis. 254 A. M. HENSSEN AND P. W. JAMES ” oe : Nee . : a . ¢ 4 ff r, v be fe pee , 4 Fig. 23. Anatomy in Steinera radiata (both holotypes, BM). A S. radiata subsp. radiata, L.s. of thallus. B S. radiata subsp. aucklandica, L.s. of thallus. C-E S. radiata subsp. aucklandica, squash preparations showing spores and paraphyses; C Thickened apex of the ascus indicated by arrow; D Asci with deformed spores; E Submoniliform paraphyses. F S. radiata subsp. radiata, squash preparation of asci. Scale = 20 um. Typus: New Zealand: Auckland Islands, Auckland Island, Mount Eden. Ad saxa basaltica ferro abundantia in uliginosis inter caespites Chionochlorae antarcticae cum Argopsis megalospora, 320 m, 31 December 1962, P. W. James NZ 858/2 (BM—holotypus; CHR, MB—isotypi). Similar to subsp. radiata in habit and anatomy, differing in the narrower, ellipsoid—clavate asci, 55-60 8-12 wm, and ellipsoid spores (9:5—) 12-14 (-15) 3-4-5 (-5) wm. Paraphyses often simple, less branched at the apices, and apical cells more noticeably swollen, up to 9 wm, than in subsp. radiata. Pycnidia numerous, inapparent, of the Umbilicaria type (see p. 000), 200150 wm, outline rather irregular, convoluted, lined with short-celled conidiophores. Conidiogenous cells mainly producing conidia terminally. Conidia 5x2, rod shaped. Specimens seen: New Zealand: Auckland Islands, Auckland Island, Mount Eden, P. W. James NZ 858/2 (type collection); Auckland Island, Mount Eden, in small declivities on basalt rocks, with Argopsis megalospora, 340 m, 1 January 1963, P. W. James NZ 863/1 (BM). Observations: Steinera radiata is a very distinctive species, forming neat, rosette-shaped thalli with numerous emergent ascocarps, which at first are deeply immersed in the thallus. The species differs from THE LICHEN GENUS STEINERA 235 Fig. 24 Habit photograph of Steinera radiata subsp. aucklandica (holotype. BM). Scale = 2 mm. other taxa in the genus in not having Nostoc as its phycobiont; the identity of the alga is, however, not known for certain but the presence of basal and intercalary heterocysts suggests that it belongs to the Rivulariaceae, even though the distal ends of the filaments are not tapered as is commonly found in this family. In section, the apothecial margin of S. radiata is unusually (for the genus) well organized, and the ascocarps have a tendency toward the horizontal spread (Figs 10A-B, 11) which is characteristic of Coccocarpia and Peltularia. Steinera radiata subsp. aucklandia is closely related to subsp. radiata. The two taxa are only * separated by minor differences in the character of the spores, asci, and paraphyses (Fig. 13D-E). These differences may reflect the long isolation of the two subspecies. Steinera radiata occupies a rather anomalous position in the genus. It differs from the other species in the simple spores, well developed excipulum proprium, low hymenium, lateral expansion of the apothecium, and the different phycobiont, all characters this species has in common with the monotypic genus Peltularia. Nevertheless, the placoid habit and the more or less homoiomerous organization of the thallus justify its inclusion in Steinera. 8. Methods Unless otherwise stated the photographs of anatomical structures were prepared from freezing microtome sections mounted in lactophenol-cotton blue. All measurements are made from either squash preparations or microtome sections mounted in lactophenol-cotton blue. 9. Acknowledgements These studies were supported by a grant from The Deutsche Forschungsgemeinschaft (A.H.) and the Royal Society of New Zealand (P.W.J.). The valuable assistance of Mrs G. Traute (Marburg) in preparing the sections and plates, Miss K. Kavanagh (BM) in providing assistance with the latin diagnoses, Mr J. R. Laundon for discussions on nomenclature, and Miss D. Stephenson for help with typing, is gratefully 256 A. M. HENSSEN AND P. W. JAMES acknowledged. The curators of the herbaria at CHR, H, UPS, and W are thanked for their assistance in locating material. We are indebted to Dr D. J. Galloway for permission to describe Steinera polymorpha from his collections, for his ecological data, and for his comments on the manuscript. 10. References Crombie, J. M. 1875. New lichens from Kerguelen land. J. Bot., Lond. 13: 333-335. Dodge, C. W. 1948. Lichens and lichen parasites. B.A.N.Z. Antarct. Research Exped. 1929-31, Reports B, 7: 1-276. — 1971 [‘1970’]. Lichenological notes on the flora of the Antarctic continent and the subantarctic islands IX—XI. X New taxa and little known species from the alpine areas of New Zealand and the subantarctic islands. Nova Hedwigia 19: 453-490. Henssen, A. 1963. Eine Revision der Flechtenfamilien Lichinaceae and Ephebaceae. Symb. bot. upsal. 18: 1-123. — 1969. Die Entstehung des Thallusrandes bei den Pannariaceen (Lichenes). Ber. dt. bot. Ges. 82: 235-248. — 1974. New or interesting cyanophilic lichens II. Lichenologist 6: 106-111. —— 1975. Ontogenesis of the ascocarp in the lichen genus Steinera. Abstract of the Seventh International Botanical Congress 1: 61. Leningrad. — 1979. Problematik der Gattungsbegrenzung bei den Lichinaceen. Ber. dt. bot. Ges. 92: 483-506. — 1981. Hyphomorpha als Phycobiont in Flechten. Pl. Syst. Evol. 137: 139-143. — & Jahns, J. M. 1973 [‘1974’]. Lichenes, eine Einfiihrung in die Flechtenkunde. Stuttgart. — 1981, in co-operation with Keuck, G., Renner, B., & Vobis, G. The Lecanoralean centrum. Jn D. R. Reynolds (Ed.), Ascomycete Systematics. The Luttrellian Concept: 138-234. New York. Keuck, G. 1977. Ontogenetisch-systematische Studie tiber Erioderma im Vergleich mit anderen cyanophilen Flechtengattungen. Biblthca lich. 6: 1-175. Schwendener, S. 1869. Die Algentypen der Flechtengonidien. Basle. Tuckerman, E. 1875. Lichens of Kerguelen’s land. Bull. Torrey bot. Club 6: 57-59. — 1877. Observationes lichenologicae, no. 4. Observations on North American and other lichens. Proc. Am. Acad. Arts Sci. 12: 166-185. Vobis, G. 1980. Bau and Entwicklung der Flechten-Pycnidien und ihrer Conidien. Biblthca lich. 14: i-v, 1-141. & Hawksworth, D. L. 1981. Conidial lichen-forming fungi. Jn G. T. Cole and B. Kendrick (Eds), The Biology of Conidial Fungi: 245-273. New York, London, & San Francisco. Zahlbruckner, A. 1906. Die Flechten der Deutschen Siidpolar-Expedition 1901-1903. Jn E. von Drygalski (Ed.), Deutsche Stidpolar-Expedition 1901-1903 8: 19-55. Berlin. — 1926. Lichens (Flechten). B. Spezieller Teil. Jn A. Engler and K. Prantl (Eds), Die natiirlichen Pflanzenfamilien 8: 61-270. Leipzig. British Museum (Natural History) Seaweeds of the British Isles Volume I 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 0 565 00781 5 1977 Approx. £12. Titles to be published in Volume 10 Taxonomic studies in the Labiatae tribe Pogostemoneae. By J. 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. 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