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Heish deeds eae) Da nenstre Via tes 323: mak, Sa NS Rees: a Sea ps SRS S See ~ SS ee Coser : pS Ee = a, — Bein ‘. s 21; ae ms, Garesh ee eset ie ay ; et 2 aA ts = a abe SS “ +! . cts Sonat et ytates sc ae Nn & Wi 7989 Bulletin of the _ British Museum (Natural History) The algae of Lightfoot’s Flora scotica Peter S. Dixon Botany series Vol 11 No 1 25 August 1983 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, ete England. is World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.), . seat DS oad } © Trustees of the British Museum (Natural History), 1983 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 11 No 1 pp 1-15 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 25 August 1983 , The algae of Lightfoot’s Flora scotica [F exe Peter S. Dixon |. Department of Ecology and Evolutionary Biology, University of California, Irvine, California : 92717, U.S.A. Ay Contents MNS Ee sss ays eu ain enone dp svsloas Sssadd vias ow shines ae oeuig gens on sae peNwaa ave saaNTavo-ateNte Reached 1 UTES ES eA Gane et Vie eRe RENE a Eye ter UN Tee PRONE TEES Crane eae ees 1 Pegi ais Gt LipnUOol S MELO OL WOM. o, 225.5555 segs. cose snes sty ei xen des ap bec eieeds 2s er sevnionres Zz Py Ucacons OF] iehtloot's NEW algae. isics. access ctascanee tele ve sp esi ercnntyevaet clesedcirubee 4 Ro USVSSMS DUN DUTCUS 20.5 orca ccaas cases trek one dein casa toes sac esata cee cAcaene SeBacen er diliessoomeerrs 4 De COMIELUA CONT AL OSE yah cup clon ts oe oaven a eRe aver culty wales Fee 5 Ber KONI ER VECOPGUINA Peres CAR hs Con Crip ee CUE NIE RR TeCCe Ee Laue eee 5 WC OMIOEVAUUDHGNE a0 cikic ocean sited ee cer cae e aan one one ret na Oe euaanni eee het 5 DS OPEL WIT CQUASELL ONG ee cnccoe teens eee UA i A Ons RE NT: fi De OMT ET UA NOMI OSE: criss enn ores aeet aaEE RTE oun oat SIO IA Lage Ys RUM MES ea aN od Sun ARNE UST D Dy ORFET VO VETUCIN GIG 1500882 cven or dover coc teeeninnctat ah Goa tunnee ade asset Dy wars Beeaneeeanae 8 Be MECIAS CHLGIV IL OUUS 2 Sas acuatlie sate oe vacetunonoteccee ear min dacacece genders werand tee eome. Tae: 8 Pr TCHS ITINGUISs occ ISs cay ayaa tee Nate tnih tas er Na ean Suan ag ae ee anna 8 BO. PCS HETCIDONS 5 cosas 4 sansces sais ing Gane out ORDA PLU be one teak Cae aera aoe ete e 9 he PCUS POLYSCIIOS 25 ce anes Siades aide cbt taut Aaa u adn ctouaeN ata FYerkSiigs a Fis guinkor sate 9 iN gs oh To EW V0) 1 4 Saga Rae ein Der ey eer aE a ea hee nqmrsn ea Rest otS Wyn Vn e Meee Bere re op eer ny A 9 PS PUCUSTEDONS credo seas ieee on cen aw ie sua eaorannd toe cone ate ces aes eees tot tare ce ee oah eco ee 10 BAS VCUSVETUEUIAIUS SoU cr rscae ee sen tech ctvaze see ee ere ean sh ee eee Side Le eCae aie ee e btn 10 Moe OVE COPE ete eas es ee cote aaa ee ee eos ose Ie nee acme ne ees 11 DONOHUE irc Bt BENS PELs CI Ty SO Me atls ARON CNS GO Dal 11 MRE ELAS OE Tel Ps 177 as AR ee OR OE RS ECeR RAR TATU ica Coe oe nr err errr 12 ES CLV TONIANG coco c te erie arte ear tian Weems peiean eek Arar ew tenia uy oneNlee ea hawats 13 PCR MO WIC RCINONES: 66 20 Gone nese cns toes xa trelgu ius vbotim laure anenecateraumetestasuernsueseeuees 13 ROSCOE S osc dois oe. vpctee cau anc cindy sae aeT ohne dae tmcees DUR ena OSaRES tar eek acumutees UnkeaeG 13 Synopsis Lightfoot’s Flora scotica (1777) is of importance as it was the first flora of the northern parts of the British Isles to use Linnaean binomials. Although Lightfoot left no statement of his method of work, deductions can be made from a comparison between his publication and his algal herbarium, facilitating typification of the 18 new binomials for which Lightfoot previded diagnoses. The correct name for the alga currently known as Polyneura gmelinii is P. laciniata (Lightf.) P. Dixon, comb. nov. Introduction The Flora scotica of the Reverend John Lightfoot (1735-1788) is of importance as it was the first flora of the northern parts of the British Isles in which Linnaean binomials were used. The flora treated all groups of plants, and is of considerable importance phycologically because of the 18 species of algae for which new binomials were published. Several of these provide basionyms of names in the current checklist (Parke & Dixon, 1976). The flora represented the results of a tour of Scotland made in 1772 in the company of Thomas Pennant, who has given a detailed account (Pennant, 1776). Pennant encouraged Lightfoot to write Flora scotica, and the work was subsequently printed and published at Pennant’s expense. The date of publication has been the source of some confusion, but Price (1968) has analysed the causes of this and concluded that the flora was published in 1777. It contains 35 plates, of which eight are of algae. Of the algal plates, most were prepared by Sowerby, although one (pl.XX VIII) is attributed to Robertson and another (pl.X XXIII) is anonymous. Bull. Br. Mus. nat. Hist. (Bot.) 11 (1): 1-15 Issued 25 August 1983 y) P. S. DIXON The history of the Lightfoot Herbarium after the death of its collector in 1788 is complicated; Britten (1915) and Dixon (1959) have described the sequence of ownership. As a result of the work of Britten, part of the Lightfoot Herbarium was discovered at the Saffron Walden Museum in 1913, and transferred to the Royal Botanic Gardens, Kew (K) in 1915. The need to locate early algal collections for the preparation of Seaweeds of the British Isles led to a general survey of herbaria in 1957-60, and it was discovered in the course of this that no algae, fungi, or lichens had been involved in the transfer of the Lightfoot Herbarium from Saffron Walden to K. Search had shown in 1957 that there were then about a dozen Lightfoot algal specimens in the general algal herbarium at K, and a further six specimens at the British Museum (Natural History) (BM). Detailed consideration of the history of the Lightfoot Herbarium suggested that the missing portions must still be housed at Saffron Walden Museum and, indeed, the algae were found there in an attic in 1958 (Dixon, 1959). Portions of the Lightfoot Herbarium (comprising the lichens, fungi, and the genus Byssus) are still missing, not having come to light in the searches of 1958. The Lightfoot algae were transferred on permanent loan from Saffron Walden Museum to K in 1958, and subsequently (with other algal material) to BM in 1970 (Brenan & Ross, 1970). In 1791 Goodenough received permission from Queen Charlotte (wife of King George III), the then owner of the Lightfoot Herbarium, to examine it. Finding it ina damaged condition, he suggested that it be looked over by a competent authority and Smith was selected for this task. Smith made extensive use of the Lightfoot Herbarium in his various publications on phanero- gams, but he appears to have paid little attention to the algal sheets as no annotations in his hand have been detected. Goodenough, however, examined some algal portions of the Lightfoot Herbarium in detail, and in his treatment of the genus Fucus (as then understood) made frequent reference to Lightfoot material (Goodenough & Woodward, 1797). Goodenough relabelled many of the folders and rearranged the material of Fucus, but did little or nothing with the genera Conferva and Ulva. It is obvious that Goodenough was permitted to keep specimens (see p. 10, under Fucus repens), and he incorporated them into his personal herbarium, which was transferred to K in 1880. Thus, the few Lightfoot algal specimens prior to 1958, together with an unknown number of Lightfoot specimens of groups other than algae received from Goode- nough, possibly accounted for early opinions that the Lightfoot Herbarium was at K. Following the discovery of the Lightfoot algae at Saffron Walden, each of the original flimsy Lightfoot species folders was placed in a strong manila folder for added safety but no rearrangement was undertaken. Thus, the algae of the Lightfoot Herbarium are in as near original state as possible, apart from the rearrangement of Fucus undertaken by Goodenough. Since the transfer of the Lightfoot algae to K and thence to BM, several authors (Chapman, 1972; Dixon, 1960, 1962; Norton & Burrows, 1969; Prud’homme van Reine, 1972) have typified Lightfoot binomials on the basis of the material now at BM. The algal descriptions given by early authors, such as Hudson and Lightfoot, 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. In the case of Lightfoot, these were not stated explicitly although they can to some extent be deduced. Herbaria are cited by their official abbreviations: 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; K: The Herbarium, Royal Botanic Gardens, Kew; OXF: Fielding-Druce Herbarium, The University, Oxford. Analysis of Lightfoot’s method of work Lightfoot left no statement of his method of work. As with his contemporary Hudson (Irvine & Dixon, 1982), it is possible to analyse the text and deduce the philosophy and practice used. In the case of Lightfoot the situation is simpler than for Hudson because much of the algal ALGAE OF LIGHTFOOT’S FLORA SCOTICA 3 herbarium is intact and comparisons can be made with the published text. A complete catalogue has been prepared of the Lightfoot algal herbarium to facilitate such comparisons. The textual arrangement adopted in Flora scotica was similar to the earlier publications of Linnaeus (1753, 1763, 1767, 1771) and Hudson (1762). For each species, a polynomial diagnosis in Latin was provided, with the specific epithet placed in the margin. In many cases, the Lightfoot polynomial was taken with little or no modification from Linnaeus or Hudson. The diagnosis was followed by a list of synonyms, both pre-Linnaean and binomial. Lightfoot differed from Hudson in that he quoted Linnaean and Hudsonian synonyms exclusively in binomial form, whereas Hudson used the polynomial form most frequently (Irvine & Dixon, 1982). Hudson considered quotation of a synonym with illustration of much value as a means of interpretation, and Lightfoot did this even more extensively, citing previously published illustrations of Morison (1699), Réaumur (1712), Oeder (1766-1771), Gmelin (1768), Gerard (1636), Gunnerus (1766, 1772), Buxbaum (1728), Micheli (1729), and Dillenius (1742). In connection with the last author, Lightfoot (1777: xv) acknowledged that he had consulted the actual herbarium, whereas there is no direct evidence that Hudson had done so (Irvine & Dixon, 1982). Presumably, Lightfoot also examined the Morison herbarium at Oxford (OXF) at the same time, although he does not mention this. One significant difference between the work of Hudson and Lightfoot is that the latter gave extensive discussions of the occurrence, use, and systematic relations of all species in English, rather than Latin, a practice which he justified in the preface (Lightfoot, 1777: xi). Lightfoot gave new binomials to three types of entries: 1. Incases such as Conferva equisetifolia (p. 17), Fucus verticillatus (p. 10), and Ulva laciniata (p. 12), Lightfoot did not cite any synonyms at all, and he regarded these as new species. 2. In cases such as Conferva confragosa (p. 5), Fucus polyschides (p. 9), and Ulva crispa (p. 11), Lightfoot cited one or more pre-Linnaean polynomials in synonymy, so that the Lightfoot binomial was based on a combination of his own material (if, in fact, he had any) and the material on which the pre-Linnaean polynomial was based. Such a binomial validates a species as ‘new’ by modern standards, but Lightfoot did not consider it so because he was not discriminating between pre-Linnaean polynomials and binomials. 3. In cases such as Conferva corallina (p. 5) and Fucus nereideus (p. 9), Lightfoot cited one or more binomials in synonymy, so that his names are superfluous and illegitimate (Art. 63 of Stafleu et al., 1978). Understanding these categories facilitates typification of the Lightfoot binomials. Cataloguing the algal herbarium showed that Lightfoot had specimens from other parts of the British Isles numerically more extensive than those from Scotland. Some of these had been collected personally on one or other of the many tours which Lightfoot made to various parts of the British Isles (cf. Riddelsdell, 1905), while others had been received from correspondents. Very few of the specimens in the Lightfoot Herbarium are dated, and information as to the place of collection is usually scanty. At least half the specimens bear no indication of the place of collection at all, and in many other cases it consists only of an indication of the county from which the specimen was obtained. Dates of collection are also rare. Because of these difficulties it is often not possible to establish whether a specimen was collected on the Scottish tour or, if not, whether the specimen was in Lightfoot’s possession prior to that tour or incorporated after- wards. In the introduction to Flora scotica, Lightfoot acknowledges that he received considerable help in the course of his tour from various Scottish botanists, who contributed information, and, in a few cases, specimens. From the text of Flora scotica, and from annotations on algal specimens in the Lightfoot Herbarium, it would appear that Thomas Yalden ‘a late student of physic at Edinburgh’ was the most prolific contributor of Scottish specimens. The Lightfoot Herbarium contains numerous specimens which are referred to unpublished binomials, particularly from Ellis or Solander. It is not clear whether the specimens in question were received from Ellis or Solander, or merely referred by Lightfoot to an unpublished species 4 P. S. DIXON name emanating from Ellis or Solander. There is no evidence in the form of annotations to suggest that they were identified by either Ellis or Solander. Sir Thomas Frankland, who had worked with Hudson (see Irvine & Dixon, 1982), was probably the most prolific of all Lightfoot’s contributors. Despite the fact that they were contemporaries, there is no evidence of any direct collaboration between Lightfoot and Hudson. For example, there are no specimens annotated by Hudson in the Lightfoot Herbar- ium, although several are annotated by Frankland as having been ‘named by Hudson, from his own mouth’. Moreover, the only references to Flora scotica (Lightfoot, 1777) are in the appendix to the second edition of Hudson’s Flora anglica (Hudson, 1778), and not in the main text. Obviously, Flora scotica was published before Hudson was aware of its contents, by which time the second edition of Flora anglica had been set. A significant discovery arising from this investigation is that certain species treated in Flora scotica apparently do not occur in Scotland. An example of this is Fucus nereideus (see p. 9). In other cases, such as Fucus polyschides (see p. 9), the Scottish origin of the material used in the original treatment is suspect, although absolute proof is lacking. Presumably, Lightfoot was travelling light on his tour of Scotland and collecting relatively little material. He based the inclusion of these entities on memory rather than on a specimen in hand. This would account for some curious discrepancies in geographical distribution noted during the preparation of forthcoming parts of Seaweeds of the British Isles. Typifications of Lightfoot’s new algae The following notes refer to the typification of new binomials coined by Lightfoot, listed alphabetically, initially by genus and then by species. 1. Byssus purpurea Lightfoot (1777: 1000) Lightfoot described Byssus purpurea on the basis of material collected ‘Upon the base of the abbot Mackinnon’s tomb, in the ruin’d abbey at J. Columb-kill’ [= Iona, Argyll, Scotland]. He also cited a reference to an illustration by Micheli (1729), with a query ‘Michel. Gen. p.211.n. 13. tab. 90. f. 2?’. As indicated previously, the fungi, lichens, and the genus Byssus of the Lightfoot Herbarium are still missing, not having been found during the searches at Saffron Walden in 1913 and 1958. Neither has any Lightfoot specimen referred by him to Byssus purpurea been located in any other herbarium. Therefore, as indicated in Dixon & Irvine (1977), typification of B. purpurea Lightf. on the basis of material is not possible at present. Furthermore, the citation of the reference to Micheli (1729) with a query means that B. purpurea Lightf. cannot be typified by his material. Specimens were obtained by various collectors from the type locality within a few years of Lightfoot’s visit. Although subsequent collection from a type locality may or may not represent an accurate re-collection of the original material, the present species does represent a special case as the type locality is unique: Reconstruction of the abbey at Iona during the present century involved re-roofing the chancel and cleaning the interior, which was in ruins at the time of Lightfoot’s visit. The late Dr K. M. Drew visited the cathedral in 1948 in an effort to establish the identity of Byssus purpurea Lightf., but there was then no trace of algae on the tomb, which is in the re-roofed part. She was able to collect material corresponding to Lightfoot’s alga from a wall between the refectory and the cloisters in the still unreconstructed part of the abbey (Drew, personal communication). Byssus purpurea Lightf. is the basionym of Audouinella purpurea (Lightf.) Woelk. This species occurs in various habitats in the British Isles (Dixon & Irvine, 1977) and is extremely. polymorphic. The alga described by Lightfoot represents the low-growing stunted form found on maritime rock above high water: the species develops to much greater size on intertidal rocky shores. ALGAE OF LIGHTFOOT’S FLORA SCOTICA 5 2. Conferva confragosa Lightfoot (1777: 976) The Lightfoot treatment of Conferva confragosa is as follows: ‘CONFERVA filamentis mucosis simplicibus equalibus violaceis. — a Linn@o non descripta. (Dillen. musc. 15. t. 2. f. 4.)’, with the further comment ‘We observ’d it upon the rocks in the waterfalls on the mountain of Goatfield; in the isle of Arran, &c.’ The Lightfoot treatment is based therefore on material personally collected (if not actually preserved) and on the entity described previously by Dillenius (1742). The Lightfoot Herbarium does not have a folder annotated as Conferva confragosa. No Lightfoot specimen so annotated has been found elsewhere in the Herbarium or in any other herbarium. The Dillenian Herbarium at OXF contains a specimen annotated ‘Conferva mucosa confrago- sis rivulis innascens’ which represents the Dillenian material of the entity referred to by Lightfoot. This material can therefore be regarded as lectotype of Conferva confragosa: it was selected as such by Drouet (1968) and referred by him to Microcoleus irriguus (Kiitz.) Drouet, a blue-green alga. 3. Conferva corallina Lightfoot (1777: 988) The Lightfoot treatment of Conferva corallina is as follows: ‘CONFERVA filis geniculatis dichotomis. Syst. nat. edit. 13. p. 818. CONFERVA corallinoides. Sp. pl. 1636. (Dillen. musc. 33. t. 6. f. 36. et ejus varietas minor, t. 6. f. 37. CONFERVA geniculata. Ellis. Philos. Transact. vol, 57. t. 18. fig. f. F. opt.’ Conferva corallina Lightf. is thus a superfluous and illegitimate name, antedated by the two previously-described species cited in the original treatment (Conferva corallinoides Linnaeus, 1753 and Conferva geniculata Ellis, 1767), and it must be typified from the type materials of the ons name. All binomials refer to the alga known currently as Griffithsia corallinoides (L.) atters. 4. Conferva diaphana Lightfoot (1777: 996) Lightfoot described Conferva diaphana as a new species, in his sense of that term, with neither pre-Linnaean nor binomial synonyms cited. He stated that it occurred ‘Upon rocky shores, in basons of water left by the tides, and often adhering to Fucuses’. The Lightfoot Herbarium contains a single folder annotated ‘Conferva diaphana’ in Light- foot’s hand which contains six specimens (or packets containing specimens); no Lightfoot specimens have been discovered elsewhere. From the annotations on these six specimens it would appear that one was collected at Bognor (Sussex), two at Scarborough (Yorkshire) by Frankland, while two are completely unannotated. The sixth specimen is annotated ‘Conferva Diaphana. FI: Caledon: & Ellis & Solander’ in Lightfoot’s hand, but it is not clear whether this annotation implies that this specimen served as the basis for the description of Conferva diaphana in Flora scotica, or whether it is a specimen collected subsequently outside Scotland and merely identified by Lightfoot as belonging to the species which he had previously described. This specimen has been accepted provisionally as the lectotype of C. diaphana Lightf., and a series of illustrations of it is presented (Figs 1 & 2). Conferva diaphana is the basionym of Ceramium diaphanum (Lightf.) Roth, and its attribu- tion has been the subject of great controversy (cf. Du Rietz, 1930; Kylin, 1909; Petersen, 1908, 1911; Sjdstedt, 1928; Svedelius, 1927, 1929). The often polemic literature has resulted from the considerable unrecognized morphological plasticity in the genus Ceramium, and a failure to follow correct typification procedures. The illustrations of the material accepted as the pro- visional lectotype of Conferva diaphana Lightf. should help to resolve problems with respect to the application of that name. Although it is unfortunate that the evidence is not indisputable that the specimen selected was used as the basis for the original diagnosis, that specimen was at least referred by the original author to the entity under discussion. 6 P. S. DIXON 25 um ; ( : ) A Oy p| ‘BeCase RAB! Fig. 1 Provisional lectotype material of Conferva diaphana Lightf. A. Apex. B. Portion of axis, 20 segments behind the apical cell. C. Portion of axis, 80 segments behind the apical cell. bh ocd e / a Q No, 9 SS ; d re . ‘ & e g & jes nage: : : 100 um : DOL ¢ Os ey a (C: AAT ai? " ; a Lae ae r 5 . ‘ 3 ie 006 \) 0 0 i 0 % ) 9 Fig. 2 Provisional lectotype of Conferva diaphana Lightf. A cortical band in a mature portion of an axis. ALGAE OF LIGHTFOOT’S FLORA SCOTICA # 5. Conferva equisetifolia Lightfoot (1777: 984) Lightfoot described Conferva equisetifolia as a new species, in his sense of that term, with neither pre-Linnaean nor binomial synonyms cited. He stated that it occurred ‘In the Frith of Forth, but rare. A specimen was communicated by Mr. Yalden. The same we also found on the coast of Cornwall, and at the rocks call’d the Needles, near the isle of Wight.’ No trace of the Yalden specimen has been found, either in the Lightfoot Herbarium or elsewhere. The Lightfoot Herbarium contains a single folder annotated ‘Conferva equisetifolia’, with several loose specimens and a label on which there is an illustration and a diagnosis written in Lightfoot’s hand. The diagnosis on the label is slightly different from that published, but its presence, together with the rough illustration, suggest that this was the material used by Lightfoot in his treatment. The annotation on the label concludes ‘ad littora Insula Vectis prope rupes Needles dictae. a Dom.Woods Clerico.’ It would seem appropriate to consider this material as being of type status and a lectotype has been selected from it. The material is referable to the alga known currently as Halurus equisetifolius (Lightf.) Kiitz. Although the attribution of Conferva equisetifolia Lightf. has never been questioned by subsequent authors, it is extremely doubtful if a specimen of Halurus equisetifolius could have been collected in the Firth of Forth. Hooker (1821) and Greville (1824) both accepted the occurrence of the species in that area on the basis of Lightfoot’s statement, although strong doubts were expressed by Harvey (1846). Present knowledge indicates that Halurus equisetifo- lius occurs as far north as Argyll on the west coast of Scotland, but that it is absent from east Scotland, having its northern limit in the North Sea in Yorkshire. 6. Conferva nodulosa Lightfoot (1777: 994) The Lightfoot treatment of Conferva nodulosa is as follows: ‘CONFERVA filamentis genicula- tis ramosissimis, articulis nodulosis, fructificationibus tuberculatis lateralibus. -CONFERVA elongata ? Huds. Fl. Angl. 484. n. 25. (Dillen. musc. 35. t. 6. f. 38’. He further states that it occurs ‘On the sea shores frequent, often adhering to Fucuses’, but cites no specific locality. It is therefore evident that Conferva nodulosa Lightf. is based on material observed or collected personally, together with the Dillenian illustration and/or material. The punctuation of Lightfoot’s treatment is such that the query could be interpreted as applying only to Conferva elongata Huds., or to both that and the Dillenian reference, as the original description of C. elongata (Hudson, 1762) cites the same Dillenian reference. As Lightfoot had examined the Dillenian Herbarium while Hudson apparently had not, the most appropriate interpretation of Lightfoot’s treatment, and of his expression of doubt, is that he was using his knowledge of the Dillenian Herbarium to interpret C. elongata Huds. As discussed elsewhere (Irvine & Dixon, 1982) there are grounds for thinking that Hudson included several discordant elements in his initial concept of C. elongata. The Lightfoot Herbarium contains several specimens of C. elongata which confirm the current application of that epithet to a species of Polysiphonia, and one of these is annotated ‘Hudson named this for Mr Frankland at different Times’. The Lightfoot Herbarium contains nine folders annotated as ‘Conferva nodulosa’ in Light- foot’s hand, and a single unannotated folder containing further material identified by Lightfoot. These folders contain numerous specimens which all appear to have been collected in England (Sussex, Hampshire, Dorset, and Cornwall); there is no sign of any specimen indisputably collected in Scotland. No specimens referred by Lightfoot to C. nodulosa have been detected in other herbaria. All specimens are referable to the genus Ceramium and most to the alga known currently as C. rubrum (Huds.) Agardh, a specimen of which has been accepted provisionally as the lectotype of Conferva nodulosa Lightf.; it should be appreciated that this specimen was collected in Sussex, not Scotland. From this it might seem that Conferva nodulosa Lightf. might simply be dismissed as a, synonym of C. rubra Huds., the basionym of Ceramium rubrum (Huds.) Agardh. Conferva rubra Hudson (1762: 486), however, is a superfluous and illegitimate name because of a reference in the synonymy in the protologue to Fucus cartilagineus Linnaeus (1753: 1161), the basionym of Plocamium cartilagineum (L.) P. Dixon. Thus, an alternative name must be found 8 P. S. DIXON for the alga known currently as Ceramium rubrum, one of the best-known binomials used in the red algae. Consequently, accurate typification of taxonomic synonyms published shortly after Conferva rubra Huds. is of great importance. 7. Conferva verticillata Lightfoot (1777: 984) Lightfoot described Conferva verticillata as ‘species non descripta’, i.e. as a new species, with neither pre-Linnaean nor binomial synonyms cited. He stated that it occurred ‘Amongst the sea rocks, in basons of water left by the tides, in the Frith of Forth, and many other places, not unfrequent’. The Lightfoot Herbarium contains several specimens, of which two small packets are particularly relevant; no material has been detected in any other herbarium. The two packets are annotated as follows: (a) ‘my Conf: verticillata Fl: Scot: Examine & compare it with C: spongiosa’, in Lightfoot’s hand. (b) ‘my Conf: spongiosa FI: Scot: I described it as it appear’d thro’ a small Magnifier, but I now think it verticillata’, in Lightfoot’s hand. Both specimens are of the alga known for many years as Cladostephus verticillatus (Lightf.) Agardh, although recently Prud’>homme van Reine (1972) reduced it in status to C. spongiosus f. verticillatus (Lightf.) P. van Reine. The first specimen (a) listed above is selected here as the lectotype of Conferva verticillata Lightf. The lectotypification by Prud’homme van Reine (1972) does not precisely indicate which of the Lightfoot specimens is being selected. 8. Fucus endiviifolius [‘endiviefolius’| Lightfoot (1777: 948) Lightfoot described Fucus endiviifolius as a new species, in his sense of that term, with neither pre-Linnaean nor binomial synonyms cited. He stated that it had been collected ‘In the Frith of Forth, and on the coast of Jona’ and provided an illustration (Lightfoot, 1777: pl.XXXII f, g) prepared by Sowerby. The Lightfoot Herbarium contains two folders labelled by Lightfoot as Fucus endiviifolius, one of which contains two specimens collected by Frankland at Scarborough with no indication of relevance to the original treatment. The second folder contains a single mounted specimen (for which the published illustration is a perfect mirror image) together with a diagnosis and instructions to the artist. These are as follows: ‘The Fructifications are visible near the Margins. Draw this of the natural Size, & put it in a Plate with another small one’. This specimen is clearly the holotype of F. endiviifolius and it is referable to the alga known currently as Cryptopleura ramosa (Huds.) Kylin ex Newton. Because of the excellent illustration there have been no uncertainties as to the attribution of Fucus endiviifolius Lightfoot during the past two centuries. Although it is not possible to be certain whether the specimen was collected at one or other of the localities cited, or from elsewhere, this is ofittle importance, since Cryptopleura ramosa is of widespread occurrence throughout the British Isles. 9. Fucus ligulatus Lightfoot (1777: 946) Lightfoot described Fucus ligulatus as a new species, in his sense of that term, as he was unable to refer it to either a pre-Linnaean polynomial or a binomial name. He stated that it occurred ‘In the Frith of Forth, about New-Haven, and other places, but not common’, and provided an illustration (Lightfoot, 1777: pl.XXIX) prepared by Sowerby. The Lightfoot Herbarium does not now contain any folder annotated as Fucus ligulatus; no relevant material occurs in other folders nor has any Lightfoot material been discovered elsewhere. In the absence of material, F. ligulatus Lightf. must be typified by the illustration mentioned above (cf. Chapman, 1972), which is of the alga known currently as Desmarestia ligulata (Lightf.) Lamouroux. ALGAE OF LIGHTFOOT’S FLORA SCOTICA 9 Nomenclatural studies of the genus Desmarestia (e.g. Pease, 1917, 1920; Setchell & Gardner, 1924, 1925; Chapman, 1972) have failed to appreciate that Fucus ligulatus Lightf. is an illegitimate name because it is a later homonym of F. ligulatus S. Gmelin (1768), which is referable to the alga known today as Calliblepharis jubata (Gooden. & Woodw.) Kitz. The next epithet for the alga known currently as Desmarestia ligulata is herbacea, derived from Fucus herbaceus Hudson (1778: 582). Although initially described by Hudson independently of F. ligulatus Lightf., the synonymy of these two entities was accepted by Hudson (1778, Appendix: 662) thus rendering F. herbaceus superfluous (Art. 63.1). Subsequently, Turner (1809) de- scribed independently another species of Desmarestia under the name Fucus herbaceus which is thus an illegitimate name because it is a later homonym of the illegitimate Fucus herbaceus Hudson. Lamouroux (1813) transferred F. herbaceus Turn. to Desmarestia in his original treatment of that genus so Desmarestia herbacea Lamouroux (1813: 45) must be regarded as a new name (Art. 72, Note 1). In the same way Desmarestia ligulata Lamouroux (1813: 45) must also be regarded as anew name. It is the oldest name available for the alga under discussion, and lectotypified by the illustration of Fucus ligulatus provided by Lightfoot (1777: pl.X XIX). 10. Fucus nereideus Lightfoot (1777: 956) Lightfoot described Fucus nereideus on the basis of material collected ‘In the Frith of Forth and other places, but not common’, but also made reference to two previously-published binomials, F. sericeus S. Gmelin (1768) and F. spinosus S. Gmelin (1768). Fucus nereideus Lightf. is therefore a superfluous and illegitimate name, which must be typified from the Gmelin material of these two species. Although both have been considered frequently to be referable to the genus Gelidium (cf. J. Agardh, 1851), this is unlikely in the case of F. sericeus as there are no species of Gelidium reported from its place of collection (Kamtschatka). Other workers (cf. Harvey, 1849) have regarded F. sericeus as a species of Ptilota, and this is a more likely attribution. However, Gmelin material is not available for further study at the present time. The material in the Lightfoot Herbarium identified as Fucus nerideus is of Pterocladia capillacea (S. Gmelin) Bornet & Thuret, a species restricted in the British Isles to southern and western shores. This indicates once again that the material of the Lightfoot Herbarium could not have originated in the Firth of Forth (Dixon, 1960). 11. Fucus polyschides Lightfoot (1777: 936) Lightfoot described Fucus polyschides on the basis of material collected ‘Upon the sea-shores . . ., but not common, as at J-Columb-kill [= Iona, Argyll, Scotland], &c’ and comments further that ‘We have observ’d the same also in South Britain, upon the coast of Cornwall, where it grows so large that a single specimen is sometimes a load for a man’s shoulders’. He also refers to the illustration by Réaumur (1712) of F. arboreus polyschides (from which, no doubt, the epithet was derived) and the illustration of Fucus palmatus given by Gmelin (1768) ‘sed non omnia synonyma’. Gmelin referred the alga figured and described to F. palmatus L. (= Palmaria palmata (L.) Kuntze), but it is clearly conspecific with that described subsequently by Lightfoot. The Lightfoot Herbarium contains two folders, both of which are annotated in Goodenough’s hand ‘Fucus bulbosus Huds. — [Fucus] polyschides Lightfoot’, each containing a single loose specimen. Associated with one specimen is a label annotated in Lightfoot’s hand ‘Cornwal Stalks F: polyschides.’, while the labels with the other are annotated, also in Lightfoot’s hand: ‘Cornwal a new Species’ ‘My F. polyschides or furbellowed Fucus’. Fucus polyschides is the basionym of Saccorhiza polyschides (Lightf.) Batters. Norton & Burrows (1969) accepted the argument (Dixon, 1960) that not all algae described in Flora scotica were based on Scottish material, and typified Fucus polyschides by the second specimen mentioned above, collected in Cornwall, England. 12. Fucus prolifer Lightfoot (1777: 949) Lightfoot described the entity as ‘F. frondibus submembranaceis dichotomis catenato-proliferis, 10 P. S. DIXON apicibus dilatatis bifidis’ with the questioned attribution to it of two previously-published entities ‘FUCUS crispus ? Syst. nat. edit. 13. p. 815. et Huds. Fl. Ang. p. 472 ?’, together with a reference to an illustration by Buxbaum (1728: pl. 60, fig. 2). The species was said to occur ‘Upon the shores of the western coast, adhering to shells and stones’ and Lightfoot provided an illustration (pl.XXX), prepared by Sowerby, showing a whole plant at natural size and two fragments magnified. No material referred by Lightfoot to Fucus prolifer occurs anywhere in the Lightfoot Herbarium and none has been discovered elsewhere. Lightfoot achieved some notoriety for copying illustrations apparently without acknowledgement (cf. Price, 1968), although the magnified illustrations (which are original) suggest that he must have had access to material of F. prolifer which is now lost. It was suggested (Dixon, 1962) that Fucus prolifer Lightf. should be lectotypified by the Lightfoot illustration, and no additional information has come to light subsequently to change this view. Because of the excellent illustration provided by Lightfoot, there has been little uncertainty over the attribution of Fucus prolifer Lightf. to the alga known variously as Phyllophora epiphylla (Mill.) Batters, P. rubens Grev., or correctly P. crispa (Huds.) P. Dixon (Dixon, 1962). Lightfoot’s F. prolifer is a later homonym of F. prolifer Forskal (1775), which is a species of Caulerpa. 13. Fucus repens Lightfoot (1777: 961) Lightfoot described Fucus repens on the basis of material which ‘grows upon the sea-rocks, which are wash’d by the waves, in many places, particularly near Musselburgh, &c’, and he also refers to a Dillenian illustration (Dillenius, 1742) ‘Dillen. musc. 50. t. 10. f.9. A. B.C. D. bon.’. The Lightfoot Herbarium includes three folders containing several specimens, and a further specimen originating from Lightfoot has been detected at BM-K. One specimen in the Lightfoot Herbarium is enclosed in a packet annotated ‘Fucus repens of Yalden near Musselborough. It seems to be that Tremella described by Dillen: t. 10 f. 9 sed Queere’ in Lightfoot’s hand and ‘Fucus repens I gathered this near Muselborough what do you call it it creeps along on the Stones which it covers’ in an unknown hand, possibly that of Yalden. The Dillenian entity to which reference is made is his ‘Tremella marina caespitosa, segmentis tenuibus’ (Dillenius, 1742: 50). All the specimens identified in the Lightfoot Herbarium as Fucus repens and the Dillenian alga are referable to the species of Catenella which has been known by various names during the present century: C. opuntia (Gooden. & Woodw.) Grev., C. repens (Lightf.) Batters, and correctly, C. caespitosa (With.) L. Irvine. The specimen described above, possibly collected by Yalden, has been selected as the lectotype of Fucus repens Lightt. A Lightfoot specimen of Fucus repens, once at K and now at BM, was received at the former as part of the Goodenough collections. It is associated with a label annotated ‘Her Majesty is very glad that this little Specimen may be of Some use to Dr. Goodenough, and desires him to Keep it. Her Majesty will also give commission to Mr. Agnus to Search for the fucus that is wanting, in case it had got among other things’. As mentioned previously, Goodenough worked on the Lightfoot Herbarium while it was in the possession of Queen Charlotte. It is curious, therefore, that Goodenough & Woodward (1797) should have described as Fucus opuntia the same entity described previously by Lightfoot as F. repens, because, in this case, Goodenough had not only examined the Lightfoot Herbarium but actually had a portion of the Lightfoot material in his possession. 14. Fucus verticillatus Lightfoot (1777: 962) Lightfoot described Fucus verticillatus as a new species, in his sense of that term, with neither pre-Linnaean nor binomial synonyms cited. He stated that ‘We found it sparingly upon the rocks of the Little Isles of Jura’, and provided an illustration (Lightfoot, 1777: pl.XXXI) prepared by Sowerby. The Lightfoot Herbarium contains several specimens, one of which resembles the illustration ALGAE OF LIGHTFOOT’S FLORA SCOTICA 14 (Lightfoot, 1777: pl.XXXI) except that the artist omitted some of the major axes. This specimen is with two other specimens in a folder with a loose label. The specimen is annotated ‘Fucus verticillatus. Fl. Scot.’, while the label reads ‘Fucus (verticillatus) frondibus tubulosis subarticu- latis ramosis ramis verticillatis subulatis setaceo-ligulatis. In this the Branches are verticillate as in Horse-tail or Equisetum. The Fructifications are red Spots in the Substance of the Leaves near their Summits, but these I fear cannot be shown. Let him draw it just as it appears, of the Size of Nature’, both in Lightfoot’s hand. The other specimens in the Lightfoot Herbarium are annotated as having been collected in Yorkshire or Cornwall, but there is no real evidence to indicate that the illustrated specimen was collected in Scotland. As the illustration in Flora scotica is clearly based upon it, although admittedly as a somewhat simplified representation, it would seem best to select this specimen as the lectotype of F. verticillatus. The specimen is of the alga known currently as Chylocladia verticillata (Lightf.) Bliding. One curious aspect is the Goodenough & Woodward (1797) were unable to locate the type material stating (p. 208) ‘We have received specimens of this plant . . . under the name of F. verticillatus of Mr. Lightfoot. It is a matter of great regret to us that we could not meet with his original specimen.’ A later comment (p. 233) states that ‘Mr. Lightfoot’s F. verticillatus unfortunately had been mislaid, so that no opportunity offered of proving that species from his original specimen’. That F. verticillatus Lightf. was the oldest name for the common European species of Chylocladia was not appreciated for many years, the alga being known generally as C. kaliformis (Gooden. & Woodw.) Hook. The priority was first accepted by Bliding (1928), and again independently by Papenfuss (1947). 15. Ulva cornuta Lightfoot (1777: 972) The Lightfoot treatment of Ulva cornuta is as follows ‘ULVA subrigida cornuta terrestris. — (Dillen. musc. 52. t. 10. f. 13.)’. He further comments ‘It is about an inch long, sometimes two or three; divided deeply, without order, into various acute horned segments; the edges turn’d inwards, otherwise a flat membrane. It is of a pale-green color, and substance somewhat rigid’. The diagnosis given by Lightfoot differs somewhat from that given by Dillenius (1742) which is ‘Tremella terrestris cornuta’. The Lightfoot Herbarium does not now contain any folder annotated as Ulva cornuta and no relevant specimen has been found in any other herbarium. A specimen in the folder annotated ‘Ulva montana’ in Lightfoot’s hand may be relevant, however. This consists of a minute fragment of material in a packet annotated ‘No 1 this I had from Scotland for the Tremella verrucosa. I am doubtful Please to dilute it and try what you can make of it’, in an unknown hand, and ‘Ulva cornuta. FI: Scot: but possibly may be a Sponge or Zoophyte’, in Lightfoot’s hand. The Lightfoot annotation suggests that this is a specimen received after the publication of Flora Scotica, and not part of the material on which the original treatment was based. Identification of this material proved to be impossible. The material in the Dillenian Herbarium at OXF referred to by Lightfoot is also unidenti- fiable. It consists of a scrap of material which Batters (in Druce & Vines, 1907) interpreted as ‘possibly a fungus’; personal investigation suggested that Batters’ interpretation was probably correct. Thus, one element cited by lightfoot in the protologue of Ulva cornuta is unidentifiable, as is also the specimen now in the Lightfoot Herbarium which may or may not be the material referred to by Lightfoot. Neither of the specimens bears any close resemblance to the comments made by Lightfoot, cited above, which if anything suggest that Ulva cornuta Lightf. could have been a lichen. It is possible that after the publication of Flora scotica the original material of U. cornuta was moved by Lightfoot within his herbarium from the algae to the lichens, which are still missing. 16. Ulva crispa Lightfoot (1777: 972) Lightfoot described Ulva crispa as follows: ‘ULVA tenera crispa terrestris. — Raii synop. p. 64. 12 P. S. DIXON n. 12. (Dillen, musc. 52. t. 10. f. 12. A. B. C. D.)’, with the further comment ‘This is found lying upon the ground in shady places, at the foot of walls and houses’. Lightfoot’s treatment was based therefore on his own material and on the entities described previously in Ray (1724) (which was in fact prepared by Dillenius) and by Dillenius (1742). The Lightfoot diagnosis differs slightly from the diagnoses of both Ray (‘Ulva terrestris tenerrima, viridis crispa’ and Dillenius (‘Tremella terrestris tenera, crispa’). The Lightfoot Herbarium does not now contain any folder annotated as Ulva crispa. A specimen which appears to be the original Lightfoot material of U. crispa occurs in the Herbarium in the folder annotated Ulva montana. This specimen is enclosed in a packet annotated ‘No. 22. In moist shady places on the North side of Walls where no Grass grows, generally’, in an unknown hand and ‘Ulva terrestris tenerrima viridis crispa. R. Syn: 64. not mentioned by Linn: or Huds: This of Ray may be no other perhaps than varietas 8 of Ulva Lactuca. Hud:’, in Lightfoot’s hand. On the reverse of the packet is the annotation ‘This is a thin membranaceous substance, & I think it mostly is where people Pisses agst. Walls.’, in an unknown hand. This material is of the alga known currently as Prasiola crispa (Lightf.) Kitz. No other material of Ulva crispa originating from Lightfoot is known to exist elsewhere at the present time. Examination of the Dillenian Herbarium at OXF reveals that there is a relevant specimen filed in the herbarium of the Historia muscorum. This was identified by Batters (in Druce & Vines, 1907) as Prasiola crispa and personal examination confirms this determination. From the original Lightfoot treatment, and from the material in the Lightfoot Herbarium, it would appear that Ulva crispa was based principally on Lightfoot’s own material, even though it is known that Lightfoot had consulted the Dillenian Herbarium. The slight changes in diagnosis between Lightfoot’s treatment and those of Ray and Dillenius also support this view. Conse- quently, the specimen in the Lightfoot Herbarium has been selected as the lectotype of Ulva crispa. 17. Ulva laciniata Lightfoot (1777: 974) Lightfoot described Ulva laciniata as a new species, in his sense of that term, with neither pre-Linnaean nor binomial synonyms cited. He stated that ‘We gathered it upon the coast of Iona’. The Lightfoot Herbarium contains one folder annotated ‘Ulva laciniata’ in Lightfoot’s hand which contains a single specimen and a separate label. The label is annotated ‘Ulva laciniata. I-columb-kill.’, in Lightfoot’s hand. The specimen bears a lengthy annotation in Lightfoot’s hand, with instructions to his artist, as follows: ‘Ulva (laciniata) frondibus planis purpureis, apice dilatitatis laciniatis undulatis. With a Microscope you may see the Seeds lodged in the Substance of the Membrane near the Edges, appearing like fine Grains of a red Powder. These I fear cannot be express’d. Draw the Plant of the Size of Nature just as it appears.’ This specimen bears a close resemblance to the illustration published by Lightfoot (1777: pl.XXXIII), prepared by an unknown artist. No relevant material has been detected in any other herbarium. The specimen on which the published illustration was based is considered the holotype of Ulva laciniata Lightf. The identity of Ulva laciniata Lightf. has been the subject of much confusion. It appears that Hudson (1778) was the principal cause in that in the appendix to the second edition of Flora anglica he made the comment (Hudson, 1778: 652) ‘Obs. an varietas Ulvae umbilicalis’. Subsequently, C. Agardh (1824: 190) listed Porphyra laciniata (Lightf.) Agardh as the first species of his newly created genus Porphyra. Later authors followed this attribution without question until Drew (1955: 3) suggested that Ulva laciniata might be a member of the Delesseriaceae, and rejected its application in Porphyra. Following the discovery and initial examination of the alga in the Lightfoot Herbarium the opinion expressed by Drew was confirmed (Dixon, 1959). Detailed examination of the type material of Ulva laciniata Lightf. showed that it belongs to the alga known as ‘Nitophyllum gmelinii Grev.’ or ‘Polyneura gmelinii (Lamouroux) Kylin’, the ALGAE OF LIGHTFOOT’S FLORA SCOTICA 13 basionym of which is Delesseria gmelinii Lamouroux (1813: 124). Since Lamouroux cites Fucus palmetta Gmelin (1768) as asynonym, Delesseria gmeliniiis illegitimate, having the same type as the former (see Silva, 1952, and Guiry, 1977). However, the species described by Gmelin has been assigned elsewhere, so that the alga which has been called Nitophyllum or Polyneura gmelinii is without a name. Ulva laciniata Lightf. provides an epithet for the taxon, and its correct name therefore becomes Polyneura laciniata (Lightfoot) P. Dixon, comb. nov. (= Ulva laciniata Lightfoot 1777: 974). 18. Ulva montana Lightfoot (1777: 973) Lightfoot described Ulva montana as ‘ULVA plana coriacea terrestris sanguinea’ as a new species in his sense of that term, with neither pre-Linnaean nor binomial synonyms quoted. He stated that it occurred in various localities: ‘It grows upon the ground amongst the grass and moss on the sides of the mountains in the isle of Skye, and in Ross-shire on the side of a mountain call’d Cail-veg, between Loch-broom and Lead-beg, on the western coast, and doubtless in many other such-like places’. The Lightfoot Herbarium contains a folder annotated Ulva montana in Lightfoot’s hand which contains clumps of material, a label, and two packets which are not relevant to U. montana (see Ulva crispa and Ulva cornuta). The label is annotated ‘Mountain Dulse Skye. Ulva montana. Fl. Cal.’ in Lightfoot’s hand: the specimen has been shown to Dr Francis Drouet and has been identified by him as Anacystis montana (Lightf.) Drouet & Daily. The publication in which Ulva montana Lightf. was transferred to Anacystis (Drouet & Daily, 1952) made no mention of type material, although the subsequent, more detailed, treatment (Drouet & Daily, 1956) indicated that the type (of unspecified status) occurred at K, prior to the discovery of the Lightfoot Herbarium at Saffron Walden and its transfer to K. Search of the BM-K collections has failed to find the material referred to by Drouet & Daily (1956). Under these circumstances, it would seem best to regard the specimen in the Lightfoot Herbarium as the lectotype of Ulva montana Lighttf. Acknowledgements I wish to thank Mrs L. M. Irvine for helpful discussions, and Dr D. M. John and Mrs J. Moore for assistance in the search for Lightfoot material of Ulva montana at BM. 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Ceramium-studies. I. Remarks on Danish species of Ceramium. Bot. Tidsskr. 31: 97-105. Price, J. H. 1968. Bibliographic notes on works concerning the algae II. The Flora Scotica of John Lightfoot (1777; 1789). J. Soc. Biblphy nat. Hist. 5: 57-67. Prud’homme van Reine, W. F. 1972. Notes on Sphacelariales (Phaeophyceae) II. On the identity of Cladostephus setaceus Suhr and remarks on European Cladostephus. Blumea 20: 139-144. Ray, J. 1724. Synopsis methodica stirpium britannicarum . . . 3rd ed. [xvi] +482+[30] pp. Londini. Réaumur, R. A. F. 1712. Sur les fleurs et les graines de diverses plantes marines. Hist. Acad. r. Sci. Mem. Math. Phys. 1712(2): 21-44. Riddelsdell, H. J. 1905. Lightfoot’s visit to Wales in 1773. J. Bot., Lond. 43: 290-307. Setchell, W. A. & Gardner, N. L. 1924. Phycological contributions, VII. Univ. Calif. Publs Bot. 13: 1-13. — & 1925. The marine algae of the Pacific coast of North America. Part III. Melanophyceae. Univ. Calif. Publs Bot. 8: 383-898. 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: 241-323. Sjéstedt, L. G. 1928. Review of some dubious Swedish Ceramium types, their classification and ecology. Acta Univ. lund. I1, 23(12): 1-52. ALGAE OF LIGHTFOOT’S FLORA SCOTICA 15 Stafleu, F. A. et al. (Ed.) 1978. International code of botanical nomenclature . . . xiv+457 pp. [Regnum veg. 97]. Utrecht. Svedelius, N. E. 1927. The seasonal alternation of generations of Ceramium corticatulum in the Baltic. A contribution to the periodicity and ecology of the marine algae. Nova Acta R. Soc. Scient. upsal. Vol. Extr. Ord. (15): 1-28. 1929. The seasonal alternation of generations of Ceramium corticatulum. Some anticritical remarks. Svensk bot. Tidskr. 23: 366-387. Turner, D. 1809. Fuci. . . 2. [2]+162+[2] pp. London. ee eS eC <=> ell ig I El he ale 6 tee al ee ae ee — rir ote ry j f. , 4 i 7 z “i OS ed ig ye eee ne LON a eee rene British Museum (Natural History) 1881-1981 Centenary Publications 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. In the companion volume, The Evolving Biosphere, museum scientists have chosen an evolutionary concept—speciation, coevolution, biogeography etc. and related this to the group of animals or plants in which they are specialising. Thus beetles and birds exemplify sympatric and allopatric speciation, butterflies mimicry and certain fishes explosive evolution. In both volumes the text is supplemented by over one hundred specially commissioned pieces of two-colour artwork. These two books will be invaluable to all sixth-form and undergraduate biology and geology students. The Evolving Earth: 276 X 219 mm, 280pp, 138 line illustrations, 42 halftones The Evolving Biosphere: 276 X 219 mm, approx. 320pp, 133 line illustrations Published: May 1981 Co-published by the British Museum (Natural History), London and Cambridge University Press, Cambridge. Titles to be published in Volume 11 The algae of Lightfoot’s Flora scotica. By Peter S. Dixon. A taxonomic study of the lichen genus Micarea in Europe. By B. J. Coppins. The hepatics of Sierra Leone and Ghana. By E. W. Jones and A. J. Harrington. Studies in the Corallinaceae with special reference to Fosliella and Pneophyllum in the British Isles. By Y. M. Chamberlain. Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, Dorchester Cee oe : Bulletin of the _ British Museum (Natural History) _ A taxonomic study of the lichen genus Micarea in Europe Brian John Coppins : Botany series Vol 11 No 2 29 September 1983 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), 1983 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 if vs : RB? ; Ki f 7 O COM dA : BRE | £7 9CF 1983 As RR prireeres Nae fo ISSN 0068-2292 Ny. 4/4 “SALES oF rice oe, OW Botany series g; rato we Vol 11 No 2 pp 17-214 Ry ; USA tere) So British Museum (Natural History)) Cromwell Road eager ee ee Issued 29 September 1983 A taxonomic study of the lichen genus Micarea in Europe Brian John Coppins \, \ Royal Botanic Garden, Inverleith Row, Edinburgh, EH3 5LR Contents a ee Nae sal let ASN TPCT EG aT Tine Oy ie A le TOS IER Se Ak tr SER ORB Sp ORE AN ROSS 18 RuINLORCay PACK REOUNG {cca straits snse ra5esc.ssniitycd 1 Remetesne caste boeciveesanee ere neviae veue eeu 19 MECMIE ACC AE coc tec crc cenage recs oacae occ eet cc tetech pip tacuwaeehnctoree toate oe noe ei eee 19 OLA dds HESAOR Rare oe eA Pee EAMETS TS AY CCCORT PETES Seo UTC COLIN A te ee ER RED 20 tg EA Set ark Ara ba Sy (nt op ee SE MeN at PROS AEP eg Wey Wane Sup Nncen atin ee 20 PS ee cates cee a a etter Suen sure eed ac eta De asin atg RR pr aaL sec einrs dtaneataseeent 2h BI SUUCIES or ye trek coe Lay nies oral costa ooo cd cna ae aR a ool eda Bae FETS 21 BABIN WRICTOSCODY 153 cock figlen Merce ac cok anion be Uk case ee een acme cin ER AUTOS KEE G Ss oes acoA 21 PRUE AA VOT CHLOMALORE AD AY. oot st ys 43 csv eae ose ie Oa pa cenk Sree aac ate stata oan EY Zi MINOTAUR Coun tc sp ent corey eater aeec a ReC ere ae e gine pom ue Raa a eee sohocen trae ree es 21 ook 2) 10s 81-4 JEN Wi EER AERA HAA LEER AM ne ST rh pO ne TAR SAE Sep eh ep OPE ds eae ei TAR 22 CLL) Se ke eee eae BROS iA Antenne fear toe Ae Mnary Cee Ener MORE Sing ARMED OA MRO tafe bh Ze PEVEOUION IS atic coca nate ene les tonite atau deg a eid kat ROO CE rete scanner nts 25 ROOM AIOGIA Syn rce cee eeretia oy scene ues eee he ee nach aor cena Te Re ee ae 25 PLDOINCCIAT EXLCTMATEALUTOS os ocois vehiaws nvnass ont cresoee eens saad na ns meseestn yee ee Maes 26 PIO A INTENNG EOALOLCS itech rs era nat yts.619 ce, pap Aey borne seas done Sareea hae vee 30 Pepe (COMOIALS AUS) aac sess earn? cles rerieta) na eenewaies os ax siontate aa eesk rien 63 SEDGE 29.2 o ecto guceantosc11 sama e see APCL SL Atd Com akeah pote ORTaT PIReN ag de Lette boas ae 84 MAGE TE SUDSEANCOS hacer none ian in cae Cite een coe Vee cier i (4a thats aw eI Rd ad 84 BEL CR RM reed Te EER ECR COTE ETT BURNET RC pene eee EE RICE EY Reine Peer eres 87 TOMOU Soares cg ek hae ds SoS ee nd tosh otte SU DNEN ER coe anos Nasu AU ne oa) de 88 RSE NO TU NDILALS 5 tert case oct ccc ahalaeg eee coe ee eee ne Tae maar ee eee aren 88 PRSCINE NADIA Soest e trey ioe cte demain cctuehen irate aaNet ee ee ean A own ei acre 90 Crs alas! 1h (a Male ye net seme ie LALOR Eto ene WRU ERRORS GNIS Ragen tect nnn saa Meese MACE Pe ee aNE OT to 93 | Ur a DNR Oi trees Aah aah Pet ete ESRD DEPED Ne Were ene pan ee BEAT ete pee ar ena eMC Tae 93 | TCs) «a eae a eel tue Ren CLG RER GS nines mae TEED AEE reenter TRC rn ee ere tro R ET ee cee ae 94 NRE lex seria verse ane tg eA rata day euae on ada meu snr delete cand tay poets ene 96 ME POUR OTC aioe Noe ae Seda weenie ud Sali AN Seen eee A asdceaice saneeh 96 MELLEL DONCTIC COMSICE TAN OUS 55 carers saat, ysecceans eee aha aa oe ds sO ete ed RIE APOTIOTICCONGKMET AHIOUIS 52045 5 1h vege cris usa au Abaha se sub dd gSeee- osey halen tecanaan terahenet? 98 FRIDARENCTIC COMSIGCELANONS: «caccoas: srsqcascuydareeecs cre Mopsuenaa ail 4 ceddevereny oes eomeueanee: 99 a eo, [A ae CP eee CEE ES SONA ae Rarer Ter erin na Ferre er. Cr er red TERN Aref 100 ATMO 1G RE Vere WICHUNCAUONS oxociyficacnian iia onere meaner nee cna ee eee eres 100 WEY TS EMO AIS SDOIES <5 conrss ckeah vas ee eee a yaees ua Tana eee Tan we ea tetunnnccor een dies 101 Key to European species that may occur without apothecia..................scceeee sence ees 107 SMR EMROS SSRN ieee res res tite aie ed MEd A OLIN gh ome bar ine mae tbs enead daliceebe es scene 108 Br RUC ed Naieae ck cee cae Ripe PERT eet rae eh ane et anien Se way seit ears peede tes 108 INE IGOOSIF ES cal pei Tig es Scns ets Sets we ceive oe eesigts Me navn ndatyes okaes7 se eure em ee 110 ie WEAR THA pein iat du es dalek Gi Ses kU Os 14 OC teu Daly PSE SE MORN PIER GERS OSL eT Mr ah Neate 112 CPD) RE np 1, 1 RO ats ot ae ae hae nen Oe Pree Meer eee eMart rye any NOP Eee err Tet 114 ad SY Ss Vr Tale ee ee GA MEY PN OODLE PLES Pee Tr IE EENE A eR ETOP CES TE 117 ONY RL 2007 COAL lie PTS eh a eA OP ne heey fee Oe yee nery He nnLS 118 RAIS CUACPEOE saccade} asain ticddsiens 224 ae dae soos dower cea ta toncd i eaoe beth ge Seee 121 BE COAL GRIN occa ae Gari an oa 8 Ge aie Koi eee eee SERIO Uh biehy'eneaa aoe en EMER 124 SEAT CT ONSIPIOR sia c FL 4s vas Ss cg acV oi eou Meosnes sone eeaae e RUSKIN TTR SH Ge wenn ces sea bene Uae 125 UES SPOT Te Sea eer SOAS RCRD AEC Beat Roti, ete eatin Te eanertiy PPAR Pr TONS Er Teter ery eores 126 Bull. Br. Mus. nat. Hist. (Bot.) 11 (2): 17-214 Issued 29 September 1983 18 BRIAN JOHN COPPINS DIE CTCF ONG 0c ONS a ach piucttnn vdeeip Oe GTR Ao es Sova thew si vedadoy rok e sd eed Tee Hee TEO OT 127 TD FIVE: CLC TILSEO as, Seed cs oath ale ccteonn aah cea ua oo taaasanhan eutade eas be eee ene TEAR Vie ne 131 5 ae So 7771 7) REM AeS eReRO PP aa ner Prep nmney iN ye Tera Rone Peart Swain Joe URE 134 1G, HM SIGDUNOSONG clio 2c anchors tice ssoneoooen tides Gaok arate ORE TiAak Gate et 134 i BE OR LETT Te 1! RNS Roi Ren ee TeON CATR OSE RANE Pe mEL IS On COCHIN? J MOREE OME Sree 135 AGS OTICTOSSAIG Or eas iced peas FU, Ste OORT ous CRIN ed Be 137 1 RRR) Be TT TAS: IRE ENA RCE ARETE ED CETTE Peete TER TRUER RS Amn AILS BRS MDDS at ra! ese BIO E 138 Ah BI sald hg 01.17 Rae Nr Ore nat er mPa BN. noire Pas ARR SOE II Fina are rk Re ee 140 UD ME ONTO ce Sic osapieeusticd diese bsues tiie caemubedn deh ee Ts ake sce eons tin eee ee Lea 142 BA LI LEEATOUL oie Barn tes ee ee sla SRR elt Teel Te ia 147 PANY! ES TCT) 177 RARER ROpRRe ee ECR PEET] ater try Pete are SAE ER Tae Aer ae Ok EOC MR ols Own ot 148 PODS OE elt 1 PE RRA EID PE NPE PE ENE SEER AE ERC NV BAPE «PEO REE Ne] OE nee NR 150 a Sat NE IMCUIOWLG aos hice. eo nt ie hed foe ce em ees SAT eh Coed dasa aeiene 154 PINS) Bp To] ils bd * Qe oe ESE PPAR IATA RE REO RO Lie eC OPE eae AM AEN ap Se aE Se 155 ZIM IGIONOOOUE socaioenes ys oasrs ba det este Sey ee TR a 156 OS IVE THISEL Ge eH os fence hac ilc e p0 a ak a ae EES Soc oga a Kane bes UN Oe 158 LET cm WVENTIMENT AL oot cic ban cco doe. neon eeu nete ok OER REL Hee RPT TEC ed ees donee 160 ee AE TURN TLOCONIIG 5 juianie Sergi Sho, olex tee Ronn cae nies Shraneee ieee oa oi Ran octey Sei ees eens 161 Doe WE: TQ OLB a ioc crosaiemaa tinny inc sai mmepsauh wie Grace lave pire cwinke eutend pone cada emne eae 163 SS PIM TESCO ONE ood ae ea 2a Soa acannon ids Whee aaa 165 Bde WE SOUV ACER 85 oe os caesar ae ea ete dace eae ak, enna ee cava 167 Ly AN MOSIDONSIE eas ie Se tse che as adda asa BERGE Tew ieee ob chase La tad dda nae remeca giana 169 Doe Ms POUOCOIID Bicdeaais cise ss tata dceoug en Gannon MeO TaREE ah ae Pea eRw eRe 169 SH AVE IT ASULA ens 5 cakeng selvig sc teen pre tuwiii oid Ties aadcsrete sacea ane eC ALS pokes raiaeedl ead 173 DOs VAD VOMGIOPNON EG: Kaas oeliccntns teen Mis oh a ick ah wees eon aad een Cos bc hoeete Ngataee 179 SOS DA TRO UGPONG 2.51552 neve ni 1s press ener snwcon ade feast fee Vibdias lah ctres euesasn.s 181 Dl eda CR TEDLAND (yr, Sed Saath er aac tad tase Ciel ce cae pe lah ary on Ve 182 DOr WV SUDICDTOSU Rc. serie oe hora an eclods os ending bios sar atoeea en ICOM RGR ER Lo 182 IVs SULDFULOPN: bs be os a8 gas oars oh wee ace acces nome aiagmocuae san wied ob seus e wae aTER eee Leal 183 PUI SU UI OIGSCOIES ne tesla biita Ss ee ee erate ene ee aE 185 BL OVE OV IV ICOLG cuea ade shone ane gohasdl seg stor Nanaimo teat aE eee 186 Ee cE SV IUOINCOLAGS Scions ook peta tanner ndeee, ryan aan eee OCP ree Rcanicn hetioanhav kee SOI 188 AS AIA OPIATES cree Se ae oe a eS es ald epee CORO A ete duseak HemoCue edldde ah Bene coe, Sa 190 Mae IVE SEO OTCUN GLC lec pre rn tc a ES SEN TRINA SLE te aaa Pay Un oe cope 192 bs ate ANF g 705.11 Weep a MON EMO EE ONT Win chee ME Pa GREET Urs cate cD Rae's PARTS ny AP ence eR Cres 194 | Bp. (ej | Fra Pra eb, C: Be ren Ca ne ERED SN BRD Se NEBR RES COME EEE oct Nn koe a RSE A Re 196 INGGK TO EXSICCATAC elaine rk ae cet px wee da Rass one sak ae See SaeN es COCK Se Reed 203 PLEKNOW LC CREM ICIS sc, coy pose nd visas peed ened So eres cas ae pe ra, gee een ia 206 | Fos a) 12 EE OR a ote DR Ane a Ar UREN AOS Neen PTET as ROR D NIU GAN AS Cp LY dre POMP MULT MERRT UA, sf 207 TION oes) eae ces Sk Cevamtcos phe toe SET Coe LT A er aN 210 Synopsis A taxonomic revision is presented for the lichen genus Micarea in Europe, with special emphasis on those species occurring in the British Isles. In brief, this genus is here circumscribed to include most crustose lichens with lecideine (biatorine) apothecia, a poorly defined excipulum (sometimes absent altogether) of radiating paraphysis-like hyphae, a non-granular epithecium, Lecanora-type, 8-spored asci, simple to multiseptate, hyaline ascospores, and a ‘grass-green’ phycobiont usually of the so-called ‘micareoid’ type. Comparisons and possible relationships with similar genera are discussed. Noteworthy discoveries made during this study include the finding of cephalodia in three species, dimorphic paraphyses in several species, and a wide array of anamorphic forms, with three species each found to have three conidial states. Type studies have been made for nearly all names known, as well as those suspected, to be referable to Micarea in its present, wider concept. Forty-five species are recognised, of which 31 are confirmed from Britain. Seven species are new to science: Micarea adnata, M. curvata, M. hedlundii, M. muhrii, M. nigella, M. olivacea, and M. myriocarpa Vézda & V. Wirth ex Coppins. Several described species are included in Micarea for the first time, and additional name changes are required for nomenclatural reasons; nine new combinations result: M. assimilata (Nyl.), M. crassipes (Th.Fr.), M. elachista (Kérber) Coppins & R. Sant., M. globulosella(Nyl), M. intrusa(Th.Fr.) Coppins & Kilias, M. lignaria var. endoleuca (Leighton), M. melaenida (Nyl.), M. melanobola (Nyl.), and M. subviolascens (Magnusson). Several taxa are ex- LICHEN GENUS MICAREA IN EUROPE 19 cluded from the genus and the new combinations, Psilolechia clavulifera (Nyl.) and Bacidia prasinata (Tuck.), are proposed. Keys for the identification of all the accepted European taxa are given. The taxo- nomic parts are preceded by an outline of the historical background to the study of Micarea, and details of materials and methods employed in this study. Detailed accounts of the morphology, chemistry, and ecology in the genus are provided, and a discussion of distributions is supported by maps for the British taxa. All Micarea species occur on acidic, nutrient poor substrata, and most are confined to cool-temperate, boreal, or oceanic regions; a few occur in arctic-alpine areas but the genus is poorly represented in dry, lowland, Mediterranean regions. Prior to this study, most of the species were little-known or misunderstood; clarification of their taxonomy has been achieved by paying particular attention to their anamorphic states, chemistry (including pigmentation), and detailed anatomy. Consideration of the distribution and ecology of the species has proved invaluable in ordering the taxonomic chaos which pre- viously surrounded the notoriously variable species of the genus. Historical background Lecideaceae Until recently the circumscription of the Lecideaceae (and of the genera within it) had changed little from that adopted by Zahlbruckner (1926). It included most lichens with the following combination of characters: a crustose to squamulose thallus, a ‘grass-green’ phycobiont (excluding Trentepohlia and Phycopeltis), + immersed to sessile, disc-like apothecia without a thalline margin, mainly colourless spores, and an absence of parietin (or related pigments) and (or) polarilocular spores. The principal genera in the family (e.g. Lecidea, Catillaria, Bacidia, Biatorella, Mycoblastus, Lopadium, Bombyliospora, and Toninia) were separated mainly on the basis of spore characters, i.e. size, septation, and number per ascus. This classification largely ignored many features which (according to modern mycological concepts) now merit careful consideration, although they were used to varying degrees for the delimitation of species or infrageneric categories above this rank. In brief, these features involve the structure of asci, excipular and hypothecial tissues, paraphyses and anamorphs, ontogeny, finer aspects of thallus structure, and nature and location of pigments and lichen substances. In addition, investigations of the phycobiont(s) and considerations of ecology and phytogeography often provide valuable supplementary information. However, the use of some of these features in an attempt to define more natural genera is not a purely recent phenomenon. Several lichenologists working in the 1850s and 1860s made bold attempts in this direction. With regard to the Lecideaceae s. lat., two lichenologists deserving special mention are G. W. K6rber (who introduced Lecidella, Lopa- dium, Pyrrhospora, Schaereria, Schadonia, and Steinia) and A. B. Massalongo (who introduced Catillaria, Psilolechia, Sarcosagium, Scoliciosporum, Strangospora, and Toninia). In the latter half of the 19th century lichenology came under the almost monarchical influence of William Nylander, whose simplistic generic concepts gained precedence over the more far-sighted works of Kérber, Massalongo, and others. From the 1870s right up to the 1950s there were few attempts to reassess the generic concepts of Nylander or the slightly more complex, but no less artificial, system of Zahlbruckner. Between about 1929 and 1954 the French lichenologist M. G. B. Choisy resurrected many of the old and more or less forgotten genera, and created several new ones (e.g. Haplocarpon [= Huilia], Hypocenomyce, Trapelia, and Tremolecia). Unfortu- nately, Choisy’s works made little impact at the time and it was not until the mid-1960s that lichenologists began to look more carefully at the delimitation of genera. Recent investigations have led to the reinstatement (although often with emendations) of many of these genera and many new genera have had to be described (e.g. Fuscidea, Herteliana, Melanolecia, Trapeliop- sis, Tylothallia, and Vezdaea). Most are included in the key to European lichen genera in Poelt & Vézda (1981). Despite the many advances made during the last 15 years, it will be several decades before a reasonably natural generic classification within the Lecideaceae s. lat. is achieved. The enormity of the task can be appreciated from the fact that Zahlbruckner, (1921-40) accepted no less than 1450 species in the genus Lecidea alone! In addition to the high number of taxa involved, further problems arise from the locating and obtaining on loan suitable (including type) material, and the many difficulties in observing and interpreting many microscopical, morphological, and ontogenetic features. 20 BRIAN JOHN COPPINS Micarea The genus Micarea was first validly described in 1825 by Elias Fries in his Systema orbis vegetabilis (see p. 96), and was placed in his ‘Tribus Collemaceae’ on account of its rather gelatinous thallus, although he noted that it had Lecidea-like apothecia. The generic name was little used by most 19th century lichenologists, although it was accepted with the single species (M. prasina) by a few such as K6rber (1855). Towards the end of that century, J. T. Hedlund submitted his doctoral thesis to the University of Uppsala. In this work (Hedlund, 1892) he adopted and emended the genus to include 20 species, and his circumscription of the genus is essentially the same as that accepted by Vézda & Wirth (1976) and myself, although many species have since been added. Hedlund’s sagacious work was evidently too revolutionary for his time, and it did not achieve international recognition. It seems that Hedlund was disillusioned and had difficulties in finding a position. He turned to horticulture (especially dendrology) and then became an authority on Sorbus. Although Hedlund’s obvious talents were not lost to botany, they were sadly lost to lichenology, especially the study of microlichens. Vézda & Wirth (1976) slightly expanded Hedlund’s concept of Micarea by including Lecidea sylvicola and related species (all without ‘micareoid’ algae), and also Bacidia beckhausii. Apart from the exclusion of B. beckhausii, my own concepts are much the same, although I have emended the genus very slightly so as to include species such as Lecidea (Helocarpon) crassipes and Catillaria intrusa. The first species of Micarea to be described was Lecidea [Micarea] lignaria Ach. (1808). By 1850 only seven of the currently accepted 45 European species were validly published. At about this time lichenologists began to make use of better quality microscopes and the additional characters they revealed. By the end of the nineteenth century 34 of the accepted species had been described. The European Micarea flora was increased by only one species (Lecidea subviolascens) between 1900 and 1960. In 1961 Vézda published Bacidia [Micarea] subleprosu- la, and as a result of the present studies (some in collaboration with Mr P. W. James, Dr A. Vézda, and Dr V. Wirth) a further nine species have been described. In the future a few additional species will probably be added to the European flora, but the greatest expansion within the genus will come from the study of extra-European collections; indeed, many European, undescribed, and described (in other genera) species are already known to me from parts of the world outside Europe. Materials During the course of this study I have examined about 3,000 specimens (c. 1800 from the British Isles) attributable to Micarea, plus a further c. 500 specimens which have been referred to other genera. Material has been received on loan from (or studied in) the following institutional herbaria: ABD, ANGUC, BEL, BERN, BG, BM, BON, C, DBN, DEE, DUKE, E, G, GLAM, GZU, H, HAMU, HBG, HFX, IMI, K (now in BM; cited as BM ex K), L, LD, LIV, LSR, M, MANCH, NMW, NWH, O, S, STD, STU, SUN, TUR, U, UPS, VER, WCR, WIS, WRSL,; abbreviations according to Holmgren et al. (1981). In addition, numerous specimens have been received on loan from private herbaria. From the British Isles these are: Dr H. J. M. Bowen (Oxford), Dr R. W. M. Corner (Penrith), Mr I. P. Day (Carlisle), Dr U. K. Duncan (Arbroath; lichen herbarium recently gifted to E), Dr A. Fletcher (Leicester), Mr V. J. Giavarini (Parkstone, Dorset), Dr O. L. Gilbert (Sheffield), Mr R. Gomm (Taunton), Rev. G. G. Graham (Hunwick, Co. Durham), Mr A. Henderson (Leeds), Dr C. J. B. Hitch (Saxmund- ham, Suffolk), Dr P. D. Hulme (Aberdeen), Dr A. R. Pentecost (Royal Tunbridge Wells), Dr F. Rose (Liss, Hampshire; many specimens now in BM), Dr M. R. D. Seaward (Bradford), Mr J. F. Skinner (Southend-on-Sea), Dr P. B. Topham (Dundee), Mr R. G. Woods (Newbridge on Wye, Powys); and from elsewhere in Europe: Dr J. Hafellner, Dr H. Mayrhofer, and Prof. Dr J. Poelt (all Graz, Austria), Mr L.-E. Muhr (Karlskoga, Sweden), Dr A. Vézda (Brno, Czechoslo- vakia), and Dr V. Wirth (Ludwigsburg, W. Germany). When studying a group of much misunderstood lichens it is a rewarding exercise to investigate folders of other superficially similar (but strictly unrelated) taxa, especially those that occur in LICHEN GENUS MICAREA IN EUROPE 21 similar habitats. During this study numerous specimens of Micarea have been found in folders of widely misinterpreted (‘dustbin’) names, such as ‘Bacidia sphaeroides’, ‘Catillaria erysiboides’, and ‘Lecidea vernalis’. Copies of the list of specimens examined have been lodged at BM, DBN, E, GZU, M, NMW, UPS and US. Methods Field studies I have attempted to observe as many as possible of the accepted species in their natural habitats because such experience is invaluable for the appreciation of environmentally controlled variation and ecological requirements. For this end I have made field studies in most parts of mainland Britain, and also S.E. Ireland, Denmark (N. Jylland), mid- and N. Sweden. In addition, I have collected Micarea specimens during earlier expeditions to France (Bretagne) and western Ireland. I have been successful in finding two-thirds of the European species, and all but two (M. assimilata and M. subleprosula) of the 31 British species. Light microscopy Observations and measurements of external features were mostly made at X50 using a Vickers stereomicroscope equipped with a measuring eyepiece. Internal features were investigated with a Wild M20 microscope which was fitted with a drawing tube and a 2-5 x adapter for the drawing of spores and conidia etc. Most sections were made by hand with a razor blade, but some were cut by a freezing microtome. Sections were usually mounted in water, followed by (or directly in) 10% or 50% KOH(K), domestic bleach(C), 50% HNO3, or ammoniacal erythrosin (0-5g erythrosin in 100 ml 10% ammonia solution), but other mountants such as cotton-blue in lactophenol (LCB), congo red, and alcian blue were also used at times. Tests for amyloid reactions were made by mounting directly in Lugol’s iodine solution (1g iodine and 2g potassium iodide in 300 ml distilled water), or in this solution following treatment with 10% KOH. More permanent preparations were made by ringing mounts in LCB with nail varnish, or by mounting in polyvinyl alcohol (PVA) or cotton blue in PVA (see Omar et al., 1979). For further details on techniques see also ‘Guide to keys and identifications’ (p. 100). Thin-layer chromatography The t.l.c. techniques employed were those described by Walker & James (1980) which are based on the standard method of Culberson (1972). For the purposes of routine analysis only two of the three basic solvent systems were found to be necessary (i.e. H.E.F. and T.D.A.). Nomenclature In the list of synonyms for a given species each entry begins with the oldest valid name (basionym) and is followed with later combinations included in Hedlund (1892), Smith (1911, 1926), and James (1965a), and some others which have important nomenclatural implications. Zahlbruckner (1921-40) and Lamb (1963) should be consulted for additional combinations. The entries are listed chronologically, except for entirely invalid (or illegitimate) and misapplied names which are included at the end of the lists. Abbreviations of authors are according to Hawksworth (1980); those of journals to the third [1980] edition of Serial Publications in the British Museum (Natural History) Library; and those of books to Hawksworth (1974) or Stafleu & Cowan (1976-81). Apart from those newly described in recent years, few names in Micarea have been formally typified (according to Art. 7 of the Code); all cited lectotypes and neotypes are selected in this work, unless otherwise indicated. The nomenclature of lichens not treated in detail mainly follows Hawksworth et al. (1980), but a few later changes are used. eae. BRIAN JOHN COPPINS Morphology Thallus Thallus structure in Micarea is crustose and basically simple, but nevertheless, encompasses much interspecific and intraspecific variation. For the purposes of explanation thallus morphol- ogy can be roughly divided into five types. [Note: the discussion of the first type includes some general features applicable to all types. ] (i) Areolate-type (Fig. 1A—C) The term areolae (pl.) is used here to describe discrete, rounded (when viewed from above), flattened, or more usually convex (sometimes + globose) portions of the thallus which have developed directly from the prothallus lying in or on the substratum. These should be distinguished from the (often angular) segments derived from the cracking of a previously continuous crust. The areolae in Micarea mostly range from the 0-06 to 0.2 mm diam, but are characteristically larger in certain species. Thalli comprising smaller, discrete, granular, soredia- like structures (goniocysts) are dealt with in the next category (ii). Species that have well-developed, convex areolae sometimes appear + squamulose, e.g. M. assimilata, M. incrassata, M. lignaria (as in type of Lecidea trisepta var. polytropoides), M. melaenida, and M. subviolascens. Catillaria zsakii (a synonym of M. melaenida) was placed in the squamulose genus Toninia Massal. by Lettau, but its thallus is not truly squamulose and it was excluded from Toninia by Baumgartner (1979). In M. subviolascens (and to a lesser extent in some other species) the areolae become confluent and the resultant crust becomes secondarily cracked into ‘islands’ (each containing several primary areolae), thus giving the thallus a rather squamulose appearance. The areolae in most species are whitish or pale to medium grey, and often tinged dull greenish, bluish or brownish. The areolae of M. lignaria var. endoleuca are characterically ivory or cream-yellow. More vivid yellow, citrine or orange thallus colourations are not known in Micarea. A few whitish, arachnoid, prothalline hyphae are sometimes visible in specimens with scattered areolae, but a thick prothallus as found, for example, in some species of Phyllopsora (Swinscow & Krog, 1981) is unknown in Micarea. The thallus in most Micarea spp. is effuse and wide-spreading, and even when thalli form small patches amongst other lichens, a delimiting hypothallus, such as found in many species of Fuscidea and Lecidella, is never formed. Small thalli forming + circular patches amongst other lichens, suggesting parasitic behaviour, are characteristic of M. intrusa (p. 140). In vertical section the areolae usually lack a well-defined cortex, but their surface is sometimes covered by a thin (c. 2-10 wm thick), hyaline amorphous layer which does not dissolve in concentrated KOH (Fig. 1A, C). Such a layer is found in the areolae of, e.g. M. alabastrites, M. assimilata, M. cinerea, M. elachista, M. incrassata, M. lignaria, M. peliocarpa, M. subnigrata, M. subviolascens, and M. ternaria. Areolae with a similar external appearance, but without an amorphous covering layer, can be seen in well-developed specimens of, for example, M. denigrata, M. nitschkeana, M. globulosella, and sometimes M. melaena (Fig. 1B). The thallus of these species (and to a lesser extent that of M. elachista) is often invaded and disrupted by the dematiaceous hyphae of a fungal parasite(s), and non-lichenized algae, resulting in a dark, often blackish, scurfy crust. This phenomen on is best exemplified by M. melaena, which in Britain is rarely found with a healthy, well-developed thallus. By contrast, during my excursions in Sweden I found it to be rarely parasitised. Susceptibility to invasion by parasites or opportunists may be related to climatic factors, being increased in the British Isles by the overall weather and warmer conditions. It would appear, therefore, that the amorphous covering layer (never present in M. melaena) is an effective barrier against potential invaders, but whether or not this is the reason for its evolution is a matter for speculation. The nearest approaches to the development of a true cortex are found in M. elachista and M. melaenida. Sections of healthy areolae show an outer layer (c. 10-12 wm and 12-20 um thick respectively) of compacted, evacuated hyaline hyphae (Fig. 1C). The amorphous covering layer of M. elachista often partially disintegrates causing the areolae to exhibit a white-pruinose appearance, a feature not noted in any other species of the genus. LICHEN GENUS MICAREA IN EUROPE 23 | _] q Fig. 1 Some thallus types in Micarea. A, areolate-type without cortex but with amorphous covering layer (e.g. 4. lignaria). B, areolate-type without cortex or amorphous covering layer (e.g. M. denigrata). C, areolate-type with both cortex and amorphous covering layer (M. elachista). D, goniocysts (M. prasina). Scale = 50 wm. The outermost hyphae of areolae are frequently coloured or surrounded by a pigment, usually that which is found in the upper hymenium of the apothecia, or the pycnidial walls, of the given species; for example, the dilute olivaceous, K+ violet pigment in M. denigrata, M. elachista, M. globulosella, M. nitschkeana, and M. subviolascens, the green, K—, HNO3+ red pigment in M. cinerea, M. lignaria, M. peliocarpa, M. sylvicola, and M. ternaria, and the brown, K—, HNO3— pigment in M. subnigrata. The intensity of pigmentation is much dependent on exposure to light and pigment may be entirely absent in shade forms. Species that always lack pigment in their apothecia and pycnidia (e.g. M. alabastrites, M. pycnidiophora, amd M. stipitata) similarly lack pigment in the thallus. In the areolae of most species the algal layer is in direct contact with the substratum. However, a white medulla, devoid of algal cells, may be formed in the larger areolae of some species, e.g. M. incrassata, M. intrusa, M. lignaria, and M. subnigrata. 24 BRIAN JOHN COPPINS (ii) Goniocyst-type (Fig. 1D) Several species have a finely granular thallus composed of discrete, + globular structures, mostly c. 12-40 wm diam. These ecorticate granules consist of clustered algal cells intertwined and surrounded by short-celled hyphae, and are never protected by an amorphous covering layer. They are often seen to have short protruding hyphae, but they never have distinct spines as found in some species of Vezdaea (Poelt & Débbeler, 1975). They are similar to soredia, but because they are the main component of the thallus and not derived from specialised structures (soralia) or eroding or disintegrating parts of a different thallus type (see (iii) below), the term goniocyst (Ozenda, 1963) seems the most applicable for them. In M. prasina, M. hedlundii, and M. melanobola the thallus is composed entirely of goniocysts, the first of which appear to arise directly from the prothalline hyphae; further development is presumably by a process of division or budding from existing goniocysts. M. botryoides has a ‘primary’ thallus of flattened granular-areolae and in a few specimens I have noticed goniocysts developing from these areolae, which then become obscured as the gonio- cysts proliferate. However, in most specimens of M. botryoides no areolae can be seen; either they have become obscured or, perhaps in some cases the thallus develops as goniocysts from the outset. In M. prasina and M. melanobola the outermost hyphae of superficial (exposed) goniocysts are often surrounded by the K+ violet pigment which also occurs in the upper hymenium of these species. When well developed, thalli composed of goniocysts have a + gelatinous appearance when moist; this was the reason why Elias Fries originally placed the genus Micarea in his ‘Tribus Collemaceae’. The thallus of M. synotheoides consists of small dark granular-areolae and is somewhat intermediate between the ‘areolate-type’. The thallus of M. myriocarpa sometimes has a scurfy appearance but is often organized into small (10-15 wm diam) goniocyst-like granules. (iii) Sorediate-type This is represented by the apparently closely related M. leprosula and M. subleprosula. Their thalli are essentially of the ‘areolate-type’ but their areolae lack an amorphous covering layer and are very fragile (easily broken by touching with the point of a needle), often breaking down or eroding to form irregularly shaped, pale green or yellowish green soredial granules, c. 20-50 wm diam. (iv) Smooth- or scurfy-type Included here are thalli which, although developed on the surface of the substratum, are not organized into discrete areolae or goniocysts. The thalli may be smooth, continuous to rimose, or irregularly scurfy-granular. Such thalli may be formed by species that normally have well defined areolae (e.g. M. lignaria and M. melaenida) or, by lignicolous species whose thalli are often endoxylic (e.g. M. anterior, M. nigella, and M. olivacea). The thalli of M. bauschiana and M. lutulata vary from smooth or rimose, to scurfy-granular, and lack areolae, whereas the similar M. lithinella, M. sylvicola, and M. tuberculata sometimes produce areolae. Another two species whose thalli fall within this category are M. adnata and M. turfosa. (v) Immersed-type The thallus of many lignicolous species is indistinct and developed below the surface of the substratum, which may have a bleached appearance (cf. M. muhrii). An endoxylic thallus is occasionally formed by species that more characteristically have superficial areolae, e.g. M. cinerea, M. denigrata, M. lignaria, M. muhrii, and M. peliocarpa. Conversely, M. melaeniza and M. misella are usually endoxylic but forms with areolae are sometimes encountered. The thallus in M. anterior, M. nigella, and M. olivacea is often endoxylic, but a thin + smooth crust is sometimes formed on the surface of the lignum. The thalli of M. contexta, M. eximia, and M. rhabdogena are invariably endoxylic. Sections of the substratum show small goniocyst-like clusters (c. 15-40 wm diam) surrounded and inter- connected by pigmented hyphae. As in the relevant examples in thallus types (i-iii), and in M. turfosa in group (iv), the pigment involved is that found in the upper hymenium or pycnidial walls of the given species, viz.: dark green and K— in M. contexta and M. eximia, and olivaceous and K+ violet in M. rhabdogena. LICHEN GENUS MICAREA IN EUROPE Is) The species which at times grow on bryophytes usually have a superficial thallus, but even so, the thallus is often, in part, endocuticular. When on bark Micarea thalli are generally epiphloeodal, but at least partially endophloeodal thalli have been seen in M. cinerea and M. peliocarpa. Phycobionts Among the 45 species of Micarea accepted in this revision three types (? genera) of ‘grass-green’ phycobiont can be distinguished by LM observations on thallus squashes. Unfortunately the identities of these algae are unknown and they await critical studies by an algologist. In addition, two genera of blue-green alga may be involved in the formation of cephalodia (q.v.). The commonest alga in Micarea is that found in the type species (M. prasina) and was discussed at some length by Hedlund (1892, 1895). In the course of this study this alga is hereafter referred to as ‘micareoid’ (Fig. 2). Its cells are fairly regular in appearance, + globose, thin-walled and c. 4-7 wm diam. Reproduction within the thallus appears to be by a process of cell division in which the protoplast divides in two, followed by the laying down of a dividing wall which joins up with existing wall of the parent cell (Fig. 2A). No internal divisions into aplanospores as found in, for example, Myrmecia and Trebouxia, have been observed. Contact between the fungal hyphae and the algae cells is by intracellular haustoria (Peveling, 1974) which are readily seen at X 1000 (mounted in 10% KOH, followed by ammoniacal erythrosin). The haustoria may be peg-like (Fig. 2B, D-E), swollen to become clavate (Fig. 2G) or capitate (Fig. 2C, H), or spreading into a foot (Fig. 2F). In collapsed algal cells (Fig. 2H) the haustoria are intensively stained. A characteristic feature of the micareoid alga and its relationship with the mycobiont is that a hypha frequently becomes closely aligned along the dividing line of two separating algal daughter cells (Fig. 2D—-E); the hypha concerned is often seen to penetrate one or both of the cells by haustoria. The second algal type is found in Micarea sylvicola and its presumed relatives M. bauschiana, M. lutulata, and M. tuberculata. The cells are thin-walled and irregular in size and shape, varying from 5-12 «m diam when globose to up to c. 15X10 wm when + ellipsoid. Iam undecided as to their mode of reproduction within the thallus. The large number of cells at the lower end of the size range is suggestive of aplanospore formation, but I have never observed a mother-cell undergoing division. Haustorial penetrations by the mycobiont hyphae have been detected, but they are never as distinct as those involving micareoid algae or the phycobiont of M. intrusa (see below). The third algal type is confined to M. intrusa, and has large globose cells, 7-21 wm diam, with thick hyaline walls c. 1-24m thick. Cell division within the thallus seems to be by a process of the division of a mother-cell into three or four daughter-cells (i.e. ‘protococcoid’ division). Many of the cells are clearly seen to be deeply penetrated by a haustorium (Fig. 55). This phycobiont looks very similar to that of Scoliciosporum umbrinum and may well be identical with it. Cephalodia For reviews of the morphological, taxonomic, and physiological aspects of cephalodia see Jahns (1974), James & Henssen (1976), and Millbank (1976). Cephalodia have been found in three species of Micarea: M. assimilata, M. incrassata and M. subviolascens. The first two species have thalli composed of verrucose areolae with a micareoid phycobiont. However, sections of their thalli reveal the presence of + globose structures (c. 200-600 zm diam) containing a blue-green alga of the genus Nostoc. In many cases these structures are visible externally and closely resemble the areolae except that they are brown, and usually darker, in colour. Internally they consist of numerous ramifying fungal hyphae (presum- ably belonging to the Micarea) and dense masses of Nostoc cells which have lost their normal (when free-living) filamentous arrangements; and I am in no doubt that these structures are cephalodia. I am less certain of the status of the more loosely organized clusters of Stigonema which are sometimes associated with the same two Micarea species, and also M. subviolascens. However, the Stigonema filaments are, at least partially, disrupted and fungal hyphae are present. 26 BRIAN JOHN COPPINS Fig.2 ‘Micareoid’ phycobiont of Micarea alabastrites and its relationship with mycobiont hyphae; see text for further details. Scale = 10 um. The three above-mentioned Micarea species are morphologically closely related and occur in exposed arctic-alpine situations. In such habitats the formation of cephalodia with a blue-green alga(e) which has the ability to fix atmospheric nitrogen, is of undoubted nutritional benefit to the mycobionts. Most morphological and physiological studies concerning cephalodia have involved macrolichens. Reports of cephalodia in crustose lichen are relatively few, although some are listed by James & Henssen (1976). Additional examples in the Lecideaceae include Huilia aeolotera, H. elegantior, H. panaeola (Hertel, 1977), and Lecidea pallida (Fries, 1874). Apothecia: external features As a rule the apothecia in Micarea are small to medium sized, convex to + globose or tuberculate, and usually immarginate. However, this generalization encompasses considerable variation even within a single given species. With regard to shape, some species (e.g. M. denigrata and M. prasina) are very variable with apothecia. that may be shallow-convex to convex-hemispherical (Figs 3A—-B, 4A-B) and finally + globose (Fig. 3D) or tuberculate (Fig. 3C). Apothecia of other species may be less variable: those of M. alabastrites and M. cinerea are broadly convex to hemispherical, sometimes tuberculate, but never + globose; at the 27 LICHEN GENUS MICAREA IN EUROPE ‘unl QZ *xoidde = a]e9g *(viuvUsy 'W JO SuLo}) ayeydys “q *(vuvUsy "W) WnHindIoxa poxoayal YIM ssoqgo]s ‘q ‘aye[NdIOqn} wiNnid9y}0de nq ‘ONIP “DQ “pexeye: wnjndioxe pue popuedxe wintdeyjode yng ‘oyIp ‘g *(vdiv20nad ‘W) wn -ndioxa padojaaap-[[aM YIM XOAUOD ‘Y *(SHeWIWIeISeIP-IuIas) Dasvo1py UI sadAy winisayjode aui0g ss ¢ “BI BRIAN JOHN COPPINS 28 ‘un yz ‘xoidde = a[e9g ‘(vJDSsp49U1 “PF JO SULIOJ DWOS) WINTIdyYJOdAY SUTJOOL, YJIM X9AUOD ‘¢ *(Sadissp49 PF) WN{NdIdx9 podojaasp-|]oM YIM 9}eIIdNs ‘gq (‘vyauzYyII) “PW) wIN| -ndioxa ynoyyIM ‘dJeupe pure XdAUOd SY ‘(ONRWUWeIZeIP-IWas) Vasvo1PF UI sodA} UINIDOYIOde sUIOS =f “BI ase . N =p. % Ths ai) 2 ' a ai LICHEN GENUS MICAREA IN EUROPE 29 other extreme those of M. contexta, M. eximia, and M. myriocarpa are + globose from the beginning and often tuberculate. The range of variability for most other species falls somewhere between these last two extremes. Apothecia are usually sessile (or partly hidden by adjoining areolae or goniocysts) but occasionally, as with M. lignaria, M. botryoides, and M. ternaria, they may be turbinate or short-stalked (due to vertical elongation of excipular and (or) hypothecial tissues), but only in M. crassipes and an undescribed species from New Zealand is this a constant feature (Figs 3E, 4B). Apothecia are usually immarginate, but in those species with a well differentiated excipulum (e.g. M. cinerea, M. denigrata, M. muhrii, M. peliocarpa, and M. ternaria), a faint marginal zone is sometimes apparent, especially in young apothecia. This zone is rarely raised above the level of the disc, but it is often paler in colour (especially when wet) and slightly more glossy. Even in M. crassipes, the only species whose apothecia regularly exhibit a distinctly raised marginal rim, this feature is soon occluded as the disc expands and becomes convex. The dimensions of apothecia in Micarea fall mainly within the range 0-1-0-6 mm diam. In some species (e.g. M. denigrata, M. peliocarpa, and M. prasina) there is considerable variation in apothecial size, but in others it may be less so. Characteristically small apothecia (<0-3 mm diam) are found in, for example, M. contexta, M. eximia, M. melaeniza, M. misella, M. myriocarpa, and M. rhabdogena. Species often with large apothecia, sometimes growing to a diameter of 0-7 mm (or even up to 1 mm), include M. assimilata, M. cinerea, M. incrassata, and M. ternaria. These dimensions relate to ‘normal’ convex to globose apothecia; however, larger sizes are often attained when the apothecia become tuberculate (Fig. 3C). The apothecia of M. tuberculata are usually less than 0-3 mm diam, but when tuberculate may reach 0-5 mm diam. Some other examples with equivalent dimensions are: M. anterior (0-3 mm, 0-6 mm), M. bauschiana (0-4 mm, 0-7 mm), M. botryoides (0-25 mm, 0-5 mm), M. cinerea (0-7 mm, 1-3 mm), M. elachista (0:3 mm, 0-8 mm), M. lutulata (0-4 mm, 0-8 mm), and M. sylvicola (0-5 mm, 1-2 mm). The apothecia of M. adnata are nominally c. 0-2-0-4 mm diam but they often coalesce and become somewhat tuberculate, producing large nodulose clusters up to about 2 mm across. The apothecia of certain species (M. adnata, M. alabastrites, M. pycnidiophora, and M. stipitata) are always devoid of pigments and hence whitish or pallid in colour and + translucent when wet. Conversely, those of certain other species are heavily pigmented and invariably blackish and opaque when wet (e.g. M. assimilata, M. contexta, M. crassipes, M. eximia, M. incrassata, M. intrusa, M. melaena, M. melaenida, M. melaeniza, M. nigella, M. olivacea, M. subviolascens, and M. ternaria). Black apothecia are also characteristic of M. lignaria, M. misella, and M. turfosa, but bluish grey or brown shade-forms have been encountered. The apothecia of a number of species (notably M. bauschiana, M. denigrata, M. peliocarpa, and M. prasina) commonly occur as forms ranging from whitish or pallid, to blue-grey or pale-brown (sometimes white/blue-grey or white/brownish piebald), through dark grey or dark brownish grey, to black. The consequent array of colour forms, combined with much variability with regard to size and shape of apothecia, thallus development and substrata, has resulted in a large synonymy for each of these species! In such variable species apothecial colour is largely dependent on the light regime experienced by individual apothecia. This can be demonstrated in the field by observing the species in situations where large changes in light intensities occur over short distances (a few centimetres or less). Such a situation is portrayed in Fig. 5, by an example of M. bauschiana growing all over the exposed parts of a flat stone lodged in the dry underhang of a bank, alongside a woodland road. Species such as M. curvata, M. elachista, M. myriocarpa, M. osloensis, and M. subnigrata have brown or brown-black apothecia due to the presence of a brown epithecial pigment. The external brownish colour may also result from the combination of a colourless (or + so) hymenium and epithecium and a dark brown hypothecium (e.g. in forms of M. lutulata and M. muhrii). However, brown apothecia as sometimes encountered in, for example, M. bauschiana and M. denigrata, often indicate the presence of a dematiaceous hyphomycete (? Bispora sp.) parasitizing the apothecial tissues. Apothecia which appear blue-grey or blue-black contain a green (K—) epithecial or hymenial pigment (e.g. in M. bauschiana, M. peliocarpa, and M. sylvicola), or similarly located high concentrations of the olivaceous (K+ violet) pigment (e.g. in 30 BRIAN JOHN COPPINS i r oS | | / V4 q My if 7 ie ai AW iZ ROAD Be EO a ye Fig. 5 Sketch demonstrating a situation which M. bauschiana shows variation in the colour of its apothecia, from pallid (A), through pallid/brownish or pallid/blue-grey (B), to blue-black (C). Site: roadside bank in woodland in lower Glen Roy, Westerness (cf. Coppins 3492/3, E); the bank is c. 1m high). M. denigrata, M. nitschkeana, and M. prasina. | know of no Micarea that has apothecia possessing pruina or, exhibiting a bluish (caesious) ‘bloom’ when wetted. Such features are due to the existence in the epithecium of minute colourless (or pale straw when dense) granules that dissolve in K, and are characteristic of several species often misidentified to Micarea, e.g. Bacidia beckhausii, Lecanora symmicta agg., Lecidea caesioatra, L. turgidula, and Scoliciospor- um pruinosum. Apothecia: internal features All tissues of the apothecia are bound by a weak gel-matrix which + dissipiates in 10% KOH to clearly reveal the component parts (i.e. hyphae, paraphyses, and asci). In some species the apices of the paraphyses and/or the hyphae of the hypothecium are additionally cemented by dense pigment deposited in the matrix and/or on the hyphal walls, thus tending to impair their separation in K. The presence of the gel-matrix serves to separate species of Micarea from some superficially similar species of Vezdaea (Poelt & Débbeler, 1975). Hymenium and epithecium The term ‘epithecium’ is here employed in its broad sense to refer to the upper part (usually c. 3-15 ym) of the hymenium where this differs in appearance from the remaining lower part(s). LICHEN GENUS MICAREA IN EUROPE 31 This difference can be due to the much branched and entangled apices of the paraphyses, or (and) deposits of pigment in the gel-matrix, or on (or in) the walls of the paraphyses. The upper layer of the hymenium differentiated in such ways is perhaps more correctly termed the ‘epihymenium’ (Poelt, 1974a) or ‘pseudoepithecium’ (Korf, 1973). In its strict (original) sense an ‘epithecium’ refers to a layer of branches of paraphyses that overtop the asci. Such a layer is approached in some species of Micarea and is best exemplified by M. contexta and M. melanobola. In Micarea the difference between an ‘epihymenium’ and an ‘epithecium’ is not clear-cut, and so I have chosen to employ the latter term which has been long and widely employed in lichenology. The height of the hymenium in Micarea species is sometimes difficult to measure accurately because the hymenium often merges rather imperceptibly into the hypothecium. Measurements are most accurately made by mounting thin sections in Lugol’s iodine, in which the strongly amyloid (dark blue) hymenium contrasts strongly with the non-amyloid (or + so) hypothecium. Even when accurately determined the height of the hymenium is rarely a useful character in distinguishing similar species. The overall range in height is about 23-90 wm, and in most cases is within 35-50 xm. Shallow hymenia (rarely exceeding 35 xm) are characteristic of many of the species with small spores, viz.: M. hedlundii, M. melanobola, M. misella, M. myriocarpa, M. nigella, M. olivacea, M. osloensis, M. rhabdogena, and M. tuberculata. Likewise, tall hymenia are characteristic of species with large spores, and the tallest hymenium (65-90 xm) belongs to the species with the largest spores, namely M. subleprosula. The height of the hymenium sometimes increases with the age of the apothecium, and this is particularly true in M. bauschiana and M. sylvicola: small, yet mature, apothecia usually have a hymenium c. 40 wm tall, but as the apothecium enlarges and increases in convexity, the hymenium often increases in height up to about 60 zm; at the same time the paraphyses appear to ‘stretch’ and become thinner (especially in the lower part of the hymenium). The hymenium in Micarea never contains hyaline, crystalline inclusions that adhere to paraphyses (as found in some species of Lecidella, and many Graphidaceae) or numerous oil droplets (as found in several species of Buellia and Caloplaca). Epithecial granules that dissolve in K and often give the apothecia a pruinose appearance are also never found in Micarea. Minute granules of blue-violet (K+aeruginose) pigment, like those found in Lecidea hypnorum, are occasionally observed in the hymenium of M. contexta and M. lignaria. The colour of the hymenium and epithecium is an important character in the identification of Micarea species. A discussion of the various pigments involved is given in the sections on ‘chemistry’. The hymenium may be dilutely and + evenly coloured throughout, or the colouration may be more intense in the upper part. In many species the hymenium is often intersected by dark vertical streaks, a feature usually due to dense pigment surrounding individual, or small fascicles of stout paraphyses (see p. 61). Asci The asci of Micarea species are clavate or cylindrical-clavate in shape and belong to the Lecanora-type of Honegger (1978). When mounted in Lugol’s iodine (following treatment in 10% KOH) the ascus is seen to have a non-amyloid wall surrounded by an amyloid outer-layer (‘fuzzy coat’) and an internal staining amyloid apical dome (Fig. 6). Micarea has sometimes been placed in the Arthoniaceae (e.g. Lamb, 1953) but the ascus of members of that family does not have an amyloid outer-layer and the apical dome is not deeply amyloid. In a few species of Arthonia and Arthothelium I have observed a faint bluing in the part of the apical dome immediately adjacent to the ocular chamber. In some Arthonia species a tiny amyloid ring is apparent above the apex of the ocular chamber, and, despite statements to the contrary by Eriksson (1981), such a ring is found in the asci of A. radiata (type species of Arthonia Ach.) and A. fuscopurpurea. The same type of ring was also described for Bryostigma leucodontis by Poelt & Débbeler (1979). Such aspects of ascus structure in the Arthoniaceae clearly merit more detailed investigation. The ascus structure in Micarea supports my opinion that the genus should be placed in the Lecideaceae s. str., at least for the time being. The asci of all European Micarea species are 8-spored. The North American Lecidea populina 32 BRIAN JOHN COPPINS Fig. 6 Ascus apex of Micarea alabastrites in optical section (LM); mounted in Lugol’s iodine following pre-treatment in 10% KOH. A, young ascus. B, mature ascus. ac, apical cushion; aw, ascus wall; d, apical dome (tholus); oc, ocular chamber; ol, outer layer of ascus wall. Shading indicates intensity of amyloid reaction; note that in reality the apical cushion and ocular chamber are probably completely non-amyloid. Scale = 10 um. Mill. Arg. has 16-spored asci and was transferred to Micarea by Anderson & Carmer (1974). Unfortunately the type material of L. populina has been out on loan from H and not available to me during the course of this study. However, the species apparently occurs in Xanthorion communities and, if this is so, then it is unlikely to be a Micarea. Spores The wide variety of spore types found in Micarea can be seen in the selection of spores from all the European species illustrated in Figs 7-33. A few species (e.g. M. assimilata, M. contexta, and M. lithinella) have spores that are + consistent in size, shape, and septation, but for many other species (e.g. M. anterior, M. botryoides, M. denigrata, M. prasina, and M. turfosa) these characters can be very variable even within the same ascus. The smallest spores are found in M. myriocarpa (5:5-8-5X1-5—2-5 ym) and the largest ones are found in M. subleprosula (40-60 x 5—6:6 wm). Spores may be simple or up to 7-septate, the variations within these limits depending on the species. Spores with more than 7-septa are very rare, although a few 9-septate spores have been observed in M. subleprosula, and a collection (Coppins, 1834) of M. synotheoides has spores with up to 11 septa. Among the great range of spore shapes encountered, some may be broadly ellipsoid (M. subnigrata and M. intrusa), but regularly globose spores do not occur in Micarea. Healthy spores are always thin-walled and colourless. However, the walls of old spores trapped within the hymenium sometimes become slightly thickened and pale straw coloured, or they may become impregnated with hymenial pigment. Spore walls always appear smooth (LM at X 1000) and are never surrounded by a gelatinous epispore. The cytoplasm of spores, asci, and ascogenous hyphae in M. intrusa is sometimes a dilute orange, turning purple-red in K. LICHEN GENUS MICAREA IN EUROPE ai Fig.7 A, M. adnata (E-holotype). B, M. anterior (H-NYL 21655 — lectotype). Scale = 10 wm. 33 34 BRIAN JOHN COPPINS at so sate Fig.8 MM. alabastrites. A, (H-NYL 18656 — holotype). B, (Coppins 2588, E). Scale = 10 um. LICHEN GENUS MICAREA IN EUROPE 35 L j v q Fig.9 M. assimilata. A, (H-NYL 16556 -lectotype). B, (Norway, Oppland, Fogstuen, 1857, Lindsay, E). Scale = 10 wm. 36 BRIAN JOHN COPPINS D0QQU 0OSORC oR egE L = | i] 1 Fig. 10 A, M. bauschiana (M - lectotype). B, M. botryoides (Coppins 8429, E). Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE a) aoe ees j t Fig. 11 M. cinerea (Coppins 2533, E). Scale = 10 wm. 38 BRIAN JOHN COPPINS Ono OU UU COE | j ' 1 Fig.12 A, M. contexta (S—lectotype). B, M. crassipes (Vézda Lich. Sel. 11, BM). C, M. curvata(WRSL- holotype). Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE 39 DOGO HYG ta df JURCHE i 1096 Fig. 13 M. denigrata. A, (UPS —lectotype). B, (H —lectotype of Lecidea hemipoliella). Scale = 10 um. Ls U 40 BRIAN JOHN COPPINS (Hoe tet ae O0000UG | Ws J t t Fig. 14 A, M. elachista (L-—lectotype). B, M. eximia (S — lectotype). C, M. hedlundii (UPS — holotype). Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE 41 Fig. 15 M. globulosella. A, (S — lectotype). B, (Czechoslovakia, Slovakia, Vysoké Tatry, 1879, Lojka, BM). Scale = 10 um. 42 BRIAN JOHN COPPINS Wises IAG ne On je =i i] 1 Fig. 16 M. incrassata. A, (S— holotype). B, (Kerguelen, 1875, Eaton, E). Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE 43 = & Fig. 17 M. intrusa. A, (UPS — holotype). B, (BM — isotype of Lecidea melaphana). C, (Norway, Hordaland, Fjell, 1980, Skjolddal, BG). Scale = 10 wm. 44 BRIAN JOHN COPPINS JOG ab ees | ‘ q Fig. 18 M. leprosula (UPS — lectotype). Scale = 10 um. LICHEN GENUS MICAREA IN EUROPE 45 if mK oe ae T q Fig.19 M. lignaria. A, (H-ACH 265 — lectotype). B, (TUR-VAINIO 21274 - lectotype of Lecidea trisepta var. polytropoides). Scale = 10 um. 46 BRIAN JOHN COPPINS JUQUQOD: 00000 - QO0000 L ] r a | Fig.20 A-B, M. lithinella; A, (M-—lectotype); B, (Germany, Heidelberg, Zwackh, H-NYL 19191). C, M. lutulata (Wales, Pembroke, Tycanol, 1980, James, BM). Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE 47 p9668808 a qT Fig.21 A, M. melaena (UPS -—lectotype of Lecidea milliaria var. turfosa). B, M. melaeniza (S— holotype). C, M. melanobola (H-NYL 21614 - lectotype). Scale = 10 um. 48 BRIAN JOHN COPPINS -— - = Fig. 22 M. melaenida. A, (H-NYL 18843 - holotype). B, (Fl. Hung. exs. 714, BM —topotype of Catillaria zsakii). C, (WRSL - lectotype of Catillaria schumanii). Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE 49 JOOUUOI > 000000): weeCHERE [= J rat | Fig. 23. A, M. misella (H-ACH 52-holotype of Lecidea resinae subsp. globularis). B, M. muhrii (E — holotype). C, M. myriocarpa (hb Wirth 6085 — holotype). Scale = 10 um. 50 BRIAN JOHN COPPINS 0000) L j ' 1 Fig. 24 A, M. nigella (E — holotype). B, M. nitschkeana (Coppins 2426, E). C, M. olivacea (BM — holotype). Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE 51 sBg8 SHAG Fig. 25 M. peliocarpa. A, (H-NYL 18716 — isotype of Lecidea violacea). B, (England, New Forest, Great Wood, Bramble Hill Walk, 1970, Coppins et al., E). Scale = 10 um. 52 BRIAN JOHN COPPINS Fig. 26 M. prasina. A, (UPS -lectotype). B, (H-NYL 19056 — lectotype of Lecidea subviridescens). Scale = 10pm. LICHEN GENUS MICAREA IN EUROPE 53 968800 | ote JO08GE | JOGG86 L at] t q Fig. 27M. prasina. A, (H-NYL 21604 — lectotype of Lecidea prasiniza). B, (GZU — holotype of Micarea polytrichi). Scale = 10 um. 54 BRIAN JOHN COPPINS JU L ] v | | a Oa Fig. 28 A, M. pycnidiophora (E — holotype). B, M. osloensis (UPS — holotype). Scale = 10 um. LICHEN GENUS MICAREA IN EUROPE 55 HO0N0UDOYL es es | Fig. 29 A, M. rhabdogena (BM - isolectotype). B, M. stipitata (E- holotype). Scale = 10 wm. 56 BRIAN JOHN COPPINS 7 ae Fig. 30M. subleprosula (Muhr 4380, E). Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE Sy, rs 4 Fig. 31 A, M. subnigrata (H-NYL 19136 — lectotype). B, M. subviolascens (Havaas Lich. Exs. Norv. 139, BG). C, M. sylvicola (UPS — lectotype). D-E, M. synotheoides; D, (H-NYL 19101 — lectotype); E, (Coppins 3257, E). Scale = 10 wm. Fig. 31 D-E on following page LICHEN GENUS MICAREA IN EUROPE 59 [as ] t J Fig. 32. A, M. tuberculata (O - lectotype). B, M. ternaria (Thomson 9188, DUKE). Scale = 10 um. 60 BRIAN JOHN COPPINS Fig. 33 M. turfosa. A, (VER - holotype). B, (Vézda Lich. Sel. 1135, BM). Scale = 10 um. LICHEN GENUS MICAREA IN EUROPE 61 Paraphyses The paraphyses in Micarea are characteristically thin and branched. When measured (in 10% KOH, or ammoniacal erythrosin) at the mid-hymenium they may be very thin and only about 0-6-1 um wide (e.g. M. anterior, M. botryoides, M. contexta, M. eximia, M. lithinella, M. misella, and M. prasina), or thin and c. 1-1-5 wm wide (e.g. M. adnata, M. cinerea, M. denigrata, M. intrusa, M. muhrii, and M. peliocarpa), or relatively stout and about 1-5—1-8 xm wide (e.g. M. assimilata, M. incrassata, M. lignaria, M. osloensis, and M. subnigrata); these measurements relate to paraphyses not coated in pigment. In old, much expanded, apothecia the paraphyses sometimes appear ‘stretched’ and thinner than normal (especially in the lower half of the hymenium); this phenomenon has frequently been observed in collections of M. bauschiana, M. sylvicola, and M. lignaria. In many species the paraphyses gradually widen towards their apices, but the apices are never regularly clavate or capitate. This widening is often enhanced by the deposition of closely adhering pigment which sometimes gives the appearance of a ‘hood’ (e.g. M. melaena); such coatings or hoods can be detached by gently boiling and then tapping sections or squash preparations in 50% KOH. Ina few cases (e.g. M. melanobola) the pigment cannot be separated in this way and appears to be located within the walls of the paraphyses. Paraphyses with dark pigmented apical ‘caps’, like those found in Catillaria s. str. (Killias, 1980: 253), Buellia, and many species of Lecanora, are not known in Micarea. In all species of Micarea a large proportion of the paraphyses are branched, even if the branching is mainly confined to the epithecium. Species with sparingly branched paraphyses include M. assimilata, M. incrassata and M. lignaria. Anastomozing paraphyses have been observed in all the species, but often the anastomoses are + confined to the lower third of the hymenium. The degree of branching and anastomosing is difficult to quantify for the practical purposes of identification, but this character can be useful when comparing collections micro- scopically. A similarly difficult character is the relative abundance of the paraphyses. The two extremes can be referred to as: (a) ‘numerous’ — large in number and immediately discernible when observing a mount in 10% KOH at 400; (b) ‘scanty’ — few in number and not immediately obvious when observed in the same way. In hymenia with scanty paraphyses the ‘extra space’ is taken up by hymenial gel or a higher proportion of asci, or a combination of both. The situation in most species lies somewhere between the two extremes. Furthermore, the relative propor- tions of hymenial gel and asci to paraphyses sometimes increases as the apothecium expands (cf. the above example of M. bauschiana, M. sylvicola, and M. lignaria). An accurate assessment of the abundance of paraphyses is not usually essential for the routine identification of Micarea species, although it can be helpful when comparing difficult, convergent forms of M. denigrata and M. misella (see couplet 11 of the main key). In addition to the ‘normal’ paraphyses described above, the hymenium of several species (e.g. M. bauschiana, M. botryoides, M. eximia, M. nigella, M. sylvicola, and M. tuberculata) contain scattered individuals, or small fascicles, of rather stout paraphyses. These ‘paraphyses’ are about 2-3 wm wide and usually more distinctly septate than normal paraphyses. Furthermore (especially in species with a dark hypothecium), they are often coated in pigment throughout their length (Fig. 34), with the result that the hymenium is seen to be intersected by dark vertical streaks. They are mainly found in species with ‘scanty’ paraphyses, and appear to extend deep into the hypothecium. The elucidation of their true status and function awaits detailed ontogenetic studies, but it is possible that they have a strengthening, spacing or protective function during the maturation of the hymenium from the primary corpus. Hypothecium The area of tissue lying below the hymenium, or between the hymenium and the excipulum (if present), is generally referred to by lichenologists as the ‘hypothecium’. In many groups of lichenized discomycetes (including the Lecideaceae) this area can be divided into an upper, usually narrow, layer containing mainly ascogenous hyphae, and a lower, often much deeper layer of structural tissue (hyphae gelatinised to various extents, according to genus or species). These two layers are often of different colour or colour intensity. Where the two layers are 62 BRIAN JOHN COPPINS t 4 Fig. 34 Two types of paraphyses in Micarea tuberculata. A, normal, non-pigmented paraphyses. B, stout, pigmented paraphyses. Scale = 10 wm. distinct they have been referred to as the ‘subhymenium’ and ‘hypothecium’ respectively (e.g. Hertel, 1977b). The term ‘hypothecium’ in this context has been substituted by many students of non-lichenised discomycetes by the term ‘medullary excipulum’ (Korf, 1973), and was called the ‘ental excipulum’ by Eckblad (1968). In Micarea the two layers are well-differentiated in only a few species (e.g. M. crassipes); and in the present work ‘hypothecium’ refers to all ascocarp tissue lying below the hymenium, apart from the excipulum (where present). This same terminology was adopted in the recent study of Catillaria by Kilias (1981). The hypothecium in Micarea is composed of deeply staining (e.g. in LCB and ammoniacal erythrosin), swollen-celled ascogenous hyphae (c. 2-5 wm wide) and moderately staining, slender ‘structural’ hyphae (c. 0-7-2 wm wide; slightly varying in width according to species), embedded in a gelatinous matrix. In species with a colourless or pale hypothecium the matrix is + dispersed in K and the hyphae are then clearly visible (400). In some species with a coloured hypothecium the pigment may be + evenly distributed through the matrix and the hyphae are similarly distinct in K (e.g. M. eximia, M. olivacea, and M. turfosa). However, in the majority of species with a darkly coloured hypothecium (e.g. M. assimilata, M. botryoides, M. melaena, M. myriocarpa, M. nigella, M. sylvicola, and M. tuberculata) the pigment, although present in the matrix, exists as a strongly adhering coat around the hyphae, such that the hyphae appear thick-walled and broad (c. 1-5-4 wm). The pigment coatings tend to bind the hyphae together so that they are often indistinct in K. For the routine identification of Micarea species it is not essential to know the location of pigment (i.e. + evenly distributed through the gel-matrix versus LICHEN GENUS MICAREA IN EUROPE 63 tightly bound to hyphae) but such knowledge is sometimes of supplementary value, e.g. when comparing M. eximia versus M. nigella and M. olivacea versus M. tuberculata. A more important diagnostic character in Micarea is the colour of the hypothecium in water mounts, and the corresponding colour changes obtained by the addition of KOH and HNO;3. A discussion of the pigments involved is given under ‘chemistry’. The height of the hypothecium (in vertical section) is largely dependent on the overall size and (especially in species with a poorly developed excipulum) convexity of the apothecium. The measurements given in the species descriptions relate to normally developed, non-tuberculate apothecia. This character is often very variable for a given species and consequently of little diagnostic value when comparing closely similar species. One exception to this is the case of M. contexta (20-90 um) versus M. melaena (80-160 pm). In Lugol’s iodine the hypothecial tissues are non-amyloid, although there is sometimes a faint bluing in the vicinity of ascogeneous hyphae (especially in the upper part of the hypothecium). Excipulum The size and distinctiveness of the excipulum (‘ectal excipulum’) in Micarea varies greatly according to species and the age of the apothecium. In species such as M. cinerea, M. crassipes (Fig. 4B), M. peliocarpa, and M. ternaria the excipulum is sufficiently well developed that their young apothecia are often weakly or distinctly (M. crassipes) marginate in outward appearance. However, even when well developed and initially distinct, the excipulum may become reflexed and + occluded as the apothecium expands and increases in convexity (Fig. 3A—B) or becomes tuberculate (Fig. 3C). In many species the excipulum is always extremely reduced or absent (Fig. 4A). When present, the excipulum is composed of outwardly radiating branched and anastomosing hyphae that + separate in K. The hyphae closely resemble paraphyses, but are usually more dense and more richly branched. With markedly convex apothecia it can be difficult to distinguish between reflexed portions of the hymenium and what might be an excipulum. In such cases the excipulum (if present) can be identified in good thin sections by the absence of asci and a negative (non-amyloid) reaction to Lugol’s iodine. The excipulum often differs in colour or colour intensity from the hymenium, although a similar colour difference may sometimes be shown by reflexed parts of the hymenium. In M. crassipes and rare forms of M. lignaria (‘f. gomphillacea’) the excipular and hypothecial tissues become vertically extended to form a stipe (Figs 3E, 4B). Anamorphs (conidial states) With a few noteworthy exceptions, such as Lindsay (1859, 1872) and Gliick (1899), the conidial states (anamorphs) of lichenized fungi have received little detailed attention from taxonomists. Several recent monographic studies have shown that anamorphs can provide useful characters at various hierarchial levels of classification and the reader is referred to Vobis (1980) and Vobis & Hawksworth (1981) for further background information on the conidial states of lichens. Within the genus Micarea there is a diverse array of anamorphic forms, possibly unrivalled by any other genus of lichens, except perhaps for some of the genera in the Asterothyriaceae (Vézda, 1979). Information gained from the study of anamorphs has proved invaluable to me for the delimitation of species in Micarea; indeed, several species frequently occur without apothecia but with numerous pycnidia, such that a detailed knowledge of the latter is often essential for their identification (see ‘key to species without apothecia’). Conidiomata The conidiomata are usually pycnidial, and are globose, ovoid, doliiform, or ceriberiform in shape. They may be immersed (or partly so) within the thallus or substratum, sessile, or borne on stalks (pycnidiophores). When stalked, the pycnidia are usually + doliiform and the stalk-tissue is comprised of loosely interwoven hyphae bound by a gel matrix which is often pigmented. In addition, the ‘stalk-part’ often includes effete pycnidia (Fig. 35B—D). The stalks are sometimes branched due to the simultaneous development of two (or more) pycnidia at the 64 BRIAN JOHN COPPINS Table 1 Conidial states (anamorphs) found in European species of Micarea. Conidium — type Stalked Micarea micro- meso- macro- pycnidia adnata + ~ alabastrites anterior assimilata bauschiana botryoides + + cinerea + + contexta = ete crassipes + curvata denigrata te + elachista + + eximia + + + +++4++ globulosella Ge hedlundii incrassata + intrusa leprosula lignaria var. endoleuca lithinella lutulata + melaena + + melaenida + melaeniza melanobola misella muhrii myriocarpa + nigella + + + +++ +++ + i+ nitschkeana + olivacea osloensis peliocarpa + te prasina EF pycnidiophora rhabdogena Ss Stipitata subleprosula subnigrata os + subviolascens sylvicola synotheoides + ternaria tuberculata turfosa + +++4++4 +++ Total (species) Zo 28 9 9 apex of stalk tissue (Fig. 35F) or at the apex of an old pycnidium (Fig. 35D). Stalked pycnidia are apparently rare in other genera of crustose lichens (or related fungi). However, I have seen branched pycnidiophores associated with lignicolous Chaenothecopsis spp. growing on the moribund thalli and ascocarps of Chaenotheca spp. (several collections in E). LICHEN GENUS MICAREA IN EUROPE 65 Fig. 35 Micarea botryoides, conidiomata: diagram showing the various means of ‘stalk’ formation and branching (see text for further details). Scale = 100 um. The pycnidia of species which consistently lack pigment in their apothecia (e.g. M. adnata, M. alabastrites, M. pycnidiophora, and M. stipitata) correspondingly lack pigment in their wall tissues. In species with coloured apothecia the pycnidial walls are usually (at least in part) coloured also. For a given species the pigment involved is usually that found in the hymenium (or epithecium) of the apothecia, but in a few cases (e.g. M. lutulata) it is the pigment found in the hypothecium which is involved. The link between apothecial and pycnidial pigmentation helps distinguish pycnidia belonging to a Micarea from any intermixed pycnidia belonging to a non-lichenized or parasitic fungus. It further helps to distinguish the pycnidia belonging to the different species of Micarea in mixed populations; for example, the stalked pycnidia of M. misella (walls olivaceous, K+ violet) are often found intermixed with those of other species such as M. anterior (walls reddish brown, K—) and M. melaeniza (walls dark olivaceous, K—). When a pycnidium is deeply immersed in the thallus it is often only the uppermost part around the ostiole which is coloured, and likewise, if the pycnidium is semi-immersed it is often only the upper, exposed part which is coloured. With the exception of the species that are characteristi- cally devoid of pigment and some forms of other species in deep shade, the walls of sessile or stalked pycnidia are coloured more or less throughout. As mentioned above, the conidiomata of Micarea species are pycnidial, but there is one exception to this. M. adnata has two anamorphic states: one in which the conidia (mesoconidia) are produced internally in immersed pycnidia; and another where the conidia (macroconidia) are borne externally on cushion-like sporodochia which resemble small apothecia. This latter state could be considered to be a hyphomycetous anamorph but comparative ontogenetic studies are required to investigate the theoretical possibility that these sporodochia have evolved from pycnidial conidiomata by an exertion of the ‘hymenium’, and a thickening of the subtending wall-tissue to form a supporting cushion (Fig. 36). Unfortunately this hypothesis is not supported by intermediate forms in other species, although the pycnidia (with filiform or curved-hamate macroconidia) of, for example, M. cinerea and M. peliocarpa often have wideiy 66 BRIAN JOHN COPPINS Fig. 36 Suggested pathway for evolution of a sporodochium (D, as in M. adnata) from a pycnidium (A). The step AB can be seen in the macroconidial conidiomata of e.g. M. cinerea and M. peliocarpa, and the mesoconidial conidiomata of e.g. M. denigrata and M. sylvicola. No structure equivalent to ‘C’ is known from Micarea. gaping ostioles which eventually expose the ‘hymenium’. The often dubious distinctions between pycnidia, acervuli, and other types of conidiomata are discussed by Nag Raj (1981). Although the sporodochial anamorph of M. adnata has unusually large, oblong-ellipsoid conidia and the longest conidiogenous cells known in the genus, there seems to be no fundamental difference in these features from those in other Micarea anamorphs, and the mode of conidiogenesis is apparently the same in all cases (see below). Conidia Vobis & Hawksworth (1981) estimate that perhaps as many as 8000 species of lichenized fungi have conidial anamorphs, and that in most cases a given species has only one conidium type. However, it is becoming increasingly evident that a large number of lichens (especially crustose lichens) have two conidium types, although many of the examples have not yet been notified in the literature. Some examples from my own studies include Anisomeridium biforme, A. Juistense, Catillaria globulosa, Lecania cyrtellina, Lecanora quercicola, and Opegrapha niveoat- ra. Hedlund (1895) found M. denigrata and M. prasina to each have two conidium types and during the present study 16 of the 45 European species of Micarea have been found to have two conidium types. Even more surprising has been the discovery that three species (M. denigrata, M. nitschkeana, and M. lignaria) each have three conidium types (Figs 42, 45), and to my knowledge these are the first reported instances of fungi (whether lichenized or not) with more than two pycnidial (coelomycetous) anamorphs. There are, however, a few species of sooty LICHEN GENUS MICAREA IN EUROPE 67 mould (e.g. in the Metacapnodiaceae) which have three hyphomycetous anamorphs (Hughes 1972, 1976). The main distinguishing features of the three conidium types found in Micarea are as follows (see Figs 37-52 for example): Microconidia, narrowly cylindrical or narrowly fusiform, aseptate, eguttulate and not constricted in the middle, mostly in the range 3-8 x0-5-1 wm; produced in small immersed or + sessile pycnidia, mostly c. 20-60 wm diam. Mesoconidia, variable in shape, e.g. cylindrical, oblong, ovoid-oblong or obovoid-oblong, aseptate, often biguttulate and (or) slightly constricted in the middle; produced in small to large, immersed, sessile or stalked pycnidia. All conidia found in stalked pycnidia are included here. Macroconidia, mostly filiform or curved and often septate, or helicoid and septate (M. subnigrata (p. 183)); produced in medium-sized to large (often up to 200 wm diam or more) pycnidia. Also included here are the large, aseptate, eguttulate, oblong-ellipsoid conidia produced by the sporodochia of M. adnata. In all cases no single pycnidium has been found to contain more than one conidium type. A few specimens of M. denigrata and M. nitschkeana have been found with all three anamorphs on the same thallus; but such occurrences are rare and it is more usual to find just one or two of them. A few species (e.g. M. assimilata, M. crassipes, and M. incrassata appear to have only one anamorphic state, with conidia somewhat intermediate between microconidia and mesoconidia as defined above; thus the assignment of their conidia to one or other of these types (Table 1) must be considered tentative. The distinction between the three conidium types (especially micro- and mesoconidia) is most obvious when two or three of them occur on the same thallus. The use of the terms ‘micro-’, ‘meso-’ and ‘macroconidia’ are here applied solely to the conidium types found in Micarea-— they may not necessarily be analogous to their use in other lichenized or non-lichenized fungi. Table 1 indicates the conidium type(s) known for each of the European species of Micarea, and from this a numerical summary of the various known combinations is as follows: Combination of conidium types Number of species (anamorphs) (holomorphs) Micro- + meso- + macro- 3 Micro- + meso- 10 Micro- + macro- 5 Meso- + macro- 1 Micro- only 6 Meso- only 14 Macro- only 0 Anamorph(s) unknown 6 The role of each of the conidium types is as yet unknown. Vobis (1977) obtained successful germination and subsequent growth of mycelium from the macroconidia of Lecanactis abietina, but was unable to germinate the microconidia of the same species. His results suggest that the former may well act as asexual propagules, and the latter as spermatia in sexual reproduction. It seems most likely to me that the microconidia of Micarea species are spermatia. As to whether or not they are essential components of the sexual reproductive process is much less certain; 21 of the treated species are not known to produce microconidia. The role of conidia as asexual propagules is still a matter of much debate and speculation among lichenologists, but such a role (at least in certain cases) can be inferred from the fact that a few crustose species (e.g. Lecanactis subabietina and an undescribed Bacidia) are not known to have apothecia or vegetative diaspores (i.e. soredia, isidia, etc.) but always have numerous pycnidia. To such examples can be added the numerous crustose lichens whose apothecia are known, but which commonly occur as sterile populations with numerous pycnidia, e.g. Anisomeridium juistense (macroconidial state), Arthonia phaeobaea, A. spadicea, Bacidia arnoldiana, B. carneoglauca, B. trachona, Cliostomum graniforme, C. griffithii, Lecanactis abietina, Lecidea erratica, Opegrapha niveoatra, O. vermicellifera, and O. vulgata. This last group also includes Micarea botryoides, M. denigrata, M. misella, M. pycnidiophora, and M. stipitata, all of which commonly occur with abundant mesoconidia-containing pycnidia and few (if any) mature apothecia. 68 BRIAN JOHN COPPINS ok “AC i Fig. 37 M. adnata (E—holotype), macroconidia and conidiogenous cells. Scale = 10 um. LICHEN GENUS MICAREA IN EUROPE 69 — Fig. 38 A, M. botryoides (Coppins 8429, E), mesoconidia and conidiogenous cells. B, M. melaeniza (S - holotype), mesoconidia. C-D, M. anterior Sweden, Angermanland, Langsele, 1892, Hedlund, S); C, mesoconidia; D, microconidia. Scale = 10 um. BRIAN JOHN COPPINS 70 ‘WM CT = 9[B9S ‘S][99 SNOUSSOIpIUODS puke eIpluOsOIORW ‘(q ‘E¢ggz suiddo)D) vasauld ‘Pw 6¢ “Sl LICHEN GENUS MICAREA IN EUROPE a | 000 0008 0000 0000 pjo0000ot t— \ UM: Fig. 40 A-B, M. contexta (S —lectotype); A, mesoconidia; B, microconidia. C, M. eximia (Malme Lich. suec. exs. 26, C), mesoconidia. D, M. bauschiana (Coppins 4111, E), microconidia and conidiogenous cells. Scale = 10 wm. a BRIAN JOHN COPPINS ail o— ISO , wil Fig. 41 A-B, M. incrassata, microconidia; A, (S — holotype); B, (Kerguelen, 1875, Eaton, E). C, M. crassipes (Vézda Lich. Sel. 11, BM), (?)mesoconidia and conidiogenous cells. Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE v8: I j | oat Fig.42 M. denigrata (Coppins 1888, E), conidia with conidiogenous cells; A, macroconidia; B, mesoconi- dia; C, microconidia. Scale = 10 wm. 74 BRIAN JOHN COPPINS JOO 00 NN). «» Ode 1000 . 0000 0) 000 0000, VN. Fig.43 A-B, M. globulosella (Czechoslovakia, Slovakia, Vysoké Tatry, 1879, Lojka, BM); A, mesoconi- dia; B, microconidia. C, M. globulosella (S — lectotype), mesoconidia. D-E, M. synotheoides (Coppins 2942, E); D, mesoconidia; E, microconidia. Scale = 10 wm. a LICHEN GENUS MICAREA IN EUROPE 2 A000 a0a ce aml a be j0 UY Fig. 44 A, M. hedlundii (UPS — holotype), mesoconidia and conidiogenous cells. B, M. /utulata (Wales, Pembroke, Tycanol, 1980, James, BM), mesoconidia and conidiogenous cells. Scale = 10 wm. 76 BRIAN JOHN COPPINS ae C Fig. 45 M. lignaria. A, macroconidia (Coppins 4658, E). B, mesoconidia (Coppins 8436, E). C, microconidia (Coppins 8952, E). Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE we) is fz at N00000000 » Fig. 46 A, M. melaena (Coppins 6041, E), macroconidia. B—C, M. melanobola (H-NYL 2164 - lectotype); B, mesoconidia; C, microconidia. Scale = 10 wm. 78 BRIAN JOHN COPPINS i 00000 Ay Fig. 47 A, M. olivacea (BM — holotype), mesoconidia and conidiogenous cells. B, M. nigella (E - holotype), mesoconidia and conidiogenous cells. C, M. myriocarpa (Coppins 8939, E), mesoconidia and conidiogenous cells. Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE 79 Fig.48 MM. peliocarpa (England, New Forest, Great Wood, Bramble Hill Walk. 1970, Coppins et al., E). A, macroconidia and conidiogenous cells. B, microconidia and conidiogenous cells. Scale = 10 wm. 80 BRIAN JOHN COPPINS )0- QM N00) UND Fig. 49 A-B, M. prasina (GZU — holotype of M. polytrichi); A, mesoconidia; B, microconidia. C, M. prasina (Coppins 8009, E), mesoconidia. D, M. prasina (Coppins 2835, E), microconidia. Scale = 10 pm. LICHEN GENUS MICAREA IN EUROPE 81 Fig.50 M. subnigrata (Coppins 8417, E). A, macroconidia and conidiogenous cells. B, microconidia and conidiogenous cells. Scale = 10 wm. 82 BRIAN JOHN COPPINS Fig. 51. A, M. sylvicola (Muhr 2579, E), mesoconidia and conidiogenous cells. B, M. tuberculata (O - lectotype), mesoconidia and conidiogenous cells. Scale = 10 wm. LICHEN GENUS MICAREA IN EUROPE 83 NOUO ION: SOUCOUOOU » 10000000000 . JO0000U Fig. 52. A-B, M. ternaria, mesoconidia; A, (H-NYL 18682 — holotype); B, (Thomson 9188, DUKE). C, M. aff. ternaria (Fair Isle, 1976, Duncan, BM), mesoconidia. Scale = 10 wm. 84 BRIAN JOHN COPPINS In the case of M. denigrata it is a general rule (but with exceptions) that when on worked wood (fence-posts, garden furniture, window frames, etc.) its thallus has numerous pycnidia (with mesoconidia) and often few (if any) mature apothecia. On natural substrata (e.g. fallen tree trunks in old woodlands) pycnidia with microconidia or macroconidia are more prevalent, although they are usually relatively fewer in number and associated with numerous apothecia. It seems highly likely that the success of M. denigrata as a primary coloniser of newly available substrata is largely due to the successful role of its mesoconidia as asexual propagules. Its microconidia probably function as spermatia, but the function of the macroconidia is less obvious, although I suspect that they, like the mesoconidia, act as asexual diaspores (perhaps in a different way). Another observation pertinent to this discussion is that I have frequently observed the meso- and macroconidia of Micarea species being extruded through the ostioles of the pycnidia as white mucilaginous blobs; such phenomena are usually seen after periods of wet weather. It is tempting to suggest that these extrusions facilitate dispersal beyond the limits of the parent thallus by the action of rain or passing arthropods and mollusca. Dispersal by invertebrates may be of especial importance for species such as M. botryoides, which occur in sheltered situations rarely subjected to rain-wash. I have only rarely observed microconidia being extruded as white blobs, thus suggesting that there is no need for them to be dispersed beyond the limits of the parent thallus. Indeed, if microconidia do function as spermatia, and if the mycobiont of the Micarea is homothallic, there would be no requirement for them to be dispersed more than a few millimetres. Conidiogenous cells | The conidiogenous cells of Micarea species are always phialidic, although in many cases the phialides are seen to undergo ‘percurrent proliferation’ (Figs 42B, 44B). The conidiogenous cells belong to Types I or II of Vobis & Hawksworth (1981) and arise from the inner wall of the pycnidium (or on the outer surface of the sporodochium in M. adnata). Their subtending cells are never sufficiently regular in shape for them to be termed ‘conidiophores’. In several species whose pycnidial walls are intensely pigmented the bases of the conidiogenous cells are often similarly pigmented (Figs 47B, 51B). The nearest approach to the differentiation of wall-tissue from a conidiogenous layer is found in the thick-walled pycnidia of M. elachista. There is much variety in the shape of conidigenous cells (e.g. ampulliform, doliiform, lageniform, cylindrical) and there is sometimes much variation within a single pycnidium (Figs 42B, 44A). Conidiogenous cells with long cylindrical necks often have swollen bases (Figs 38A, 47B, 51B). Although critical observations and measurements have not been made for all species, the size and shape of conidiogenous cells have rarely been found to be useful characters for the separation of closely similar species. However, one exceptional case is that of saxicolous forms of M. olivacea (Fig. 47A) versus M. tuberculata (Fig. 51B), in which the conidiogenous cells of the latter are much longer. Chemistry The discussions below are presented under two sub-headings. The first deals with ‘lichen substances’, which are readily extractable in acetone and identifiable by thin-layer chromatogra- phy (t.1.c.; see ‘Methods’). The second part deals with pigments that cannot be analysed in this way; their chemical nature is at present unknown and they can only be characterised by their colour in water and subsequent reactions with reagents such as K and HNO3. Lichen substances Prior to the present studies there is little evidence in the literature to suggest that species of Micarea contain any lichen substances. However, there are a few hints given in some early descriptions: for example, Leighton (1879: 362) gives ‘K+ yellow, C+ orange-red’ reactions for Lecidea milliaria [Micarea lignaria], which probably relate to his specimens of the var. endoleuca. The only reported t.l.c. analysis of a Micarea appears to be that of Huneck & LICHEN GENUS MICAREA IN EUROPE 85 Table2 Chemical content and corresponding spot test reactions of European species of Micarea known to contain lichen substances. +, present; +, presence variable and sometimes absent; ?, present occa- sionally as trace amounts or contaminant; R, red; Y, yellow; fY, faint yellow; O, persistent orange; ( ) reactions not obtainable in many collections; *, including 3 or 4 accessory substances. See eS 2 2 FA ieee acs ic s fe) S = = os iS Bo eee Fe ek we ees Apothecia 3 S c § 2 8 & Thallus reactions reactions os 3 Fi » a. oo ar =4 K CRC” Pp G alabastrites 35 = R = R cinerea + = R = R curvata ate — R = R denigrata 1: = R = R globulosella ak use R aes R hedlundii ? = <= re = leprosula =e ee — R R R lignaria + = = R oa var. endoleuca + ty O _ = melaena + = R = oe misella as — (R) = (R) nitschkeana + —= R = R peliocarpa a — R _ R prasina s. str. ? 2 i a rae = prasina s. lat. 2 fe ms a= a — prasina s. lat. ¢ + = nae = — pycnidiophora + _ = eS R subleprosula + — R Y a Follmann (1972) who detected an unknown substance [= argopsin] in Bacidia [Micarea] lignaria [var. lignaria]. During my investigations a third of the 45 species of Micarea found in Europe have been shown to contain lichen substances (Table 2). Of the remaining 30 species the following have not been analysed by t.l.c. because the available specimens were too meagre: M. contexta, M. intrusa, M. lithinella, M. melaeniza, M. melanobola, and M. rhabdogena; however, spot tests on the thalli and apothecia in these specimens are all negative and the presence of lichen substances is unlikely, but not impossible. Eight principal lichen substances are so far known in Micarea: alectorialic acid (including three or four accessory substances — ? derivatives), a depside of the B-orcinol series; argopsin, a depsidone of the B-orcinol series; gyrophoric acid, a tridepside of the orcinol series; two xanthones (not yet positively identified); and three unknown substances each found in different chemical races of M. prasina. The relative positions of these substances on chromatograms using solvent systems TDA and HEF are shown in Figs. 53 and 54. Argopsin (1’ — chloropannarin) is found in M. lignaria var. lignaria and M. leprosula. It was originally isolated from Argopsis friesiana Mill. Arg. (belonging to the Stereocaulonaceae), and discussed at some length by Huneck & Lamb (1975). During the present investigation, and miscellaneous studies in collaboration with Mr P. W. James, it was found that this, then unknown, PD-+ red substance in M. lignaria was identical to that in Lecidea efflorescens (Hed1.) Vainio and Phyllopsora rosei Coppins & P. James. A sample of P. rosei was sent to Dr J. A. Elix who identified the unknown substance as argopsin. This substance was found in several other species of Phyllopsora by Swinscow & Krog (1981), who referred to it as ‘albicans unknown 2’. The two unidentified xanthones are found together in M. lignaria var. endoleuca; more ample specimens of this variety are required for further chemical studies. The most commonly encountered lichen substance in Micarea is gyrophoric acid, one of the 86 BRIAN JOHN COPPINS F F A Ga TDA ah ee K L 5k age av, S B 4a-@ 5 os z Al Q Q ED aL E2 Ww) Ww hh oO room 4 ~‘J ioe} oO Fig. 53 Diagram of chromatogram in solvent system TDA. 1, Control (Parmelia acetabulum plus Cladonia subcervicornis). 2, M. peliocarpa.3, M. subleprosula. 4, M. lignaria. 5, M. leprosula. 6, M. lignaria var. endoleuca. 7, M. prasinas. lat. 8, M. prasinas. str.9, M. prasina s. lat. A, atranorin, B, norstictic acid. C, fumarprotocetraric acid. D, gyrophoric acid. E, alectorialic acid, and accessory substances (E', E”). F, argopsin. G and H, xanthones. J, prasina unknown A. K, prasina unknown B. L, prasina unknown C. A re F 77, @ @- & , , L E @®m SS El D @e @ c> Q) 5 fr D E1 D B 4+ © oe | E2 3+ © Cc 2k me L n n r 1 1 4 re | 1 2 3 4 5 6 7 8 fe) Fig. 54 Diagram of chromatogram in solvent system HEF. Explanation as for Fig. 53. LICHEN GENUS MICAREA IN EUROPE 87 most widely occurring compounds among lichen genera. It has been found in 11 Micarea species, although its presence in M. curvata has only been inferred from spot tests, the solitary specimen (holotype) being too small for t.l.c. analysis. In addition to these occurrences, gyrophoric acid is frequently detected in trace amounts in chromatograms of other taxa, especially M. prasina s. ampl. The fact that Micarea species often grow intermixed with other lichens, including several that contain gyrophoric acid (e.g. Lecidea icmalea and L. granulosa agg.) raises the question as to whether these trace amounts represent its presence as an accessory substance or as a contaminant. This is difficult to answer. Gyrophoric acid is certainly found in the M. prasina complex, as it has been detected in large amounts in the type material of Lecidea levicula Nyl. from Cuba. (Further studies are required to establish the taxonomic status of L. levicula.) Well-developed specimens of M. denigrata, M. nitschkeana, and M. melaena have an areolate thallus containing readily detectable quantities of gyrophoric acid. However, the thallus of these species is sometimes scurfy-granular and blackish due to disruption by invading dematiaceous fungi and non-lichenized algae, and gyrophoric acid is produced in very low amounts or is apparently absent altogether (not detectable by t.].c.). The most surprising result of the chemical studies in Micarea was the discovery of three unknown, but very distinctive, compounds in M. prasina. The three compounds can be characterised thus (UV at 254 my): prasina unknown A. TDA 5: HEF 6, UV+ blue-white. After H,SO, and charring: UV+ dull orange-red, dull orange in daylight. prasina unknown B. TDA 5: HEF 6, UV + blue (less bright than ‘unknown A’). After H2SO,4 and charring: UV + vivid citrine-yellow, yellow (without orange tinge) in daylight. prasina unknown C. TDA 4-5: HEF 6-7 (slightly higher than ‘unknowns A and B’), UV+ grey or pale mauve (TDA) or + colourless (HEF). After H,SO, and charring: UV + violet-blue, + colourless in daylight but turning a pale pinkish-lilac after several weeks. In all the numerous European and North American specimens of M. prasina s. ampl. examined so far, these compounds have never been found in combination, and M. prasina exists as three distinctive chemical races (see taxonomic account of M. prasina for further discussion). None of the three compounds can be identified in the tabulations of Culberson (1972), and they have never been encountered in other genera during the numerous and diverse investigations by Mr P. W. James (pers. comm.). Samples of specimens containing ‘unknowns A and B’ are currently being studied by Dr J. A. Elix. The identification of lichen substances (even if only by spot tests) is essential for distinguishing sterile specimens of M. leprosula from M. subleprosula, and for separating the two varieties of M. lignaria; it is also of great value in the routine identification of many other species (see ‘Keys to species’). The three European races of M. prasina are not distinguishable by spot tests, but are readily identified by t.l.c. Pigments The nature of pigments and their location (especially within apothecia) are of prime importance in the delimitation of species within many lichen genera, particularly those included in the Lecideaceae, and Micarea is no exception. Little is known of these acetone insoluble pigments and a detailed knowledge of their structure and biogenesis would be invaluable for the evaluation of their taxonomic significance. On the basis of the colours seen in water mounts and the colour changes brought about by the application of a strong alkali (KOH) and a strong acid (HNO3) at least eight pigments can be recognized in Micarea. A very provisional summary of these pigments is given below: Pigment A. Green or aeruginose, K— or + green intensifying, HNO3+ red; in various tissues (according to species) of M. assimilata, M. bauschiana, M. cinerea, M. intrusa, M. lignaria, M. melaena, M. peliocarpa, M. sylvicola, and M. tuberculata. Pigment B. Purple, K+ green, HNO3+ purple-red; mostly in the hypothecium of M. assimilata, M. contexta, M. crassipes, M. eximia, M. melaena, M. nigella, and M. sylvicola (rare forms). Pigment C. Purple, K+ purple intensifying (sometimes partly dissolving into solution); occasionally in 88 BRIAN JOHN COPPINS the epithecium (M. melaenida and forms of M. crassipes and M. melaena) but mainly in the hypothecium (M. assimilata, M. crassipes, M. melaena, M. melaenida, and M. subviolascens). Pigment D. Olivaceous, K+ violet (also C+ violet), HNO3+ red; in various tissues (but rarely in the hypothecium, and then in low concentrations) of M. denigrata, M. elachista (mostly in pycnidia), M. globulosella, M. hedlundii, M. melanobola, M. misella, M. nitschkeana, M. prasina, M. subviolascens, and M. synotheoides. Pigment E. Fuscous brown, K+ dissolving into solution, HNO3-—; in the epithecia of M. elachista and M. rhabdogena. Pigment F. Brown (sometimes slightly tinged reddish), K— (not dissolving), HNO3— or + orange- brown; in various tissues (according to species) of M. botryoides, M. curvata, M. incrassata, M. lutulata, M. muhrii, M. myriocarpa, M. osloensis, M. subnigrata, and M. turfosa. It is quite possible that more than one pigment is involved here. Pigment G. Dilute yellowish, K+ purple (oily droplets), HNO3—; in goniocysts and sometimes the lower hymenium and hypothecium of M. hedlundii. Pigment H. Pale yellowish orange, K+ purple, HNO3~—; in cytoplasm of some ascogenous hyphae, asci, and spores of M. intrusa. This is possibly similar (or identical) to pigment G. Pigment complexes involving mixtures of pigments A, B, and C are often found (especially in the hypothecium) in M. assimilata, M. contexta, M. crassipes, M. melaena, M. subviolascens, and M. sylvicola; see the individual species accounts for further details. The existence of such complexes suggests that pigments A, B, and C are closely related chemically. Pigment A is probably identical to that found (usually in the epithecium) in a large number of lichens, especially in Lecidea s. lat., Catillarias. lat., Bacidias. lat. and Lecanora s. lat. Pigment C is probably of rare occurrence outside Micarea, but I know of it in the epithecium of Bacidia beckhausii and in several species of Pertusaria (e.g. P. hymenea and P. oculata). I have not encountered pigment E in any other lichens, but it should be compared with the pigment(s) responsible for the ionomidotic reaction found in several genera of non-lichenized discomycetes (Korf, 1973). Pigment F is probably of wide occurrence, but there may be several pigments that impart a brown or ‘melanized’ appearance in tissues. The dark violet-blue, K+ aeruginose granules occasionally seen in the hymenium of M. contexta and M. lignaria (and also the apothecia of Lecidea hypnorum and Dactylospora lobariella), is probably related to (if not the same as) pigment B, and it may be the same as the epithecial pigment found in Bacidia absistens, Mycoblastus fucatus, and Schaereria tenebrosa. Ecology No detailed ecological studies have been made in connection with this primarily taxonomic study of Micarea, and much of the discussion given below is based on floristic notes and casual field observations. For most amateur and professional lichenologists alike, the species of Micarea are little known, poorly understood, and much overlooked in the field. For this reason the following account is more of a guide to collectors, rather than an ecological dissertation. Additional ecological notes are also included in the taxonomic accounts of each of the Micarea species treated. Syntaxonomic nomenclature follows James et al. (1977). General habitats Micarea species occur in a wide range of habitats but are confined to substrata with a low pH (below c. pH 5) and generally avoid nutrient enriched situations. Thus, they are absent from limestone rocks, tops of bird-perching stones, seashores, and basic bark. There are a few partial exceptions, for example M. denigrata can occur on wooden fencing and other timber in farmyards, M. prasina can occur on soil and debris among rocks in the upper seashore, and M. lignaria has been found on exposed limestone growing over mats or cushions of moribund bryophytes from which free calcium ions have presumably been leached out. The bark of Acer and Ulmus normally has a pH which is too high for species of Micarea (with the exception of M. prasina in some situations), but in areas heavily polluted by sulphur dioxide and its derivatives (e.g. West Yorkshite Conurbation), bark pH can be substantially lowered (Gilbert, 1970), thus providing a suitable substratum for species such as M. melaena (q.v.) and M. botryoides (q.v.). LICHEN GENUS MICAREA IN EUROPE 89 The principal habitats (I-XI) in which Micarea spp. are found are given below; each is provided with a list of the species which can be expected to occur in it (allowing for climatic and phytogeographical variations). Species given in square brackets [ ] are rarely and unusually found in the given habitat, and those prefixed with a dagger (+) are not known in this habitat from the British Isles. I. Sheltered, dry underhangs, usually in valley woodland or narrow ravines, including dry undersides of up-ended tree root systems in woodland; growing on rock, loose stones, consolidated soil, and exposed roots. M. bauschiana M. myriocarpa M. botryoides M. sylvicola M. lignaria M. tuberculata M. lutulata II. Exposed parts of sheltered rocks, usually in woodland; more frequently wetted than I; growing directly on rock. M. botryoides M. sylvicola +M. curvata [M. cinerea] M. lignaria s. lat. [M. denigrata] M. lithinella +[M. muhrii] M. melaena [M. nitschkeana]* M. olivacea [M. prasina] M. peliocarpa a, stones amongst Calluna in heathland. Ill. Over bryophytes on rocks, boulders, old stumps, or fallen trees in woodland at low altitudes (500 m in the British Isles). M. adnata M. lignaria s. lat. M. botryoides M. melaena M. cinerea M. peliocarpa M. leprosula M. prasina [M. stipitata] IV. Exposed, hard siliceous rocks; growing directly on rock. M. intrusa +M. subviolascens M. lignaria M. aff. ternaria® M. subnigrata b, coastal districts only. ¥. Over bryophytes or peaty debris on exposed turf or on, or amongst, rocks and boulders in open situations in upland, montane, or ‘arctic’ districts. M. assimilata M. peliocarpa +M. crassipes M. subleprosula M. incrassata +M. ternaria M. leprosula M. turfosa M. lignaria s. lat. [M. cinerea] M. melaena VI. Over bryophytes or plant debris on the ground in old dunes, disused lead and zinc mines, sea-cliffs, or by woodland tracks. M. botryoides M. prasina M. leprosula [M. denigrata] M. lignaria s. str. +/M. misella] M. peliocarpa VII. | On bare mineral soil at low altitudes. +M. melaenida [M. lignaria] +M. osloensis [M. prasina] [M. leprosula] VIII. Corticate trunks of healthy trees; usually in woodland; sometimes overgrowing bryophytes (*). *M. alabastrites *M. pycnidiophora *M. cinerea *M. stipitata 90) BRIAN JOHN COPPINS +M. elachista *M. synotheoides +M. globulosella [M. botryoides] M. melaena [M. denigrata] +M. melanobola [M. leprosula] *M. peliocarpa [M. lignaria] *M. prasina [M. nitschkeana] IX. On attached twigs of trees or large shrubs, or thin stems of small shrubs; in woodland, heathland, or scrub. +M. cinerea M. peliocarpa M. lignaria s. str. M. prasina M. nitschkeana [M. denigrata] X. On timber (i.e. worked wood), e.g. fencing, garden furniture, old window frames, and shingles. M. cinerea M. nitschkeana M. denigrata M. peliocarpa +M. elachista [M. globulosella] M. lignaria s. str. [M. leprosula] M. melaena [M. sylvicola] M. misella XI. Directly on lignum of old stumps and decorticate trunks. M. adnata M. misella M. alabastrites +M. muhrii +M. anterior M. nigella M. cinerea M. nitschkeana +M. contexta M. olivacea M. denigrata M. peliocarpa +M. elachista M. prasina +M. eximia +M. rhabdogena +M. hedlundii [M. botryoides] M. lignaria s. lat. [M. leprosula] M. melaena [M. myriocarpa] +M. melaeniza [M. synotheoides] From the above lists it will be seen that several species (e.g. M. lignaria, M. melaena, M. peliocarpa, and M. prasina) inhabit a very wide range of habitats and substrate. At the other extreme there are many species which are much more restricted e.g. M. tuberculata (1), M. subnigrata (IV), M. assimilata (V), M. melaenida (VIII), M. pycnidiophora (VIII), and M. anterior, M. contexta, M. eximia, and M. melaeniza (XI). In most of the communities in which they occur Micarea species are usually of minor importance with regard to cover values. However, there are some exceptions such as the Micareetum sylvicolae association of underhangs and exposed tree root-system (see below), the community dominated by M. prasina on trunks in dense conifer plantations, some lignicolous assemblages on fallen trunks, old stumps, and worked wood. Micarea nitschkeana on Calluna twigs and litter in some lowland heaths, M. melaena on sandy or peaty soil in some heathlands and moorlands, and M. lignaria on the ground in some old lead mine workings. Specific habitats Deciduous (broad-leaved) woodland In Britain the genus Micarea is best represented in terms of number of species per site in the mature, + natural woodland (both deciduous and coniferous) on acid soils, in areas with a high annual rainfall (at least 1000 mm distributed over at least 160 ‘wet days’; see Coppins, 1976). The best examples of ‘Micarea-rich’ deciduous (broad-leaved) woodlands are found in Wales, the English Lake District and the west of Scotland north of the Clyde Estuary. The Micarea species found in them on trees and stumps are included in lists HII, VII, and XI above, except that M. misella, M. nigella, and M. olivacea are primarily species of coniferous woodlands. The occurrence of Micarea spp. on bark (or over bryophytes thereon) is favoured by leaching, and is further favoured, to the detriment of more basicolous lichens, by the effects of ‘acid rain’ LICHEN GENUS MICAREA IN EUROPE 91 (Anon, 1980; Fowler et al., 1982). Acid rain effects, resulting from the pollution emitted from the industrial areas of west-central Scotland (Clydeside and Glasgow conurbation), are prob- ably the explanation for the prevalence on the trunks of mature trees (e.g. Quercus, Alnus, Betula, and Fraxinus) of such species as M. alabastrites, M. cinerea, M. peliocarpa, M. stipitata, and M. synotheoides in the deciduous woodlands of the Cowal Peninsula in southern Argyll- shire. The communities in which these species occur are probably referable to the Parmelietum laevigatae, and in areas subjected to ‘acid rain’ it appears that these communities have to some extent, and in certain situations (especially on Quercus and Fraxinus), replaced the more basicolous communities of the Lobarion pulmonariae. In the deciduous woodlands of drier parts of the British Isles (annual rainfall of <1000 mm over <160 ‘wet days’) Micarea species are less prevalent on the trunks of healthy trees, and M. prasina is the only species commonly encountered, although M. nitschkeana can often be found on twigs and thin branches. However, M. peliocarpa is occasionally found growing amongst (but not on) mosses on the bark of old Quercus trunks in ancient woodlands, especially in the New Forest, Hampshire. Similar occurrences of M. peliocarpa are known from the old oak-woods of north Jylland in Denmark, and north-west France. Coniferous woodlands The formerly extensive native Caledonian pine-forest of Scotland is now reduced to a few scattered relics totalling about 10050 ha (26000 acres), of which only about 1620 ha (4000 acres) is represented by dense stands of Pinus sylvestris (Steven & Carlisle, 1959; Goodier & Bunce, 1977). These woodlands cannot be described as ‘natural’ because they have all been managed commercially to some extent during the last 200 years. Nevertheless, most of them contain many old trees, together with standing and fallen decorticate trunks, and are rewarding sites for the seeker of Micarea species. From the analysis of various vegetational and edaphic characteristics, the Scottish native pinewoods can be divided into four ‘site types’ (Bunce, 1977). The Micarea species recorded by me from some of the pinewoods in three of these site types are listed in Table 3. This indicates a poor Micarea flora for the eastern site type which is probably due to their lower rainfall. However, it is in these eastern (Speyside and Deeside) pinewoods where special efforts should be made to search for the group of lignicolous species (M. anterior, M. contexta, M. eximia, M. melaeniza, M. muhrii, and M. rhabdogena) which are so far known only from the coniferous forests of Scandinavia (especially mid-Sweden). To these can be added the more widely distributed M. elachista and M. hedlundii. An ever increasing proportion of British woodland (especially in Northumbria and Scotland) is composed of densely planted evergreen conifers such as Picea sitchensis, P. abies, Pinus sylvestris, and to a lesser extent other species including Abies grandis, A. procera, Pinus contorta, and Pseudotsuga menziesii. In some old plantations (70+ years) in the central and western Scottish highlands species of Micarea (including M. alabastrites, M. cinerea, M. peliocarpa, M. stipitata, and M. synotheoides) may colonise the trunks of mature trees, particularly in less-shaded situations by forest tracks and fire-breaks. However, most of the commercial forests contain closely planted trees less than 30 years old, and the light regime in such forests is so poor that it + prohibits the development of both a ground flora and an epiphyte flora of tree boles. The most successful lichen in these deeply shaded situations is the ubiquitous M. prasina which occurs in them as a form with a thinly developed thallus and small, whitish (translucent when wet) apothecia; in this form it is easily overlooked as a thin dark green algal-scum covering trunks and fallen branches. Old stumps in dense plantations sometimes also support M. melaena and the newly described M. nigella (q.v.). Larix is another frequently planted conifer, but because it is deciduous, its trunks are soon colonised by photophilous communities of the Pseudevernietum furfuraceae and Usneion barbatae. These communities are dominated by foliose and fruticose lichens, thus leaving little available space for crustose species, although species such as M. melaena, M. peliocarpa, and M. prasina are sometimes present in small quantities. On the other hand, the long slender twigs of Larix are commonly colonised by M. nitschkeana, often accompanied by Scoliciosporum chlorococcum, so forming communities referable to the Bacidietum chlorococcae. 92 BRIAN JOHN COPPINS Saxicolous habitats Of the 45 European species of Micarea, 21 have been found growing directly on rock (lists I, II, and IV), although the genus is poorly represented in the main saxicolous lichen alliances of exposed siliceous rocks (e.g. Lecideion tumidae, Rhizocarpon alpicolae, and Umbilicarion cylindricae; list 1V). As saxicoles Micarea species are more prevalent in sheltered ravines and woodlands. In dry underhangs in rock faces, steep banks, and below overhanging trees, up to seven species (list I) may be found in the ombrophobous, aerohygrophilous Micareetum sylvicolae. The species in this community grow on rock, loose or lodged stones, exposed roots, consolidated soil, and encroaching dry mats of bryophytes. Associated lichens from other genera may include Coniocybe furfuracea, Enterographa hutchinsae, Melaspilea subarenacea, Microcalicium arenarium, Opegrapha gyrocarpa, O. zonata, Porina chlorotica, P. lectissima, Psilolechia clavulifera, and P. lucida. This community is usually well defined, but in some situations it intergrades with other assemblages of shaded rocks such as the Lecideetum lucidae, Coniocybetum fururaceae, Opegraphetum horistico-gyrocarpae, and the Racodietum rupestris. A few Micarea species (list II) may be found on the upper sides of stones, boulders, and rocks where they are subjected to direct wetting by rain, or by drips from the overlying tree canopy. In most cases these are incidental occurrences of species more characteristic of other substrata. However, such situations may be the normal habitat of the little known M. curvata and M. lithinella. In northern and western Britain several Micarea species occur on mossy rocks. The communi- ties involved are often difficult to place, some belonging to bryophyte-dominating syntaxa, others to normally epiphytic communities (e.g. Parmelietum laevigatae), or to chomophytic communities dominated by bryophytes and Cladonia species which, in turn, are often invested by a gelatinous algal scum. Terricolous habitats Areas of lowland heathlands where the peaty or sandy soil has been laid bare of tall vegetation by erosion, disturbance, or burning are colonised by several species of algae bryophytes and lichens. The lichens involved often include species such as Lecidea icmalea, L. uliginosa, L. oligotropha (rare in Britain), Baeomyces rufus, B. roseus, and some Cladonia spp. Micarea species are not usually involved in such communities, although M. melaena may at times be present, occasionally achieving local dominance. In mature Callunetum in the heathlands of eastern England and Jylland (Denmark) M. nitschkeana is sometimes encountered in abund- ance on litter, as well as on the thin attached twigs of the Calluna. Acidic soils contaminated by heavy metals (especially lead) are often rather bare with an open vegetation, and Micarea species (especially M. lignaria) may occur in quantity growing over moribund bryophytes and plant debris. Soil, plant debris, and moribund bryophytes amongst coastal rocks are often colonized by M. prasina (q.v.). M. melaenida (q.v.) is found exclusively on consolidated fine grained (argillaceous) mineral soils. The terricolous species encountered in upland, montane, or arctic regime are given in list V; the reader is referred to the individual species accounts for further details. Man-made substrata A detailed treatment of the lichens of man-made substrata is given by Brightman & Seaward (1977). Several Micarea species occur on worked wood (list X), but M. denigrata (q.v.) is by far the most successful species. M. denigrata has also been found on pieces of hardboard lying in a dune slack at Tentsmuir in Fife. Old sackcloth and other fabrics lying on the ground in old lead-mine workings are frequently colonised by lichens, including M. lignaria. A more surpris- ing find was that of M. nitschkeana, growing with Scoliciosporum umbrinum, on a small plastic carton in a heathland on the Isle of Wight. Further field studies will undoubtedly extend this list of artificial substrata. Micarea denigrata and M. prasina are rapid colonisers of newly available substrata, and have been collected respectively on dead culms of Cladium mariscus and Phragmites australis in natural habitats. Both of these reeds are sometimes used as thatch, and a close inspection of thatched roofs ought to reveal the presence of Micarea species. LICHEN GENUS MICAREA IN EUROPE 93 Habitats in lowland pastoral areas The lowlands of eastern England, the English south midlands and parts of eastern Scotland contain few ‘natural’ habitats suitable for Micarea species. In addition, these same areas suffer, at least to some extent, from the effects of incoming pollution (especially sulphur dioxide and its derivatives) and indigenous pollution in the form of agricultural biocides and fertilisers. Nevertheless, the diligent collector will usually discover suitable niches where Micarea species are to be found. The most commonly encountered member of the genus in lowland agricultural and suburban districts is M. denigrata which can grow on worked wood in a wide range of situations, and on old tree stumps in hedgerows. M. prasina can be found on trees and shrubs in small woodlands planted as fox coverts or for the rearing of pheasants. It may also be found on sheltered stems of shrubs in waste ground, marginal land, or gardens, where M. nitschkeana may also occur. A major refuge for wildlife (including lichens) in lowland Britain is provided by churchyards (Anon, 1973, 1978). Although records of Micarea species are rather few, M. denigrata can often be found on timber and, more rarely, on siliceous memorial stones; and M. lignaria and M. peliocarpa have been recorded on the siliceous stone-work of old walls and memorials. Distribution Britain Maps 1-28 present the distributions of all the Micarea species occurring in the British Isles, with exception of M. lithinella which is known from just one 10 km grid square (44/86). It is seen from these maps that records for most species are concentrated in the western and/or northern districts. This is partly due to the fact that suitable habitats are most numerous in these areas, and it is probable that several species (e.g. M. bauschiana, M. lignaria, M. melaena, M. peliocarpa, and M. sylvicola) would be more evenly distributed if there were more available habitats in the lowland areas of the south and east. It is impossible to discuss in isolation the effects of climate or substrate availability on plant distributions. Substrate availability is largely dependent on topography and geology, but the physical and chemical nature of an existing substratum can be much influenced by the prevailing climate. For example, the bark of trunks of mature Quercus in high rainfall districts tends to be more leached, soft, and friable (i.e. more suitable for Micarea spp.) than in low rainfall districts. As a second example the species of the Micareetum sylvicolae (see p. 92) are ombrophobous (avoid frequent direct wetting), but are also aerohygrophilous (require + constant high humidity). In western Britain the high rainfall, high incidence of cloud cover (i.e. low duration of possible sunshine), and incoming moist air-stream from the Atlantic on the prevailing westerly winds result in high levels of relative humidity (and low levels of saturation deficit) more or less throughout the year (see Climato- logical Atlas of the British Isles, London: HMSO (1952)). Under such conditions a well developed Micareetum sylvicolae can be found on underhangs in a wide range of lowland habitats. Eastern districts generally have a lower rainfall and lower incidence of cloud cover resulting in lower levels of relative humidity, such that the Micareetum sylvicolae tends to be more confined to narrow river valleys with a long history of continuous tree cover. A western bias to the present day British distribution of many lichens is often due to the fact that the eastern side of the country is generally that most affected by industrial and urban development, intensive agricultural practices, and the resultant forms of air pollution (Hawksworth, Coppins & Rose, 1974; Coppins, 1976). A climatically determined western distribution, attributable to the General Western Group of Coppins (1976), is shown by Micarea adnata, M. alabastrites, M. cinerea, M. lignaria var. endoleuca, M. stipitata, and M. synotheoides. These species have a eu-Atlantic or sub-Atlantic European distribution, and are confined to areas with an annual rainfall of over 800 m (mostly over 1000 mm) distributed over at least 160 wet days [‘wet day’ = period of 24 h in which 1 mm (or more) of rain is recorded]. The genus Micarea is poorly represented in southern (Mediterra- nean) Europe, and it is not surprising that few British species exhibit a marked southern tendency. The best example of such a species is M. pycnidiophora, which is mainly confined to 94 BRIAN JOHN COPPINS Table 3 Distribution of Micarea species in some of the site types of Scottish native pinewoods. SITE. TYPE SW C E Annual rainfall (mm) 2000-2800 1300-1800 760-1150 lignaria var. endoleuca + ct a Stipitata + — =e adnata + a ae alabastrites + fe = lignaria var. lignaria + + = synotheoides a + + melaena + = a peliocarpa ~ + + prasina + + + cinerea ad ab: 12. nigella — + = nitschkeana — es a misella — ae Es Site types are err to Bunce (1977). The woods studied (with 10 km grid-references) are as follows. South-western (SW): Barisdale (18/80); Black Mount Woods (27/24 & 27/34). Central (C): Black Wood of Rannoch (27/55); Glen Affric (28/12 and 28/22); Glen Moriston (28/31); Glen Strathfarrar (28/33); Guisachan Forest (28/22). Eastern (E): Abernethy Forest (38/01); Rothiemurchus Forest (28/90). the New Forest, and its British distribution reflects its overall southern eu-Atlantic distribution, although it appears to avoid areas with an extremely high rainfall. A markedly eastern distribution is not clearly exhibited by any of the well known British species, although M. nitschkeana appears to occur with greater frequency and in greater abundance outside of the predominantly western areas with a very high rainfall. Amongst the species of low altitude habitats (mostly <500 m) a distinctly northerly distribu- tion (not related to high rainfall) is best exemplified by M. misella. The arctic-alpine species of Micarea have a northern distribution, and are apparently confined to Scotland. One possible exception is the little known M. subleprosula with its single British locality in north Wales. However, this species has probably been overlooked, and I believe it to be just a matter of time before it is recorded from Scotland. The most widely and evenly distributed species of Micarea in Britain are M. denigrata and M. prasina; both are efficient colonisers of a wide variety of newly available substrata, even in areas of suburban development and intensive agriculture. The dearth of records of common species in Ireland is due to under-recording. Europe An accurate assessment of distribution patterns exhibited by such little known and poorly understood organisms as members of the genus Micarea is likely to be an impossible task for many years. Distributions can be easily misinterpreted due to uneven’ recording. Many large areas may be underworked (or not studied by all) by collectors familiar with the group concerned, and other areas may be, by contrast, intensively studied. Consequently, when analysing the resultant records there is always the danger of artificially creating ‘centres’, ‘headquarters’ or trends of distribution. This state of recording is applicable at the present time to Micarea species, which are best known in mainland Britain, the southern halves of Norway, Sweden, and Finland, southern Bavaria and the Schwarzwald (Black Forest), Austria, the Italian Tirol, and the Sudety and Carpathian Mts of Czechoslovakia. It is probably safe to generalise that these areas have, coincidentally, the highest concentrations of Micarea species. However, field studies are required to determine which Micarea species are present in other potentially rich areas which are as yet little studied. These areas include Bretagne, south-east France, the Pyrénées, Cantabrian Mountains, northern Apennines, Dinaric Alps, southern Carpathians, Transylvanian Alps, Balkan Mountains, and the Rhodope Mountains. LICHEN GENUS MICAREA IN EUROPE 95 The identification (and subsequent annotation and cataloguing) of a single specimen of Micarea (which more often than not is fragmentary or in poor condition) can be very time consuming, and often involves lengthy microscopical examination. The unfortunate conse- quence of this is that I have not been able to examine a large number of potentially available specimens from several major institutional and private herbaria. Despite the above problems and shortcomings, I have tentatively attempted below to assign the species of Micarea to some general distributional types. In most cases I have paid little attention to purely literature sources of records because I have found that many such reports, even those by lichenologists whose works I hold in the highest esteem, can be unreliable in respect of current species concepts. A. Eu-Atlantic — mainly confined to oceanic areas including west Norway, the British Isles, western France, and Macaroenesia. M. alabastrites M. Stipitata M. nigella M. subnigrata M. olivacea +M. subviolascens *M. prasina (with ‘unknown C’) M. synotheoides *M. pycnidiophora *with southern tendency fwith affinity to group G B. Sub+Atlantic — as above, but also present in high rainfall (annual rainfall of >800 mm) areas of central Europe (Alps, Carpathians, etc.). M. adnata M. lignaria var. endoleuca M. cinerea i. Boreal — mainly confined to Fennoscandia, but absent or rare from west Norway. M. anterior M. melanobola M. contexta M. muhrii M. eximia M. osloensis M. melaeniza M. rhabdogena D. Boreal-continental — present in Fennoscandia and central Europe, but rare or absent from west Norway and western Britain. M. elachista M. lithinella M. hedlundii *M. melaenida *but known from western France E. Continental — known only from central Europe. M. curvata F. Montane (arctic-alpine) — at high altitudes in montane regions of Britain and central Europe, but sometimes at low altitudes in Fennoscandia. M. assimilata M. subleprosula M. crassipes M. turfosa M. incrassata G. Arctic — mainly confined to within the Arctic Circle, but possibly extending southwards along the Atlantic coast. M. ternaria H. Widely Distributed — known from British Isles, Fennoscandia, central Europe, and often else- where. More records will probably reveal distinct phytogeographical tendencies for some of the species included here. M. bauschiana M. melaena M. botryoides +M. misella M. denigrata M. myriocarpa tM. globulosella M. nitschkeana M. intrusa tM. peliocarpa M. leprosula *M. prasina M. lignaria var. lignaria M. sylvicola M. lutulata M. tuberculata *races containing ‘prasina unknowns A and B’ twith possible affinity to group D twith possible affinity to group B 96 BRIAN JOHN COPPINS World The present study is confined to species of Micarea that occur in Europe. However, some extra-European specimens have been examined in connection with nomenclatural matters, and others have been examined incidentally. According to current information the genus is best represented in Europe, but this may, or may not, be true. Micarea is well represented in North America from where I have seen 12 species (all European taxa: M. crassipes, M. denigrata, M. globulosella, M. lignaria s. str., M. melaena, M. misella, M. nitschkeana, M. peliocarpa, M. prasina, M. sylvicola, M. ternaria, and M. turfosa); this list will undoubtedly be much extended in the near future. From other regions I have seen (but not necessarily critically examined) specimens of Micarea from Japan, Borneo, Tasmania, New Zealand, South Africa, South America (Brazil), and the Antilles (Cuba). It is likely that species of Micarea are to be found in most temperate and boreal regions, as well as in many tropical regions (especially mountainous areas). The numerous collections that I have received from New Zealand include several undescribed taxa but they also include at least one European species, Micarea peliocarpa. Three additional European species known from the southern hemisphere are M. incrassata (Ker- guelen), M. lignaria (Brazil), and M. misella (Brazil). The genus Micarea Micarea Fr. Syst. orb.: 256 (1825). — Micarea Fr. emend Hed. in Bih. K. svenska VetenskAkad. Handl. II, 18(3): 27 (1892). Lectotype: Micarea prasina Fr. (see note below). Helocarpon Th. Fr., Lich. arctoi: 178 (1860); Nova Acta R. Soc. Scient. Upsal. II, 3: 278 (1861). Type species: Helocarpon crassipes Th. Fr. [= Micarea crassipes (Th. Fr.) Coppins]. Stereocauliscum Nyl. in Flora, Jena 48: 211 (1865). Type species: Stereocauliscum gomphillaceum Nyl. [= Micarea lignaria (Ach.) Hedl.] Micarea sect. Bryophagae Poelt & Débbeler in Bot. Jb. 96: 337 (1975). Type species: Micarea polytrichi Poelt & Débbeler [= Micarea prasina Fr.]. Note. The name Micarea was first validly published in December 1825 by Fries (op. cit.), although it was twice mentioned by the same author earlier in the same year. In Sched. crit. lich. suec. part 3, fasc. 4, page 21 (pre 7 May) it appeared in a note under the entry for Biatora fuliginea. However, it was not accepted in that work, and must therefore be considered invalid according to Art. 34.1. The second appearance was in Stirp. agri. femsion. page 37 (June 1825) in the form of the combinations ‘Micarea fuliginea’ and ‘M. nigra’. Neither of these was accompanied by a description and the generic name was again invalidly published (Arts 32, 34.1(e)). When validly published, Micarea was introduced with four species: M. prasina, M. coccinea, M. fuliginea, and M. nigra. Kérber (1855) emended the genus to include only M. prasina and is thereby considered to have lectotypified the genus on that species. See ‘Excluded taxa’ for notes on the remaining three original names. Thallus crustose or immersed in the substratum, effuse and often wide-spreading, never bordered by delimiting hypothalline lines. Superficial thallus consisting of + spherical granules (goniocysts) up to c. 60 +m diam; or convex to subglobose areolae which in some species may dissolve into soredia: or more rarely present as a thin + smooth to rimose, or scurfy-granular crust. Thallus in section ecorticate (or weakly corticate in a few species), but sometimes (when areolate) covered by a hyaline amorphous layer, and outermost hyphae sometimes pigmented. Phycobiont ‘grass-green’; cells usually thin-walled and c. 4-7 wm (‘micareoid’), or more rarely larger (‘non-micareoid’ phycobiont types); a few species also have cephalodia containing Nostoc or Stigonema. Apothecia small (mostly <1 mm diam), whitish or variously coloured (mostly grey, dull brown or blackish), epruinose, usually immarginate or + so, adnate, sessile or rarely stipitate, convex to + globose and often becoming tuberculate. Hymenium with amyloid gel-matrix. Asci clavate or cylindrical-clavate, of Lecanora-type, 8-spored. Spores hyaline, smooth-walled, variously shaped (ellipsoid, ovoid, fusiform, or acicular), usually less than 6 wm wide, simple to multiseptate but never muriform. Paraphyses few to numerous, septate, mostly branched (especially above), often anastomosing, mostly in range of 0-7-1-7 wm wide at mid-hymenium; LICHEN GENUS MICAREA IN EUROPE 97 apices never regularly clavate or capitate and never with a dark brown apical ‘cap’, but sometimes irregularly incrassate and sometimes with thickened pigmented walls in about upper 5-15 wm; in addition, some species have a few stout paraphyses (which are c. 2-3 wm wide, and, in some cases, pigmented throughout their length) occurring as scattered individuals or small fascicles. Hypothecium (including subhymenium) variously pigmented, of interwoven hyphae that become outwardly orientated towards the hymenium, mixed with wider, short-celled ascogenous hyphae. Excipulum often absent or indistinct, if discernible then non-amyloid (or + so) and composed of radiating branched and anastomosing paraphysis-like hyphae that become distinct and + separate in K. Pycnidia often present, very varied in form from immersed to sessile or stalked, the stalks (pycnidiophores) sometimes branched. Pycnidial walls hyaline or pigmented. Conidiogenous cells ampulliform to cylindrical (sometimes with swollen base), phialidic, sometimes with 1-3 proliferations. Conidia hyaline, smooth-walled, of three basic types: (i) microconidia — + cylindrical, aseptate, eguttulate, in range 3-5-9x0-5-1 um, borne in immersed to sessile pycnidia usually heer al ae : a | ie | A | A ) Ze, ih | A 7 aa see ae 2 ie j NGS ae eq | it | | Be =a a ake rai e one ao Pia, i yD ' Map1 = Micarea adnata @ 1950 onwards © Before 1950 Pycnidia usually present, white with hyaline walls, of two types: (a) immersed in areolae, c. 100-220 um diam, ostiole often widely gaping; conidia (macroconidia) markedly curved, often sigmoid, sometimes faintly 3—-S-septate, 21-55xc. 1 wm; (b) + sessile, c. 50-100 wm diam, ostiole not, or only slightly gaping; conidia (microconidia) narrowly fusiform-cylindrical, 5-7 x0-5-0-7 wm. Chemistry: Thallus and apothecia C+ red; t.l.c.: gyrophoric acid. Observations: M. alabastrites is morphologically and chemically very similar to M. peliocarpa, and it could easily be dismissed as representing a pale, shade form of that species. Apart from the complete absence of pigment from all its tissues, M. alabastrites is subtly different from M. peliocarpa in having slightly larger apothecia and spores. Shade forms of M. peliocarpa are usually sparingly fertile with scattered apothecia, whereas the apothecia of M. alabastrites are almost invariably numerous, crowded and often confluent. My early doubts regarding the distinction of these two species were dispelled by the collection, on several occasions, of M. alabastrites with adjacent thalli of M. peliocarpa with blackish apothecia; the slight differences in the sizes of apothecia and spores were confirmed with these collections. The two species also differ somewhat in distribution and habitat; M. peliocarpa is widely distributed throughout much of Europe on a wide range of substrata (tree trunks, mossy rocks peaty soil etc.); whereas M. alabastrites has a hyperoceanic distribution and occurs on tree trunks (or on bryophytes thereon) and occasionally lignum, but apparently never on mossy rocks or peaty soil. In the recent checklist of British lichens (Hawksworth et al., 1980) I mistakenly placed 132 BRIAN JOHN COPPINS Table 4 Diagnostic features for the separation of Micarea alabastrites, M. cinerea, and M. peliocarpa. peliocarpa alabastrites cinerea Apothecia size (mm) 0-14-0-4(-0-6) 0-2-0-7 0-2-0-7 Apothecia colour pallid to black whitish or pallid pallid to black Hymenium height (um) 40-55 45-55 55-70 Spore septation (1-)3(-S) 3(-7) (3-)7 Spore length (um) (11-)15—23(-24) (16—)18—26(-29) (19-)23-34(-38) Spore breadth 3-5(-6) (4-)4-5-S(-6) 4-5-6 Macroconidia shape curved-sigmoid curved-sigmoid flexuose Macroconidia length (4m) 21—40(-S0) 21-55 50-110 Thallus colour greenish white to greenish white or greenish white to dark blue-grey pale green dark blue-grey ‘alabastrites’ as asynonym of M. cinerea. At that time I had only seen the fragmentary isotype of Lecidea alabastrites in BM; I found this to have a few 7-septate spores and considered it a juvenile, shade-form of M. cinerea. Subsequent examination of the holotype (a large, healthy specimen) proved me to be wrong, and that it actually belonged to a hyperoceanic species which I had intended to describe as new to science. M. cinerea is another close relative of M. alabastrites and is more prone to occur with + pallid apothecia than M. peliocarpa. It differs from both these species in having spores which are mostly 7-septate at maturity, and macroconi- dia which are flexuose and much longer; for a comparison of these three species see Table 4. Occasional 7-septate spores have been found in several specimens of M. alabastrites, but they never number more than two or three in each squash preparation. M. alabastrites is one of the several lichens, named ‘Bacidia (or Bilimbia) sphaeroides’ by British lichenologists. This name is based on Lichen sphaeroides Dickson, which is a species of Biatora Fr. (non Ach.) and currently known as Catillaria sphaeroides (Dickson) Schuler. The name Bacidia sphaeroides as commonly used by Scandinavian workers refers to a species (not known in Britain) which also belongs in Biatora; its correct name (basionym) is probably Bilimbia tetramera de Not. Habitat and distribution: In the British Isles M. alabastrites is mainly found in communities of the Parmelietum laevigatae association on the trunks of Quercus, Betula, and, more rarely, Alnus, Fagus, Crataegus, Ilex, Pinus, Pseudotsuga, and Juniperus. Associated lichens include Bryoria fuscescens, Catillaria pulverea, Cladonia spp., Hypogymnia physodes, Lepraria incana agg., Micarea cinerea, M. peliocarpa, M. stipitata, M. synotheoides, Mycoblastus sanguinarius, M. sterilis, Parmelia crinita, P. laevigata, P. saxatilis, Platismatia glauca, Sphaerophorus globosus, and Usnea spp. Occasionally it is found on lignum of fallen decorticate trunks, especially in the western native pinewoods; associated species include Lecidea granulosa agg., Micarea lignaria, M. peliocarpa, and Platismatia glauca. To date, it is not known to occur on mossy boulders, directly on rock, or on the ground on peaty debris, etc. Its distribution in the British Isles is correlated with areas experiencing at least 180 ‘wet days’ per annum and is referable to the ‘General Western Group’ of Coppins (1976). Elsewhere it is known from western Norway (Hordaland), the Azores (on Cryptomeria), and the Canary Islands (on Erica arborea). 3. Micarea anterior (Nyl.) Hedl. (Figs 7B, 38C-D) in Bih. K. svenska VetenskAkad. Hand. III, 18 (3): 76, 86 (1892). — Lecidea anterior Nyl. in Flora, Jena 58: 299 (1875). — Catillaria anterior (Nyl.) Zahlbr., Cat. lich. univ. 4: 29 (1926). Type: Finland, Tavastia australis, Asikkala, 1863, J. P. Norrlin (H-NYL 21655 — lectotype!; H — isolectotypes!). Micarea anterior f. diluta Hedl. in Bih. K. svenska VetenskAkad. Handl. III, 18 (3): 76, 86 (1892). Type: Sweden, Halsingland, Jarvs6, vii 1890, J. T. Hedlund (S — holotype!; UPS — isotype!). LICHEN GENUS MICAREA IN EUROPE 113 ~ i?) 1 2 3 4 y Ps = oT Se a a (s- f| | Ads 3 oat by “Ra Map2_ Micarea alabastrites @ 1950 onwards O Before 1950 Thallus effuse, endoxylic, or epixylic, then whitish and irregularly verrucose to 100 um thick, sometimes becoming secondarily rimose, but without distinct primary areolae; in section, without an amorphous covering layer. Phycobiont micareoid, cells 4-7 um diam. Apothecia numerous, few, or absent, immarginate, convex, adnate, often becoming tubercu- late, pale reddish brown, dull orange-brown or red-brown, 0-15—0-35 mm diam, or to 0-6 mm when tuberculate, Hymenium 38-50 um tall, + hyaline with irregular, dilute red-brown (K—, or dulling) blotches, especially in upper part. Asci clavate or cylindrical-clavate, 35—45 x 10-12 wm. Spores oblong-ovoid to fusiform or + bifusiform, often slightly curved, (0-)1(-2 or ?-3)-septate, 9-14(-16) x2-5—4(-4-5) wm. Paraphyses rather numerous, simple or sparingly branched, many becoming much branched above with their apices entangled, 0-6-1 um wide; apices often irregularly incrassate, hyaline and up to 1-5(—2) wm wide, or sometimes thickened with reddish brown pigment and up to 3 wm wide. Hypothecium c. 60-110 wm tall, hyaline. Excipulum thin but clearly seen in sections as a reddish brown reflexed zone; hyphae radiating, branched and anastomosing, hyaline and c. 1 wm wide, but becoming thickened with brownish pigment and up to 2 wm wide in the outer excipulum. Pycnidia usually numerous; of two types: (a) sessile on old apothecia, but more usually borne on slender stalks (pycnidiophores) which arise from the thallus or in some cases from old apothecia; 70-250 um tall and 40-70 wm diam (overall); stalks pallid but often with reddish brown blotches, glabrous, simple, or sometimes branched and bearing two or three pycnidia; pycnidia (excluding stalk) short cylindrical or slightly tapering inwards above, 40-80 um tall and 40-70 «um diam, with dark reddish brown walls, K— or dulling; conidia (mesoconidia) cylindric- 114 BRIAN JOHN COPPINS al, often faintly biguttulate and slightly constricted in the middle, 3-5—4-5x1-2-1-6 um; (b) sessile on the thallus or old apothecia, c. 30-40 wm diam, with dark reddish brown walls, K— or dulling; conidia (microconidia) narrowly cylindrical, (3-2—)3-6—4-5 x0-7-0-9 wm. Chemistry: Thallus K—, C—, PD-; sections of thallus and apothecia C—; no substances detected by t.l.c. Observations: Micarea anterior is characterised by its reddish brown apothecia, rather narrow, mostly 1-septate spores, pale pycnidiophores bearing dark brown pycnidia, and a + endoxylic, or thin and irregularly verrucose, thallus. Stalked pycnidia are a feature of several other lignicolous species (e.g. M. melaeniza, M. misella, M. nigella, and the occasionally lignicolous M. botryoides), but in those species the pycnidiophores appear black and have different pigmentations when examined microscopically. The pycnidiophores of M. hedlundii are brownish but distinctly tomentose, and those of the normally corticolous M. pycnidiophora and M. stipitata are entirely hyaline. M. adnata has its mesoconidia produced in immersed pycnidia and has macroconidia borne on small apothecia-like sporodochia; further dis- tinguishing features from M. anterior include a + waxy appearance of the whole plant, mostly paler apothecia, and relatively broader spores with rounded apices. Richly fertile specimens of M. anterior could be confused with Catillaria erysiboides (q.v.) but that species has marginate young apothecia, short ovoid spores, and a non-micareoid phycobiont. Habitat and distribution: M. anterior is found on the rather soft and smooth lignum of decorticate trunks of Picea and Pinus, and associated with such species as Cetraria pinastri, Cladonia spp., Hypogymnia physodes, Lecidea icmalea, Lepraria incana agg., Micarea adnata, M. contexta, M. misella, M. prasina, Parmeliopsis ambigua, P. hyperopta, Pertusaria pupillaris, and the non-lichenised Cryptodiscus pallidus. It is a rare or overlooked species, apparently confined to Scandinavia, from where it is known from several localities in middle and southern Sweden, and south-west Finland. Exsiccata: Malme Lich. Suec. 22 (M, S). 4. M. assimilata (Nyl.) Coppins, comb. nov. (Fig. 9; Map 3) Lecidea assimilata Ny\., Lich Scand.: 221 (1861). Type: Norway, Nordland, Helgeland, M. N. Blytt (H-NYL 16556 — lectotype!). Lecidea assimilata a. [var.] irrubata Th. Fr., Lich. Scand. 2: 522 (1874); nom. inval. (Art. 26). Thallus growing on bryophytes, plant debris or sandy soil, composed of confluent, irregular, convex-verrucose, sometimes flattened and subeffigurate areolae that are often intermixed with cephalodia. Areolae white or brownish white, matt or sometimes glossy, 0-08—-0-4 mm diam; in section, sometimes with a hyaline amorphous covering layer up to 7 wm thick, outermost hyphae hyaline; a + algal-free medulla sometimes differentiated in large areolae. Thallus hyphae c. 1-8-3 wm diam. Phycobiont micareoid, cells 4-7 wm diam. Cephalodia often present, irregularly globose and hidden amongst the areolae, sometimes visible externally as brown areolae-like structures, 0-2-0-4 mm diam; containing Nostoc, cells 3-5 wm diam. Less often present are irregular, rather loose clusters (? cephalodia) of Stigonema. Apothecia numerous, immarginate, convex to subglobose, matt or + glossy (subnitid), 0-3-0-8 mm diam, sometimes forming tuberculate clusters up to 1-5 mm diam. Hymenium, 45-50 ym tall; upper part (epithecium) dark aeruginose, olivaceous or brownish-green, K—, HNO3+ red; mid-hymenium dilute greenish; lower hymenium dilute greenish and K—, or dilute purplish and K+ purple intensifying or K+ sordid green. Asci clavate, 45-50 x 12-14 wm. Spores oblong-ellipsoid to oblong-fusiform, O(-1)-septate, (10—)12-16(-19)x3-5 gm. Paraphyses numerous, simple below, but often forked or branched above, 1-5—1-7(—2) wm wide, sometimes widening above to 3 wm; apical walls hyaline although surrounded by densely pigmented gel-matrix. Hypothecium c. 150-400 um tall, dark purple brown, K+ purple intensifying or (especially in upper part) K+ dark green; all parts HNO3+ purple-red; hyphae interwoven but LICHEN GENUS MICAREA IN EUROPE 115 + vertically orientated near the hymenium, c. 1-7-2-7 wm wide, embedded in a densely pigmented matrix; ascogenous hyphae c. 1-7—2-7 wm wide, embedded in a densely pigmented matrix; ascogenous hyphae c. 2-5-5 wm wide. Excipulum reflexed, mottled reddish brown or sordid olivaceous; hyphae radiating, branched and anastomosing, 1-5—2 um wide. Pycnidia rare, semi-immersed to sessile, black, 40-100 zm diam; walls dull reddish brown, or dull olivaceous in part; conidia (? microconidia) cylindrical, 6-9 1-1-5 wm. Chemistry: Thallus K—, C—, KC—, PD-; t.l.c.: no substances. Observations: Micarea assimilata is characterized by its conspicuous, whitish, verrucose areolae which are often intermixed with brown cephalodia, rather large, convex, black apothecia, green (K—) epithecium, dark purple-brown hypothecium and predominantly simple, large spores. It is apparently closely related to M. crassipes, M. incrassata, M. melaenida, and M. subviolascens, all of which share rather stout, simple or sparingly branched paraphyses, rather stout excipular hyphae, dark hypothecia and an absence of lichen substances. In many ways M. assimilata is identical to M. incrassata and both occur in similar habitats; indeed the latter has often been considered a variety (var. infuscata) of the former (e.g. Anderson, 1964; Hertel, 1977). However, M. incrassata differs in several respects, the most important of which is its quite different, red-brown hypothecium which lacks any trace of purple pigmentation in water, K or HNO. In addition, it has an often darker thallus, less prominent apothecia and a generally higher proportion of septate spores. M. subviolascens is similar in appearance to M. assimilata, but grows on rock and has a green, K+ violet epithecial pigment. M. melaenida differs from the last three species in its comparatively small apothecia and less conspicuous thallus, absence of green pigmentation in apothecial tissues, preponderance of 1-septate spores, shorter conidia, and confinement to argillaceous, or fine-grained, mineral soils. M. crassipes occurs in much the same habitats as M. assimilata and the two species have been much confused. However, M. crassipes is easily distinguished by its apothecia which are thinly marginate (at least when young) and markedly constricted below, often being turbinate or even short-stipitate; also by its thallus which is composed of small granular-areolae which sometimes proliferate to give it an isidiose appearance. In section, the apothecia of M. crassipes display a well-developed excipulum and a distinctly two-zoned hypothecium. Pycnidia are rare and difficult to find in members of the M. assimilata group, and have not yet been found at all in M. subviolascens. The remaining species appear to have one conidium type only; although the conidia are rather large, they are probably microconidia. However, the relatively shorter and broader conidia of M. crassipes may be better described as mesoconidia. Cephalodia are here reported from M. assimilata and M. incrassata, apparently for the first time. In both cases the blue-green alga concerned is Nostoc, which loses its filamentous form. In addition, irregular clusters of partially disrupted Stigonema filaments are sometimes present amongst the areolae of M. assimilata, M. incrassata, and also M. subviolascens. Hyphae, presumably belonging to the lichens, are seen to ramify through these clusters; but whether or not these loosely organised structures can be considered to be cephalodia is a problem requiring further detailed anatomical and experimental investigation. Cephalodia or cephalodia-like structures have not been encountered in any other Micarea species. The closely related Lecidea limosa Ach. and L. stenotera (Nyl.) Nyl. have often been confused with M. assimilata and M. incrassata, but they can be distinguished by their pale hypothecia, paraphyses which are individually supplied with a dense, hyaline, gelatinous sheath, and more organised excipular structure. Together with Catillaria contristans (Nyl.) Zahlbr. they probably represent a distinct genus within the Lecideaceae, although perhaps not far removed from Micarea. In British herbaria (at least) there has been much confusion between M. assimilata and Lecidea hypnorum Lib. The latter has marginate, brown-black apothecia, simple paraphyses, 0(-3)-septate spores which become straw coloured and finely warted with age, a reddish brown excipulum composed of radiating, stout, pachydermatous, heavily conglutinated hyphae, c. 3-5 um wide, and bears little resemblance to a Micarea. In addition, the hymenium, hypothe- cium and excipulum of L. hypnorum (and the related L. berengeriana (Massal.) Th. Fr., and L. sanguineoatra auct.) often contain minute granules of a dark violet (K+ aeruginose) pigment. 116 BRIAN JOHN COPPINS More understandable cases of mistaken identity involve Lecidea caesioatra Schaerer, which bears a close superficial likeness to M. incrassata. It can be distinguished from this Micarea by its dark blue-grey apothecia which have a caesious bloom when wet, excipulum of conglutinated (in K) hyphae, and non-micareoid phycobiont with cells c. 8-12 wm diam. Habitat and distribution: M. assimilata grows on decaying bryophytes and plant detritus on the ground or on soil accumulation in rock crevices in arctic (i.e. low altitudes at high latitudes) or montane situations. The realisation that this species belongs in Micarea came so late in the present study, that I have been able to examine only a limited amount of herbarium material. To date, I can confirm its presence in northern Scandinavia and the single outlying locality in Scotland. Specimens purporting to be Lecidea assimilata from the Alps and other central European mountains have proved to be M. crassipes, M. incrassata, or ‘Lecidea’ species not referable to Micarea. Material of ‘Lecidea assimilata’ from Greenland distributed as Hansen Lich. Groenland Exs. 172 and 247 belongs to Lecidea stenotera and L. limosa respectively. The Scottish locality is Ben Lawers in Perthshire, a mountain long renowned for its arctic-alpine flora (Anon, 1972; James, 1965b). M. assimilata was collected there by W. L. Lindsay in 1856. Recent collections from Ben Lawers, and other Scottish localities, are referable to other species, such as Lecidea caesioatra, L. hypnorum, L. limosa, and Micarea incrassata. Exsiccata: Krypt. Exs. Vindob. 2268 (BM, H). Malme Lich. Suec. 216 (H). 0 1 2 4 Pade iz & o ‘ Le He @ * So Sea vale ; —yasmeires — _ we 4. pe © uy is i SS ae ee. eee : Ese Map3_ Micarea assimilata ® Before 1950 + Micarea incrassata @ 1950 onwards LICHEN GENUS MICAREA IN EUROPE 117 5. Micarea bauschiana (KOrber) V. Wirth & Vézda (Figs 5, 10A, 40D; Map 4). in Vézda & V. Wirth in Folia geobot. phytotax., Praha 11: 95 (1976). — Biatora bauschiana Korber, Parerga lich.: 157 (1860). — Lecidea bauschiana (K6rber) Lettau in Hedwigia 55: 28 (1914). Type: Germany, Baden-Wiirttemberg, Baden-Baden, on the way to Yburg, on porphyry, 1859, Bausch, Rabenh. Lich. Eur. 648 (M - lectotype!, sel. V. Wirth & Vézda (loc. cit); M—isolectotype!); additional isotype material distributed as Arnold Lich. Exs. 120 (BM!, M!). Lecidea infidula Nyl. in Flora, Jena 51: 475 (1868). Type: Jersey, C. Larbalestier (H-NYL p.m. 5413 - holotype!). Lecidea lynceola Th. Fr., Lich. Scand. 2: 561 (1874). Type: Norway, Christiana [Oslo], Tveten, 20 v 1868, N. G. Moe 257 (UPS — holotype!). Lecidea semipallens Nyl. in Flora, Jena 59: 234 (1876). Type: Ireland, West Galway, Lough Inagh, 1875, C. Larbalestier (H-NYL 19399 — lectotype!; isolectotypes: BM ex K!, H-NYL 19402!). Lecidea dilutiuscula Nyl. in Flora, Jena 59: 308 (1876). Type: England, South Devon, near Buckfastleigh, H. B. Holl (BM - lectotype!; H-NYL 10754 — isolectotype!). Lecidea rusticella Ny}. in Flora, Jena 61: 245 (1878). Type: Ireland, West Galway, Connemara, Tullywee Bridge, 1876, C. Larbalestier (H-NYL 20206 — holotype!; topotypes [‘1878’]: BM!, BM ex K!). ? Lecidea callicarpa Larbal. ex Leighton, Lich. Fl. Br., ed. 3: 266 (1879). Type: Ireland, West Galway, Glencorbot near Kylemore, 1877, C. Larbalestier (not seen; possibly in BM (incl. BM ex K) but not traced). ?Lecidea semipallens var. obscurior Lang ex Havaas in Bergens Mus. Arb. 1935(2): 27 (1935). Type: Norway, Dalsb¢, 1903, J. J. Havaas (not seen; not traced in BG or H). ?Catillaria microspora Maslova in Ukr. bot. Zh. 30(5): 665 (1973). Type: USSR, Regio Volhyniensis, distr. Ljuboml., in pineto haud procul pag. Kamenca, ad saxa granitica, 11 vi 1969, W. R. Maslova (KW — holotype, not seen; but a slide prepared from the holotype by H. Kilias examined). Thallus thin (<100 um thick), slightly uneven, continuous or finely rimose, usually becoming scurfy-granular and then often thicker (toc. 300 um thick), but never forming discrete areolae or goniocysts, greenish grey, grey-green or pale grey-brown, sometimes ‘oxydated’ and then pale ochraceous or ferrugineus; often obscured by a pale green (white in old herbarium specimens) covering of free-living algae. Thallus in section, ecorticate, without an amorphous hyaline covering layer; outermost hyphae hyaline and without surrounding pigment. Phycobiont not micareoid; cells thin-walled, + globose, S—12 wm diam. Apothecia numerous, immarginate and convex-hemispherical from the start, mostly becom- ing subglobose or tuberculate, 0-2-0-3 mm diam, or up to 0-7 mm diam when tuberculate; very variable in colour, from pale bluish grey (sometimes pallid in part and then ‘piebald’) to dark bluish grey to black, shade forms pallid or pale brown; brown colouration often due to the infestation of ascocarp tissues by a dematiaceous hyphomycete (?Bispora sp.). Hymenium (35—)40-50(-60) wm, sometimes hyaline throughout (shade forms), more usually dilute green- ish, olivaceous or aeruginose and often more intensely pigmented in the upper part, K—, HNO;+ red. Asci cylindrical-clavate, 35-509-12 wm. Spores ellipsoid or ovoid-ellipsoid, simple, 6-5—10-52-5—4 wm. Paraphyses rather scanty, of two types: p.p. evenly distributed, sparingly branched and anastomosing below, septa indistinct, 0-8—1-2 wm wide, often very thin (‘stretched’) in the lower hymenium of old, much expanded apothecia; apical parts usually branched and sometimes incrassate (to 2 wm wide); p.p. fewer in number, as scattered individuals or in small fascicles, straight, simple or occasionally forked above, distinctly septate, stout, c. 1-5-2 wm wide; apices swollen (up to 4 um wide) and often cemented together (when in fascicles) by deeply pigmented matrix. Hypothecium c. 80-150 ym tall, hyaline, or dilute greenish or dilute brownish olive (K—, HNO3+ reddish) in upper part; hyphae interwoven, but becoming vertically orientated toward the hymenium, hyaline, c. 1-2 wm wide; pigment when present confined to the gel-matrix and not closely adhering to the hyphae; ascogenous hyphae with swollen, cells c. 2-5 zm wide. Excipulum not evident even in sections of young apothecia; part of the hymenium is often reflexed to below the hypothecium to give the appearance of an excipulum. Pycnidia often present, immersed, c. 40-80 wm diam; walls greenish K—, HNOs, + reddish, or rarely hyaline (shade forms); conidiogenous cells irregularly subcylindrical, 6-9 1-1-4 um, 118 BRIAN JOHN COPPINS often with a swollen base up to 3 wm wide; conidia (microconidia) cylindrical, eguttulate, 4-6x0-8-1 um. A few collections with pallid apothecia have associated pycnidia, c. 60-100 wm diam, with hyaline walls and containing (often biguttulate) mesoconidia 2:8—4-3 x 1-1-3 wm (see ‘Observations’ below). Chemistry: All parts K—, C—, KC—, PD—; no substances detected by t.l.c. Observations: Micarea bauschiana is closely related to, and often confused with, M. sylvicola (q.v.), and the two species often grow together. M. bauschiana is notoriously variable in the colour of its apothecia, but the full range of variation from pallid to blackish can be seen over small distances (a few centimetres) in a single population (Fig. 5: p. 30) and the variation appears to be solely phenotypic. When growing on iron-rich rocks the thallus of M. bauschiana is often oxydated. In a few specimens (including types of Lecidea rusticella) this oxydation has extended to the apothecia in which finely granular, ferruginous material has been deposited (especially in the epithecium and hypothecium); such specimens have been confused with M. /utulata and species of Protoblaste- nia. A few collections from Austria and Scotland (considered here as M. cf. bauschiana) have entirely pallid apothecia which are accompanied by pycnidia containing mesoconidia (see description above). Such pycnidia have not been detected in specimens of M. bauschiana with: pigmented apothecia, and it is possible that the former collections represent a distinct taxon. However, one of these collections (Austria, Steiemark, Graz, Schéckl N of St Radegund, 1978, Poelt (GZU) also has typical (for M. bauschiana) microconidia-containing pycnidia, and it may be that there is some environmental control (e.g. a response to very low light intensities) involved in the initiation of the mesoconidial anamorph. Further careful field observations and laboratory studies are required to establish the status of these seemingly anomalous collections. Forms of M. bauschiana (s. ampl.) with pallid apothecia should be compared with the much rarer M. lithinella (q.v.) which can be distinguished by its micareoid phycobiont. In addition, M. lithinella seems not to occur in the ombrophobous, aerohygrophilous Micareetum sylvicolae, and is a rather ombrophilous, substratohygrophilous species of damp, shaded rocks or stones. However, I must stress that this ecological interpretation of M. lithinella is based on the limited data available from the herbarium specimens and requires confirmation by field studies. Material recently distributed as ‘M. bauschiana’ in Hertel Lecid. exs. no. 54 is not this species. It has weakly marginate apothecia due to a well developed excipulum, and a large-celled (9-17 wm diam) phycobiont, the cells of which are deeply penetrated by distinct haustoria (as in M. intrusa). It is probably an undescribed species of Micarea and will be treated in a later publication following more critical studies. Habitat and distribution: M. bauschiana is a faithful member of the Micareetum sylvicolae and grows on rocks, stones, roots, and consolidated soil in dry underhangs in woodlands or sheltered valleys. In the British Isles M. bauschiana is the commonest member of the M. sylvicola group. It is particularly common in the north and west but also occurs in suitable situations (e.g. dry stones in sandy banks in woodlands) in the lowlands of south-west England. It seems to be widely distributed in Europe but I have not seen enough specimens to make an assessment of any distributional tendencies. It occurs in the Azores and the Canary Islands, but I have not seen any additional material from outside Europe. Exsiccata: Arnold Lich. Exs. 120 (BM, M), 1233 (M). Johnson Lich. Herb. 434 p.p., 504 (HAMU). Larbal. Lich. Herb. 68 (BM), 305 p.p. (LD). Rabenh. Lich. Eur. 648 p. max. p. (M). Zwackh Lich. Exs. 279A-B, 594A-B, 595 (M). 6. Micarea botryoides (Nyl.) Coppins (Figs 10b, 35, 38A; Map 5) in D. Hawksw., P. James & Coppins in Lichenologist 12: 107 (1980). — Lecidea apochroeella var. botryoides Nyl. in Flora, Jena 50: 373 (1867). — Lecidea botryoides (Nyl.) Zahlbr., Cat. lich. univ. 3: 740 LICHEN GENUS MICAREA IN EUROPE 119 1 Map4 _ Micarea bauschiana @ 1950 onwards © Before 1950 (1925). Type: Finland, Tavastia australis, Lammi, Evo, Lapinkallio, 1866, J. P. Norrlin 404 (H-NYL 20685 p.p. — lectotype!; H —isolectotype!). Note: In the protologue Nylander states ‘ad lignum putridum’ but this is clearly an error (see Vainio 1934: 361). The lectotype has been selected from a packet in the Nylander Herbarium containing two pieces of rock — one with M. lutulata, the other (lectotype) with the apothecia and characteristic stalked pycnidia of M. botryoides. It is possible that Nylander’s diagnosis was based in part on the apothecia of M. lutulata, which bear many similarities to those of M. botryoides. Because Art. 70 of the ICBN is no longer applicable Lecidea apochroeella var. botryoides cannot be rejected as being based on discordant elements; consequently the part containing M. /utulata is excluded, and that part including stalked pycnidia (as described in Nylander’s diagnosis) is chosen as the lectotype. Thallus effuse, thin, more rarely developing into a thick, loose crust up to c. 0-4 mm thick, scurfy-granular, pale to dark dull green, dark olive-green, or dull greenish black, sometimes whitish buff when on dry, deeply shaded rocks. When very thin the thallus consists of flattened granules, c. 20-50 wm diam., which are often dispersed and interconnected by white, arachnoid, prothalline hyphae. Thickening of the thallus is caused by the production of goniocysts (c. 16-30 sum diam), the first of which appear to arise by budding from the primary granules; when thick the thallus has a + gelatinous appearance when wet. Phycobiont micareoid, cells 4-7 wm diam. Apothecia usually few, more commonly absent, immarginate, at first convex-hemispherical, soon becoming + globose and much constricted below (sometimes short-stipitate), often becoming tuberculate, black, or dark brown (in deep shade), matt, 0-1-0-25 mm diam, or to 0-5 mm when tuberculate. Hymenium 25-35 um tall, hyaline or sometimes tinged dilute brown (K—, HNO;—) or dilute olivaceous (K—, or K+ green intensifying, HNO3+ red) in places, but 120 BRIAN JOHN COPPINS always with numerous dark brown (K+ olivaceous tinge, HNO3+ reddish tinge) vertical streaks. Asci cylindrical-clavate or clavate, 20-35 x 7-9 um. Spores ovoid, oblong-ellipsoid or oblong-ovoid, straight or slightly curved, 0-1(-3)-septate, 8-13(-16) x2-3-3-7(-4) um. Para- physes rather scanty, of two types: p.p. evenly distributed, flexuose, simple or sparingly branched, sometimes anastomosing, thin, 0-7—1 wm wide, sometimes widening above to 1-7 um, walls hyaline throughout; p.p. fasciculate, simple or sparingly branched, stout, c. 2-2-5 wm wide, sometimes widening above to 3-5 wm, coated + throughout by dark brown pigment. Hypothecium c. 60-120 um tall, dark reddish brown, K— or dulling, HNO3— or red tinge slightly intensifying; hyphae coated with dark brown pigment, c. 2-3 wm wide, interwoven but becoming vertically orientated towards the hymenium and sometimes continuing into it as stout, fasciculate, pigmented paraphyses; ascogenous hyphae similarly pigmented, with short, swollen cells to 5 wm wide. Excipulum indistinct, sometimes evident in young apothecia as a reflexed reddish brown zone concolorous with, or slightly paler than, the hypothecium; hyphae radiat- ing, branched and anastomosing, c. 1-1-5 4m wide, their walls sometimes with a thin coating of brown pigment. Pycnidia always present and numerous, sessile or, more usually, distinctly stalked, black, 50-400 um tall (including stalk) and 40-90 wm diam; stalks simple, or branched and bearing up to six pycnidia; the ‘stalk-part’ below the current conidia-producing pycnidia often includes old pycnidia (see Fig. 35); the stalk and pycnidia are usually black but in extreme shade forms the stalk tissue may be + colourless, contrasting with the dark brown or blackish, current and old pycnidia. In microscope preparations (at x 400): pycnidiophore tissue dilute to dark fuscous or reddish brown, K— or dulling, HNO3~— or red tinge slightly intensifying; pycnidial wall dark greenish brown, K— or K+ green intensifying HNO3+ red. Conidiogenous cells + cylindrical or elongate ampulliform, often with swollen base which is thickened with brownish pigment, often with one or two percurrent proliferations, 3-5—7-5 x 1-1-4 um, base sometimes swollen to 2-5 wm wide. Conidia (mesoconidia) + cylindrical, often biguttulate, sometimes slightly constricted in the middle, 3-5—4-8x 1-1-5 um. Chemistry: Thallus K—, C—, PD—; sections of apothecia and thallus C—; no substances detected by t.l.c. Observations: Micarea botryoides is easily recognised by its numerous, black, stalked pycnidia, whose walls are dark greenish brown (K— or K+ green intensifying), and its normal occurrence on substrate other than lignum. It is very similar to the rare M. melaeniza, but that species has shorter conidia, smaller, simple spores, and is apparently confined to lignum. M. misella often has stalked, black pycnidia, but is usually lignicolous. However, it does occasional- ly occur on decaying bryophytes where it could be confused with M. botryoides. In such instances their separation is easy because the pycnidia of M. misella contain an olivaceous or dull brownish pigment that turns violet in K. When on rock the apothecia of M. botryoides could be confused with those of M. lutulata, but the latter species has smaller, simple spores, a non-micareoid phycobiont, and immersed pycnidia. When on soft lignum it should be compared with M. nigella which is superficially identical, but has a purple-brown (K+ green) pigment in its apothecia and pycnidia. Sterile forms of M. botryoides have puzzled lichenologists for many years, being dismissed with such remarks as ‘indeterminate pycnidia’ or ‘fungus’. In 1867 Leighton distributed sterile material of M. botryoides in his exsiccate (no. 388), as ‘Lecidea sabuletorum var. milliaria (Fr.), spermagonia’, evidently believing it to be the pycnidial state of Micarea lignaria, the apothecia of which occur on some of his specimens. The identity of these pycnidia remained a mystery until my discovery of fertile material in Scotland in 1976 and my subsequent examination of the type material of Lecidea apochroeella var. botryoides in 1979. Habitat and distribution: M. botryoides is usually found as a constituent of the Micareetum sylvicolae in dry underhangs, growing on solid rock, loose stones, consolidated soil, tree roots, and loose mats of, moribund bryophytes; but it also occurs on rock, stones, and decaying bryophytes in wetter shaded situations. There are a few collections made from the soft, LICHEN GENUS MICAREA IN EUROPE 12 "93 aie 0 1 2 4 0 BY i. By ee Se HX Beals Bd af t Py : 0. ft Cr) se 4 Vel aro ba lie’ ae i 8 ie nee 8 ¢ Paice ee : rg Py, s ies \—qasmeres— ge we OKs i" ra) if Miles Cee 2 e . ‘ F AS PTA \ ae ee ome ee | “E| Li + % “ a, 3 ee 4 TF FRC 4 f % < a oe | 4 LY 2 sper] = 2 al a @ Sica : Al le ; aan Caee, ane oa ah a pe | ; ! z : Z 4 Map5 Micarea botryoides @ 1950 onwards O Before 1950 crumbling lignum of old stumps, but I have not seen it on the firmer, + smooth lignum favoured by M. melaeniza and M. misella. In the environs of some industrial regions (e.g. the West Yorkshire conurbation) it has been found on bark at the bases of trunks of Acer and Betula in sheltered woodlands. M. botryoides is a widespread species, especially in western and upland regions of Britain, and is certainly much more common than current records would suggest. Several of these records result from accidental gatherings, being subsequently identified on samples of other members of the Micareetum sylvicolae. This gives an indication as to how often this species must have been overlooked. It is little recorded outside Britain, and to date I know it only from Norway (S6r-Tr6éndelag and Hordaland), Finland (Tavastia australis), and north-west France. Exsiccata: Leighton Lich. Brit. 388 (BM, BON, DBN, E, M). 7. Micarea cinerea (Schaerer) Hed. (Figs 11, 39; Map 6) in Bih. K. svenska VetenskAkad. Handl. III, 18 (3): 81, 93 (1892). — Lecidea cinerea Schaerer, Lich. Helv. spic., sect. 3: 156 (1826). — Bacidia cinerea (Schaerer) Trevisan in Linnaea 28: 293 (1856). — Bilimbia cinerea (Schaerer) Kérber, Parerga Lich.: 164 (1860). Type: Switzerland, ‘ad infimos Abietum truncos, in sylva Konitz’, L. E. Schaerer (G — holotype!; M — ? isotype!). Lecidea sphaeroides var. albella Schaerer, Lich. Helv. spic., sect. 4-5: 165 (1833). Type: Switzerland: ‘Schweiz’ [on conifer bark], L. E. Schaerer (M — neotype!). See note (i). Biatora delicatula Korber, Denkschr. Feier ihres fiinfzigjih. Best. herausg. Schles. Gesellsch. vater. Kult., 122 BRIAN JOHN COPPINS Breslau: 233 (1853). — Bilimbia delicatula (K6rber) Korber, Syst. lich. Germ: 212 (1855). Type: lectotype as for Lecidea sphaeroides var. albella Schaerer. See note (ii) below. Bilimbia cinerea f. hypoleuca Stizenb. ex Arnold in Flora, Jena 58: 598 (1864). — Micarea cinerea f. hypoleuca (Stizenb. ex Arnold) Hedl. in Bih. K. svenska VetenskAkad. Handl. U1, 18 (3): 82, 93 (1892). Type: Germany, Bayern, near Eichstatt, Affenthale, on young Picea abies in a wood, viii 1864, F. C. G. Arnold (M - lectotype!). Notes. (i) No material labelled Lecidea sphaeroides var. albella could be traced in G (Geissler, in litt.) and an undated specimen in M is selected as neotype. If it can be proven that this specimen was collected before 1833, it should be regarded as a lectotype. (ii) No material referred to in the protologue of B. delicatula has beén traced in L, WRSL or elsewhere. However, K6rber gives ‘Lecidea sphaeroides o albella Schaer. Enum. 193.’ as a synonym. This citation refers to Schaerer (1850) and is an indirect reference to Lecidea sphaeroides var. albella Schaerer (1833). Consequently I have typified both names with the same specimen in M. Thallus effuse, sometimes partly immersed in the substratum (especially when on lignum), more usually developed on the surface of the substratum as rounded, shallow-convex, hemis- pherical or + globose areolae. Areolae scattered to + contiguous, smooth, greenish white to blue-grey, or more rarely becoming dark grey, often dark coloured on upper surface but greenish white below, c. 40-160 wm diam or up to 300 wm diam if containing a pycnidium of the macroconidial anamorph. Areolae in section, ecorticate but with a hyaline amorphous covering layer c. 2-5 wm thick; outermost hyphae often with grey-green to aeruginose walls, K—, HNO3+ red. Phycobiont micareoid, cells 4-7 wm diam. Apothecia usually numerous (see ‘observations’), adnate, plane to convex, sometimes becoming tuberculate, sometimes with an indistinct margin that is + flush with the level of the disc, pale leaden-grey to grey-black (margin often paler), or ivory-white or pallid in shade forms, (0-2)0-3-0-7 mm diam or up to 1-3 mm when tuberculate. Disc matt and finely roughened, but margin smooth and often + glossy. Hymenium 55-70 um tall, hyaline, but usually olivaceous or aeruginose (K—, HNO3+ red) in upper part (epithecium). Asci clavate 50-65 x 15-20 um. Spores fusiform, often slightly curved, (3—)5—7-septate, (19-)23-34(-38) x4-5—6 wm. Para- physes numerous, branched and often anastomosing, 1-1-4 um wide; apices often more richly branched and entangled, often slightly incrassate to c. 1-8 wm, or 2:5 wm due to thickening by greenish pigment. Hypothecium c. 40-70 um tall, hyaline; hyphae interwoven, c. 2-4 wm wide. Excipulum well developed, hyaline or pale straw, or richly branched and anastomosing hyphae, c. 1-1-5 wm. Pycnidia frequently present, of two types: (a) immersed in areolae, white or faintly greenish around the ostiole, 160-300 wm diam, ostiole often widely gaping; conidia (macroconidia) + straight or flexuose, filiform, 9-17-septate, 50-110Xc. 1 wm; (b) + sessile, white, c. 40-70 wm diam, ostioles not, or only slightly, gaping; conidia (microconidia) narrowly fusiform-cylindrical (3-8-)4-5 x0-5—0-7 wm. Chemistry: Thallus and whitish apothecia K—, C+ red, PD—; apothecia in section C+ red; t.l.c.: gyrophoric acid. Observations: M. cinerea is very variable with regard to the colour of its apothecia and thallus; as with M. peliocarpa this variation is due to the amount of green pigment produced in response to environmental factors, especially light. Morphologically and chemically M. cinerea is closely allied to M. alabastrites and M. peliocarpa; the diagnostic features of each of these three species are compared in Table 4. Forms of M. cinerea with blackish apothecia can be confused with M. lignaria (including var. endoleuca) but on close inspection the latter will be seen to differ in its + globose apothecium with a poorly differentiated excipulum (in vertical section), usually brownish or greenish upper hypothecium, less strongly branched paraphyses and chemistry. M. cinerea is usually well fertile with numerous apothecia, but sterile forms with numerous macroconidia-containing pycnidia have been encountered; the macroconidia are about twice as long as those found in M. peliocarpa and M. alabastrites and so are readily identifiable (see ‘Key to species without apothecia, pp. 107-108). LICHEN GENUS MICAREA IN EUROPE 123 Habitat and distribution: In the British Isles M. cinerea is mostly found in communities of (or closely akin to) the Parmelietum laevigatae on the trunks or over bryophytes thereon of Quercus, Betula and, less often, Alnus, Corylus, Fraxinus, Ilex, Sorbus, Larix, Pinus, and Pseudotsuga. Associated species in these communities include Catillaria pulverea, Cladonia coniocraea, C. macilenta, C. squamosa, Haematomma caesium, H. elatinum, Lecideaicmalea, Lepraria incana, Micarea alabastrites, M. stipitata, M. synotheoides, Mycoblastus sterilis, Parmelia laevigata, P. saxatilis, Platismatia glauca, Stenocybe septata, Thelotrema lepadinum, Trapelia corticola ined. Frullania tamarisci, Lejeunea ulicina, Scapania gracilis, Dicranum fuscescens, and Hypnum cupressiforme. Less commonly it is found in the lignum of fallen, decorticated trunks. Occur- rences on other substrata are rare although I have found it on one occasion growing directly on rock (Coed Hafod in Denbigh) where it was associated with Trapelia involuta on the top of a dry-stone wall in an oak-birch wood. In addition, it was found on mosses on epidiorite at an altitude of 1000 m in the Ben Alder range of Inverness-shire by Dr O. L. Gilbert; the specimen is sterile but has pycnidia with the characteristic macroconidia. In Britain M. cinerea usually occurs at lower altitudes (mostly below 300 m), and exhibits a distribution pattern attributable to the General Western Group of Coppins (1976). However, there are two outlying easterly localities (in North Northumberland and East Perth) both of which are sheltered, more or less undis- turbed, valley woodlands. From outside Britain I have seen material of M. cinerea from Hordaland and Rogaland in western Norway, the east Sudety and Vysoké Tatry of Czechoslovakia, southern Germany, and the Swiss, Austrian and Italian Alps. Most collections from southern Germany and the Alps are iis OR, ZL 7 » 7. 4 ae ! t 6 - 6 no $ 2 5 5 Us . KG 5 6 a lia 3 \ 3 J i ae x 2 va 4 2 ie arg Pe ee aa Faas, ae t 2 a 2 3 4 5 6 a Map6 Micarea cinerea @ 1950 onwards O Before 1950 124 BRIAN JOHN COPPINS from the trunks of conifers (Picea, Pinus, and Abies), but they also included specimens from the trunks of A/nus and Betula, lignum of stumps, fallen trees, and old fence posts. In addition, it was collected several times on thin twigs of Picea by F. Arnold and his contemporaries, and it is possibly of interest to note that I do not know it as an inhabitant of twigs in north-west Europe. Unlike M. alabastrites and M. peliocarpa, M. cinerea is not known from Macaronesia, and I have not seen any material of it from outside Europe. Exsiccata: Arnold Lich. Exs. 548 (M), 549 (BM ex K, M). Arnold Lich. Mon. 47 (BM ex K), 115 (BM ex K,M, MANCH), 116 (BM ex K, M). Britz. Lich. Exs. 846 (M). Hepp Flecht. Eur. 21 p.p. (BM, M). Lojka Lich. Hung. 60 (M). Vézda Lich. Sel. 1087 (BM, S). Zwackh Lich. Exs. 898 (M). 8. Micarea contexta Hed. (Figs 12A, 40A-B) in Bih. K. svenska VetenskAkad. Hand. II, 18 (3): 83, 96 (1892). — Catillaria contexta (Hedl.) Zahlbr, Cat. lich. univ. 4: 35 (1926). Type: Sweden, Halsingland, Ovanaker, 1891, J. T. Hedlund (S - lectotype!). Thallus effuse, endoxylic, inapparent or evident as a slight bleaching of the wood, consisting of scattered, small, rounded granules (c. 15-25 wm diam) buried between the wood fibres; external hyphae of granules with dark green walls, K—, HNO3+ red. Phycobiont micareoid, cells 4-7 wm diam. Apothecia numerous, immarginate, + globose from the start, often becoming tuberculate, black, matt, 0-1-0-2 mm diam, or to 0-3 mm diam when tuberculate. Hymenium 35-45 pm tall; upper part (epithecium) dark green, K— or green intensifying, HNO3+ purple-red; remaining (lower) part + hyaline or dilute greenish with dark green, vertical streaks, sometimes with a few, minute, purple-violet (K+ aeruginose) granules. Asci clavate, 35-40 10-14 um. Spores ovoid or oblong-ovoid, upper cell broader than the lower and with a more rounded apex, 1-septate, or rarely with a thin, additional septum in the lower cell, 7—13(—14) x (2:3-)3-4-5 wm. Paraphyses rather scanty, of two types: p. max. p. branched and anastomosing, haline, thin, 0-6-0-8 wm wide, apices often with thickened, pigmented walls and then to 2 wm wide; p. min. p. scattered or in small fascicles, simple, with thickened, pigmented walls throughout and 1-5—2 wm wide, apices sometimes widening to 3 wm. Many apices overtopping the asci to form a + distinct epithecium. Hypothecium 20-90 um tall, dark green or dark purple (colours often intermixed), K+ dark olive-green or aeruginose, HNO3+ purple-red; hyphae interwoven, hyaline or thickened with pigment, 1-2 wm wide; ascogenous hyphae with swollen cells to 4 wm wide. Excipulum indistinct, sometimes evident as a narrow, dark greenish zone, c. 5—12 wm wide, forming a lateral border to the reflexed edge of the hymenium. Pycnidia usually present but inconspicuous, immersed between the wood fibres or emergent to sessile, black, with dark greenish walls, K — or green intensifying, HNO3+ purple-red; of two types (a) c. 40 wm diam; conidia (mesoconidia) cylindrical, often faintly biguttulate 3-8-4-7X 1-3-1-8 wm; (b) c. 20-40 wm diam; conidia (microconidia) narrowly cylindrical, 4-5 x 0-8-1 wm. Chemistry: Apothecia sections C—; material insufficient for analysis by t.l.c. Observations: Micarea contexta is characterised by its endoxylic thallus, very small, black, + globose apothecia, dark green epithecium, dark greenish or purplish hypothecium, ovoid, 1-septate spores, thin hyaline paraphyses and an absence of stalked pycnidia. It is apt to be confused with diminutive, immature forms of M. melaena, but that species has more numerous paraphyses, longer microconidia, and a superficial, granular thallus which contains gyrophoric acid when in a healthy condition. When mature, M. melaena has more robust apothecia and larger, 3-septate spores. M. eximia can be distinguished from M. contexta by its more brightly coloured epithecium and narrower, + fusiform spores; and M. nigella can be distinguished by its simple spores and stalked pycnidia. M. olivacea differs in having more numerous paraphyses which are broader (1-1-2 wm) when hyaline, a complete absence of purple pigmentation in its apothecia, and relatively narrower, + oblong spores. LICHEN GENUS MICAREA IN EUROPE 125 Habitat and distribution: M. contexta occurs on conifer lignum, probably in sheltered, woodland situations. Species associated with M. contexta on the specimens examined include Arthonia helvola, Cetraria pinastri, Cladonia spp., (scattered squamules), Hypogymnia phy- sodes, Lecanora symmicta agg., Lecidea efflorescens, L. pullata, Micarea anterior, M. misella, Parmeliopsis ambigua, and P. hyperopta. The known distribution of M. contexta is restricted to middle Sweden, from where it was found in several localities by Hedlund. Supposed collections from outside Sweden seen by me belong to M. melaena. Nevertheless, M. contexta should be sought for in other regions, especially those containing naturally occurring coniferous forests. Exsiccata: Malme Lich. Suec. 28 (M, S). 9. Micarea crassipes (Th. Fr.) Coppins, comb. nov. (Figs 4B, 12B, 41C) Helocarpon crassipes Th. Fr., Lich. arctoi: 178 (1860); and in Nova Acta R. Soc. Scient. Upsal. III, 3: 278 (1861). — Lecidea crassipes (Th. Fr.) Nyl. in Flora, Jena 45: 464 (1861). Type: Norway, Finnmark, Aldjok, 1857, Th. M. Fries (S —lectotype!; isolectotypes: BM!, BM ex K!, LD, UPS!). Lecidea crassipes f. moriformis Th. Fr., Lich. Scand. 2: 520 (1874). Type: Norway, Finnmark, Masdy, 16 vii 1864, Th. M. Fries (UPS — lectotype!). Lecidea crassipes f. pulverula Th. Fr., Lich Scand. 2: 520 (1874). Type: Norway, S6r-Tréndelag, Oppdal hd., Dovre, Kongsvoll, Hégsnyta, 11 viii 1863, Th. M. Fries (UPS — lectotype!). Lecidea hypopodia f. subassimilata Ny\. in Bull. Soc. linn. Normandie, IV, 1: 242 (1887). Type: U.S.A., Alaska, Bering Strait, St Lawrence Island, 1879, E. Almquist (H-NYL 20915 — holotype!). Thallus growing on bryophytes or plant debris, rarely spreading on to sandy soil, composed of small, + globose areolae. Areolae cream-white, pale grey-brown, sometimes ash-grey in part, matt, rather fragile, 0-07-0-2 mm diam; the larger areolae sometimes budding to produce smaller, secondary granules thereby giving the thallus an isidiose appearance. Areolae in section without a hyaline covering layer; thallus hyphae entirely hyaline, c. 2-3 ~m wide. Phycobiont micareoid, cells 4-7 wm diam. Apothecia numerous, sessile but (in section) markedly constricted below, or turbinate, or short-stalked, 0-2-0-6 mm diam; disc matt, at first plane with a thin, often slightly raised, concolorous or slightly paler, often + glossy, margin, 0-02—0-05 mm wide; disc later expanding, becoming convex and excluding the margin. On rare occasions apothecia become tuberculate and up to 1-4 wm diam. Hymenium 45-50 wm tall; upper part (epithecium) dark green or olivaceous and K+ green-intensifying, or purple-brown and K+ purple-intensifying; remaining (lower) part dilute purplish, K+ purple-intensifying or K+ sordid-green. Asci clavate, 40- 48x11-14 wm. Spores 0(-1)-septate, fusiform-ellipsoid, (9-)10-17(—21) x(2:5-)3-+4:5 wm. Paraphyses numerous, simple, or sparingly branched above, sometimes anastomosing, 1-1-5 pm wide, sometimes widening above to 2-5 wm; apical walls hyaline but often surrounded by dense pigment. Hypothecium dark, variably pigmented: upper part (c. 30-60 wm) dark purple-brown, K+ dark olive-green, or K+ purple-intensifying in part or in whole; lower part (including ‘stipe’) paler and mottled, brown- or vinose-red and K—, or with purple tinge and K+ purple-intensifying; hyphae in upper part tightly interwoven but becoming vertically orientated towards the hymenium, c. 1-5-2 wm wide, embedded in a densely pigmented gel-matrix, intermixed with ascogenous hyphae c. 2-5-4 xm wide; hyphae in lower part loosely interwoven, becoming outwardly directed laterally (i.e. as they approach the excipulum), c. 1-1-5 um wide, embedded in a gel-matrix that swells and partially dissolves in K. Excipulum well developed, and conspicuous in young mature apothecia, c. 50 wm wide laterally; innerpart concolorous with upper hypothecium; outer part dilute to dark olivaceous, K+ green-intensifying; hyphae radiating, branched and anastomosing, c. 1-1-5 wm wide. Excipular and hypothecial tissues usually extended vertically downwards to form a ‘stipe’ (Fig. 4B). Pycnidia rare, sessile, black, c. 60-150 wm diam, ostiole sometimes gaping; walls c. 10— 17 wm wide, green-black (K— or K+ green-intensifying) in upper parts, changing to dark red-brown (K—) below; conidiogenous cells elongate-ampulliform, strongly tapered towards 126 BRIAN JOHN COPPINS the neck, 6-9 x 3-4 wm, neck 1-1-3 wm wide; conidia (? mesoconidia) cylindrical, 4-5—5-7 x 1-2- 1-5 um. Chemistry: Thallus K—, C—, KC—, PD-; t.l.c.: no substances. Observations: The combination of a finely granular (sometimes isidiose) whitish to grey, muscicolous thallus and turbinate or short-stalked, marginate (when young), black apothecia make this an easy species to recognise, even with a hand lens. Internally, the apothecia have a well differentiated excipulum and a distinctly two-zoned hypothecium. This structure, which is rather ‘advanced’ for a Micarea, has led me to deliberate on the merits of the monotypic genus Helocarpon Th. Fr. However, characters such as the basic arrangement, organisation and form of apothecial hyphae (including paraphyses), ascus structure, morphology of pycnidia, and thallus structure (including phycobiont), are still those of a Micarea. The vertical extension of excipular and hypothecial tissues to produce a ‘stipe’ is not unique to M. crassipes, and is found in some specimens of M. lignaria and M. ternaria, and in an undescribed species from New Zealand (North Island: Trounson Kauri Reserve, on hepatics on rotten stumps 1967, D. M. Henderson (E)). This last species closely resembles M. crassipes (e.g. apothecia marginate when young, apothecial pigmentation, thallus type) but has broader, ellipsoid or ovoid, 1-septate spores (13—-16-5 5-5-7 ym) and rather sparse paraphyses. Overall considerations of morphology and ecology lead me to assign M. crassipes to the group including M. assimilata, and it is to the account of that species the reader is referred for further discussions. Habitat and distribution: M. crassipes grows over bryophytes and plant debris on the ground and amongst rocks in northern Fennoscandia and at high altitudes (probably above 1000 m) in the Alps and other central European mountains. From Norway and Sweden I have seen several collections from north of c. 63°N and in Russia it occurs on the Kola Peninsula. In Finland it extends to the southernmost parts (Karelia australis, c. 60°30'N). In central Europe it occurs in the western Sudety (Krkono&e, at 1000 m) of Czechoslavakia, and southwards its range extends east to the Alps (Austrian Tirol, and Dolomiti in north Italy) and west to the Transylvanian Alps (Muntii Retezat, at c. 2000-2500 m) of Romania. From outside Europe it has been collected on St Lawrence Island in the Bering Strait; it is quite likely to have a circumpolar distribution. Exsiccata: Arnold Lich. Exs. 556 [A] (BM ex K, H, H-NYL 16578), 556B (BM ex K, H, H-NYL p.m. 5105), 1121 (BM ex K). Fellman Lich. Arct. 165 (BM ex K, H, H-NYL 16570). Malme Lich. Suec. 362 (BM). Norrlin and Nyl. Herb. Lich. Fenn. 194A, B(H). Vézda Lich. Bohem. 282(LD). Vézda Lich. Sel. 11 (BM). 10. Micarea curvata Coppins, sp.nov. (Fig. 12C) Thallus probabiliter albido-griseus vel brunneo-griseus, granuloso-verruculosus. Algae cellulis 4-7 wm diam. Apothecia immarginata, convexo-hemisphaerica mox tuberculata, atrobrunnea, 0-2-0-5 mm diam, aut ad 0-65 diam ubi tuberculata. Hymenium c. 60 wm altum, + hyalinum, cum vittis verticalibus, pallide fuscis; parte summa (epithecio) pallide fuscis, K—, Ascosporae fabiformes vel valde curvatae, (0-)1-septatae, 9-11-72-5-3-8 jum. Paraphyses ramosae et anastomosantes, graciles, c. 0-8-1 wm latae, apicibus haud incrassatis et haud pigmentiferis. Hypothecium pallidum. Excipulum reflexum, pallidum, margine externo pallide fusescenti. Pycnidia ignota. Thallus et apothecia in sectione C+ rubra. Typus: Germania, Guestphalia, Gravenhorst, in muro lapideo ad lapidem arenarium, leg. Th. R. J. Nitschke (WRSL -holotype!). Thallus probably grey or brownish grey and granular-verrucose, but difficult to interpret in the single specimen in which the thallus is invaded by foreign algae and dematiaceous hyphae. Phycobiont micareoid, cells 4-7 wm diam. Apothecia convex-hemispherical to tuberculate, immarginate, dark brown, 0-2-0-5 mm diam, or to 0-65 mm diam when tuberculate. Hymenium c. 60 wm tall + hyaline with pale fuscous brown vertical streaks; upper part (epithecium) pale fuscous brown, K—, HNO3. Asci clavate, c. 45-50 10-13 xm. Spores fabiform or distinctly curved, (0—-)1-septate, 9-11-7X2-5-3-8 wm. LICHEN GENUS MICAREA IN EUROPE 127 Paraphyses branched and anastomosing c. 0-8-1 wm wide, not swollen or pigmented at apices. Hypothecium pale, tinged dilute straw-brown. Excipulum soon reflexed, internally pale with straw-brown tinge, becoming slightly darker (pale fuscous) towards the outer edge; hyphae radiating, branched and anastomosing, c. 0-8-1 wm wide. Pycnidia not found. Chemistry: Sections of thallus C+ red, PD—; sections of apothecia C+ red; probably containing gyrophoric acid but material insufficient for analysis by t.l.c. Observations: M. curvata is characterised by its distinctly curved, 1-septate spores, fuscous brown pigment in the apothecia which is unchanged by K or HNO3, and C+ red reactions (? gyrophoric acid) of apothecia and thallus in sections. It is reminiscent of M. subnigrata but that species has uncurved, ellipsoid spores and all parts C—. In outward appearance M. curvata is similar to some forms of the almost ubiquitous Scoliciosporum umbrinum, with which it could be mistaken in the field. Habitat and distribution: M. curvata is known only from a single collection on sandstone in the Wroclaw Herbarium, which is labelled ‘?Biatora nov. sp. Auf einer Steinmauer bei Gravenhorst/Westfalen leg. Nitschke’ (label kindly transliterated by Prof. Hertel). No recognis- able associate species are present on the small specimen although there are a few, rounded, white areolae (C+ red) probably belonging to Trapelia coarctata s. ampl. From the appearance of the specimen it occurred in a humid, sheltered, possibly shaded or north-facing situation. 11. Micarea denigrata (Fr.) Hed. (Figs 1B, 13, 42; Map 7) in Bih. K. svenska VetenskAkad. Hand. I11, 18 (3): 78, 89 (1892). — Biatora denigrata Fr. in K. svenska VetenskAkad. Handl. 1822: 265 (1822). — Catillaria denigrata (Fr.) Boistel, Nouv. Fl. Lich. 2: 199 (1903). Type: Sweden, Smaland, ‘Femsjé in parietibus vetustus’, E. M. Fries (UPS - lectotype! [t.l.c.: gyrophoric acid]). Lecidea anomala f. pyrenothizans Nyl., Lich. Scand.: 203 (1861). — Micarea denigrata f. pyrenothizans (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Handl. M11, 18 (3): 78, 89 (1892). Type: Finland, Nylandia, Helsingfors [Helsinki], on lignum, 1860, W. Nylander (H-NYL 21665 — lectotype!). Lecidea parissima Nyl. in Crombie in J. Bot., Lond. 9: 178 (1871), andinJ. Linn. Soc. Bot. 11: 484 (1871). Type: England, Middlesex, Hendon, on old pales, 1870, J. M. Crombie (H-NYL 21659 — holotype!; isotypes: BM!, BM ex K!). Lecidea spodiza Ny). in Flora, Jena 57: 9 (1874). Type: Scotland, Perthshire, near Killin, on old worked conifer lignum, 1873, J. M. Crombie (H-NYL 21734 — lectotype!; BM — isolectotypes!). Lecidea hemipoliella Nyl. in Flora, Jena 58: 11 (1875). — Micarea hemipoliella (Nyl.) Vézda in Vézda & V. Wirth in Folia geobot. phytotax., Praha 11: 100 (1976). — Micarea denigrata f. hemipoliella (Nyl.) Hed. in Bih. K. svenska VetenskAkad. Handl. III, 18 (3) 78, 89 (1892). Type: Finland, Tavastia australis, Evo, + smooth bark of Alnus, 1873, J. P. Norrlin (H — lectotype!, isolectotypes: H(<3)!, H-NYL 21691!). Lecidea discretula Ny]. in Flora, Jena 58: 444 (1875). Type: Romania, ‘Circa Thermas Herculis in Banatu (Hung.), supra Daedaleam in trunco quercino’, 1874, H. Lojka (H-NYL 21693 — holotype!). Lecidea aniptiza Stirton in Rep. Trans. Glasgow Fld Nat. 4: 85 (1876). Type: Scotland, Perthshire, Killiecrankie, on lignum, 1875, J. Stirton (GLAM - lectotype!; BM — isolectotype!). Lecidea denigrata var. submisella Nyl. in Vainio in Medd. Soc. Fauna Fl. fenn. 3: 112 (1878). — Catillaria synothea f. submisella (Nyl.) Blomb. & Forss., Enum. Pl. Scand.: 92 (1880). Type: Finland, Tavastia australis, Korpilahti, on lignum, 1874, E. A. Lang [Vainio] (H-NYL 21661 — lectotype!). Lecidea spodiza f. ecrustacea Lamy in Bull. Soc. bot. Fr. 25: 440 (1878). Type: France, Puy-de-Dome, Monte Dore, near Cascade du Queureilh, on lignum of conifer trunk, 31 vii 1878, Lamy (H-NYL 21731- lectotype!). Lecidea praeviridans Ny\., Suppl. Lich. env. Paris: 5 (1879). — Biatorina praeviridans (Nyl.) Boistel, Nouv. Fl. Lich. 2: 194 (1903). — Catillaria praeviridans (Nyl.) Zahlbr., Cat. Lich. univ. 4: 64 (1926). Type: France, Haute Loire, Saugues, on Pinus bark, 18 -, A. Boistel (H-NYL 19221 — lectotype! [t.l.c.: gyrophoric acid]; BM — isolectotype!). Lecidea denigrata f. sublivescens Ny]. in Flora, Jena 64: 539 (1881). Type: Romania, Maramures, ‘prope 128 BRIAN JOHN COPPINS balneum kabola Pojana, com. Marmaros in Hung.’, on bark of Pinus sylvestris, H. Lojka 4455, ‘Lich. Hung. exs. (ined.) n. 305. ad int.’ (H-NYL 11663 — holotype!). Lecidea denigrata f. pseudoglomerella Harm. in Bull. Séanc. Soc. Sci. Nancy. I, 33: 58 (1899 [‘1898’]). - Catillaria denigrata var. pseudoglomerella (Harm.) Boistel, Nouv. Fl. Lich. 2: 199 (1903). Type: France, Meurthe-et-Moselle, La Malgrange, on oak posts [lignum], 8 v 1894, J. Harmand, Lich, Loth. 838p.p. (ANGUC - lectotype! [t.l.c.: gyrophoric acid]; DUKE — isolectotype!). Catillaria prasina f. longior Erichsen in Schr. naturw. Ver. Schleswig-Holst. 22: 101 (1937). Type: West Germany, Schleswig-Holstein, Lauenberg, Sachsenwald, Rev. KI. Viert, on roots of old Fagus, 2 xii 1934, C. F. E. Erichsen (HBG — holotype!). Micarea denigrata var. friesiana Hedl. in Bih. K. svenska VetenskAkad. Hanadl. III, 18 (3): 78, 89 (1892); nom. inval. (Art. 26). Micarea denigrata var. friesiana f. vulgaris Hedl. in Bih. K. svenska VetenskAkad. Handl. III, 18 (3): 79, 90 (1892); nom. inval. (Art. 26). Micarea andesitica Vézda in Poelt & Vézda, Bestimmungsschl. europ. Flechten. Erganzungsheft I: 160 (1977); nom. nudum (Art. 32). Spec. orig.: Hungary, Matra, Matrafiired, Pipis-Legy, 380 m, on andesitic rock, 17 xi 1976, A. Kiszely & A. Vézda (hb Vézda!). ?Lecidea hemipoliella* semialbula Stirton in Rep. Trans. Glasgow Fld Nats. 4: 89 (1876). — Biatorina synothea var. semialbula (Stirton) A. L. Sm., Monogr. Br. Lich. 2: 122 (1911). Type: Scotland, Sutherland, Altnaharra, on lignum, Stirton (not seen; not traced in BM or GLAM). ?Catillaria synothea f. major B. de Lesd., Rech. Lich. Dunkerque, Suppl. 1: 118 (1914). Type: France, Nord, Dunkerque, Ghyvelde, on piece of wood (Pinus) on the sand ina dune, B. de Lesdain (not seen). ?Catillaria synothea f. fusca B. de Lesd., Rech. Lich. Dunkerque, Suppl. 1: 118 (1914). Type: France, Nord, Dunkerque, St-Pol, on a stake, B. de Lesdain (not seen). Lecidea synothea auct., non Ach. (1808). See note below. Note: Lecidea synothea Ach. in K. svenska VetenskAkad. Handl. 1808: 236 (1808). From the comments on Acharian specimens by Hedlund (1892: 91) and my examination of Acharian material in BM, it is clear that this name was based on material of the common and widely distributed species generally known as Bacidia umbrina (Ach.) Bausch or Scoliciosporum umbrinum (Ach.) Arnold (basionym: Lecidea umbrina Ach, Lich. Univ.: 183 (1819)). In the 19th century ‘synothea’ was commonly applied to Micarea denigrata. Despite Hedlund’s comments, this misapplication has been continued by some authors, even in recent years (e.g. Ozenda & Clauzade, 1970: 402). To avoid the confusion that would arise if ‘umbrina’ were replaced by ‘synothea’, a formal proposal to reject the name Lecidea synothea Ach. (and combina- tions from that name), under the provision of Art. 69.1, will be presented elsewhere. Thallus effuse and often widespreading, sometimes endoxylic but usually developing on the surface of the substratum (especially in the vicinity of apothecia) as convex to irregularly subglobose areolae. Areolae 60—200 xm diam, greenish white to greenish grey, matt; in section without an amorphous covering layer, external hyphae hyaline but surrounding gel matrix often with dilute olivaceous, K+ violet pigment. Thallus sometimes scurfy and dark grey-brown to blackish, owing to invasion by dematiaceous fungi and non-lichenized algae. Phycobiont micareoid, cells c. 4-7 wm diam. Apothecia usually present and numerous (see ‘observations’ below), scattered to confluent, broadly convex to subglobose, sometimes tuberculate, 0-15—0-5 mm diam, or to 0-6 mm when tuberculate; immarginate, or sometimes young apothecia with an indistinct, shallow marginal rim, paler than the disc; disc pallid to brown or piebald (shade forms), more usually dark grey or black, matt. Hymenium (25—)30-40 um tall, dilute olivaceous or dull brownish, K+ violet; pigment often concentrated in upper part (with lower part + hyaline), and confined to the gel-matrix. Asci clavate, 28-369-12 um. Spores oblong-ellisoid, oblong-ovoid, fusiform or bacilliform, often slightly curved, (0—)1-septate, upper cell usually slightly shorter and broader than the lower, (7—)9-16(—18) X2-3-3(—3-5) wm. Paraphyses numerous, branched and some- times anastomosing, 1—-1-5(-1-7) wm wide; apices scarcely wider, and never with closely adhering pigment. Hypothecium 60-110 um tall, hyaline or very dilute yellowish straw; hyphae c. 1-1-5 ym wide, interwoven or some vertically orientated in upper part, intermixed with short-celled, ascogenous hyphae hyphae c. 2-4 wm wide. Excipulum indistinct, but usually evident in sections of young, shallow-convex apothecia, hyaline; hyphae radiating, branched and anastomosing, hyaline, c. 1-1-5 wm wide. Pycnidia usually present and numerous, immersed in the thallus or substratum (endoxylic LICHEN GENUS MICAREA IN EUROPE 129 forms), sometimes emergent, hyaline grey or black; walls hyaline but usually dilute olivaceous or brownish, and K+ violet in the upper (exposed) parts; in endoxylic forms, walls olivaceous (K+ violet) throughout; of three types: (a) 60-150 wm diam, ostioles eventually gaping; conidia (macroconidia), curved or hamate, (1—)3-septate, 12—24xc. 1 wm; (b) 80-160 wm diam, ostioles often gaping: conidia (mesoconidia) short cylindric or obovate, sometimes faintly biguttulate, 2-8-4-5(-5) x 1-2-1-8 wm often extruding from the ostioles as a conspicuous white blobs; (c) 30-50 gm, ostioles not gaping; conidia (microconidia) narrowly fusiform or bacilliform, (4-5)5—7-5 x0-7-0-8 wm. Chemistry: Thallus K—, PD—; sections of thallus and apothecia C+ orange-red, rarely C— (also, parts with dull olivaceous pigment, C+ violet). If thallus is heavily parasitised and scurfy the C+ orange-red reaction may be difficult to obtain and gyrophoric acid may not be detectable by tke, Observations: Micarea denigrata is a common and extremely polymorphic species exhibiting a wide range of genotypic and phenotypic variation. Its thallus can vary from being completely endoxylic to forming a thick, granular-verrucose, areolate crust. The colour of the apothecia varies from pallid, through grey or brown, to black in a + direct relationship with exposure to light. Specimens may be abundantly fertile with numerous, large, mature apothecia. On the other hand, they may be sterile with just a few, scattered immature apothecia (or even none at all) but with numerous pycnidia containing mesoconidia; alternatively the latter may be replaced by pycnidia containing microconidia and, or, macroconidia. Indeed, all possible combinations of anamorphic states have been found and a few collections (e.g. Coppins 1888 and 8384) have all three states, plus apothecia, on the same thallus. Spore length is usually in the range of 9-14 wm, but extreme forms are known in which the range is 8-10 wm, or 12-18 um. Gyrophoric acid is usually present in the thallus and/or the apothecia (and easily demonstrated by tests with C), but in some specimens its concentration is low and only detectable with certainty by t.l.c.; in a few diminutive, endoxylic forms I have been unable to detect gyrophoric either by a C test, or byt.l.c. Inshort, M. denigrata exhibits extreme variation. Despite this, with care and patience the species is rarely difficult to separate from other, similar species. The biggest problems I have experienced are where the decision has to be made between what could be a diminutive form of M. denigrata with small spores, or a form of M. misella with an unusually high proportion of 1-septate spores and without its characteristic stalked pycnidia (that contain mesoconidia). Such decisions are settled by the careful observation and measurement of paraphyses, and microconidia (see couplet 11 of the main key and Table 7). Shade forms of M. denigrata with brown, + globose apothecia have been confused with M. elachista, but the latter has a brown (K+ dissolving) epithecial pigment, a more complex thallus structure (with a ‘cortex’ and amorphous covering layer), distinctive pycnidia, and always lacks gyrophoric acid (sections of thallus and apothecia C—). Forms of M. denigrata with a scurfy-granular thallus can be similar to M. prasina, but microscopic examination will show the thallus appearance to be caused by its disruption by invading foreign fungi and algae, and not by the presence of the small + discrete granules (goniocysts) characteristic of M. prasina. Furthermore, M. prasina never gives C+ orange-red (gyrophoric acid) reactions, and it usually has broader, rarely curved spores with more rounded apices. The closest relative to M. denigrata is M. nitschkeana, which is + identical with regard to thallus structure, pycnidial types, apothecia structure, pigmentation, and chemistry. Morphologically the only important difference is that the mature spores of M. nitschkeana are 3-septate and more consistently curved (cf. Figs 13 and 24B). In addition, the two species differ in their preferance of substrata: M. denigrata favouring lignum or dead bark of old tree trunks or stumps, and M. nitschkeana favouring corticate twigs and small branches of trees and shrubs. However, there is some degree of overlap as M. nitschkeana is occasionally found on the lignum of fence posts, but, on the other hand, I have never encountered M. denigrata on attached living, corticate twigs. In the field, well developed specimens of M. denigrata sometimes have a superficial resemblance to M. cinerea, M. lignaria, and M. peliocarpa, but these three species have K— hymenia and larger, 3 (or more)-septate spores. Diminutive endoxylic forms of M. denigrata are indistinguishable in the field from the several, 130 BRIAN JOHN COPPINS predominantly lignicolous species with small, black apothecia, but all such species (except M. misella as discussed above) appear very different when examined microscopically. Habitat and distribution: M. denigrata is commonly found on the lignum of fallen trunks and old stumps of broad-leaved and coniferous trees, especially in rather open situations at the edges of woodlands, woodland glades, and hedgerows, etc. The communities in which it occurs are difficult to define, but to give some indication the following list of associated species has been made from collections from fallen decorticate trunks of Pinus in the native pine-woods of Scotland: Buellia griseovirens, Cladonia coniocraea, C. macilenta, Hypogymnia physodes, Lecanora expallens, L. symmicta agg., Lecidea aeruginosa, L. icmalea, Micarea peliocarpa, Mycoblastus sterilis, Parmelia saxatilis, Platismatia glauca, Pseudevernia furfuracea, Xylo- grapha abietina, and X. vitiligo. M. denigrata is a very successful primary coloniser of untreated timber-work. It also occurs on the same substrate whose preservative has lost its effectiveness, or whose painted surface has flaked off. On such substrata M. denigrata has been found in a wide range of situations, e.g. gates, fence posts and rails, garden furniture, picnic tables and seats, window frames of greenhouses, wooden roof tiles (shingles), telegraph poles, and the woodwork of old carts and farm machinery. When on such worked wood it often forms almost pure stands, and is often present as a form with few apothecia but numerous pycnidia containing mesoconidia; the conidia often extruding as white blobs easily visible to the unaided eye or through a X10 lens. Lichens associated with M. denigrata on worked wood in the British Isles include Buellia punctata, Cyphelium inquinans, Hypogymnia physodes, H. tubulosa, Lecanora conizaeoides, L. f 0 2 4 max i Nn w oa A Map7 Micarea denigrata @ 1950 onwards O Before 1950 LICHEN GENUS MICAREA IN EUROPE 131 dispersa agg., L. piniperda, L. pulicaris, L. saligna, L. symmicta agg., L. varia, Lecidea aeruginosa, L. icmalea, Mycoblastus sterilis, Parmelia sulcata, Scoliciosporum chlorococcum, S. umbrinum, Strangospora moriformis, and Thelocarpon laureri. Further evidence of the pioneering abilities of M. denigrata comes from the find (Coppins 1888) of it on hardboard lying in a dune-slack, and the observations by Poelt (1977; as M. hemipoliella) on its colonisation of dead leaves of Cladium mariscus in a fen in Bavaria. Other reports on soft vegetable matter are wanting, although I have found it on decaying mosses (with Cladonia chlorophaea agg.) on the slope of a stable sand dune on Holy Island in Northumber- land (Coppins 4456). M. denigrata is occasionally found on the rather dry and loose bark of old trees (e.g. Acer pseudoplatanus, Alnus, Betula, Castanea, Sambucus, Ulmus, and Pinus), usually towards their bases; however occurrences on + smooth bark are very rare (e.g. type material of Lecidea hemipoliella, on Alnus in southern Finland). To my knowledge M. denigrata never occurs on the healthy, living twigs of trees, or shrubs (see ‘observations’ above). There are two British collections on shaded sandstone, one from Fife in a conifer plantation, the other from Yorkshire in a ditch embankment; associated species included Lecidea granulosa agg., L. icmalea, Micarea peliocarpa, and Parmeliopsis ambigua. In addition, it has been found growing with Psilolechia lucida on the east-facing vertical side of a tomb in a Suffolk churchyard. M. denigrata is widely distributed in mainland Britain, but has not yet been reported from Ireland. In mainland Europe it is again widespread although I have not seen material from north of about 67°N; its range extends into eastern Europe and the Balkans and continues to the Caucasus. I have examined North American material from Newfoundland, Washington (state), and Colorado, thus suggesting a widespread distribution in that subcontinent. Exsiccata: Anzi Lich. Ital. 256 (BM). Arnold Lich. Mon. 46 (BM ex K). Britz. Lich. Exs.: 464 p.p. (H). Fries Lich. Suec. 98 (E, UPS). Harm. Lich. Loth. 838 (ANGUC, DUKE). Hepp Flecht. Eur. 14 (E). Johnson Lich. Herb. 373 (BM). Krypt. Exs. Vindob. 3153 (BM, BM ex K), 3651 (BM, BM ex K, M), 4858 (GZU). Kutak Lich. Bohem. 205 (O), 516 (O), 517 (O). Malbr. Lich. Norm. 387 (M). Malme Lich. Suec. 145 (M, S). Migula Crypt. Germ. 132 (BM, E, MANCH). Mougeot & Nestler Stirpes Crypt. 1329 p. min. p. (BM). Norrlin & Nyl. Herb. Lich. Fenn. 177 (BM, H), 745 (BM, H). Poelt Lich. Alp. 22 (BM, M, WIS). Rabenh. Lich. Eur. 626 (BM, BM ex K, E). Samp. Lich. Port. 132 (LD). Vézda Lich. Sel. 1430 (BM). Weber Lich. Exs. 73 (E, DUKE, M, NMW, WIS). Zwackh Lich. Exs. 394 (BM ex K, H). 12. Micarea elachista (K6rber) Coppins & R. Sant., comb. nov. (Figs 1C, 14A) Biatora elachista Kérber, Parerga lich.: 159 (1860). — Catillaria elachista (K6rber) Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 455 (1934). Type: Germany, Baden-Wiirttemberg, Heidelberg, on old trunk of Castanea sativa, W. E. von Ahles (L 910, 138-100-lectotype!; L 910, 138—32-isolectotype!) See note below. Lecidea anomala *L. glomerella Nyl., Lich. Scand.: 203 (1861). — Biatorina glomerella (Nyl.) Arnold in Flora, Jena 53: 474 (1870). — Catillaria glomerella (Nyl.) Th. Fr., Lich. Scand. 2: 578 (1874). — Micarea glomerella (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Handi. III, 18 (3): 75, 85 (1892). Type: Finland, Ostrobottnia australis, Botom, 1859, A. J. Malmgren (H-NYL 19123 — lectotype!). Lecidea poliococca Ny\., Lich. Scand. : 203 (1861). — Catillaria denigrata f. poliococca Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 460 (1934). — Catillaria synothea var. poliococca (Ny1.) Erichsen in Annls mycol. 41: 205 (1943). Type: Sweden, Uppland, ‘Upsalia’, in silva ‘Parken’, ad pinos decorticatos’, 1852, W. Nylander (H-NYL 19144 — holotype!). Lecidea sororians Nyl. in Flora, Jena 58: 445 (1875). — Bacidia sororians (Nyl.) H. Olivier in Bull. Géogr. bot. 21: 168 (1911). Type: Finland, Tavastia australis, Korpilahti, near Raianlahti, on rock with Stigonema sp., 1873, E. A. Lang [Vainio] (H-NYL 17234 — holotype!). Lecidea glomerella f. simplicata Nyl. in Vainio in Medd. Soc. Fauna Fl. fenn. 10: 28 (1883). Type: Finland, Tavastia australis, Evo, ‘supra truncum pineum’, 1874, J. P. Norrlin, Norrlin & Nyl. Herb. Lich. Fenn. 314 (H — lectotype!; isolectotypes: BM!, H! M!). Lecidea glomerella var. poliococcoides Vainio in Medd. Soc. Fauna Fl. fenn. 10: 29 (1883). Type: Finland, Karelia borealis, Lieksa, Vieki, on burnt lignum, 1875, E. A. Vainio (TUR-VAINIO 22326 — holo type!). 132 BRIAN JOHN COPPINS Catillaria elachista var. carbonicola Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 458 (1934). Type: Finland, Tavastia australis, Korpilahti, ‘hiiltyneella kannolla’, 1873, E. A. Lang [Vainio] (H-NYL 19143 - isotype!). Notes on the typification of Biatora elachista K6rber: In the protologue to B. elachista, Kérber (loc. cit.) gives the following habitat and locality information: ‘On alten Striinken der Castanea vesca bei Heidelberg von Hrn. v. Zwackh und Ahles aufgefunden.’ Material borrowed from the K6érber Herbarium (L) consisted of four specimens: i. bei Heidelberg, Zwackh, annotated: ‘Biatora elachista Kbr. nov. sp.’ (L 910, 138-101). ii. Heidelberg, Ahles (L 910, 138-100). iii. identical label as ii. (L 910, 138-32). iv. Forstrevier Goleow bei Rybnik, iii 1872, B. Stein (L910, 138-298). (i-iii) are on Castanea lignum and (iv) is on conifer lignum. The most obvious choice for lectotype is (i) but it is not a Micarea and does not conform to the usual interpretation of the name (e.g. Vainio, 1934: 45; Ozenda & Clauzade, 1970: 401). I do not know the identity of this species, but it may be close to Lecidea hypopta Ach.; a brief description follows: Thallus probably mostly endoxylic but obscured by a pale farinose crust of a non-lichenised alga. Apothecia numerous, reddish brown to brown-black, epruinose 0-15—0-3 mm diam. Young apothecia with a pale margin which is soon excluded as the disc expands and becomes convex. Epithecium reddish brown, turning olivaceous brown in K. Hymenium 30-35 um tall. Paraphyses simple or forked in their upper part, c. 1-1-5 wm wide; upper 5S—15 wm often with pigmented walls and up to 2-5 wm wide. Asci clavate 28-35 x 10-11 wm, 8-spored. Spores simple or 1-(rarely 2-) septate oblong-ellipsoid, straight or slightly curved, contents often becoming brown, 8-13 x3—4 wm. Excipulum hyaline within, reddish brown at edge, often with a penetrating algal layer c. 25 wm wide. Algal cells c. 7-13 wm diam. Pycnidia not found. Specimens (ii) and (iii) are presumably those that K6rber attributed to Ahles in the protologue and are therefore syntypes. Both belong to the species generally known as Catillaria elachista (K6rber) Vainio, and it is with these that the name is lectotypified here. Specimen (iv) is not a syntype and does not belong to either of the above taxa. It has small, brown thinly white-pruinose apothecia, a dark reddish brown hypothecium and small, ellipsoid spores, 7—8 x 2-5-3 ym, and is referable to Lecidea apochroeella Nyl. Thallus effuse, superficial, consisting of dispersed to continuous, convex to subglobose areolae; areolae greenish white or whitish grey, sometimes tinged grey-brown or olivaceous, occasionally dark brown (when on burnt lignum), matt, sometimes + white-pruinose, c. 0-08-0-16(-0-:25) mm diam. Areole in section (Fig. 1C) with a c. 10-12 wm tall, hyaline or greyish (pigment in gel-matrix, K+ violet), algal-free ‘cortex’, composed of interwoven, hyaline hyphae (c. 1-5 wm wide) that separate in K; outer surface of cortex sometimes bound by a hyaline, amorphous, densely gelatinised layer (‘epicortex’), c. 3-5 um tall. This organised structure with a ‘cortex’ and ‘epicortex’ is often disrupted by the invading torulose hyphae of a dematiaceous hyphomycete. Phycobiont micareoid, cells 4-7 wm diam. Apothecia usually numerous, immarginate, convex to + globose, often becoming tuberculate, dark brown to brown-black, matt, (0-08-)0-12-0-3 mm diam, or to 0:8 mm diam when tuberculate. Hymenium 30-40 um tall. Upper part (epithecium) usually well defined, up to 12 wm tall, dark fuscous-brown; pigment concentrated into dense amorphous clumps, but dissolving and fading into solution (without changing colour) in K, HNO3— and not dissolving; epithecium of young apothecia sometimes Kf+ violet due to the additional presence of the dull olivaceous, K+ violet pigment which occurs in greater concentrations in the pycnidia. Remain- ing (lower) part of hymenium, hyaline with dilute yellowish brown vertical streaks. Asci clavate, 23-35 x 10-12 xm. Spores fusiform, oblong-fusiform or ovoid-oblong, often slightly curved, mostly 0-1-septate and slightly constricted at the septum, (9—)11—15(-19) x2-3-5 wm; occa- sionally becoming 2- or 3-septate and up to 20-24 um long; old spores sometimes with brownish contents. Paraphyses numerous, hyaline throughout, 0-8-1 wm wide in mid-hymenium but gradually widening above to 1-7(—2) wm; sparingly branched and sometimes anastomosing, but becoming richly branched above where their entangled apices, together with the brown pigment, form a + well delimited epithecium. Hypothecium 60-150 um tall, pale, tinged dilute yellowish brown, K—, HNO3~-; hyphae hyaline, 1-1-5 wm wide, interwoven, becoming + vertically orientated towards the hymenium; ascogenous hyphae with swollen cells, mostly 2— LICHEN GENUS MICAREA IN EUROPE 133 4 wm wide. Excipulum poorly developed and much reflexed, sometimes evident as a narrow, pale fuscous-brown zone; hyphae hyaline, radiating, branched and anastomosing, 1-1-5 wm wide. Pycnidia usually present and numerous, developing from within areolae but soon becoming emergent and sometimes + sessile; of two types: (a) c. 100-200 um diam, grey-brown but usually paler or whitish around the ostiole, surface smooth and + glossy; ostioles distinct and c. 20 wm diam, sometimes gaping and to 50 wm diam; wall not continuing below the base, laterally c. 23-40 um wide, dilute olivaceous or brownish, K+ violet, and formed of interwoven hyaline hyphae (c. 1-5 wm wide) that + separate in K; conidiogenous cells cylindrical, occasionally percurrently proliferating, c. 4-7x1-2 wm; conidia (mesoconidia) + cylindrical, sometimes slightly wider at proximal end, often faintly biguttulate, 3-5—4-5 x 1-3-1-7(—2) wm. (b) similar in appearance and structure to above, but smaller and 60-100 wm diam; conidiogenous cells cylindrical, c. 3-6X1 um; conidia (microconidia) narrowly cylindrical, (4-)4-5—6(-6-5) x 0-7-1 wm. Chemistry: Thallus K—, C—, KC—, PD-—; sections of apothecia C—; no substances detected by t.l.c. Observations: The combination of + globose areolae, + globose, brown apothecia, dense brown (dissolving in K) epithecial pigment, C— hymenium, fusiform, 1—3-septate spores and distinctive, + glossy pycnidia usually make this species easy to identify. Some confusion has been made with shade forms of M. denigrata, but these can be distinguished by their rather adnate and often larger apothecia, C+ orange-red (gyrophoric acid) hymenium and thallus, never glossy, thin-walled pycnidia, and, when present, curved macroconidia. M. elachista is closely related to M. rhabdogena (q.v.), which differs in having an endoxylic thallus, smaller, mostly simple, spores, and black pycnidia. The areolae of M. elachista sometimes have a white-pruinose appearance: the ‘pruina’ resulting from the partial disintegration of the thin ‘epicortex’. Habitat and distribution: M. elachista is found, often in the company of Parmeliopsis spp., on the lignum (rarely bark) of partially or wholly decorticate trunks or large stumps of old trees, especially Castanea, Pinus, and Quercus. Most collections from France and Germany were made from Castanea, whereas those made from Scandinavia were mostly from conifers. It occasionally occurs on burnt or charred stumps and my collection (Coppins 6017) from Sweden, on Pinus, was accompanied by Chaenotheca ferruginea, Hypocenomyce friesii, Lecidea granu- losa agg., Micarea melaena, and Parmeliopsis spp. It seems to be rare on worked timber, but was collected on old fence-posts in Bavaria by Arnold. I have seen only one saxicolous gathering of M. elachista, i.e. the holotype of Lecidea sororians from southern Finland, with which occurred colonies of a Stigonema sp., a few lobes of a brown Parmelia (? P. verruculifera) and, according to Nylander, Biatorella torvula (not present in the existing material). This specimen is very small and in poor condition, and the few spores seen appeared to be abnormally developed, 1—3-septate, 15-24x2-2:5 um; other features of the thallus and apothecia agree with M. elachista (pycnidia were not found) and I think it is most unlikely to represent a distinct taxon. M. elachista appears to avoid the more strongly oceanic areas of Europe, and is known from mid-Sweden, southern Finland, France (Haute Vienne, the Massif Central and the Pyrénées), southern Germany (Hessen, Baden-Wirttemberg and Bavaria), and the Austrian Tirol. Its presence in Britain is a possibility and it should be sought for, especially in the central Highlands and east Scotland, and the Welsh border counties. Exsiccata: Arnold Lich. Exs. 1471 (BM ex K, M). Arnold Lich. Mon. 246 (BM ex K, M). Malme Lich. Suec. 21 (M, S). Norrlin & Nyl. Herb. Lich. Fenn. 314 (BM, H, M); 724 (BM). Vézda Lich. Sel. 1134 (BM). Zwackh Lich. Exs. 122 (H-NYL 18828, M). 134 BRIAN JOHN COPPINS 13. Micarea eximia Hed. (Figs 14B, 40C) in Bih. K. svenska VetenskAkad. Hand. III, 18 (3): 80, 84, 95 (1892). — Catillaria malmeana Zahlbr., Cat. lich. univ. 4: 56 (1926); nom. nov.; non Catillaria eximia Malme. Type: Sweden, Dalarna, Alvdal, Hallstugen, vi 1891, J. T. Hedlund (S — lectotype! [t.l.c.: no substances]; S — isolectotype!; material distributed as Malme Lich. Suec. Exs. 26 is possibly part of this collection). Thallus effuse, endoxylic, of minute clusters (c. 15-40 wm diam) of the phycobiont amongst the wood fibres, with intertwining hyphae that often have green walls, K—, HNO3+ red. Phycobiont micareoid, cells 4-7 wm diam. Apothecia numerous, immarginate, + convex-globose from the start, often becoming tuberculate, black, matt or slightly glossy, 0-1-0-2 mm diam, or to 0-4 mm diam when tuberculate. Hymenium 30-45 um tall; upper part bright to dark aeruginose, K— or dulling, HNO3+ purple-red; remaining (lower) part + hyaline with aeruginose vertical streaks. Asci clavate, c. 30-40x11-12 um. Spores oblong-fusiform, sometimes slightly curved, 9-14 (—16) x 1-8—2:5 wm. Paraphyses rather scanty, branched and anastomosing, thin and 0:7-0-8 wm wide, but upper c. 5-17 wm with thickened, pigmented walls and then 2-3 um wide; usually a few paraphyses have pigmented walls throughout their length and are about 2 wm wide in the middle hymenium. Hypothecium 45-120 um tall, dilute reddish- or purplish brown, sometimes with darker blotches, K+ dull greenish, HNO3+ purple-red, pigment confined to gel-matrix; hyphae interwoven, hyaline, c. 1-1-5 wm wide; ascogenous hyphae with swollen cells up to 4 wm diam. Excipulum indistinct, sometimes evident as a dark green or brownish (K+ olivaceous) reflexed zone with branched, radiating, pigmented hyphae c. 1-5-2 wm wide, Pycnidia numerous but inconspicuous, usually immersed between surface wood fibres, but sometimes + emergent, black, c. 35-80 wm diam; walls dark green, K—, HNO3+ purple-red, composed of tightly bound, pigmented hyphae, c. 2 wm wide. Conidia (mesoconidia) cylindri- cal, sometimes biguttulate, 3-9-5-5 x 1-1-4 um. Chemistry: Sections of apothecia and thallus C—; no substances detected by t.1.c. Observations: Micarea eximia is characterized by the combination of an endoxylic thallus, small, black, + globose or tuberculate apothecia, bright green upper hymenium, reddish or purplish brown (K+ green) hypothecium, simple or 1-septate, oblong-fusiform spores, and an absence of stalked pycnidia. It is most likely to be confused with M. contexta and M. olivacea. The former has broader, ovoid spores and a darker hypothecium; and the latter has a less brightly coloured hymenium, a darker olivaceous hypothecium, more numerous and broader paraphyses, shorter and slightly broader spores with rounded apices, and shorter mesoconidia. M. nigella differs from M. eximia in having a purplish brown upper hymenium, a darker hypothecium (hyphal walls pigmented), ellipsoid to oblong-ovoid, simple spores, and stalked pycnidia. The spores of M. eximia are similar to those of M. denigrata, but that species has an olivaceous, K+ violet pigment in its upper hymenium and pycnidial walls, and a + hyaline hypothecium. Habitat and distribution: M. eximia is a rare or overlooked species of conifer lignum, known only from middle Sweden and northern Finland. Associated species on the specimens examined include Bacidia retigena, Calicium glaucellum, Cetraria pinastri, Cladonia spp., Lecidea pullata, Micarea misella, Parmeliopsis aleurites, P. ambigua, P. hyperopta, and Xylographa vitiligo. 14. Micarea globulosella (Nyl.) Coppins, comb. nov. (Figs 15, 43A—B; Map 24) Lecidea globulosella Nyl., Lich. Jap.: 69 (1890). — Bacidia globulosella (Nyl.) Zahlbr., Cat. lich. univ. 4: 202 (1926). Type: Japan, Yokohama, on bark, 1879, E. Almquist (S—lectotype!; isolectotypes: H-NYL 17412! and 17413!, S!). Thallus effuse, of scattered or, more usually, + contiguous areolae. Areolae convex, whitish or grey, not gelatinous when wet, 40-150 wm diam; in section without an amorphous covering LICHEN GENUS MICAREA IN EUROPE 135 layer, outer hyphae sometimes olivaceous (or brownish) and K+ violet. Phycobiont micareoid, cells 4-7 wm diam. Apothecia numerous, immarginate, convex and adnate to subglobose, sometimes becoming tuberculate, greyish or brownish black, sometimes pale grey or pallid (shade forms), 0-1-0-3 mm diam. Hymenium 35-40 um tall, dilute olivaceous or dilute olive-brown, K+ violet. Asci clavate, 30-35 x9-12 um. Spores fusiform-acicular or + rod-shaped, slightly curved or + straight, 0-3(—6)-septate, 13-26x(1-5—)2-2:5(-3) wm. Paraphyses numerous, branched and sometimes anastomosing, c. 1 wm wide, sometimes widening to 1-5 wm towards their apices; apical walls hyaline. Hypothecium 50-70 um tall, hyaline. Excipulum indistinct, evident in some sections as a narrow, reflexed, lateral border to the hymenium, hyaline or dull olivaceous (then K+ violet); hyphae radiating, branched and anastomosing, c. 1 ~m wide. Pycnidia frequent but inconspicuous, immersed or emergent, whitish to dark grey; walls olivaceous or dull brownish, K+ violet (especially around the ostioles), sometimes + hyaline. Pycnidia of two types: (a) immersed within large areolae, sometimes emergent, 60-100 um diam; conidia (mesoconidia) + cylindrical, 3-6-5-3 x 1-1-4 um; (b) similar but smaller, c. 30-40 pum diam; conidia (microconidia) narrowly cylindrical, 3-8-5 x0-8—1 wm. Chemistry: Thallus C+ red, K—, PD—; apothecia sections C— or C+ red (also C+ violet due to olivaceous pigment); t.l.c.: gyrophoric acid. Observations: M. globulosella is closely related to M. denigrata and M. nitschkeana but can be distinguished by its longer, almost acicular, or rod-shaped spores. My early suspicions that M. globulosella could be a long-spored variant of M. nitschkeana were removed by the examination of microconidia, those of M. nitschkeana being significantly longer (mostly 5-5—7 wm). Long, curved macroconidia, as found in M. denigrata and M. nitschkeana, have not yet been found in M. globulosella, but they should be sought for in additional material. See under M. synotheoides for differences from that species and Bacidia beckhausii. The name Micarea bacidiella (a synonym of B. beckhausii) was mistakenly applied to M. globulosella by Vézda & Wirth (1976) and Poelt & Vézda (1977). Habitat and distribution: M. globulosella is a rare but widespread species, so far known to me from Wales, Sweden, Finland, south-east France, Bavaria (Allgau), Czechoslovakia (High Tatra), Canada (Quebec), and Japan (Honshu). It seems to prefer conifer bark, but at the Welsh locality it occurred on the top of an old gate in a wooded valley, and was accompanied by Lecanora piniperda, Lecidea aeruginosa, and L. icmalea. The specimen from Allgauer Alpen is associated with Graphis scripta, Menegazzia terebrata, Stenocybe major, and Frullania sp. on Abies; that from Sweden is on Picea with Cetraria pinastri and Ptilidium pulcherrimum; and those from Czechoslovakia are on Picea, with Cetraria chlorophylla, Hypogymnia physodes, and Ptilidium pulcherrimum. At the French locality it was growing on an old basidiome of the polypore Daedaleopsis confragosa on Pinus hapelensis. It seems to require humid conditions in old forest situations, but has a more widespread distribution than M. synotheoides which is strongly oceanic. M. globulosella may have a greater tolerance to cold winter temperatures, such as occur in the central European montane regions. Exsiccata: Rasanen Lichenoth. Fenn. 426 (hb Vézda). 15. Micarea hedlundii Coppins, sp. nov. (Figs 14C, 44A) Thallus effusus, olivaceo-viridis, subtiliter granulosus ad 0-4 mm crassus, constatus ex goniocystis; goniocystae c. 20-40 wm diam, omnes cum pigmento flavo-brunneolo, K+ purpureo-violaceo et oleoso. Algae cellulis 4-7 ~m diam. Apothecia vulgo pauca vel etiam nulla, immarginata, convexo-hemisphaerica mox tuberculata, pallida vel griseo-fusca demum obscure fusca, 0-15—0-5 wm diam. Hymenium c. 35 wm altum, + hyalinum vel p.p. dilute olivaceo-brunneolum, K+ violaceum. Ascosporae ellipsoideae, ovoideae vel oblongae, simplices interdum 1- septatae, 6-5—10(—12) x2-3+4 wm. Paraphyses aliquantum paucae, ramosae, c. 0-7—1(-1-5) um latae, apicibus vix incrassatis, incoloratis, Hypothecium pallidum. 136 BRIAN JOHN COPPINS Excipulum paulum evolutum. Pycnidia numerosa, conspicua, stipitata, 0-1—-1-0 mm alta, 0-07-0-14 mm diam, cum stipibus simplicibus vel ramosis, toto griseo-brunneo vel roseo-brunneo cum tomento exili albido. Conidia oblongo-ellipsoidea vel oblongo-ovoidea, (4—)4-5—5-5(—6) x 1:3-1-7 wm. Typus: Norvegia, Opplandia, par. Ringebu, ad oriento-boreo-orientum ex Ringebu, ad Sgraa, inter Nyhamnsbekken et Ulveslabekken, alt. 400 m, ad truncum decorticatum, 25 viii 1979, leg. L. Tibell 8657 (UPS — holotypus). Thallus epixylic or over bryophytes on lignum, effuse, dull olive-green, consisting of fine granules (goniocysts) forming a loose crust to c. 0-4 mm thick. Goniocysts c. 20-40 wm diam, the centre of each with a yellow-brown or dull orange pigment which in K appears as purple-violet oily droplets. Phycobiont micareoid, cells 4-7 wm diam. Apothecia few or sometimes absent, immarginate, convex and soon becoming tuberculate, pallid to grey-brown or dark brown, 0-15—0-5 mm diam. Hymenium c. 35 wm tall, + hyaline with vertical streaks of straw-brown or pale olive-brown, K+ violet, C+ violet, HNO3+ reddish; lower hymenium sometimes with dull orange, K+ purple-violet (oily droplets) pigment. Asci clavate, c. 30X12 wm. Spores ellipsoid, ovoid or oblong, simple, a few sometimes 1-septate, 6:5—10(-12) X2:3-4 wm. Paraphyses rather scanty, branched and sometimes anastomosing, 0-7-1(-1-5) um wide, apices not swollen or pigmented. Hypothecium c. 70-100 um tall, hyaline, or with dull orange, K+ purple-violet (oily droplets) pigment in upper part; hyphae interwoven, becoming vertically orientated towards the hymenium, c. 1-1-7 wm wide, intermixed with swollen short-celled ascogenous hyphae. Excipulum indistinct and soon reflexed, sometimes discernible in young apothecia as a narrow, hyaline, non-amyloid zone; hyphae radiating, branched and anastomosing, c. 0-8-1 wm wide. Pycnidia numerous and conspicuous, stalked with one or to about five borne terminally on simple or branched pycnidiophores. Pycnidiophores (including pycnidia) grey-brown or pinkish brown, covered with a thin whitish tomentum, c. 0-1-1 mm tall and 0-07-0-14 mm diam; lower parts of pycnidophores often covered in goniocysts. Pycnidiophore and pycnidial wall tissues composed of hyaline hyphae bound by a dilute reddish brown matrix reacting K+ violet or violet-brown and HNO3+ reddish; surface of pycnidiophores and pycnidia with protruding, slender, flexuose, hyaline tomental hyphae c. 0-7-1 xm wide. Conidiogenous cells ampulliform to + cylindrical, 5-10Xc. 1-5 wm, often swollen at base to 2:5 wm wide. Conidia (mesoconidia) oblong-ellipsoid or oblong-ovoid, (4~)4-5—5-5(—6) x 1-3-1-7 wm. Chemistry: Thallus C—, PD—; apothecia sections C— (but with C+ violet pigment); no substances or traces of ‘prasina-unknown B’ (? contaminant) detected by t.l.c. Observations: Micarea hedlundii is readily identified by its finely granular, darkish green thallus with distinctly stalked, pinkish brown, tomentose pycnidia. Several other Micarea species have stalked pycnidia but in none of them are they tomentose. Another unique feature of M. hedlundii is the dull orange pigment, present in the goniocysts and sometimes in the lower hymenium and upper hypothecium, which appears as purple-violet oily droplets in K. Unlike the olivaceous K+ violet pigment found in this species and others (e.g. M. denigrata, M. nitschkeana, and M. prasina), it remains unchanged in C and 50% HNO3. M. hedlundii is unlikely to be confused with any other Micarea, but is probably closely related to M. prasina. M. anterior has reddish brown, stalked pycnidia but they are glabrous and produce shorter conidia; in addition its apothecia and pycnidia are completely devoid of K+ violet pigments. The chemistry of M. hedlundii is problematical. Of the three specimens tested by t.l.c., the type from Norway contains no detectable substances, but the two specimens from Austria (GZU) and Germany (hb Poelt) appear to contain small amounts of ‘prasina-unknown B’. This substance is known elsewhere only in the type race of M. prasina and it is possible that its detection in the aforesaid samples is due to contamination by that species. However, the production of ‘prasina-unknown B’ by M. hedlundii is a possibility meriting further study, especially as M. hedlundii and M. prasina appear to be closely related to one another. New and carefully collected specimens of M. hedlundii are required to resolve this problem. Habitat and distribution: M. hedlundii occurs on old stumps (? mainly of conifers) in woodlands. Few associated lichens are present on the specimens examined, mostly just a few LICHEN GENUS MICAREA IN EUROPE P37. scattered squamules of Cladonia spp. and fragments of Lepraria spp, although one of the Norwegian collections includes a Chaenothecopsis. M. hedlundii seems to be a very rare species, but is known from scattered localities in Norway, Sweden, Germany (Bayern), Austria (Steiermark), and possibly Switzerland. Etymology: This new species is named in honour of Johan Teodor Hedlund (1861-1953) in recognition of his pioneering study of Micarea included in his doctorate thesis for the University of Uppsala (Hedlund, 1892), a work that has been a constant inspiration during my own studies. 16. Micarea incrassata Hed. (Figs 4C, 16, 41A—B; Map 3) in Bih. K. svenska VetenskAkad. Handl. II, 18 (3): 82, 94 (1892). Type: Austria, ‘supra muscos in summo jungo Kraxentrag circa Brenner, Tirol, c. 2800 m. alt.’, A. Minks (S— holotype!). Lecidea assimilata B. [var.] infuscata Th. Fr., Lich. Scand. 2: 522 (1874). — Lecidea assimilata f. infuscata (Th. Fr.) Vainio in Medd. Soc. Fauna Fl. fenn. 10: 85 (1883). Type: Norway, Sér-Trondelag, Oppdal hd., Dovre, Kongsvoll, Hégsnyta, 17 viii 1863, Th. M. Fries (UPS — lectotype!; UPS — isolectotype! [t.l.c.: no substances]). Thallus growing on bryophytes, plant debris or sandy soil, composed of confluent, convex- verrucose areolae that are intermixed with cephalodia. Areolae dull grey-white, or grey-brown to dark grey, matt, 0-08—0-3 mm diam; in section, usually without a hyaline amorphous covering layer (but sometimes seen in sections of young areolae), hyphae in outer c. 10 wm often with light brown walls. A + algae-free medulla often differentiated. Thallus hyphae c. 1-8-3 wm wide. Phycobiont micareoid, cells 4-7 wm diam. Cephalodia often present, irregularly globose and hidden amongst the areolae but sometimes visible externally as brown areolae-like structures, 0-2-0-6 mm diam; containing Nostoc, cells 3-5 xm diam. Less often present are irregular, rather loose clusters (? cephalodia) of Stigonema. Apothecia numerous, immarginate, convex, + adnate and often partly immersed by sur- rounding areolae, black, matt, 0-3-0-8(—1) mm diam, sometimes forming irregular tuberculate clusters up to 1-2 mm diam. Hymenium 45-50 wm tall; upper c. 10 wm (epithecium) dark aeruginose or olivaceous, K—, HNO3+ red; remaining (lower) part dilute greenish or hyaline. Asci clavate, 45—48x11-14 um. Spores ellipsoid, oblong-ellipsoid, oblong-ovoid or oblong- fusiform, 0-1(—2)-septate, (10—)12-17x4~4-8 wm. Paraphyses numerous, simple below, some- times forked above, sometimes anastomosing, (1—)1-5—2 4m wide, sometimes widening above to 3 wm; apical walls hyaline although surrounded by dense pigment in the surrounding gel matrix. Hypothecium c. 150-400 um tall, sometimes ‘rooting’ to the base of the adjoining areolae, dark red-brown, without a purple tinge, K—, HNO3+ bright orange-brown; hyphae interwoven, or + vertically arranged in upper part, c. 1-7—-2-5(-3) wm wide, surrounded by densely pigmented matrix; ascogenous hyphae c. 2-5-5 wm wide. Excipulum indistinct, sometimes evident as a reflexed, dull olivaceous or reddish brown zone; hyphae radiating, branched and anastomosing, c. 1-2 wm wide. Pycnidia rare, immersed to sessile, black, 30-60 4m diam; walls dark green above, changing to reddish brown at base; conidia (? microconidia) cylindrical or cylindrical-fusiform, 6-9 x 1- 1-3 wm. Chemistry: Thallus K—, C—, PD—; t.].c.: no substances. Observations: See M. assimilata. Habitat and distribution: M. incrassata occurs in much the same habitats as M. assimilata but appears to be more widely distributed. In Europe it has an arctic-alpine distribution ranging from Spitzbergen (c. 79°N) in the high arctic, southwards to the Kola peninsula, central Norway (Opland), and central Sweden (Harjedalen). Further south it occurs in the Scottish highlands (Clova in Angus, and near Loch Merkland in East Sutherland), Denmark, and the Alps (Upper Bavaria, Austrian Tirol, and Switzerland). I have not seen it from Alaska or Canada although it probably occurs in those regions. From further south in North America it has been collected at 138 BRIAN JOHN COPPINS altitudes of c. 3400-3900 m in the Rocky Mountain National Park in Colorado by Anderson (1964). M. incrassata has been collected on Kerguelen Island in the southern Indian Ocean (48°30'S) and is so far the only arctic or arctic-alpine Micarea known to have a bipolar distribution. Exsiccata: Fellman Lich. Arct. 164 (BM ex K, H, H-NYL 16567), 166 (BM ex K, H). 17. Micarea intrusa (Th. Fr.) Coppins & Killias, comb. nov. (Figs 17, 55; Map 22) Lecidea intrusa Th. Fr. in Bot. Notiser 1867: 152 (1867). — Catillaria intrusa (Th. Fr.) Th. Fr., Lich. Scand. 2: 579 (1874). — Lecidiopsis intrusa (Th. Fr.) Zopf in Hedwigia 35: 338 (1896). — Conida intrusa (Th. Fr.) Sacc., Syll. Fung. 18: 187 (1906). Type: Finland, Tavastia australis, Mustiala, 1867, A. Kullhem (UPS - holotype!). Lecidea aphanoides Nyl. in Flora, Jena 51: 476 (1868). Type: Scotland, South Aberdeenshire, Braemar, ‘Craig Guie’, viii 1868, J. M. Crombie (H-NYL 20237 — holotype!); supposed isotype material in BM appears to be all Scoliciosporum umbrinum (Ach.) Arnold. Lecidea melaphana Ny). in Flora, Jena 52: 83 (1869). Type: Scotland, South Aberdeenshire, Braemar, ‘Craig Guie’, viii 1868, J. M. Crombie (BM — isotype!). Lecidea contrusa Vainio in Medd. Soc. Fauna Fl. fenn. 10: 29 (1883); nom. illeg. (Art. 63). Thallus occasionally effuse but more often forming small patches (to c. 1 cm diam) amongst other lichens, irregularly granular-verrucose and often rimose, sometimes forming discrete areolae (c. 0-1-0-3 mm diam) which may coalesce to form ‘composite areolae’ up to c. 1 mm diam and 0-5 mm tall. Thallus dark olive-grey or brownish grey; surface matt, and usually scurfy due to invasion of free-living algae. In section, sometimes with a hyaline amorphous covering layer up to 12 wm thick but this is usually disrupted by invading algae; walls of uppermost hyphae sometimes thickened with olive-green pigment, K—, HNO3+ red; phycobiont layer c. 70-100 mm thick, above a + distinct hyaline medulla. Phycobiont not micareoid; cells globose, large, 7-21 wm diam, with thick hyaline walls 1-2 wm thick, each cell deeply penetrated us haustorium of the mycobiont (Fig. 55). Apothecia numerous, immarginate, convex, adnate, black, or slightly glossy when young, 0-14-0-4 mm diam. Hymenium 40-S0 um tall; upper (c. 20 wm) part aeruginose-green, K—, HNO3+ red; remaining (lower) part + hyaline. Asci clavate, 35-45 x 14-15 um; cytoplasm of asci and spores sometimes with an orange, K+ purple-red pigment. Spores ellipsoid or ovoid-ellipsoid, occasionally oblong-ellipsoid or oblong fusiform and then sometimes slightly curved, simple or 1-septate and septum often eccentric, rarely 3-septate (7—)9-14(-17) x (4-)4-5-5-5(-6) wm. Paraphyses numerous, branched and anastomosing, c. 1-1-5(—-1-7) wm wide; apices not swollen or pigmented (green pigment confined to the surrounding gel-matrix). Hypothecium c. 110-200 mm tall, hyaline or with very faint olivaceous brown tinge in places, Fig.55 Micarea intrusa: a phycobiont cell penetrated by a haustorium of an associated mycobiont hypha. Scale = 10 um. LICHEN GENUS MICAREA IN EUROPE 139 upper part sometimes pale orange, K+ purple-red (see below); hyphae interwoven, but becoming vertically orientated towards the hymenium, hyaline, c. 1-7—2 wm wide; ascogenous hyphae short-celled, to 5 wm wide, contents sometimes with dilute orange, K+ purple-red pigment. Excipulum reflexed to below the surface of surrounding thallus, thin, dilute brownish or + hyaline in places, sometimes darkish brown at outer edge; hyphae radiating, branched and anastomosing, c. 1-5—2 wm wide. Pycnidia not seen. Chemistry: Thallus surface and sections K—, C—, KC—, PD—, I—; apothecia sections C—; not analysed by t.l.c. Observations: This species is placed in Micarea with some hesitation, owing to its unusual habitat (for a Micarea) and large-celled, thick-walled, non-micareoid phycobiont. However, much the same habitat is shared by M. subnigrata (a species with a typical micareoid phycobiont) and no other fungal characters appear to merit its exclusion. It is similar in appearance to M. subnigrata which can be distinguished by its brown epithecium and usual presence of pycnidia containing helicoid macroconidia. Superficially, M. intrusa is virtually indistinguishable from Table5 Species associated with Micarea intrusa in Norway (see text). + vertical + horizontal rocks rocks Aspicilia morioides + Caloplaca cf. lamprocheila + Candelariella vitellina Cornicularia normoerica 8 Fuscidea cyathoides F. intercincta F. tenebrica Haematomma ochroleucum Lecanora badia L. gangaleoides L. intricata ac L. polytropa L. cf. salina Lecidea fuliginosa a L. fuscoatra ate L. lactea 5 cts L. lapicida re L. leucophaea L. vorticosa + Micarea subnigrata Opegrapha gyrocarpa Pertusaria dealbescens ca P. lactea + Placopsis gelida Porina chlorotica P. guentheri Ramalina subfarinacea Rhizocarpon geographicum a R. lecanorinum 5 R. riparium subsp. lindsayanum so Rinodina gennarii Schaereria tenebrosa 2 Stereocaulon vesuvianum i Trapelia involuta af Umbilicaria spodochroa a ++ +++ +4+4+4+4++4+ +++4+4 oo 140 BRIAN JOHN COPPINS forms of Scoliciosporum umbrinum with a well developed thallus, but that species has vermiform or sigmoid-curved spores. The phycobiont in M. intrusa and S. umbrinum appears to be the same; any future critical appraisal of the delimitation of Scoliciosporum should seriously consider the generic disposition of M. intrusa. The pale orange, K+ purple-red pigment commonly found in the ascogenous hyphae, asci and spores has not been seen by me in any other Micarea, although it is conceivable that it is the same, or similar, to the pigment found in the goniocysts of M. hedlundii (q.v.). Habitat and distribution: M. intrusa occurs on hard siliceous (igneous) rocks in communities referable to the Umbilicarietum cylindricae in its broad sense (James et al., 1977). In Nylander’s protologue of Lecidea aphanoides it is said to occur on calcareous rock, but this is erroneous; application of 50% HNO; to fragments of substratum from the holotype produced no efferves- cence. The formation of small patches amongst other crustose lichens indicates that M. intrusa may be, at least facultatively, parasitic. Such suggestions have been previously propounded by Magnusson (1942), Poelt (1958), and Wirth (1973), all of whom related its behaviour to that of Lecanora intrudens and Lecidea furvella. In several collections, including the type of Lecidea intrusa, it occurs amongst the areolae of Huilia panaeola, but it is by no means confined to that ‘host’. From her studies on the island of Sotra in Hordaland, Norway, Miss L. Skjolddal informs me that M. intrusa occurs on sheltered, sunny, west or south-west facing exposures of gneiss and amphibolite, on steep surfaces or, in one case, on a + horizontal surface. Miss Skjolddal kindly provided me with a list of associated species (Table 5). M. intrusa is probably widespread in areas of western Europe with exposed, hard, igneous rocks; however, it is a very inconspicuous species and records are wanting from many potential localities. I have seen material from southern Scandinavia (several localities), North Norway (Finnmark), and the Grampian Mountains of Scotland. In addition, Wirth (1973) cites collec- tions from France (Vosges) and West Germany (Schwarzwald). Exsiccata: Norrlin & Nyl. Herb. Lich. Fenn. 182 (BM). 18. Micarea leprosula (Th. Fr.) Coppins & A. Fletcher (Figs 18, 53-54; Map 8) in Fletcher in Lichenologist 7: 111 (1975). — Bilimbia milliaria y leprosula Th. Fr., Lich. Scand. 2: 382 (1874). — Micarea violacea var. leprosula (Th. Fr.) Hedl. in Bih. K. svenska VetenskAkad. Handl. III, 18 (3): 81, 92 (1892). — Bilimbia leprosula (Th. Fr.) H. Olivier in Bull. Géogr. bot. 21: 179 (1911). — Bacidia leprosula (Th. Fr.) Lettau in Hedwigia 52: 133 (1912). Type: Sweden, Uppsala, Witulfsberg [Wittuls- berg], 26 vi 1852, Th. M. Fries (UPS — lectotype! [t.1.c.: argopsin and gyrophoric acid]). Thallus effuse, superficial, of convex to subglobose areolae c. 100—350(—400) wm diam; these usually coalesce toward the centre of the thallus, whence they may proliferate, producing secondary granular-areolae, such that the thallus becomes thicker (to c. 700 wm thick). Areolae blue-grey or, more rarely, grey-brown, matt with a minutely roughened surface with white flecks (x50 lens); lower sides of + globose areolae, and areolae in shaded situations, often unpig- mented and greenish white. Areolae fragile and often breaking down or eroding to form sorediate patches with irregularly shaped soredial granules c. 20-50 wm diam. Sections of intact areolae ecorticate and without an amorphous hyaline covering layer; outermost hyphae often coloured with greenish or brownish pigment, K—, HNO3+ reddish. Phycobiont micareoid, cells 4-7 um diam. Apothecia often absent, at first adnate but often becoming constricted below, convex to subglobose, commonly tuberculate, immarginate or rarely very faintly marginate when young, matt or slightly glossy, dark blue-grey or black, 0-15—0-4 mm diam, or to 0-7 mm diam when tuberculate. Hymenium 45-60 um tall, hyaline or dilute green, but dark green (K—, HNO3+ red) in upper part (epithecium). Asci clavate, 40-55x14-16 wm. Spores oblong-ellipsoid, oblong-fusiform or clavate-fusiform, often curved, (1—)3-septate, 14~-26(-29) x4-S(-5-5) wm. Paraphyses numerous, branched and anastomosing, (0-7—)1—1-5 u~m wide; apices often more LICHEN GENUS MICAREA IN EUROPE 141 richly branched, not or only slightly incrassate (to 1-8 ~m wide) and without closely adhering pigment (epithecial pigment confined to surrounding gel-matrix). Hypothecium c. 35-45 wm tall, but becoming much taller in markedly convex or tuberculate apothecia, dilute straw- or fuscous-brown, pigment confined to gel matrix; hyphae interwoven, but becoming outwardly orientated towards the hymenium and excipulum, c. 0-8—1-5 wm wide; ascogenous hyphae with swollen cells, c. 2-6 wm wide. Excipulum distinct in young, moderately convex apothecia, but reflexed and obscured in subglobose or tuberculate apothecia, dilute fuscous brown (K—); hyphae radiating, branched and anastomosing, c. 0-5—1-5 wm wide. Pycnidia not found. Chemistry: Thallus K—, C+ red, PD+ red; apothecia in section C+ red; t.l.c.: argopsin and gyrophoric acid. Observations: M. leprosula is characterized by its sterile or sparingly fertile, granular thallus composed of blue-grey (or brown-grey), fragile areolae which often dissolve (or become abraded) to form contrasting, yellowish green, sorediose patches. These characters are shared by the much rarer M. subleprosula, and the thalli of both react C+ red. However, M. subleprosula can be separated by the PD + yellow reaction of its thallus due to the presence of alectorialic acid. When fertile M. subleprosula is found to have much larger, mostly 7-septate spores. M. leprosula is usually distinctive in the field (although material should always be checked with M. subleprosula in mind) but can easily be overlooked when it occurs on bryophytes on shaded rocks or trees, whence the thallus often lacks its characteristic bluish tinge and the areolae are thinly scattered. The species is usually sterile but fertile specimens are occasionally encountered, especially in sheltered (but not deeply shaded) situations, such as on boulders and old walls in woodland. Hedlund (1892) regarded M. leprosula as a variety of M. peliocarpa (q.v.) and the apothecia of the two species are somewhat similar. However, those of M. leprosula exhibit several minor, yet significant, differences; spores tend to be slightly longer; apothecia tend to be more markedly convex and more frequently tuberculate; hypothecium and excipulum are dilutely pigmented with a dull brown pigment; and paraphyses never become thickened and pigmented at their apices. The two species also differ in several fundamental aspects of thallus morphology and chemistry. With regard to the production of argopsin and to the presence of a dull brown pigment in the excipulum and hypothecium M. leprosula shows some affinity to M. lignaria var. lignaria, with which it often occurs. However, M. lignaria lacks gyrophoric acid, has usually longer and more septate spores, and somewhat stouter and only sparingly branched paraphyses. The areolae of M. leprosula and M. subleprosula differ from those of such species as M. lignaria, M. peliocarpa, and M. cinerea, in lacking an amorphous hyaline covering layer (see ‘morpho- logy’). Habitat and distribution: M. leprosula is widely distributed in upland areas in the north and west of Britain, occurring at altitudes from about sea-level to at least 1100 m. It is found, usually on moribund bryophytes (e.g. Andraea spp. and Rhacomitrium spp.) on rocky ledges, boulders, and old walls, in exposed hilly areas, mountain sides or woodlands. Associated lichens noted amongst British collections include Arthrorhaphis citrinella, Baeomyces rufus, Belonia incar- nata, Cladonia cervicornis, C. coccifera, C. crispata, C. portentosa, C. squamosa, C. subcervi- cornis, Coelocaulon aculeatum, Lecidea granulosa, L. icmalea, Lepraria neglecta, Micarea lignaria, M. peliocarpa, and Vorarlbergia renitens. From Scotland I have seen two corticolous specimens, on trunks of Alnus and Betula; associated species present included Cladonia coniocraea, C. squamosa, Lecidea icmalea, Micarea melaena, and Platismatia glauca. There are two outlying localities in the lowlands of southern England. In Dorset it occurs on waste heaps of slag clay with M. lignaria (q.v.); and in Kent it was found with M. peliocarpa on wood chips lying at the side of a woodland track in a chestnut (Castanea) coppice. M. leprosula is poorly recorded outside Britain, probably because it is so often sterile. To date, I have seen one specimen from Norway (Hordaland) and several collections from mid- and southern Sweden. Further south it seems to be well represented in the Alps, and I have seen one 142 BRIAN JOHN COPPINS Fo es Fa & aR eae aaa Hea oe a hon é BY Daler a ol eile ire ee » fer , Ee : | Leh ia ee 7 Pp ; Kit Map 8 Micarea leprosula @ 1950 onwards O Before 1950 collection from Bohemia (Czechoslovakia). This Bohemian record, and another from the Black Forest (Schwarzwald) were made from exsiccate specimens (Kutak 417 and Migula 1) of M. lignaria growing on wooden roof shingles. Exsiccata: Kutak Lich. Bohem. 417 p. min. p. (0). Magnusson Lich. Sel. Scand. 208 (BM, C, GZU). Migula Crypt. Germ. 1 p. min. p. (C, E, M). 19a. Micarea lignaria (Ach.) Hed. var. lignaria (Figs 1A, 3D-E, 19, 45, 53-54; Map 9) in Bih. K. svenska VetenskAkad. Handl. II, 18 (3): 82, 93 (1892). — Lecidea lignaria Ach. in K. svenska VetenskAkad. Handl. 1808: 236 (1808). — Bilimbia lignaria (Ach.) Massal., Ric. Lich. Crost.: 121 (1852). — Bacidia lignaria (Ach.) Lettau in Hedwigia 52: 132 (1912). Type: Sweden: ‘Svecia’, on lignum, (H-ACH 265 — lectotype; BM — ? isotype!). Lecidea milliaria Fr. in K. svenska VetenskAkad. Handl. 1822: 255 (1822). Type: Sweden, Smaland, Femsj6, on lignum, E. M. Fries (UPS — lectotype! [t.l.c.: argopsin]). Lecidea geomaea Taylor in Mackay, Fl. Hib. 2: 124 (1836). Type: Ireland, South Kerry, Dunkerron, T. Taylor (BM -— lectotype!; BM ex K — isolectotype!). Bilimbia milliaria o.. [var.] lignaria** [f.] calamophila Korber, Parerga lich.: 171 (1860). Type: Germany: ‘an Schilfrohrdachern um Minster’, J. G. F. X. Lahm (L 910, 144-1685 — lectotype!; L 910, 144-1689 — isolectotype! [t.l.c.: argopsin]. Bilimbia milliaria B. [var.] saxicola Korber, Parerga lich. : 171 (1860). Type: Germany: ‘Bilimbia trochiscus Korb. prim, B. millaria 8. saxicola Kb.! Extersteine’, on sandstone, ? Beckhaus (WRSL - lectotype!). LICHEN GENUS MICAREA IN EUROPE 143 Paratype: Germany: ‘Ibbenbiiren in Westfalen, Lahm, Hb. Hepp.’, on sandstone (BM!). Stereocauliscum gomphillaceum Nyl. in Flora, Jena, 48: 211 (1865). — Micarea gomphillacea (Nyl.) Vézda in Folia geobot. phytotax., Praha 5: 321 (1970). — Bilimbia milliaria f. [as ‘f?’] gomphillacea (Nyl.) Blomb. & Forss., Enum. Pl. Scand.: 82 (1880). — Micarea lignaria [as ‘ligniaria’| f. gomphillacea (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Hand. III, 18 (3): 94 (1892). Type: Finland, Tavastia australis, Hollola, Tiirismaa, Pirunpesa, [on granite rock], 1863, J. P. Norrlin (H-NYL 40217 — lectotype! [t.1.c.: argopsin]; isolectotypes: H-NYL 40214!, 40215!, 40216!, 40218! and p.m. 631!). Lecidea sabuletorum var. milliaria f. nigrata Nyl. in Not. Sdallsk. Fauna Fl. fenn. Forh. 8: 151 (1866). Type: USSR, Kola Peninsular, Lapponia ponojensis, Ponoi, 1863, N. J. Fellman (H —holotype!; H—isotype!). See note (i) below. Lecidea milliaria var. triseptata Ny\. in Lamy in Bull. Soc. bot. Fr. 25: 441 (1878). — Bilimbia triseptata (Nyl.) Arnold in Flora, Jena 67: 572 (1884), non Bacidia triseptata (Hepp in Zoll.) Zahlbr. Type: France: Haute Vienne, ‘Sur des roches entre Chalard et St Junien, Lamy (H-NYL 18367! — lectotype, as ‘L. milliaria var. 3-septata Nyl.’). See note (ii) below. Lecidea meizospora Harm. in Bull. Séanc. Soc. Sci. Nancy II, 33 (1898): 63 (1899). — Bacidia meizospora (Harm.) Zahlbr., Cat. lich. univ. 4: 122 (1926). Type: France, Vosges, Docelles, on Calluna roots, V. and H. Claudel & F. J. Harmand (ANGUC - lectotype!). Lecidea trisepta var. polytropoides Vainio in Ark. Bot. 8 (4): 106 (1909). Type: USSR, Magadanskya Oblast (N.E. Siberia), ‘Pitlekai, peninsula Jinretlen’ [c. 67°7'N 173°24’W], on plant debris, 1878-9, E. Almquist (TUR-VAINIO 21274, fertile part — lectotype!). See note (iii) below. Patellaria milliaria var. xylophila Wallr., Fl. crypt. Germ. 1: 349 (1831); nom. inval. (Art. 26). Lecidea milliaria var. saxigena Leighton, Lich. Brit. Exs. 210 (1856); nom. nudum (Art. 32); see note under M. peliocarpa. Type: England, Shropshire, Neescliff [Nesscliff] Hill, Leighton, Lich. Brit. Exs. 210 (E — lectotype!; isolectotypes: BM! DBN! MANCH! NMW!). Notes: (i) The type material mainly consists of the black stroma of Bryomyces gymnomitrii Dobb. on Gymnomitrium sp., with only a few apothecia of M. lignaria present. (ii) Lamy (loc. cit.) gives the locality as ‘entre le Chalard et Saint-Yrieix’. The label on the lectotype specimen is entirely in Nylander’s handwriting and there may have been an error in transcription. (iii) Vainio’s diagnosis included the description of soralia. A small portion of the type specimen is a sterile sorediate thallus (C—, PD—; phycobiont cells 7-14 wm diam) but it is not a Micarea and is not part of the larger, fertile portion. The latter belongs to Micarea lignaria and is chosen as lectotype. Thallus effuse, partially endoxylic or endocuticular, but more usually developed superficially as convex to + globose areolae. Areolae c. 80—250(—300) wm diam, discrete, scattered to contiguous, sometimes coalescing to form an uneven crust, grey-white and + glossy to grey-green or bluish grey and matt, sometimes brown-grey due to the ramification through their surface of brown, toruloid hyphae (cells c. 5—74-5—7 um) belonging to an unknown fungus. Areolae in section, with a hyaline amorphous covering layer (c. 4-12 xm tall); walls of hyphae in outermost (exposed) parts often blue-green (K—, HNO3;+ red); algal layer usually in direct contact with substratum, but in larger areolae the algal layer may be c. 90-135 um tall lying above a medulla. Medulla, when present, up to c. 100 um tall, consisting of loosely interwoven hyphae c. 1-1-5 wm wide, intermixed with dead algal cells and debris derived from the substratum. Phycobiont micareoid, cells c. 4-7 wm diam. Apothecia scattered, more usually numerous and crowded, often confluent, immarginate, convex-hemispherical, soon becoming constricted at the base and hence + globose, occasionally becoming stipitate with a grey-black ‘stipe’ up to 1 mm tall (‘f. gomphillacea’), black, matt or slightly glossy, rarely blue-grey (shade forms), 0-15—0-6(-0-9) mm diam. Hymenium 50-75 wm, often not sharply delimited from the hypothecium, dilute olivaceous or aeruginose-green (K—, HNO3-+ red) in upper, and sometimes lower, part(s) (the middle part often being + hyaline); in the uppermost part (epithecium) the pigment is more dense and is present not only in the gel matrix but also in the walls of the paraphyses; in reflexed parts of the hymenium towards the base of the apothecium the green pigment is often replaced by a dilute brown pigment; minute (less than 2X1 um) granules of violet (K+ intense aeruginose) pigment are sometimes present and lightly scattered through the hymenium. Asci clavate, 45-50X11-19 wm. Spores fusiform, straight or slightly curved, (0-)3—7 septate, 16-36(—38) x4-6(—7) wm. Paraphyses numerous, simple or slightly branched above, but more richly branched in the transition zone between the hymenium and excipulum, sometimes anastomosing, 1-3-1-8 wm wide; apices often slightly 144 BRIAN JOHN COPPINS incrassate and coloured with green or greenish brown pigment and then up to 3 wm wide, usually coherent (even in K) and, together with the surrounding pigmented matrix, form an epithecium. Hypothecium c. 100-230(—350) «um tall, dilutely pigmented with pigment confined to gel matrix; upper part dull aeruginose or olive-brownish K— or + dulling, HNO3+ reddish); lower part often without greenish tinge, and then being dilute brownish or + hyaline; hyphae interwoven or some + vertically orientated near the hymenium, c. 1-1-7 4m wide; ascogenous hyphae with short, swollen cells c. 2-5—4 wm wide. Excipulum + distinct in sections of young, hemispherical apothecia, but soon becoming strongly reflexed and not sharply delimited from the hymenium, dilute brown or sometimes darkish brown along the outer edge. Hypothecial and excipular tissues sometimes elongated vertically to form a stipe (‘f. gomphillacea’). Pycnidia inconspicuous, + immersed, with walls green (K—, HNO3+ red) in exposed upper parts and + hyaline in immersed lower parts; of three types [only type (c) is common]: (a) c. 100 wm diam; conidia (macroconidia) curved or hamate, 0-3-septate, 16-22 xc. 1 wm; (b) c. 100-140 yum diam; conidia (mesoconidia) + cylindrical, oblong-ellipsoid, obovoid or oblong-obovoid, usually distinctly truncate at proximal end, often 2-3-guttulate, 4-7(—7-6) x 1:2-1-8 um; (c) c. 40-50 wm diam; conidia (microconidia) narrowly cylindrical, only faintly truncated at proximal end, eguttulate, (4-5—)5—7(-8) x0-8-1 um. Chemistry: Thallus K-—, C—, KC—, PD+ red; sections of apothecia C—, PD-; t.l.c.: argopsin. Observations: Micarea lignaria is characterized by its whitish to grey, convex to + globose areolae which are PD+ red (argopsin; but see var. endoleuca), black, markedly convex to + globose apothecia, green upper hymenium, dilute greenish or dilute olive-brownish hypothe- cium, “nd 3—7-septate, fusiform spores. When on lignum the thallus is often reduced to small scattered areolae and sometimes it is + entirely endoxylic. The apothecia are less variable in appearance, and pale (shade) forms are very rare. However, the apothecia are occasionally found (especially on rock in dry underhangs) to be stipitate with a ‘stipe’ (composed of vertically extended hypothecial and excipular tissues) up to 0-4 mm, or even 1 mm, tall (Fig. 3E). These forms have been ascribed the status of form, variety, species and even genus, viz.: Nylander’s new species and genus Stereocauliscum gomphillaceum. In the type material of this, some ‘stipes’ are extremely tall (up to 1 mm) and composed of vertically proliferating apothecia, with the apical apothecia being the youngest; the apothecia are immature (or arrested in their development) with few asci and spores. In my opinion these stalked forms of M. lignaria result from abnormal development in response to adverse environmental conditions and are, there- fore, not worthy of taxonomic recognition at any rank. M. lignaria is often confused with M. cinerea and M. peliocarpa, but these two species have usually more flattened and often paler apothecia, more richly branched paraphyses, and a hyaline hypothecium. In addition, they contain gyrophoric acid, resulting in the C+ orange-red (quickly fading), PD— reactions of their thalli and apothecia. M. ternaria (q.v.) is very similar to M. lignaria, but has more flattened apothecia with a more discernible excipulum and spores which are never more than 3-septate; furthermore, it lacks any lichen substances. Habitat and distribution: M. lignaria occurs on a wide range of substrata, but is most common in upland districts, growing over bryophytes or plant debris on siliceous rocks and walls, or on exposed peaty ground. Associated lichens found in such habitats in Britain include Arthrorhaphis citrinella, Baeomyces rufus, Cladonia coccifera, C. floerkeana, C. squamosa, C. subcervicornis, Coelocaulon aculeatum s. lat., Lecidea granulosa, L. icmalea, Lecidoma demis- sum, Lepraria neglecta, Micarea leprosula, M. peliocarpa, Ochrolechia frigida, Parmelia saxatilis, and Pseudephebe pubescens. It often grows over bryophyte mats that are heavily invaded by gelatinous algae, and in such situations in Scotland it has been found with the rare, or overlooked, Arctomia delicatula and Belonia incarnata. When growing directly on rock M. lignaria is mostly confined to sheltered, shaded situations and is sometimes present in the Micareetum sylvicolae in rock underhangs. It is usually present in abundance in the old lead and zinc mine workings of the Pennines and Scotland, where it grows over decaying bryophytes and LICHEN GENUS MICAREA IN EUROPE 145 if ty, 2 TU siside 1 Map9 Micarea lignaria var. lignaria @ 1950 onwards O Before 1950 plant debris, loose stones, pieces of timber and sack-cloth. Mr V. Giavarini has recently found it in Dorset, growing on waste heaps of slag clay, in the company of M. leprosula, Baeomyces roseus, Cladonia arbuscula, C. ciliata, C. furcata, C. portentosa, and Coelocaulon aculeatum s. lat. M. lignaria is very rare in the lowlands of south-east England, but has been found there growing directly on the rock of east or north facing sandstone walls and churchyard memorials, and also on natural sandstone outcrops of the Sussex Weald. In upland areas M. lignaria is frequently found on the exposed lignum of fallen trunks (especially of conifers) and old timberwork, with, for example, Cladonia spp., Hypogymnia physodes, Micarea peliocarpa, Mycoblastus sterilis, Ochrolechia turneri, Parmelia saxatilis, and Lecanora polytropa (worked timber). Occurrences of M. lignaria on the bark of healthy trees are rather rare, and in Britain are confined to the high rainfall areas of the north and west, where it has been collected on Alnus, Betula, Fraxinus, Ilex, Quercus, and old Sambucus. Reports of M. lignaria on mosses on limestone rocks usually result from the misidentification of Bacidia sabuletorum or Toninia lobulata. However, on a few occasions I have seen M. lignaria growing on thick bryophyte cushions or mats at high altitudes over limestone in the north Pennines, and over calcareous mica-schist in the Breadalbane Mountains; in such situations the pH and calcium content of the substratum is presumably kept low by the leaching effect of the’ high rainfall in those areas. The altitude range of M. lignaria i in Britain is from sea-level to about 1200 m (Ben Lawers), although it may well occur at higher altitudes in the Ben Nevis group and the Cairngorm Mountains. Higher altitudes are attained in the mountains of central Europe, from where it has 146 BRIAN JOHN COPPINS been collected at 2800 m in the Austrian Tirol, c. 2000 m in the Tatry mountains of Poland and Czechoslovakia, and at nearly 2500 m in the Transylvanian Alps of Romania. M. lignaria is common in northern and western Britain, but is very rare, with only a few scattered records, in lowland, southern England east of Devon. It is widely distributed throughout most of Europe, although in southern Europe it appears to be confined to the mountains: like most species of Micarea it seems to avoid the lowland Mediterranean regions. Records suggest that it is rare in arctic Fennoscandia, although I have seen material from north Finland and the Kola Peninsula. It has been found in the Azores at altitudes of about 900 m but I do not know of it from elsewhere in Macaronesia. From outside Europe I have seen material from the north coast of western Siberia, the Franconia Mountains (at c. 1450 m) of New Hampshire in the USA, and from Brazil (Serra Montiqueira, at c. 1900 m). Exsiccata: Anzi Lich. Lang. 148 (BM). Arnold Lich. Exs. 348A, B (BM ex K, L, M). Bohler Lich. Brit. 85 (E). Claudel & Harm. Lich. Gall. 43 (BM). Cumm. Dec. N. Am. Lich. ed. I: 302 (BM, C), ed. 2: 232 (M). Flotow Lich. Exs. 129A, E and 131 (UPS). Fries Lich. Suec. 29 (E,M, UPS); 98 (E). Harm. Guide 91 (UPS). Harm. Lich. Loth. 852 (M). Hav. Lich. Norv. 555 (C). Hertel Lecid. 34 (E, GZU). Johnson Lich. Herb. 453 (E). Kalb Lich. neotropici 22, 186 (hb Kalb). Krypt. Exs. Vindob. 658 (BM, BM ex K, C, M). Kutak Lich. Bohem. 417 (O). Larb. Lich. Caes. Sarg. 83 (BM). Larb. Lich. Herb. 272 (BM). Leighton Lich. Brit. 210 (BM, DBN, E, MANCH, NMW), 238 p.p. (BON, DBN, E, M, MANCH), 386 (BM, BON, DBN, E, FRS, M). Lojka Lich. Hung. 61 (BM, BM ex K, M). Malme Lich. Suec. 288 (C, M, S). Migula Crypt. Germ. 1 p.p. (BM, C, M, MANCH); 226 (BM, C, MANCH). Moug. & Nestl. Stirp. Crypt. 1430 (E). Mudd Lich. Brit. 156 (BM, E, M, MANCH), 157 (E, MANCH), 158 (BM, E, M, MANCH). Norrlin & Nyl. Herb. Lich. Fenn. 319A, B (BM, C, H). Oliv. Lich. Orne 344 (M, S). Pisat Lich. Slov. 156 (BM, M). Rabenh. Lich. Eur. 322, 603 (BM, M). Roum. Lich. Gall. 193 (BM), 232 (BM, M). Samp. Lich. Port. 147 (LD). Schaerer Lich. Helv. 196 p.p. (BM, BM ex K, E). Vézda Lich. Bohem. 133, 258 (LD, M). Vézda Lich. Sel. 516, 858, 1036, 1088 (BM). Zwackh. Lich. Exs. 121 (BM). 19b. Micarea lignaria var. endoleuca (Leighton) Coppins, comb. nov. (Figs 53-54; Map 10) Lecidea milliaria var. endoleuca Leighton, Lich. Fl. Brit., edn 3: 363 (1879). Type: Ireland, West Galway, on the Doughraugh, 1875, C. Larbalestier (BM ex K — lectotype!; BM — ? isolectotype!). Thallus identical to var. lignaria except that the areolae usually have dull yellowish (isabelline) tinge. Apothecia and pycnidia (containing microconidia) identical to var. lignaria;, pycnidia contain- ing macroconidia or mesoconidia not known. Chemistry: Thallus K— or Kf+ yellow, C+ yellow-orange (persistent), KC+ reddish orange (persistent), PD—; sections of apothecia C-; t.l.c.: xanthone(s), possibly arthothelin and thiophaninic acid. Observations (including habitat and distribution): My preliminary studies on M. lignaria found most specimens to have a C—, PD+ red thallus reaction, but a few otherwise + identical specimens from western Ireland, north Wales, and western Scotland gave C+ persistent orange, PD-— reactions. Subsequent t.1.c. analysis proved the normal form to have the then unidentified, P+ red compound found also in Phyllopsora rosei (Coppins & James, 1979), and the anomalous form to contain two xanthones (possibly arthothelin and thiophaninic acid). Xanthones often impart a yellowish tinge to lichen thalli (e.g. Lecidella elaeochroma, L. scabra, Pertusaria flavida, and P. hymenea) and this is the case for the xanthone containing race of M. lignaria; the colour difference is most evident when the two races occur together. Of all the types of names attributable to M. lignaria s. ampl., only that applicable to the ‘var. endoleuca Leighton’ was found to contain xanthones, and that name is here adopted at the same rank. Although the var. endoleuca is morphologically identical to the var. lignaria it has a sympatric but much more restricted geographical range. In the British Isles it is confined to the more oceanic districts with one rather anomalous occurrence in Surrey (Leith Hill). Furthermore, it appears to have a lower altitudinal range, with a collection at 600 m in Snowdonia representing its known upper limit in LICHEN GENUS MICAREA IN EUROPE 147 é ee ee ee eee i Bp Pry BIE: eae. ‘ ‘i A, rg c * y Se —— oa avd ie) p Oe ae EA eee ‘o. ZL eee Fie Oe Bane a | 0 0 1 4 5 6 q J Map 10 Micarea lignaria var. endoleuca @ 1950 onwards O Before 1950 the British Isles. Only four examples of var. endoleuca have been discovered from outside Britain: from Baden (Heidelberg) and Bavaria (Bayreuth) in Germany, Bern (Langnau) in Switzerland, and Trentino (Appiano) in northern Italy. It should be checked for amongst material from other regions, especially south-west Norway, Bretagne (France) and the Pyré- nées. I believe that the var. endoleuca should be retained at varietal rank. However, as the depsidone argopsin (1’-chloropannarin) of var. lignaria is probably biogenetically distinct from the xanthones of var. endoleuca, a case could be made for its recognition at species rank. A comparative example is Parmeliopsis ambigua versus P. hyperopta (see Hawksworth, 1976). 20. Micarea lithinella (Nyl.) Hed. (Figs 4A, 20A-B; Map 11) in Bih. K. svenska VetenskAkad. Hand. I11, 18 (3): 78, 97 (1892). — Lecidea lithinella Nyl. in Flora, Jena 63: 390 (1880). — Lecidea lithinella Nyl. in Flora, Jena 45: 464 (1862); nom. nudum (Art. 32). Type: Germany, Bayern, ‘Sandsteineblécke am Waldwege von Banz nach Altenbanz in Oberfranken’, ix 1860, F. Arnold 957 (M - lectotype!; H-NYL 19192 — isolectotype (fragment)!). Thallus effuse, saxicolous, often a thin indistinct crust between protruding grains of rock, but somes developing irregularly rounded, convex, more rarely subglobose, whitish areolae, c. 40-100 xm diam. Phycobiont micareoid, cells 4-6-5 wm diam. Apothecia numerous, immarginate, adnate, convex, apparently never tuberculate, pallid or 148 BRIAN JOHN COPPINS dull yellow-orange to reddish brown (never black), (0-1—)0-15—0-4 mm diam. Hymenium 35-50 pm tall, hyaline but often tinged dilute straw in upper part. Asci cylindrical-clavate, 35-50 x 8-13 um. Spores ovoid or fusiform-ellipsoid, simple, 6-5—9-5 x2-8—4 um. Paraphyses scanty, simple or irregularly forked above, 0-7-1 wm wide but sometimes gradually widening towards the apices and up to 1-5 wm wide, hyaline throughout. Hypothecium c. 50-70 wm tall, yellow-straw to dilute orange-brown, K—, HNO;~—, pigment confined to gel-matrix and hyphal walls hyaline. Excipulum absent (not detected even in sections of young apothecia). Pycnidia apparently rare, very inconspicuous, immersed, c. 40 wm diam, with hyaline walls. Conidia (microconidia) narrowly cylindrical, 4-5-7 x 0-8-1 wm. Chemistry: Thallus C—, K—, PD—; section of apothecia C—; t.].c.: not tested. Observations: M. lithinella exhibits little variation except that between, and sometimes within, collections there is an intergradation from pallid apothecia with a dull yellowish hypothecium to darker apothecia with an orange-brown hypothecium. This variation is corre- lated with differences in exposure to light. M. lithinella is easily confused with forms of M. bauschiana with pallid apothecia. Difficult specimens can be determined by a careful examina- tion of the phycobiont, which is micareoid in M. lithinella but non-micareoid (cells c. 7-13 wm diam) in M. bauschiana. M. lutulata has darker apothecia which are commonly tuberculate, a darker (opaque) hypothecium, an often greenish hymenium, and non-micareoid phycobiont. M. muhrii is normally lignicolous but has once been found on rocks; it has adnate apothecia like M. lithinella but they are usually darker and have a taller, dark reddish brown hypothecium, and usually a greenish upper hymenium. M. myriocarpa shares a similarly coloured hypothecium with M. lithinella, but has smaller, globose to tuberculate apothecia and smaller, 1-septate spores. In their keys to Micarea Vézda & Wirth (1976) and Poelt & Vézda (1977) state that M. lithinella has a K+ violet hymenium and often 1-septate spores; this is clearly an error, probably due to confusion with saxicolous forms of M. denigrata or M. prasina. Habitat and distribution: M. lithinella occurs on acidic rocks, especially hard sandstone. Associated species encountered on the specimens examined include Baeomyces rufus (parasi- tized by Thelocarpon lichenicola on the lectotype of Lecidea lithinella), Huilia crustulata Rhizocarpon obscuratum, Scoliciosporum umbrinum, Trapelia coarctata, and T. aff. obtegens. This list of associates, plus information on some of the packets, suggest that M. lithinella has mostly been found in humid situations on outcrops and boulders by woodland roads. It was collected by Arnold, Lahm and von Zwackh from several areas of Germany: Bayern, Nord- rhein-Westfalen and Baden-Wirttemberg, respectively. In Sweden it was collected in Uppland by Hedlund who (1892: 97) also cited a collection by Blomberg from S6dermanland. One British specimen has recently been found: South-east York (V.C. 61), Wharram Quarry Nature Reserve, 44/85. 65, 1969, Coppins (E). At this locality M. lithinella was growing with Scolicio- sporum umbrinum on the underside of a large flint; the upper, more exposed, part of the flint was colonized by Lecidea erratica and Rhizocarpon obscuratum. Exsiccata: Arnold Lich. Exs. 836 (BM ex K, H-NYL 19190, M). Lojka Lich. Univ. 233 (M). Malme Lich. Suec. 125 (M, S). Zwackh Lich. Exs. 590 (H-NYL p.m. 5404, M). 21. Micarea lutulata (Nyl.) Coppins (Figs 20C, 44B; Map 11) in D. Hawksw., P. James, & Coppins in Lichenologist 12: 107 (1980). — Lecidea lutulata Ny)l. in Flora, Jena 56: 297 (1873). Type: Jersey, Rozel meadow, bases of rocks, 1873, C. Larbalestier (H-NYL 10696 — lectotype!; BM — isolectotype!; M — probable isolectotype!). See note below. Lecidea laxula Ny. in Flora, Jena 58: 11 (1875). Type: Finland, Tavastia australis, Luhanka, Hietala, 1874, E. A. Lang [Vainio] 303 (H-NYL 20689 — lectotype!; H — isolectotype!). Lecidea poliodes Nyl. in Flora, Jena 58: 10 (1875). — Micarea poliodes (Ny1.) Vézda in Vézda & V. Wirth in Folia geobot. phytotax., Praha 11: 99 (1976). Type: Finland, Tavastia australis, Evo, [on schistose rock with M. sylvicola], J. P. Norrlin (H-NYL 20683 — holotype!). LICHEN GENUS MICAREA IN EUROPE 149 Lecidea antrophila Larb. ex Leighton in Trans. Linn. Soc. London (Bot.) I, 1: 242 (1876). Type: Ireland, West Galway, Mwellan near Kylemore, in the interior of a cave, 1877, C. Larbalestier (BM ex K — lectotype!; isolectotypes: BM!, BM ex K!). Lecidea paucula Nyl. in Flora, Jena 59: 573 (1876). Type: Ireland, West Galway, ‘Montagnes de Maam’ [Maumturk Mountains], iii 1876, C. Larbalestier (H-NYL 20090 — holotype!). Micarea umbrosa Vézda & V. Wirth in Folia geobot. phytotax., Praha 11: 93 (1976). Type: Germany, Baden, Siidschwarzwald, between Neuhausle and Altglashiitte near St. Margen, 830 m, 2 v 1969, V. Wirth (hb. Wirth 1609 — holotype!). ’ Lecidea granvina Vainio in Havaas in Bergens Mus. Arb. 1909 (1): 31 (1909); nom. nudum (Art. 32). Spec. orig.: Norway, Hordaland, Hardanger, Granvin, 10 xi 1900, J. J. Havaas (BG!). Lecidea demarginata auct. p.p., non Nyl.; see ‘Excluded Taxa’. Note: typification of Lecidea lutulata. Larbalestier’s gatherings in BM and H, that were collected from Rozel meadow in 1873 and subsequently labelled ‘Lecidea lutulata’, consist of M. bauschiana, M. lutulata and M. peliocarpa. The specimen H-NYL 10696 contains M. lutulata only and agrees with the original diagnosis, e.g. ‘hypothecium fusconigricans crassum’, and is therefore selected as lectotype. Thallus effuse, thin (up to 40 wm thick), + smooth or finely rimose, usually becoming thicker (up to 600 wm thick) and scurfy-granular (but never forming discrete areolae or goniocysts), pale greenish grey or grey-green, sometimes straw-coloured in part, sometimes oxydated (on ferruginous rocks); in section ecorticate and without an amorphous hyaline covering layer. Phycobiont not micareoid; cells thin-walled, + globose, c. 5-12 um diam, or ellipsoid and up to 15X8 wm. Apothecia numerous, convex-hemispherical and immarginate from the start, later becoming + globose or tuberculate, grey-brown, dark brown or blackish, always turning blackish when moistened, 0-2—0-4 mm diam, or up to 0-8 mm diam when tuberculate. Hymenium 30-40 wm tall; upper part varying from hyaline or pale fuscous-brown through olivaceous to dark aeruginose, pigment often in patches corresponding to clustered apices of stout paraphyses, K—,HNO3+ red; lower part hyaline or dilutely coloured. Asci cylindrical-clavate, c. 30-40 x7- 10 wm. Spores ellipsoid, ovoid, or sometimes dacryoid, 6—-8(—9) x2-3(-3-4) wm. Paraphyses rather scanty, of two types: p.p. evenly distributed, irregularly flexuose, simple, or forked or with short branches (especially in the upper part), sometimes anastomosing, 0-8-1(-1-5) wm wide, apices sometimes irregularly swollen to c. 2 ~m wide; p.p. fasciculate, mostly simple, stout, c. 1-5-2 wm wide, with swollen (to 3 wm), coherent, pigmented apices. Hypothecium 120-360 um tall, dark and opaque, fuscous- or reddish brown, K— or + reddish intensifying (but never with purple or greenish tinges), HNO3—; hyphae interwoven, but becoming vertically orientated towards the hymenium, c. 1-5-2 wm wide but coated with dense brown pigment and appearing 3-4 wm wide; ascogenous hyphae with swollen cells c. 2-4 wm wide. Excipulum not seen even in sections of young apothecia; the hymenium soon becomes reflexed so as to form an excipulum-like zone below the hypothecium. Pycnidia often present but inconspicuous, im- mersed, blackish, c. 80-200 um diam, ostiole often gaping; with a single + globose locule but walls often convoluted with up to 5 locules seen in a vertical section; outer walls reddish brown, K-; inner walls of secondary locules hyaline; conidiogenous cells irregularly cylindrical, often with 1-2 percurrent proliferations, 7-10X1—1-5 um, base sometimes swollen to 2-7 wm wide; conidia (mesoconidia) cylindrical, sometimes faintly biguttulate but not constricted in the middle, (3-8—)4-5S(-5-5) x0-9-1-4 wm. Chemistry: All parts K—, C—, PD-—; t.1.c.: no substances detected. Observations: The apothecia of M. Jutulata have a similar internal pigmentation to those of M. botryoides (q.v.) and M. muhrii (q.v.), but the species is most often confused with M. sylvicola. The latter shares the same habitat as M. /utulata, but can be distinguished by its hypothecium which always has a distinctly green, or rarely purplish, tinge in water mounts or KOH; in HNO; the pigment(s) turn purple-red. M. sylvicola further differs in its larger spores, taller hymenium, slightly longer and broader conidia, and presence of greenish pigment in its pycnidial walls. M. myriocarpa occurs in similar habitats to M. /utulata and has a reddish brown, but never opaque (in thin section) hypothecium. Furthermore, M. myriocarpa has smaller apothecia, narrower, often 1-septate spores, and sessile pycnidia with much shorter conidia. 150 BRIAN JOHN COPPINS By : ? e.)mhUe ae al ike oe mee ) iene ieee e ie r 0 1°) 1 2 3 rad 4 0 7 ews zB — vol afb a PY . a.” oi wy id q ¥ eS, dees i g he = g e | a ae ye iv So ae a 17 se 0) Gs 8 ‘9. ? © aw — oe Pia ) : — 0 1 : 5 6 L ) Map 11 Micarea lithinella © 1950 onwards + Micarea lutulata @ 1950 onwards O Before 1950 Habitat and distribution: M. lutulata is found on rocks and sometimes exposed roots in dry rocky underhangs, and is a characteristic species of the Micareetum sylvicolae. It is apparently widespread in northern and western Britain; and although not yet reported from the south-west peninsula of England or the north-west of Scotland, it almost certainly occurs in those areas. It appears to be little known outside of the British Isles, but I have seen material from Norway (Oppland and Hordaland), Sweden (Varmland), southern Finland, Germany (Baden- Wirttemberg), and Czechoslovakia (Bohemia: Krkono8e). Exsiccata: Arnold Lich. Exs. 409B p.p. (WRSL). Havaas Lich. Exs. Norv. 571 (BG). Larbal. Lich. Herb. 223 (BM, BM ex K). Rasanen Lich. Fenn. 672 p. p. (LD-mixed with M. tuberculata). 22. ,Micarea melaena (Nyl.) Hedl. (Figs 21A, 46A; Map 12) in Bih. K. svenska VetenskAkad. Handl. III, 18 (3): 82, 96 (1892). — Lecidea melaena Nyl. in Bot. Notiser 1853: 182 (1853). — Lecidea vernalis var. melaena (Nyl.) Nyl. in Mém. Soc. Imp. Sci. nat. Cherbourg. 3: 182 (1855). — Bacidia melaena (Nyl.) Zahlbr. in Annis mycol. 7: 474 (1909). Type: Sweden, on lignum, E. M. Fries, Lich. Suec. Exs. 212B (UPS — lectotype!; isolectotypes: C!, H-NYL p.m. 4778 [fragment]!, M!). Lecidea milliaria var. turfosa Fr., Nov. Sched. Crit. 8: 7 (1826), non Biatora turfosa Massal. Type: Sweden, [? SmAland], on peaty turf, E. M. Fries, Lich. Suec. Exs. 212A (UPS —lectotype!; isolectotypes: C!, M!). LICHEN GENUS MICAREA IN EUROPE byl Biatora stizenbergeri Hepp, Flecht. Eur. no. 504 (1860). Type: Switzerland, Rifferschweil, on dry plant debris amongst roots in peat moor, Hegetschweiler (BM -— lectotype!; isotypes distributed in Hepp. Flecht. Eur. 504: BM!, E!, WRSL!). Lecidea ilyophora Stirton in Scott. Nat. 5: 220 (1879). Type: Scotland, Perthshire, Black Wood of Rannoch, on lignum, ix 1879, J. Stirton BM — lectotype! [t.l.c.: no substances detected]). Lecidea melaena f. catillarioides Vainio in Medd. Soc. Fauna Fl. fenn. 10: 12 (1883). — Micarea melaena f. catillarioides (Vainio) Hedl. in Bih. K. svenska VetenskAkad. Handl. II, 18 (3): 83, 96 (1892). Type: Finland, Karelia borealis, Nurmes, Riihivaara, on bryophytes on rock, 1875, Vainio (TUR-VAINIO 21478 — holotype!). Lecidea melaena f. endocyanea Vainio in Medd. Soc. Fauna Fl. fenn. 10: 12 (1883). — Micarea melaena f. endocyanea (Vainio) Hedl. in Bih. K. svenska VetenskAkad. Handl. III 18 (3): 83, 96 (1892). Type: USSR, Karelia keretina, Kivakka, on mosses on rocks, 1877, Vainio (TUR-VAINIO 21476 — holo- type!). Bilimbia melaena f. aeruginosa Vainio in Acta Soc. Fauna Fl. fenn. 53 (1): 254 (1922). Type: Finland, Tavastia australis, Hollola, on lignum, 1875, Vainio (TUR-VAINIO 21461 — lectotype!). Bilimbia melaena f. epiphaeotera Vainio in Acta Soc. Fauna Fl. Fenn. 53 (1): 255 (1922). Type: Finland, Karelia borealis, Nurmes, Louhivaara, ‘Kallion syrjalla’, on bryophytes and old Cladonia squamules, 1875, Vainio (TUR-VAINIO 21472 - lectotype!). ? Bilimbia melaena var. alnicola Savicz in Izv. imp. S. Peterb. bot. Sada 14 (1, Suppl.): 53 (1914). Type: USSR (?LE); request for loan received no reply. Bilimbia milliaria y. [var.] saprophila Korber, Parerg. lich.: 171 (1860); nom. superfl. (Art. 63). Thallus effuse, superficial, of small, scattered to coherent or clustered, granular areolae (20—)25—80(—100) wm diam, sometimes forming an uneven crust up to c. 150 wm thick. Areolae very variable in colour, pale buff or pale dull green to dark grey-green; in section without an amorphous covering layer, outermost hyphae often with dark green (K—, HNO3+ red) walls. Thallus often scurfy granular and blackish (especially in damp situations) owing to its disruption by invading dematiaceous fungi and foreign algae; in dry situations (e.g. undersides of fallen trunks) it may have a whitish or pale yellowish, farinose appearance due to a thick covering of an epiphytic alga, each cell of which is coated by hyaline, crystalline incrustations. Phycobiont micareoid, cells c. 4-7 wm diam. Apothecia numerous, immarginate, convex-hemispherical to + globose, sometimes tubercu- late, black, matt or slightly glossy, 0-12-0-4 mm diam, or to 0-5 mm diam when tuberculate. Hymenium 30-40(-50) mm tall, dull to bright aeruginose or grey-blue, K— or K+ green intensifying; alternatively lower part (or rarely the entire hymenium) dilute purple-brown, K+ greenish or K+ purple intensifying; pigment(s) often more dense around the apices of the paraphyses, thus appearing as a dark ‘epithecium’ (c. 2-10 ym tall) that contrasts with the more dilute colouration of the rest of the hymenium. Asci clavate, c. 30-40-12-15 um. Spores ovoid-oblong or oblong, with rounded apices, straight, (1—)3(—5)-septate, 12-21 x 4—S(-5:5) wm. Paraphyses numerous, branched and anastomosing, sometimes a few unbranched, thin, 0-8-1 um wide; apices not swollen, or sometimes slightly incrassate and up to 1-5 xm wide, hyaline but often individually surrounded by a pigmented hood of dense gel (easily detached in mounts in K, by tapping the cover slip). Occasionally intermixed with ordinary paraphyses are a few which are simple, stout (c. 1-5—2 ~m wide) and coated with pigment throughout their length so as to appear 3-4 wide. Hypothecium c. 80-160 mm tall, dark and + black in thick sections, pigmentation variable (see ‘observations’ below), usually purple-brown and K— or K+ purple intensifying, or, outer (more rarely whole) part K+ olivaceous with central part (‘core’) remaining + purplish brown; all parts HNO3+ purple-red; hyphae c. 0-7-1-5 wm wide but their walls coated with dense pigment so that they appear to be c. 3-4 wm wide, interwoven or becoming vertically orientated towards the hymenium, sometimes a few extending into the hymenium as stout pigmented paraphyses; ascogenous hyphae similarly pigment, short-celled, c. 2-5 wm wide. Excipulum indistinct, sometimes evident in sections of young apothecia, green or purplish (usually darker than hymenium, but paler than hypothecium); hyphae radiating, branched and anastomosing, c. 0-7—1 wm wide. Pycnidia rare, black, of two types: (a) c. 100-140 um diam, sessile; walls purple-brown or dull Olive, K+ green; conidia (macroconidia) curved or hamate, 0-7-septate, 18-33 x 1-1-5 um; (b) 152 BRIAN JOHN COPPINS c. 40-60 wm diam, semi-immersed to sessile, walls green, K—; conidia (microconidia) fusiform- cylindrical, straight, 4-8—7 x0-8-1 um. Chemistry: Thallus K—, C+ red or C—, PD; sections of apothecia C—; t.l.c.: gyrophoric acid (healthy thalli), or no substances or gyrophoric acid in trace amounts (poorly developed or disrupted thalli). Observations: Micarea melaena is one of the commonest species of the genus, especially in northern Europe, and it displays a considerable amount of variation in thallus appearance and internal apothecial pigmentation. However, my examinations of over 200 specimens have revealed many intermediates between the known extremes. When well-developed, the thallus is composed of cohering, small, granular areolae and is pale buff to dull green in colour. In exposed situations the thallus often becomes a dark green-black due to the more intense pigmentation of the outermost hyphae of the granules. The appearance of the thallus can also be affected by the presence of other organisms (see description and p. 00). When on lignum, in dry exposed situations the thallus may become much reduced to a fine scattering of tiny (less than 30 pum diam), blackish granules amongst the wood fibres. Such forms proved to be very common, during my explorations in Sweden. M. melaena also exhibits much variation with regard to the colouration of its apothecial tissues (especially the hypothecium). This is due to the presence of three (probably biosynthetically related) pigments, varying in their relative amounts and distribution. These pigments, all of which react HNO3+ purple-red, can be roughly identified as pigment A: green, K+ green intensifying; pigment B: purple, K+ green; pigment C: purple, K+ purple intensifying. In the epithecium and hymenium pigment A is often found alone, but sometimes it is replaced in the lower hymenium (or in extreme cases throughout the entire hymenium and epithecium) by pigment C. Pigment C is usually dominant in the hypothecium, but is often replaced in the upper and outer parts (or more rarely + throughout) by pigment B. If thin sections of apothecia in which pigment B is dominant are mounted in K the lower central (‘core’) of the hypothecium retains a purplish tinge (the rest having turned green), indicating the presence of a small amount of pigment C. The same type of variation, presumably involving identical (or very similar) pigments, is found in several other species, e.g. M. crassipes and M. sylvicola. The pycnidia of M. melaena are very inconspicuous and apparently sparingly produced. However, a close scrutiny of about 100 specimens revealed ten collections with the microconi- dial state and only two (e.g. Coppins 6041) with the macroconidial state. A mesoconidial state was not found but may well occur. Habitat and distribution: M. melaena is widely distributed and especially common in north- west Europe. Although, for a Micarea, it is a conspicuous species, it is often overlooked as a black or charred stain on the tops of stumps and peaty turf! It is a common inhabitant of stumps and fallen decorticate trunks in upland woodlands, or in lowland woodlands on very acid, nutrient-poor soils, and can exist in shaded to extremely exposed, sun-baked situations. In upland woods it can be found on the corticate trunks of healthy trees (especially Betula, Quercus, and Pinus). In areas heavily polluted by SO, and its derivatives (e.g. West Yorkshire conurbation) it has been found on the acidified bark of Acer and Ulmus. Occurrences on decaying timber work (e.g. fence posts, gate rails and shingles) are not uncommon, especially in upland districts. Lichens associated with M. melaena on lignum and bark in the British Isles include Bryoria capillaris, B. fuscescens, Cladonia spp., Hypocenomyce scalaris, Hypogymnia pnysodes, Lecidea granulosa agg., L. icmalea, Micarea denigrata, M. prasina, Mycoblastus sterilis, Ochrolechia turneri, Parmelia saxatilis, Parmeliopsis spp., Platismatia glauca, Trapelia corticola, and Usnea spp. M. melaena is often terricolous, growing on oligotrophic, peaty soils, in montane regions or lowland heaths. In the English lowlands it is recorded from heaths in the Sussex Weald and east Norfolk. It is common on + bare peat in the Pennines and Scottish highlands, but in the latter it is usually replaced near the higher summits (over c. 1000 m) by M. turfosa. Associated lichens on peaty turf in Britain include ‘Botrydina’ (i.e. Omphalina spp.), Cladonia coccifera, C. crispata, LICHEN GENUS MICAREA IN EUROPE 153 ; ) 1 2 = 4 ‘ rg | wal ab t f° A ¥ mt Col 2 : 4 é i ated Wale a 8 i “oi a. 8 Soa ws os Sto fos APR : nae ae ° oe 3 2 awl Se PH | 6 pes ase fe ‘ 6 ~ | ee be pane an a a ea ee “el Y Ve): ° : ( if SS : eh Amey Sal oe ale ~ = Peas “at J et eect =P Lt jaan ead py Map 12 Micarea melaena @ 1950 onwards O Before 1950 C. floerkeana, C. subcervicornis, Coelocaulon aculeatum s. lat., Lecidea icmalea, Micarea leprosula, and Ochrolechia androgyna. M. melaena is often found on rocks, growing over peaty debris or decaying mats of bryophytes and Cladonia squamules, but occurrences where it is growing directly on rock are rare. To my knowledge, all such occurrences in Britain are confined to woodland situations in rather heavily polluted areas approximating to zones 3-5 on the scale of Hawksworth & Rose (1970); these areas include Leicestershire, the West Yorkshire conurbation, and Fife. M. melaena is widely distributed in mainland Britain, but is particularly common in the upland areas of the north and west. There is a dearth of records from Ireland, where it is unlikely to be rare in suitable terrain. M. melaena is known from most parts of the European mainland and is particularly common in the boreal regions of Fennoscandia and upper Bavaria and adjoining alpine districts, but this apparent tendency may, at least partially, be a reflection of the avidity of the collectors active in those areas, and the herbaria which I have studied! It has been found in the Azores on Cryptomeria and sulphur-rich soils near fumaroles, but it is so far unrecorded from the Canary Islands. From outside Europe I have seen several collections from eastern Canada and north-eastern U.S.A. Exsiccata: Anzi Lich. Exs. Ital. 259A (BM, M). Anzi Lich. Sondr. 170B (UPS). Arnold Lich. Exs. 332A-C (M). Arnold Lich. Mon. 49 (M), 248 (C, M), 249 (C, MANCH), 407 (C, M, MANCH). Britz. Lich. Exs. 946 (M). Fellman Lich. Arct. 159 (H). Flotow Lich. Exs. 129C (UPS). Fries Lich. Suec. 212A, B (C, H-NYL p.m. 4778, M, UPS). Hepp Fl. Eur. 504 (BM, E, WRSL). Johnson Lich. Herb. 376 (BM). Krypt. Exs. Vindob. 362 (BM, M). Leighton Lich. Brit. 120 (BM). Malme Lich. Suec. 27 (C, M, S). Moug. 154 BRIAN JOHN COPPINS & Nestl. Stirp. Crypt. 1329 p.p. (E). Mudd Lich. Brit. 159 (M, MANCH). Norrlin & Nyl. Herb. Lich. Fenn. 180 (BM, C, M). Oliv. Lich. Orne 237 (M). Rasanen Lich. Fenn. 963 (M). Rasanen Lichenoth. Fenn. 344 (BM). Roum. Lich. Gall. 231 (M). Schaerer Lich. Helv. 196 p.p. (E). Vézda Lich. Sel. 14 (BM, M). Zwackh Lich. Exs. 657, 675 (M). 23. Micarea melaenida (Nyl.) Coppins, comb. nov. (Fig. 22) Lecidea melaenida Ny). in Flora, Jena 48: 146 (1865). — Catillaria melaenida (Nyl.) Arnold in Flora, Jena 53: 475 (1870). Type: France, [? Eure], Perriéres, Lenormand (H-NYL 18843 — holotype!). Lecidea relicta Nyl. in Flora, Jena 48: 354 (1865) [nec Stirton (1876)]. Type: France, Calvados, Falaise, L.-A. De Brébisson (H-NYL 21099 — holotype!). Catillaria schumannii [‘Schumanni’| Stein in Cohn, Krypt. -Fl. Schles. 2 (2): 232 (1879). — Lecidea schumanni Korber, Syst. Lich. Germ.: 255 (1855); nom. inval. (Art. 34). Type: East Germany, Reichenbach, near Ernsdorf, Schumann (WRSL - lectotype!; WRSL ~ isolectotype!). Catillaria zsakii Szat. in Magy. bot. Lap. 24: 108 (1926). — Toninia zsakii (Szat.) Lettau in Feddes Reprium Spec. nov. veg. 57: 9 (1955). Type: holotype not seen (? in BP); topotypes: Hungary, ‘In argillosis natronatis prope oppidum Karcag, Comit. Jasz-Nagykun-Szolnok. Allit. ca. 80 m. s.m. Mens. Szept. 1926’, Z. Zsak & G. Timko, Fl. Hung. Exs. 714 (E! [t.l.c.: no substances], BM!, BM ex K!), and, from same locality but at ‘100 m. s.m.’ and no date, Krypt. Exs. Vindob. 3154 (BM!, BM ex K!). Catillaria schumanni var. meridionalis Roux & Vézda in Vézda, Sched. Lich. Sel. Exs. 537 (1967). Type: France, Gard, Pujaut near Avignon, alt. 100 m, ‘supra solum argillaceo-sabulosum, ad marginem Querceti ilicis’, 6 xi 1966, C. Roux, Vézda Lich. Sel. Exs. 537 (BM — isotype!). Thallus effuse, terricolous (on argillaceous soils), forming a thin + smooth, dull greenish white crust, c. 40-100 wm thick, or forming confluent, convex areolae 0-1—0-6 mm diam; crust sometimes becoming cracked so as to appear + ‘squamulose’. Thallus in section without a distinct hyaline, amorphous, covering layer, but with a hyaline ‘cortex’ 12-20 um tall, composed of compacted, evacuated hyphae (cytoplasm presumed lost owing to lack of staining by LCB or ammoniacal erythrosin); thallus hyphae 1-5—2-5(-3) wm wide; numerous algae-free hyphae penetrating the soil substratum to a depth of c. 1 mm (? or more). Phycobiont micareoid, cells 4-7 um diam. Apothecia numerous, scattered or 2—-4-confluent, immarginate, convex to hemispherical, black, matt, 0-2-0-6 mm diam; old, contorted apothecia sometimes reaching 1-2 mm diam; occasionally a fine weft of radiating, white (hyaline) hyphae is seen around the edge of an apothecium. Hymenium 35-40 um tall; uppermost c. 1-5 wm (epithecium) with dense, amorphous, purple-brown pigment, K—, HNO3+ purple-red; remaining (lower) parts dilute dull purple, K—, HNO3+ red. Asci clavate, 35-38 x 9-12 xm. Spores ellipsoid, oblong-ellipsoid or ovoid-oblong, straight or slightly curved, 0—1-septate, (7—)9-5-15 x3-4(-4-5) um. Paraphyses numerous, mostly simple below, but often branched above, sometimes anastomosing, 1-1-7 wm wide; apices (upper S—15 wm) often purple-brown pigmented and to 2:5 wm wide. Hypothecium 60-120(—200) wm tall, mottled dark purplish brown, K—, or K+ purple intensifying in part, HNO; + purple-red; hyphae interwoven but + vertically orientated in uppermost part, c. 1-1-8 pum wide, each with a densely pigmented gel sheath thereby giving a total diameter of c. 3 um; ascogenous hyphae c. 2-5-5 xm wide. Excipulum soon reflexed, but distinct in sections of young apothecia, purple-brown (usually darker than hymenium); hyphae radiating, branched and anastomosing, c. 1-3-2 wm wide, embedded in densely pigmented gel-matrix. Pycnidia often present, immersed, black, c. 50-100 xm diam; walls purple-brown, K— or Kf+ purple intensifying; conidia (? microconidia), + cylindrical, 4-8-6(—7) x1 wm. Chemistry: Thallus K—, C—, KC—, PD—-; t.l.c.: no substances. Observations: Micarea melaenida is characterized by its whitish, terricolous thallus, immar- ginate, black apothecia (with purplish pigmentation in section), numerous and rather stout paraphyses, and 0-1-septate spores. It is apparently related to M. assimilata (q.v.) and its allies. When compared with M. melaenida, forms of M. melaena with immature spores can be distinguished by their finely granular areolae, narrow and less numerous paraphyses, and usual LICHEN GENUS MICAREA IN EUROPE 155 presence of green pigments in apothecial sections (in water or K mounts). M. turfosa is similar in some ways to M. melaenida, but has a dark coloured thallus, an olivaceous hymenium, reddish brown (never purplish) hypothecium, and larger spores which are often 1—3-septate. Habitat and distribution: M. melaenida grows on, and in direct contact with, fine-grained sandy or argillaceous, mineral soils, and is not known to occur on decaying bryophytes or plant debris, etc. Unfortunately I have never seen M. melaenida in the field, and there is little ecological information to be gleaned from the literature or herbarium labels. Furthermore, the specimens examined have included few associated species, and only Cladonia sp. (squamules) and Catapyrenium lachneum have been noted. In its Hungarian locality it was said to have occurred on soil rich in precipitated salts (‘argillosis natronalis’) although no effervescence was apparent when I tested the soil with 50% HNO3. It is probable that M. melaenida is restricted to niches where the salts have been permanently leached out. A sample of soil from Fl. Hung. Exs. 714 was sent for analysis to the Macaulay Institute for Soil Research (Aberdeen) and Dr B. W. Bache of the Institute reports ‘It bears no relationship at all to a sodic soil, and is in fact fairly acid, with pH 5-2. The conductivity is fairly low (0-8 mS in 1:2 extract), so it does not contain any great concentration of soluble salts, nor does it contain any carbonate.’ The data on the herbarium labels, although limited, do indicate that it occurs at low altitudes, for example the collection from Loire Atlantiqué was made from soil on coastal cliffs; that from Gard was made at 100 m; and those from Hungary were made at 80-100 m. M. melaenida is a rare species with a rather southern distribution, and I have seen material from several localities in France (e.g. Loire Atlantiqué, Vienne, and Gard), and single localities in East Germany (Dresden) and Hungary. It is not known from Scandinavia, and its status as a British plant is doubtful. Its first British report seems to be its appearance in the checklist of James (1965a), but I have been unable to ascertain the reason for its inclusion. However, its presence in north-west France suggests that it could occur in Britain, especially along the south coast of England or in the Channel Islands. Material from South Africa and distributed as Almborn Lich. Africani no. 84 is not M. melaenida but a mixture of two, undescribed species, both of which are probably referable to Micarea. Exsiccata: Fl. Hung. Exs. 714 (BM, BM ex K, E). Krypt. Exs. Vindob. 3154 (BM, BM ex K). Vézda Lich. Sel. 537 (BM). 24. Micarea melaeniza Hed. (Figs 21B, 38B) in Bih. K. svenska VetenskAkad. Hand. Il, 18 (3): 83, 96 (1892). — Lecidea melaeniza [‘melaniza’| (Hedl.) Zahlbr., Cat. lich. univ. 3: 798 (1925). Type: Sweden, Halsingland, lignum of old pine trunk, viii 1891, J. T. Hedlund (S — holotype!). Thallus effuse, lignicolous (sometimes extending on to bryophyte or lichen thalli), endoxylic, or epixylic with whitish convex areolae; areolae c. 100-400 wm diam and up to 45 wm thick, without an amorphous covering layer. Phycobiont micareoid, cells 4-7 wm diam. Apothecia numerous, few, or absent, immarginate, at first convex-hemispherical or subglo- bose, often becoming tuberculate, black, matt, 0-2-0-3 mm diam, or to 0-5 mm diam when tuberculate. Hymenium 28-42 um tall, upper and lower parts aeruginose-green, middle part + hyaline with aeruginose vertical streaks; K— or dulling, HNO3+ red. Asci clavate, 26-35 x 10-12 um. Spores ovoid or dacryoid, simple, 5—9x2-5-3-8 wm. Paraphyses rather numerous, of two types; p.p. evenly distributed, flexuose, branched, thin, c. 0-7-1 wm wide, hyaline, but apices sometimes pigmented and then to 2-5 wm wide; p.p. scattered or in small fascicles, stout, c. 2 wm wide, each coated + throughout their length by greenish pigment. Hypothecium c. 120-200 wm tall, dark brown, sometimes with a faint reddish (never purplish) tinge, K— or dulling, HNO3-— or red tinge slightly intensifying; hyphae coated with dark brown pigment and c. 2-3 wm wide, interwoven but becoming vertically orientated towards the hymenium and sometimes con- 156 BRIAN JOHN COPPINS tinuing into it as stout pigmented paraphyses; ascogenous hyphae similarly pigmented, but with short swollen cells to 5 wm wide. Excipulum not evident in available material. Pycnidia numerous, black, sessile and c. 40-60 wm diam, or more usually borne on black stalks (pycnidiophores) and 80-300 x 40-70 um (including stalk); stalks simple, or sometimes branched and bearing up to four pycnidia; upper part (pycnidial wall) dark olive but lower part (stalk tissue) reddish brown, both parts K—, Conidia (mesoconidia) short cylindrical, 2-3— 3-6X1-1:3 um Chemistry: Thallus PD—; sections of apothecia and thallus C—; material insufficient for analysis by t.l.c. Observations: With its black, subglobose to tuberculate apothecia, black, stalked pycnidia, inconspicuous thallus, and occurrence on lignum, M. melaeniza is indistinguishable in external appearance from M. misella and M. nigella. Microscopically, M. misella can be distinguished by the olivaceous (K+ violet) pigment in its hymenium and pycnidia, and its + hyaline hypothe- cium; and M. nigra can be distinguished by the purple-brown (K+ green, HNO3+ purple-red) pigment in its hymenium, hypothecium and pycnidia. With respect to apothecial and pycnidial pigmentation and structure, M. melaeniza is almost identical to M. botryoides. However, M. botryoides is rarely lignicolous, and has larger, often septate spores, and longer mesoconidia. M. munhrii has similar apothecial pigments to those of M. melaeniza, but is readily distinguished by its larger, adnate apothecia (which often have a pale marginal rim), larger spores, well developed excipulum, and lack of stalked pycnidia. Habitat and distribution: M. melaeniza is a rare species, known only from three collections made in Sweden (Angermanland, Halsingland, and Smaland) where it occurred on the lignum of conifer trunks. Associated species identified on the examined collections include Calicium salicinum, Cetraria pinastri, Chaenothecopsis lignicola, Cladonia spp. (squamules), Dimerella diluta, Hypogymnia physodes, Lecanora symmicta agg., Micarea anterior, M. misella, M. prasina, Usnea sp., and Lophozia sp. 25. Micarea melanobola (Nyl.) Coppins, comb. nov. (Figs 21C, 46B-C) Lecidea melanobola Ny). in Flora, Jena 50: 371 (1867). — Catillaria melanobola (Nyl.) Vainio in Acta Soc. Fauna FI. fenn. 57 (2): 465 (1934). — Lecidea erysiboides * L. melanobola (Nyl.) Nyl. in Hue in Revue Bot. Courrensan 5: 103 (1886). — Catillaria 8. byssacea f. melanobola (Nyl.) Blomb. & Forss., Enum. PI. Scand. : 91 (1880). — Micarea prasina f. melanobola Hedl. in Bih. K. svenska VetenskAkad. Hand. II, 18 (3): 87 (1892). Type: Finland, Tavastia australis, Kuhmois [Kuhomoinen], on bark of Picea abies, 1866, J. P. Norrlin (H-NYL 21614 — lectotype!; isolectotypes: H-NYL p.m. 4504!, H!). Thallus effuse, thin and indistinct, of scattered to + coherent, minute olivaceous granules (goniocysts), c. 12-25 wm diam, which appear + gelatinous when moist. Outermost hyphae of goniocysts with greenish, K+ violet walls. Phycobiont micareoid, cells 4-7 wm diam. Apothecia numerous, convex-hemispherical, immarginate, dark grey to black, 0-1-0-24 mm diam. Hymenium 30-35 um tall, hyaline, but with a clearly delimited, dark green epithecium, K+ deep violet, C+ violet, HNO3+ red; pigment closely bound to (? or also in) the apical walls of the paraphyses, and also present in the gel-matrix and apical walls of the asci. Asci clavate, 30-35 9-11 wm, upper wall(s) sometimes tinged with greenish, K+ violet pigment. Spores ellipsoid, ovoid, or oblong-ovoid, with obtuse apices, 0-1-septate, 7-9-7 x2-5-3-3 wm. Para- physes scanty, branched and anastomosing, thin, 0-5—1 wm wide; apices thickened with greenish (K+ violet) pigment and up to 1-7 xm wide, often overtopping the tops of the asci. Hypothecium 25-70 um tall, hyaline or dilute dull yellowish. Excipulum indistinct, sometimes evident as a non-amyloid, narrow (c. 10-15 um), reflexed zone of radiating, branched and anastomosing hyphae, 0-5-1 wm wide. Pycnidia numerous but small and inconspicuous, sessile, black, with dark greenish (K+ violet) walls; of two types: (a) 40-50 wm diam; conidia (mesoconidia) cylindrical or ovoid- LICHEN GENUS MICAREA IN EUROPE Table 6 Diagnostic features for the separation of Micarea melanobola, M. misella, and M. prasina. ES? prasina melanobola misella Apothecia colour pallid to dark grey dark grey to black black, rarely paler rarely black Apothecia size (mm) 0-1-0-4 0-1-0-25 0-1-0-3 Location of epithecial gel-matrix walls of paraphyses + gel-matrix pigment gel-matrix Spore septation 0-1(-3) 0-1 O(-1) Spore length (um) 8-14(-17) 6-5—9-5 7-9-7 Spore breadth (um) 2-3-4(-5) 2-3(-3-7) 2:5-3:3 Paraphyses usually numerous sparse sparse Mesoconidia («~m) (3-S—)4-6X 1-2-1-7 3-3-4-5x 1-1-3 3-5-5 x 1-1-5(-1-7) Microconidia (4m) (S5—)5-5-8 x0-7-1 4-5:5x0-7-0:8 3-8-6 X0-6-0-8 Pycnidia (meso-) immersed to sessile sessile stalked Thallus type goniocyst goniocyst endoxylic or areolate Chemistry (t.1.c.) one of 3 unknowns, unknown, not tested + gyrophoric acid ?+ gyrophoric acid in trace amounts various conifer bark mostly lignum, not known on bark Substrata oblong, often biguttulate and slightly constricted in the middle, 3-3-4-5 x 1-1-3 wm; (b) 25-40 pm diam; conidia (microconidia) cylindrical or fusiform-cylindrical, 4-5-5 x0-7-0-8 um. Chemistry: Thallus and sections of apothecia C— (not red), but olivaceous pigment reacting C+ violet; material insufficient for analysis by t.l.c. Observations: Presumably on account of its finely granular thallus, greenish (K+ violet) epithecium and small, 1-septate spores, M. melanobola was considered to be a form of M. prasina by Hedlund (1892), and asynonym of M. prasina by Vézda & V. Wirth (1976). Hedlund also suggested a relationship to M. misella, and I find that in many respects it is closer to M. misella than to M. prasina. The three species are compared in Table 6; this does not include features common to all of them (e.g. presence of olivaceous, K+ violet pigment, and + hyaline hypothecia). The single feature separating M. melanobola from both the other species is the presence of closely adhering pigment on (? or in) the apical walls of its paraphyses. To this it may be possible to add some aspects of chemistry, but unfortunately there is not sufficient available material of M. melanobola for analysis by t.l.c. The principal features separating M. melanobola from M. misella involve thallus type, spore septation, insertion of mesoconidia-containing pycnidia and substrata. Similarly, those for separating M. melanobola from M. prasina involve the sizes of apothecia and spores, density of paraphyses and, above all, size of microconidia. In external appearance M. melanobola is almost indistinguishable from M. synotheoides, but the latter is easily identified by its acicular spores. Habitat and distribution: M. melanobola is only known with certainty from the type locality in southern Finland where it occurred on the bark of Picea. On the type specimens M. melanobola is accompanied by a few scattered Cladonia squamules, some small lobes of Hypogymnia physodes and Parmeliopsis ambigua, and a few small patches of Dimerella diluta, a white Lepraria sp., and Pertusaria ? pupillaris (soralia K—, PD+ red). Further well documented gatherings are required to obtain a better understanding of the ecology of this species. 158 BRIAN JOHN COPPINS 26. Micarea misella (Nyl.) Hed. (Fig. 23A; Map 13) in Bih. K. svenska VetenskAkad. Handl. II, 18 (3): 78, 88 (1892). — Lecidea anomala f. misella Nyl., Lich. Scand. : 202 (1861). — Lecidea misella (Nyl.) Nyl. in Not. Sallsk. Fauna Fl. fenn. Férh. 8: 177 (1866). Type: Finland, Nylandia, Helsingfors [Helsinki], 1858, W. Nylander (H — lectotype!). Lecidea resinae* [subsp.] L. globularis Nyl., Lich. Scand.: 213 (1861). — Lecidea globularis (Nyl.) Lamy in Bull. Soc. bot. Fr. 25: 435 (1878). — Micarea globularis (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Hand. III, 18 (3): 78, 88 (1892). Type: Sweden, ‘Arthonia turgida var. y globifera Svecia 112° (H-ACH 52 —holotype!). Lecidea melanochroza Leighton ex Crombie in J. Bot., Lond. 9: 178 (1871). Type: Scotland, Perthshire, near Loch Tummel, J. M. Crombie (BM - lectotype!; isolectotypes: BM ex K!, and distributed in Crombie Lich. Brit. Exs. 174 (BM!) [note: this example includes M. prasina and those in E and M contain M. prasina only])). Lecidea misella f. brasiliana Vainio in Acta Soc. Fauna Fl. fenn 7 (2): 57 (1890). Type: Brazil, Minas Geraes, Carassa, 1400 m, on lignum, Vainio, Lich. Brasil. Exs. 1420 (BM ex K - lectotype!; M - isolectotype!). Bilimbia melaena f. decrustata Vainio in Acta Soc. Fauna Fl. fenn. 53 (1): 255 (1922). Type: Finland, Tavastia australis, Hollola, 1871, Vainio (TUR-VAINIO 21480 — lectotype!). Lecidea asserculorum sensu Th. Fr., Lich. Scand. 2: 473 (1874), non Ach (1810). See note by Hedlund (1892: 89). Thallus effuse, often wide-spreading, usually lignicolous and endoxylic, sometimes develop- ing on the surface of the substratum as convex to irregularly subglobose, greenish white to greenish grey, + contiguous granular-areolae, c. 60—120(-150) wm diam. Areolae in section, without a well defined cortex or hyaline amorphous covering layer, but outer 5—12 wm is often tinged with olivaceous-brown pigment (K+ violet); pigment confined to gel-matrix and not adhering to hyphal walls. Phycobiont micareoid, cells 4-7 wm diam. Apothecia numerous, few or absent, scattered to crowded and confluent, immarginate, convex-hemispherical to subglobose, sometimes shortly stipitate, sometimes tuberculate, black, rarely pallid or grey-brown (shade forms), matt. c. 0-1-0-3 mm diam. Hymenium 25-36 pm tall; upper part dull greenish brown or olivaceous, K+ violet, with pigment confined to the gel-matrix; remaining (lower) part hyaline or dilute sordid greenish. Asci clavate, 25-35 x 7-10 pm. Spores ellipsoid, ovoid, or oblong-ovoid, not curved, sometimes with 1-3 guttules, simple but a few (very rarely many) sometimes 1-septate, (6-5—)7—9-5 x2-0-3(-3-7) wm. Paraphyses scanty, sparingly branched 0-5—0-8 wm wide; apical parts often more richly branched, sometimes wider (to 1-5 wm), but without closely adhering pigment. Hypothecium c. 45-70 pm tall, hyaline, or dilute olivaceous (rarely dark olivaceous: ‘f. brasiliana’) and then K+ violet; hyphae interwoven but becoming vertically orientated towards the hymenium, c. 0-7-1 wm wide; ascogenous hyphae with swollen cells, c. 2-4 wm wide. Excipulum poorly developed but often evident as a dark brownish (K+ violet) zone bordering the reflexed edges of the hymenium; hyphae radiating, branched and anastomosing, very narrow, c. 0-5—0-8 xm wide. Pycnidia usually present black, walls green-brown, K+ violet; of two types: (a) sometimes sessile, more usually borne single in the top of a black, unbranched stalk (pycnidiophore), 70-320 am tall (including stalk) and 50-100 wm wide; stalk part composed of interwoven hyphae, c. 1-1-5 wm wide, embedded in a gel-matrix containing a dull brown or olivaceous, K+ violet pigment; conidia (mesoconidia) shortly cylindrical, sometimes faintly biguttulate, 3-5— 5X 1-1-5(-1-7) xm; (b) + immersed in the substratum (lignum) or areolae, c. 40 wm diam; wall with green-brown, K+ violet pigment in upper part, but becoming + hyaline below; conidia (microconidia) narrowly cylindrical, 3-8-6x0-6—-0-8 um. Pycnidia with stalks (a) are mostly confined to forms with an endoxylic thallus. Chemistry: Sections of thallus (when superficial) K— , C+ red, PD—; sections of apothecia C+ violet (olivaceous pigment), but only rarely C+ red; t.l.c.: gyrophoric acid detected in varying amounts, sometimes absent. Observations: Typical forms of M. misella with an endoxylic thallus, numerous apothecia, and stalked pycnidia are easy to recognize. However, when it has a well developed superficial thallus LICHEN GENUS MICAREA IN EUROPE Table 7 Diagnostic features for the separation of Micarea denigrata and M. misella. 159 misella denigrata Apothecia size (mm) Thallus type Spore septation Spore length (4m) Paraphyses Hymenium Macroconidia (4m) Mesoconidia (um) Microconidia (um) 0-1-0-3 Usually endoxylic; sometimes with areolae, 60-120 wm diam O(-1) 6-5-9-5 sparse; 0-5—0-8 wm wide C+ violet; very rarely also C+ orange-red unknown 3-5-5 X 1-1-5 3-8-5 x0-6-0-8 0-1-0-5 usually with areolae, 60-200 wm diam; rarely endoxylic (0-)1 (7—)9-16(-18) numerous; 1-1-7 wm wide C+ violet; usually also C+ orange-red 12-241; curved 2-8-4-5(-S) x 1-2-1-8 (4-S—)5-7-5 x0-7-0:8 Pycnidia (meso-) sessile to stalked immersed to emergent, never stalked gyrophoric acid usually present in large amounts; very rarely not detectable Chemistry (t.1.c.) nil; but gyrophoric acid sometimes present in trace amounts, rarely in large amounts and no stalked pycnidia it can be difficult to separate from M. denigrata, especially if in the latter no conidial (or only the microconidial) state can be found. In such instances M. misella can usually be distinguished by its small apothecia, mainly simple, uncurved ascospores and sparse, thin paraphyses. The same criteria can separate forms of both species with an endoxylic thallus and lacking diagnostic macroconidial or mesoconidial states. A C+ orange-red reaction can usually be obtained in sections of the apothecia of M. denigrata, although the reaction is sometimes very weak or unobtainable in diminutive, lignicolous, epixylic forms; in M. misella I have obtained this reaction in only one or two extremely well-developed specimens. Table 7 outlines the main diagnostic features of these two species. Apart from M. misella, black, stalked pycnidia are also a characteristic feature of M. botryoides, M. melaeniza, and M. nigella; however, the pycnidial wall tissue in these species is brownish or greenish in KOH, and never violaceous as in M. misella. For differences from M. melanobola and M. prasina see Table 6. Habitat and distribution: M. misella is almost exclusively lignicolous (especially on conifer lignum). It has not been found on bark, but has been collected on Picea cones, old basidiomes of polypores (Daedaleopsis confragosa and Gloeophyllum sepiarum), and on moribund Poly- trichum spp. by woodland tracks. Most British collections of M. misella have been made from the sides and undersides of fallen decorticated trunks of Pinus, and associated species included Lecidea icmalea, Micarea prasina, Xylographa abietina, X. vitiligo, Cladonia spp. (scattered squamules), and Hypogymnia physodes. Reports on exposed roots and stones (e.g. Watson, 1930: 53) refer to other species, especially M. bauschiana. In Britain M. misella appears to be rather rare and restricted to Scotland where it is mainly found in the native pine-woods. It has been reported (as ‘Lecidea asserculorum’) from England and Wales, but all the pertinent material seen belongs to diminutive forms of M. denigrata and M. prasina, or quite different species such as M. melaena and Cliostomum griffithii. M. misella is widely distributed in Europe, especially in areas with naturally occurring coniferous forests. It is found as far north as Lycksele Lappmark and Norbotten in Sweden, and an outlying southerly locality is on the Mediterranean island of Corsica where it was collected on the rotting lignum of Eucalyptus. From outside Europe I have seen material of M. misella from Canada (Ontario and Newfoundland), and from the highlands of Minas Geraes (at 1400 m) in Brazil. Exsiccata: Arnold Lich. Exs. 626, 627 (BM ex K, M). Arnold Lich. Mon. 172, 241, 307 (BM ex K, M, MANCH). Britz. Lich. Exs. 208 (M). Claudel & Harm. Lich. Gall. 445 (BM). Crombie Lich. Brit. 174 160 BRIAN JOHN COPPINS z & 3 o a : La Fs ane 7 5 - J Map 13 Micarea misella @ 1950 onwards O Before 1950 p.p. (BM). Elenkin Lich. Ross. 189 (UPS). Krypt. Exs. Vindob. 1532 (BM, BM ex K, M), 4214 (BM ex K, M). Malme Lich. Suec. 365 (S, WIS). Norrlin & Nyl. Herb. Lich. Fenn. 744 (H). Relig. Suza. 42 (BM, GZU, WIS). Vainio Lich. Bras. 1420 (BM ex K, M), 1451 (BM). Zwackh Lich. Exs. 1085 (M). 27. Micarea muhrii Coppins, sp. nov. (Fig. 23B) Thallus effusus, endoxylicus et labem albidam efficiens, vel epixylicus aut epilithicus; thallus superficialis cum areolis convexis, albidus, c. 0-06—0.2 mm diam, interdum coalescentibus in crustam crassiusculam demum rimosam. Algae cellulis 4-7 ~m diam. Apothecia adnata, convexa vel hemisphaerica autem haud globosa, griseo-atra, aut pallida vel rufo-brunneola vel fumosa ubi umbrose, 0-15—-0-46 mm diam, immarginata sed vulgo zona marginali, complana, + incolorata. Hymenium 40-45 um altum, parte summa obscure olivacea vel griseo-viridi et K—, vel raro hyalina. Ascosporae ellipsoideae, ovoideo-ellipsoideae vel oblongo-ellipsoideae, simplices, 9-12 4-5 ym. Paraphyses aliquantum paucae, ramosae et interdum anastomosantes, 1-1-5 um latae, apices versus vulgo incrasstae ad 3-5 um latae et vulgo pigmentiferae. Hypothecium obscure rufofuscum, K—, HNO3—. Excipulum hyalinum, paulum evolutum mox reflexum. Pycnidia pauca, immersa. Conidia cylindrica, 4-6x0-8-1 wm. Thallus et apothecia K—, C—, PD-; sine materia chemica. Typus: Suecia, Wermelandia, par. Lungsund, Pungbacken, alt. c. 170 m, in loco aperto ad truncum decorticatum supra rivum, 15 vii 1980, leg. L.-E. Muhr 2851 (E —holotypus). [t.l.c.: no substances. | LICHEN GENUS MICAREA IN EUROPE 161 Thallus effuse, endoxylic and visible as a white stain, or epixylic, or rarely epilithic; superficial thallus formed of whitish, greenish-white or grey-white, convex areolae, c. 0-06—0-2 mm diam, which are especially well developed around the apothecia; areolae sometimes coalescing to form a thickish crust up to 0-2 mm thick, which may eventually become rimose. Areolae in section without a hyaline amorphous covering layer. Phycobiont micareoid, cells 4-7 wm diam. Apothecia numerous, occasionally coalescing, adnate, convex, soon becoming hemispherical, but never globose or tuberculate, grey-black, or pallid, reddish brown or brown-grey in shade forms, 0-15—0-46 mm diam; disc matt or slightly glossy; immarginate but the adnate rim is often seen as a + colourless or watery grey zone, c. 20-40 wm wide. Hymenium 40-45 um, + hyaline with upper part dark olivaceous or grey-green, K—, HNO;+ purple-red, or + hyaline throughout in shade forms. Asci cylindrical-clavate, 38-40 10-12 xm. Spores simple, often biguttulate, ellipsoid, ovoid-ellipsoid or oblong-ellipsoid, 9-12x4-5 um. Paraphyses rather scanty, branched (especially in their upper parts), sometimes anastomosing, 1-1-5 wm wide, often slightly incrassate at apices to 2 wm wide; or upper 9-15 «um of paraphyses thickened by dense greenish pigment and then to 3-5 wm wide. Hypothecium c. 70-120 um tall, dark reddish brown, K—, HNO;-—; hyphae c. 1-2 wm wide embedded in the densely pigmented matrix, interwoven but becoming vertically orientated towards the hymenium; ascogenous hyphae with short, swollen cells to 5 wm wide. Excipulum sometimes evident in sections of young apothecia as anon-amyloid, hyaline zone, but soon reflexed and obscured in older apothecia; composed of radiating, branched and anastomosing hyphae 1—1-5(—2) um wide. Pycnidia rare, sunken within the areolae, c. 40-50 wm diam; upper part of wall around the ostiole olivaceous, lower part reddish brown, all parts K— _ Conidia (microconidia) cylindrical, 4-6 0-8-1 wm. Chemistry: Thallus K—, C—, PD—; apothecia sections C—; no substances detected by t.l.c. Observations: Micarea muhrii is characterized by its convex-adnate, never tuberculate, usually dark grey apothecia, greenish upper hymenium, dark reddish brown hypothecium (without green or purple tinges, even in K), and simple spores. It recalls the mainly saxicolous M. lutilata in its pigmentation, but that species differs in having convex-subglobose, often tuberculate apothecia, smaller spores, and a larger celled phycobiont. The lignicolous M. melaeniza is another species with similar pigmentation, but has smaller, markedly convex to tuberculate apothecia, smaller spores, and sessile or stalked pycnidia. Habitat and distribution: Often in abundance on lignum of decorticate logs that lie across streams and probably become inundated at times; collected once on a periodically inundated boulder in a stream. So far, known only from Varmland in Sweden, from where it has been found in at least four localities by Lars-Erik Muhr, in whose honour I have the pleasure of naming this species. 28. Micarea myriocarpa V. Wirth & Vézda ex Coppins, sp. nov. (Figs 23C, 47C; Map 14) Micarea myriocarpa V. Wirth & Vézda in Poelt & Vézda, Bestimmungsschl. europ. Flechten, Erganzungs- heft I: 161 (1977); nom. nudum (Art. 32). — Micarea myriocarpa V. Wirth & Vézda in V. Wirth, Flechtenfl.: 341, 345 (1980); nom. nudum (Art. 32). Thallus effusus, farinoso-granulosus, tenuissimus vel crassiusculus ad 0-3 mm crassus, pallide viridis vel viridulo-fulvus. Algae cellulis + globisis, c. 4-7 wm diam, conglomeratis in granula goniocystiformes. Apothecia immarginata, primum convexo-hemisphaerica mox tuberculata, pallide vel obscure rufa, vel fusca, 0-1-0-25 mm diam. Hymenium 25-35 xm altum, + hyalinum vel dilute rufo-brunneolum, cum vittis verticalibus obscure rufo-brunneolis, K—. Ascosporae oblongae vel oblongo-ovoideae, rectae vel +’ curvatae, 5-5—8-5 x 1-5-2-5 wm. Paraphyses aliquantum paucae, dimorphae: p.p. hyalinae, laxae, simplices vel parce ramosae, interdum anastomosantes, graciles, 0-8-1-2 um latae, apicibus interdum incrassatis ad 1-8 ym latis; p.p. pigmentiferae, fasciculatae, plerumque simplices, crassae, 2-3 wm latae. Hypothecium rufo-fuscum vel armeniaco-fuscum, K—, HNO; -—. Excipulum nullum. Pycnidia pauca et inconspicuosa, + 162 BRIAN JOHN COPPINS sessilia, doliiformia, rufo-fusca, c. 25-30 wm diam. Conidia breviter cylindrica, 2-5—3-2 x 1-1-3 wm. Thallus et apothecia K—, C—, PD—; sine materia chemica. Typus: Germania: ‘Baden, Nordschwarzwald, Schurmsee bei Sch6nmiinzach bei Schwarzenberg, alt. 790 m, Tannen am Seeausfluss’, 25 v 1976, leg. V. Wirth (hb V. Wirth 6085 — holotypus!). Thallus effuse, pale green or greenish buff, scurfy farinose-granular, thin or thickish up to 0-3 mm thick. Phycobiont ?micareoid, grouped into small goniocyst-like clusters c. 10-15 wm diam; cells + globose 4—7 xm diam, or ellipsoid 5—7 X3-5—5 ym. Apothecia numerous, immarginate, at first convex-hemispherical, soon becoming + globose, often becoming tuberculate, pale to dark reddish brown, rarely brown-black, 0-1-0-15 mm diam, or to 0-25 mm diam when tuberculate. Hymenium c. 25-35 um tall, hyaline or dilute brownish, with dark brown, vertical streaks due to fasciculate pigmented paraphyses; K—, HNO; -. Asci cylindrical-clavate, 21-25 x5-7 um. Spores oblong or oblong-ovoid, straight or slightly curved, simple or 1-septate, 5-5—8-5x1-5-2:5 wm. Paraphyses rather scanty, of two types: p.p. hyaline throughout, evenly distributed, simple or sparingly branched (especially above), sometimes anastomosing, thin, 0-8-1-2 4m wide, sometimes widening towards their apices to 1-8 wm; p.p. pigmented + throughout, in small fasciculate clusters arising from the hypothecium, stout, 2-3 wm wide. Hypothecium 35-85 ym tall, reddish brown or dull orange- brown, K—, HNO;3-—; hyphae coated with dense brownish pigment and c. 2-3 um wide, interwoven but becoming vertically orientated towards the hymenium and sometimes con- tinuing into it as pigmented, fasciculate paraphyses; ascogenous hyphae similarly pigmented but with short swollen cells, to 4 wm wide. Excipulum not evident, even in sections of young apothecia. ( pe ° nN w > Fa & OR, ase ; rae ee : Z ie . Peer ail a fre : = 4 p 4 : 2 4 ae iat w Ae| |) AT | lee 3 1 5 } 3 ® cope 2 ea poee aK c ; avian Sl x ee a = b o Map 14 Micarea myriocarpa @ 1950 onwards O Before 1950 LICHEN GENUS MICAREA IN EUROPE 163 Pycnidia occasionally present but very inconspicuous, + sessile, doliiform reddish brown, c. 25-35 wm tall and 25-30 wm diam; wall reddish brown or dull orange-brown, K—, HNO3-. Conidiogenous cells + cylindrical to ampulliform, 4—5 x 1-1-2 4m but base sometimes swollen to 2 um wide. Conidia (mesoconidia) short cylindrical, sometimes faintly biguttulate, 2-5—3-2 x 1- 1-3 wm. Chemistry: Thallus K—, C—, PD—; apothecia sections C—; no substances detected by t.l.c. Observations: Micarea myriocarpa is characterized by its numerous, small, + globose to tuberculate, reddish brown apothecia, darkish (but never blackish, even in thick sections) hypothecium, and small, narrow, 0—1-septate spores. The apothecial tissues completely lack any greenish pigmentation, unlike in M. tuberculata and Psilolechia clavulifera, species of similar habitats with which M. myriocarpa has been confused. M. botryoides is another common associate of M. myriocarpa, but can be distinguished by its black apothecia, darker hypothe- cium, larger spores, and numerous stalked, black pycnidia. Habitat and distribution: M. myriocarpa is faithful to the Micareetum sylvicolae and is found especially on exposed roots in dry underhangs under trees on steep slopes in valley woodlands. In the same situations it has been collected also on loose stones and mats of dry bryophytes. An additional habitat for M. myriocarpa is the dry undersides of stumps (especially conifers), situations which are microclimatically identical to rocky underhangs. In Britain, M. myriocarpa is widely distributed, but little collected, in western and northern districts. On the Continent the species is little known, but I have seen material from western Norway as far north as Trondheim, and Varmland in Sweden; and Wirth (loc. cit.) records it from the northern Schwarzwald in Germany (the type locality). 29. Micarea nigella Coppins, sp. nov. (Figs 24A, 47B; Map 15) Thallus effusus, immersus vel tenuissimus, albidus vel viridi-griseus. Algae cellulis 4-7 wm diam. Apothecia primum subglobosa mox tuberculata, immarginata, atra, 0-1-0-3 mm diam. Hymenium 25-30 wm altum, + incoloratum, autem parte summa (epithecio) purpureo-fusca K+ olivacea. Ascosporae ellipsoideae, ovoideae vel oblongo-ovoideae, simplices 6-5—112-5—4 wm. Paraphyses aliquantum paucae, dimorphae: p.p. hyalinae, laxae, ramosae, gracillimae, 0-7-1 wm latae, apicibus vulgo leviter incrassatis ad 2 wm latis; p.p. pigmentiferae, fasciculatae, plerumque simplices, crassae, 2-2-5 ym latae. Hypothecium obscure purpureo-fuscum, K+ obscure olivaceum. Excipulum paulum evolutum mox reflexum. Pycnidia numerosa, atra, sessilia vel stipitata, interdum ramosa, c. 40-300 ym alta et 40-80 um lata, parietibus purpureo-fuscis, K+ obscure olivaceis. Conidia ellipsoidea ad breviter cylindrica, 3-4— 4-3X1-2-1-6 um. Thallus et apothecia K—, C—, PD-. Typus: Dania, Jyllandia, c. 16 km septentriones e Hobro, Rold Skov, Torstedlund Skov, ad truncum decorticatum vetustum coniferarum in sylva mixta, 8 viii 1979, B. J. Coppins 4429 (E — holotypus). Thallus effuse, endoxylic, or + epixylic as a thin whitish to pale green-grey crust, but not forming discrete granules or areolae. Phycobiont micareoid, cells 4-7 wm diam. Apothecia few to numerous, immarginate, subglobose at first, often becoming tuberculate, black, matt, 0-1-0-2 xm diam, or to 0-3 mm diam when tuberculate. Hymenium 25-30 um tall, hyaline or tinged dull purplish brown in places, often with darker, purplish brown vertical streaks; upper part (2-15 wm) irregularly pigmented purplish brown; K+ dull green, HNO3+ red. Asci clavate, 22-27X10-11 um. Spores ellipsoid, ovoid or oblong-ovoid, simple, 6-5— 12x2-5—4 um. Paraphyses scanty, of two types: p.p. evenly distributed, branched, hyaline, thin, 0-7-1 wm wide, with apices sometimes widening to 2 wm but not becoming pigmented although embedded in pigmented gel-matrix; p.p. scattered or in small fascicles, mostly simple, stout, 2—3 m wide, coated + throughout by dark pigment. Hypothecium 70-100(—160) um tall, mottled, dark purplish brown, K+ dark dull green, HNO3+ purple-red; hyphae coated with dense: pigment and c. 1-5-3 wm wide, interwoven but becoming vertically orientated towards the hymenium and sometimes continuing into it as stout, pigmented paraphyses; ascogenous hyphae similarly pigmented but with short, swollen cells, to 4 wm wide. Excipulum indistinct} 164 BRIAN JOHN COPPINS sometimes evident in young apothecia as a dark (concolorous with hypothecium) narrow zone forming an edge to the reflexed part of the hymenium; hyphae radiating, branched and anastomosing, c. 0-8-1-5 wm wide. Pycnidia numerous, black, sessile or more usually stalked, 60-300 um tall (including stalk) and 40-80 um diam; stalks (pycnidiophores) simple or sometimes branched bearing up to 4 pycnidia; stalk and pycnidial wall tissues dark purplish brown, K+ dark green, HNO3+ purple-red. Conidiogenous cells 4-4-7-6 wm tall, with a cylindrical neck 1-9-3-7 um tall and 0-8-1-4 wm wide, and with a swollen base 2:4-3(-3-7) wm wide, the wall of which is often pigmented; percurrent proliferations not seen. Conidia (mesoconidia) ellipsoid or short cylin- drical, sometimes faintly biguttulate, 3-4-4-3 x 1-2-1-6 wm. Chemistry: Thallus K—, C—, PD—; apothecia sections C—; no substances detected by t.l.c. Observations: Micarea nigella belongs to the group of lignicolous species (including M. contexta, M. eximia, M. melaeniza, M. misella, M. olivacea, and M. rhabdogena) with an endoxylic or indistinct thallus and small, + globose to tuberculate, black apothecia. It is characterized by the purple-brown, K+ green pigment in the hymenium, hypothecium and pycnidial tissues, simple spores and stalked pycnidia. It is most likely to be confused in the field with other species with stalked, black pycnidia, namely M. melaeniza and M. misella. M. melaeniza differs in having a bright green hymenium, a red-brown hypothecium that does not turn green in K, a different pycnidial pigmentation and shorter mesoconidia. M. misella has a K+ violet hymenium, a + hyaline hypothecium and an olivaceous, K+ violet pigment in the pycnidial wall tissues. Apart from their lack of stalked pycnidia, M. contexta, M. eximia, and M. ¢ = (0) 1 2 3 4 oO WE 0 Aan J) = \ { ae Cae z 7" 2 z ORS by Du Se pw ‘a =u seal 2. A 2 ‘ oe Seats CHANNEL ISLANDS “ d it BO wee PLOTTED ON Ja = UTM GRID a’, "eed N. ej A Kt a ie) {¢) L 1 2 3 4 5 6 ) Map 15 Micarea nigella@® + Micarea olivacea ® LICHEN GENUS MICAREA IN EUROPE 165 olivacea can be distinguished from M. nigella by their 1-septate spores. M. rhabdogena has a brown (K+ dissolving) pigment in the upper hymenium, and smaller spores. M. nigella could be confused with diminutive lignicolous forms of M. melaena, but that species usually has a bright green hymenium, larger and 1—3-septate spores, and never has stalked pycnidia. Habitat and distribution: M. nigella occurs on rather soft lignum of stumps of conifers and birch, in deeply shaded situations. Primarily, it appears to be an inhabitant of conifer woodlands, and has been found both in native pine forest (Black Wood of Rannoch) and mature conifer plantations. It is known from only three localities: north Jylland in Denmark, the southern central highlands of Scotland and north-east England; and, although undoubtedly a rather rare species, it has surely been much overlooked, at least in north-west Europe. 30. Micarea nitschkeana (Lahm ex Rabenh.) Harm. (Fig. 24B; Map 16) in Bull. Soc. Sci. Nancy II, 33: 64 (1899). — Bilimbia nitschkeana Lahm ex Rabenh., Lich. Europ. Exs. 583 (1861). — Micarea denigrata var. nitschkeana (Lahm ex Rabenh.) Hedl. in Bih. K. svenska VetenskAkad. Hanadl. II, 18 (3): 79, 90 (1892). — Bacidia nitschkeana (Lahm ex Rabenh.) Zahlbr. in Anniln naturh. Mus. Wien 22: 342 (1905). Type: Germany, Nordrhein-Westfalen, between Minster and Nobriskrug, on Pinus in wood, T. R. J. Nitschke, Rabenh. Lich. Eur. Exs. 583 (M — lectotype!; isolectotypes: BM!, BM ex K!, M!). Lecidea spododes Nyl. in Flora, Jena 52: 410 (1869). — Bacidia spododes (Ny1.) Zahlbr., Cat. lich. univ. 4: 151 (1926). Type: England, Hampshire, Lyndhurst, New Forest, old pales, J. M. Crombie (H-NYL 18819 — lectotype!; isolectotypes: BM!, H-NYL 18820!). Bacidia nitschkeana f. microcarpa Erichsen in Verh. bot. Ver. Prov. Brandenb. 71: 97 (1929). Type: West Germany, Schleswig-Holstein, Eckernférde, near Levensau, Felmerholz, on Picea twigs, 9 xi 1924, C. F. E. Erichsen (HBG — holotype!). ?Bilimbia spododes f. fusca B. de Lesd., Rech. Lich. Dunkerque: 200 (1910). Type: France, Nord, Dunkerque, Malo-Terminus, on dead branch of Hippophae, B. de Lesdain (not seen). ?Bilimbia spododes f. livida B. de Lesd., Rech. Lich. Dunkerque, Suppl. I: 120 (1914). Type: France, Nord, Dunkerque, Zuydcoote, on dead branch of Hippophae, B. de Lesdain (not seen). ?Bilimbia spododes var. nigra B. de Lesd., Rech. Lich. Dunkerque, Suppl. I: 120 (1914). Type: France, Nord, Dunkerque, Malo-Terminus, dunes, on dried rhizome of Ammophila arenaria, B. de Lesdain (not seen). Thallus effuse, usually forming small patches but sometimes wide-spreading, sometimes partly endoxylic but usually developed on the surface of the substratum as crowded, often contiguous, convex to subglobose areolae. Areolae especially well developed around the apothecia, dull greenish white to green-grey, surface matt, c. 40-200 um diam; in section, without a well defined cortex and not surrounded by an amorphous covering layer, outermost hyphae sometimes surrounded by dilute olivaceous, K+ violet pigment. Thallus sometimes blackish and scurfy due to disruption by invading dematiaceous fungi and non-lichenized algae. Phycobiont micareoid, cells 4-7 wm diam. Apothecia scattered or, more usually, numerous and crowded, frequently contiguous or confluent, adnate, plane to convex-hemispherical, sometimes becoming tuberculate, immargi- nate or occasionally (especially when young) with an indistinct margin that is flush with the level of the disc, grey-black to black, or rarely whitish to pale grey-brown (shade forms), sometimes paler at the margin, matt. c. 0-1-0-3 mm diam, or to 0-4 mm diam when tuberculate. Hymenium 30-40 um tall, upper part dilute olivaceous to olivaceous, K+ violet, HNO3+ red, C+ violet, also C+ orange-red throughout due to gyrophoric acid; olivaceous pigment mostly confined to the gel-matrix, only rarely (in old, dark apothecia) closely bound to the walls of the paraphyses. Asci clavate 25—409-5-11 um. Spores fusiform, mostly curved, (1—)3(-4)-septate, 10-17 (—19) x2-5-3(-3-5) um. Paraphyses numerous, branched and anastomosing 1-1-5(-1-7) wm; apices not or only slightly incrassate, and mostly without closely adhering pigment. Hypothe- cium 30-50 um tall, hyaline; hyphae interwoven, c. 1-1-5 4m wide; ascogenous hyphae with swollen cells c. 2-5 wm wide. Excipulum distinct in sections of young, + plane apothecia, but 166 BRIAN JOHN COPPINS reflexed and obscured in older, more convex apothecia; hyaline or dilute olivaceous (K+ violet) at outer edge; hyphae radiating, branched and anastomosing, c. 1-1-5 wm wide. Pycnidia indistinct but usually present, immersed in areolae; walls hyaline throughout, or olivaceous (K+ violet) around the ostiole; of three types: (a) c. 60-80 um diam, ostiole sometimes widely gaping; Conidia (macroconidia) curved or hamate, 1—3-septate, 12-26xc. 1 pm; (b) c. 50-100 wm diam, ostiole often gaping and conidia extruded as a white blob; conidia (mesoconidia) short-cylindrical or obovate-oblong, apices rounded but often distinctly truncate at proximal end, sometimes biguttulate, (3—)3-5-5(5-7) 1-1-5 um; (c) c. 40-50 wm diam, ostiole never gaping; conidia (microconidia) narrowly fusiform or bacilliform, (4-7—)5-5- 7:5X0-7-0-8 wm. Chemistry: Thallus K—, PD-—; sections of thallus and apothecia C+ orange-red (also, parts with olivaceous pigment, C+ violet); t.l.c.: gyrophoric acid. If thallus is heavily parasitized and scurfy the C+ orange-red reaction may be difficult to obtain and gyrophoric acid may not be detectable by t.l.c. Observations: M. nitschkeana is closely related, and similar in most respects, to M. denigrata (q.v.) with mainly 1-septate spores, and M. globulosella (q.v.) with + acicular or rod-shaped 0-3(—6)-septate spores. Like M. denigrata, M. nitschkeana has three pycnidial anamorphs; usually only one or two of these states are found on any given thallus, but all three states (plus apothecia) have been noted on a single specimen from Scotland (Coppins 3369). In the field M. nitschkeana can be confused with species such as M. cinerea, M. lignaria, and M. peliocarpa, but these species usually have larger apothecia and areolae, and are quite different when examined microscopically. Habitat and distribution: M. nitschkeana is widely distributed in the British Isles at low altitudes (below 300 m), and is most frequently found on twigs or small branches of various deciduous and coniferous trees, e.g. Acer, Aesculus, Alnus, Betula, Fraxinus, Prunus padus, Quercus, Salix, Sambucus, Larix, and Picea; or on the old or dead stems of smaller shrubby plants, especially Calluna and Ulex, but also Erica cinerea, Myrica, Rosa, Sarothamnus, Viburnum, and Juniperus. It has been found to be abundant on old stems and litter of Calluna in some lowland heaths in eastern England (Coppins & Shimwell, 1971), Scotland and Denmark, where it is commonly associated with Lecanora conizaeoides and Scoliciosporum chlorococum; such communities are referable to the Bacidietum chlorococcae. Other associated species on twigs and small branches include Lecanora pulicaris, L. symmicta agg., L. icmalea, Mycoblastus sterilis, Fuscidea lightfootii (western districts), Micarea prasina, Graphis elegans, Stenocybe pullatula (on Alnus) and small thalli of Hypogymnia physodes, H. tubulosa, Parmelia sub- aurifera, P. sulcata, Platismatia glauca, Evernia prunastri, and Ramalina fairinacea. It has occasionally been met with on lignum of fallen conifers (Larix and Pinus) or fence posts, habitats more characteristic of M. denigrata. Associated species on lignum include Lecanora confusa, L. symmicta agg., Lecidea granulosa agg., L. icmalea, Mycoblastus sterilis, Hypogymnia physodes, Parmelia saxatilis, and Platismatia glauca. Occurrences on rock are rare, but a few collections have been made on sandstone or flint stones in heathland, with Lecidea granulosa agg., L. icmalea, and Micarea melaena as associates. It has also been found together with Scoliciosporum umbrinum on a plastic carton in a heathland in the Isle of Wight (Brightman & Seaward, 1977). M. nitschkeana is widely distributed through much of northern Europe. In Scandinavia it has a southern distribution and I have not seen material of it from north of latitude 62°N. It usually occurs in lowland situations, but it attains higher altitudes in the Alps (920 m) and the Nizke Tatry of Czechoslovakia (1250 m). Additional phorophytes from extra-British material include Hippophae (France: dunes near Calais), Vaccinium uliginosum (Germany: Schleswig- Holstein), and Pinus mugo subsp. pumilio (heathlands of southern Germany). From outside Europe I have seen only one specimen: USA, northern California (near San Francisco), where it occurred on fence posts. Exsiccata: Arnold Lich. Exs. 217 (BM ex K, M), 503 A, B (H-NYL, M); 503 C (BM ex K, H-NYL, M), 503 D (H-NYL, M). Arnold Lich. Mon. 48 (BM ex K, M); 270 (BM ex K). Britz. Lich. Exs. 829 (M). LICHEN GENUS MICAREA IN EUROPE 167 ( : >) 0 1 2 3 4 0 aaa ae a 1 3 n & AR, CHANNEL ISLANOS PLOTTED ON UTM GRIO Map 16 Micarea nitschkeana @ 1950 onwards O Before 1950 Claudel & Harm. Lich. Gall. 89 (L,O). Harm. Lich. Loth. 853 (M). Hepp Flecht. Eur. 20 (BM, E, L, M), 21 p.p. (E). Krypt. Exs. Vindob. 1232 (M). Lojka Lich. Univ. 137 (BM ex K, M). Magnusson Lich. Sel. Scand. 340 (M). Malbr. Lich. Norm. 287 (M). Malme Lich. Suec. 25 (M, S). Rabenh. Lich. Eur. 582, 583 (BM, BM ex K, M). Rasanen Lich. Fenn. 642 (M). Zahlbr. Lich. Rar. 110 (BM). Zwackh Lich. Exs. 470 [‘bei Miinster in Westfalen’] (H-NYL 18825), 470 bis [‘Bei Delbrueck in Kreise Paderborn.’] (H-NYL 18822, M), 534 (H-NYL, M), 587 (H-NYL, M). 31. Micarea olivacea Coppins, sp. nov. (Figs 24C, 47A; Map 15) Thallus effusus, endoxylicus vel epixylicus aut epilithicus, tenuissimus, inaequalis, interdum leviter areolatus, albidus vel viridio-griseus. Algae cellulis 4-7 wm diam. Apothecia immarginata, primum convexa vel subglobosa mox tuberculata, atra, 0-1-0-3 mm diam, aut ad 0-4 mm diam ubi tuberculata. Hymenium 30-35 wm altum, dilute sordide olivaceum cum vittis verticalibus atro-olivaceis, K+ clare olivaceo-virescens. Ascosporae oblongae vel ovoideo-oblongae, rectae, (0—)1-septatae, (7—)9-12-3 x2-5— 3-5 wm. Paraphyses numerosae, dimorphae: p.p. hyalinae, laxae, ramosae et anastomosantes, graciles, c. 1-1-2 wm latae, apicibus interdum leviter incrassatis ad 2 um latis; p.p. pigmentiferae, fasciculatae, plerumque simplices, crassae, 2-3 wm latae. Hypothecium sordide atro-olivaceum vel olivaceo-fuscum, K+ virescens. Excipulum paulum evolutum mox reflexum. Pycnidia numerosa sed inconspicua, + immersa, 25-50 um diam., parietibus sordide olivaceis vel olivaceo-fuscis, K+ clare olivaceis. Conidia breviter cylindrica, 3-4-2 1-1-3 wm. Thallus et apoihecia K—, C=, PD—. Typus: Caledonia, Mull, Aros, Druimfin, in ligno duro, 15 v 1968, P. W. James (BM — holotypus). 168 BRIAN JOHN COPPINS Thallus effuse, endoxylic to epixylic, or epilithic, forming a thin, whitish grey or greenish grey, uneven, sometimes weakly areolate crust; often appearing scurfy due to invasion by foreign fungi and algae. Phycobiont micareoid, cells 4-7 xm diam. Apothecia numerous, immarginate, convex to subglobose, often tuberculate, black, matt or slightly glossy, surface minutely roughened, 0-1—-0-25 mm diam, or to 0-4 mm diam when tuberculate. Hymenium 30-35 wm tall, without a distinct upper part (epithecium), dilute sordid olivaceous with dark olivaceous vertical streaks, K+ green intensifying, HNO3+ red. Asci clavate, 35-33X9-5-11 wm. Spores oblong or ovoid-oblong, straight, (0O—)1-septate, (7—) 9-12-32-5-3-5 wm. Paraphyses numerous, of two types: p.p. hyaline throughout, evenly distributed, branched and often anastomosing, rather thin, 1-1-2 wm wide, apices sometimes widening to 2 wm and sometimes with colourless, oily, refractive contents; p.p. broad, 2-3 wm wide, mostly simple, grouped in small fascicles and often embedded in dense pigment. Hypothecium 40-70 um tall, dark sordid olivaceous or olive-brown, K+ green intensifying, HNO3+ red; hyphae embedded in greenish gel-matrix but walls not deeply pigmented, c. 1-5-2 pm wide, interwoven but becoming vertically orientated towards the hymenium and sometimes continuing in to it as fasciculate paraphyses; ascogenous hyphae with short swollen cells, to 5 wm wide. Excipulum indistinct and soon reflexed, sometimes evident in sections of young apothecia as a hyaline or dilute sordid olivaceous, non-amyloid zone c. 20 4m wide; composed of radiating, much branched and anastomosing hyphae 0-8-1-5 wm wide. Pycnidia numerous but inconspicuous, + immersed, 25—50 xm diam; walls sordid olivaceous or olive-brown, K+ green intensifying, HNO3+ red. Conidiogenous cells elongate-ampulliform to subcylindric, 4—5-5 x 1-5-3 um. Conidia (mesoconidia) short cylindrical, sometimes faintly biguttulate, 3-4-2 1-1-3 um. Chemistry: Thallus K—, C—, PD—; apothecia sections C—; no substances detected by t.l.c. Observations: Micarea olivacea differs from M. eximia in its less brightly coloured hymenium pigmentation that is not concentrated in the upper part, more numerous and broader para- physes, shorter and slightly broader spores with rounded apices, and shorter mesoconidia. From M. nigella it can be distinguished by the complete absence of purple pigmentation in water mounts, more numerous and broader paraphyses, more elongate and 1-septate spores, and + immersed (never stalked) pycnidia. M. olivacea is easily confused with epruinose forms of the common lignicole Lecidea turgidula Fr., but that species has a dark green or olive-brown excipulum of conglutinated hyphae that do not separate in K, a paler hypothecium, and a thick walled, large celled phycobiont with cells 12-16 wm diam. In addition, the conidia of L. turgidula are smaller, c. 3-3:5X1-5-1-:8 wm. When on rock M. olivacea can be confused with M. tuberculata, but the latter has a more brightly coloured hymenium and hypothecium, somewhat smaller spores, narrower asci, and a non-micareoid phycobiont with cells c. 5-10 wm when globose or up to 15X7 wm when ellipsoid. M. tuberculata has similar pycnidia and conidia, but its conidiogenous cells are more slender and about twice as long as those of M. olivacea. Habitat and distribution: M. olivacea is an apparently rare, but probably overlooked, species, being known from just two localities, both in Scotland. At the type locality it occurred on the hard lignum of a stump by a conifer plantation; the collection contains no associate species. At the other locality it occurred with Rhizocarpon hochstetteri and Baeomyces rufus on shaded rock in a mature conifer plantation. Little more can be said of its ecology until it becomes better known, but I am inclined to believe that it is essentially a lignicolous species with a preference for hard lignum. If this is true then care should be taken not to overlook it in the field for forms of the common M. denigrata with a reduced thallus. LICHEN GENUS MICAREA IN EUROPE 169 32. Micarea osloensis (Th. Fr.) Hed. (Fig. 28B) in Bih. K. svenska VetenskAkad. Hanadl. II, 18 (3): 83, 97 (1892). — Lecidea osloensis Th. Fr., Lich. Scand. 2: 524 (1874). Type: Norway, Oslo, summit of Ryenbjerget, on decaying plant debris, N. G. Moe (UPS — holotype! [t.1.c.: no substances]). Thallus effuse, indistinct, apparently of whitish grey, + convex areolae, c. 50-70 um diam, in the single specimen seen the thallus is obscured by a bleached non-lichenized alga, giving it a pale grey ‘filmy’ appearance. Phycobiont probably micareoid, cells 4-5—6-5 wm diam. Apothecia numerous, + evenly scattered, immarginate, convex to subglobose, black or brown-black, 0-1-0-26 mm diam. Hymenium c. 30 um tall; upper part (epithecium) red-brown, K—,HNO;- or red tinge intensifying; remaining (lower) part dilute reddish brown with darker, red-brown, vertical streaks. Asci clavate, 26-30 11-13 um. Spores ellipsoid or ovoid-ellipsoid, simple, 6-9:5x3—4 wm. Paraphyses rather numerous, simple below, but mostly branched towards their apices, hyaline and 1-5—-1-8 wm wide, but occasionally surrounded throughout their length by red-brown pigment and then 1-7-2 xm wide; apices sometimes swollen to 3 wm wide, surrounded by an amorphous, red-brown, pigmented matrix that does not disperse in K. Hypothecium c. 70-85 jm tall, dark reddish brown, K—, HNO;-— or red tinge intensifying. Excipulum not evident, even in sections of young apothecia. Pycnidia not seen. Chemistry: Thallus K—, C—, PD-; sections of thallus and apothecia C—; no substances detected by t.l.c. Observations: Micarea osloensis is characterized by its small, blackish apothecia with internal reddish brown pigment that does not disperse in K, simple, ellipsoid spores, and terricolous habitat. In the field it is most likely to be overlooked for Lecidea uliginosa, but that species has a well developed, pseudoparenchymatous excipulum, paraphyses with a brown apical cap, larger spores, and ‘Trapelioid’ asci which do not have a deeply amyloid tholus. Habitat and distribution: Knowledge of M. osloensis is still restricted to its type gathering from Norway. The appearance of this material suggests that it occurred in a woodland clearing on the site of an old bonfire (small amounts of ash are present), probably in a slight depression. The specimen is accompanied by the sterile thallus of Lecidea icmalea and a few plants of the moss Ceratodon purpureus. Dr H. Krog informs me that the hill Ryenberget is now well within the city limits of Oslo, and is now much affected by the results of urbanization, although not completely destroyed. It has an altitude of about 150-200 m, and was almost certainly covered in pine forest before it was engulfed by the city. 33. Micarea peliocarpa (Anzi) Coppins & R. Sant. (Figs 3A-C, 25, 48, 53-54; Map 17) in Coppins & P. James in Lichenologist 11: 155 (1979). — Bilimbia peliocarpa Anzi in Atti Soc. ital. Sci. nat. 9: 250 (1866). — Bacidia peliocarpa (Anzi) Lettau in Hedwigia 52: 133 (1912). — Micarea violacea f. peliocarpa (Anzi) Hedl. in Bih. K. svenska VetenskAkad. Hand. II, 18 (3): 81, 92 (1892); nom. superfl. (Art. 63). Type: Italy, Piemonte, Novara, ‘Sopra i muschi nei monti del Lago Maggiore (Locarno), scoperta dal padre capuccino Agostino Daldini’ (? TO, not seen: request for loan not acknowledged); an unlocalized specimen labelled ‘Bilimbia peliocarpa Anzi Neosymb. no. 44’ [reference to original protologue] in Anzi’s handwriting occurs in UPS(!) and is possibly an isotype. Biatora lignaria B. [var.] conglomerata Hepp, Flecht. Eur., fasc. 5, no. 284 (1857). — Micarea violacea f. conglomerata (Hepp) Hedl. in Bih. K. svenska VetenskAkad. Handl. U1, 18 (3): 81, 92 (1892). Type: . Switzerland, ‘bei Rifferschweil’, on Pinus bark, Hegetschweiler, Hepp Flecht. Eur. 284 (E —lectotype!; M ~ isolectotype!). Biatora lignaria var. saxigena Hepp, Flecht. Eur. fasc. 9, no. 510 (1860). Type: Germany, Hessen, ‘auf Sandsteinfelsen bei Marburg’, W. Uloth, Hepp Flecht. Eur. 510 (E — lectotype!; M — isolectotype!). See note (i) below. 170 BRIAN JOHN COPPINS Bilimbia milliaria var. lignaria *[f.| livida K6rber, Parerga Lich.: 171 (1860). Type: same as Biatora lignaria var. conglomerata Hepp. Lecidea sphaeroides var. leucococca Ny\. in Stizenb. in Nova Acta Leop. — Carol. 34 (2): 18, t. 1, f.D,47-S1 (1867). Type: Finland, Tavastia australis, Evois [‘Evo’], on lignum, 1865, J. P. Norrlin (H-NYL 18377- lectotype!; H —isolectotype!). Bilimbia violacea Arnold in Flora, Jena 53: 473 (1870). — Lecidea violacea P. Crouan & H. Crouan ex Nyl. in Flora, Jena 45: 464 (1862), non Massal. (1852). — Micarea violacea (Arnold) Hedl. in Bih. K. svenska VetenskAkad. Handl. II, 18 (3): 80, 91 (1892). Type: France, Finistére, Brest, on schistose rock, Crouan (H-NYL 18716 — isotype!). [Note: listed as ‘Bacidia violacea (Crouan ex Nyl.) Arnold’ by James (1965a); this should not be confused with Bacidia violacea (Arnold) Arnold, Flora, Jena 67: 581 (1884), which is a Bacidia s. str.] Bilimbia trisepta Hellb., Nerikes Lafflora: 77 (1871). — Biatora trisepta Naeg. ex Mill. Arg. in Mem. Soc. Phys. Hist. nat. Genéve 16: 403 (1862); as ‘Naeg. mss. ex Dr Hepp’, nom. inval. (Arts 32, 34). — Lecidea sabuletorum f. trisepta Stizenb. in Nova Acta Acad. Leop.-Carol. 34 (2): 47, t.3,f. A, 35-62 (1867); nom. superfl. (Art. 63). — Bilimbia hypnophila b. [var.] trisepta Bausch in Ver. naturw. Ver. Karlsruhe 4: 127 (1869); nom. superfl. (Art. 63). — Bilimbia sabuletorum d. [var.| trisepta Rabenh., Krypt.-Fl. Sachsen 2: 187 (1870); nom. superfl. (Art. 63). — Bacidia trisepta (Hellb.) Zahlbr. in Engler & Prantl, Nat. Pflanzenfam. 1 (1*): 135 (1905). — Micarea trisepta (Hellb.) Wetmore in Pub. Mus. Mich. St. Univ. Biol. 3: 284 (1968). Type: Germany, Hessen, ‘auf Sandsteinfelsen bei Marburg’, W. Uloth, Hepp Flecht. Eur. 510 (E— neotype!; M — isoneotype!). See note (ii) below. Lecidea hemipolioides Nyl. in Flora, Jena 56: 294 (1873). — Bilimbia hemipolioides (Nyl.) A. L. Sm., Monogr. Br. Lich. 2: 141 (1911). — Bacidia hemipolioides (Nyl.) Zahlbr., Cat. lich. univ. 4: 114 (1926). — Micarea violacea f. hemipolioides (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Handl. III, 18 (3): 80, 91 (1892). Type: Channel Islands, Jersey, Rozel meadow, on rocks, 1873, C. Larbalestier (H-NYL 18713 - lectotype!; isolectotypes: H-NYL p.m. 4613!, BM!). Bilimbia naegelii f. lapiseda Th. Fr., Lich. Scand. 2: 379 (1874). Type: Sweden, Stockholm, Lyran, on rock, 1870, S. Almquist (UPS — lectotype!). Bilimbia milliaria f. livescens Th. Fr., Lich. Scand. 2: 383 (1874). Type: Sweden, Narke, Gotlunda, ‘pa stenmuren mellan Stabacken och Hogby’, on mosses, 1863, Blomberg (UPS — lectotype!). Lecidea albidolivens Ny\. in Flora, Jena 57: 10 (1874). — Bilimbia albidolivens (Nyl.) Blomb. & Forss., Enum. Pl. Scand.: 82 (1880). Type: Finland, Tavastia australis, Padasjoki, Nyyst6la, on lignum, 1872, E. A. Lang [Vainio] (H-NYL 18775 — lectotype!; H — isolectotype!). Lecidea fraterculans Nyl. in Flora, Jena 58: 11 (1875). Type: Finland, Tavastia australis, Padasjoki, Nysst6la, on rock, 1872, E. A. Lang [Vainio] (H-NYL 18704a — lectotype!; H-NYL 18704 — isolecto- type!). Lecidea triseptatula Nyl. in Flora, Jena, 58: 361 (1875). Type: Finland, Tavastia australis, Hollola, on lignum, 1874, E. A. Lang [Vainio] (H-NYL 18709 - lectotype!; isolectotypes: H-NYL 18710!, H!). Bilimbia albicans Arnold in Flora, Jena 65: 140 (1882). Type: Germany, Bayern, Oberbayern, near Bad Télz, W side of the Blombergs, on sandstone in wood, 5 ix 1880, Arnold, Lich. exs. 837 (BM - lectotype!). Micarea violacea f. cupreola Hedl. in Bih. K. svenska VetenskAkad. Handl. III, 18 (3): 80, 91 (1892). Type: Sweden, Hialsingland, Jarvs6, on rock, viii 1891, Hedlund (S — holotype!). Lecidea triseptatuloides Harm. in Bull. Séanc. Soc. Sci. Nancy I, 33: 64 (1899). — Bacidia triseptatuloides (Harm.) Zahlbr., Cat. lich. univ. 4: 161 (1926). Type: France, Moselle, Bitche, on Pinus bark, xii 1893, Abbé Kieffer (ANGUC - holotype!). Micarea violacea f. exigua Hedl. in Bih. K. svenska VetenskAkad. Hand. III, 18 (3): 81, 91 (1892); nom. inval. (Art. 26). Notes: (i) ‘saxigena’. This epithet has been applied at intra-specific levels to saxicolous forms of M. lignaria and M. peliocarpa. Its first valid publication was by Hepp (loc. cit.) as ‘Biatora lignaria (Ach.) var. saxigena (Leight) Hepp’ with ‘Lecidea milliaria (Fries) v. saxigena, Leight Lich. Brit. Exs. 210’ given asa synonym. All examples seen by me of Leighton Lich. Brit. 210 (issued in 1856) are M. lignaria; whereas all examples of Hepp Flecht. Eur. 510 are M. peliocarpa. Hepp’s illustration and description of spores are more applicable to M. peliocarpa than M. lignaria. The original valid introduction of the epithet ‘saxigena’ should, therefore, be typified on an example of Hepp FI. Eur. 510 and attributed to Hepp alone. Theoretically ‘saxigena Leighton’ could have been validated for saxicolous forms of M. lignaria by providing a description, excluding other names or specimens that would render it superfluous, and See homonymy with previous combinations of ‘saxigena Hepp’; but I have not found such a case in the iterature. (ii) ‘trisepta’. This epithet first appeared as ‘Biatora trisepta Naeg. mss. ex Dr Hepp’ in the protologue of LICHEN GENUS MICAREA IN EUROPE 171 Patellaria salevensis Mull. Arg. (= Bacidia salevensis (Mill. Arg.) Zahlbr.). Miller presumably regarded Naegeli’s manuscript name as a synonym of his new species. B. salevensis (type in BM!) is a limestone species and is not a Micarea. As indicated by James (1965: 103) the first legitimate publication of the epithet ‘trisepta’ for a Bacidia-like species is that by Hellbom (loc. cit.). By including the epithet in Bilimbia as ‘B. trisepta (Naeg.)’, Hellbom was clearly using it in its traditional sense and not introducing it as an entirely new name. There is no indication that any of the authors using the epithet ‘trisepta Naeg.’ had seen Naegeli’s original material, and no such material has been found amongst Hepp’s collections at BM. Because the original material is probably no longer in existence, neotypification is required. The epithet ‘trisepta’ has been a long source of confusion, being used by various authors for several species, especially Micarea lignaria, M. melaena, M. nitschkeana, M. peliocarpa, and M. ternaria. However, it is clear from the descriptions and cited specimens in Stizenberger (/oc. cit.), Bausch (loc. cit.), Hedlund (1892), and Vainio (1922) that the epithet ‘trisepta’ has most commonly been applied to the species now called Micarea peliocarpa. Because ‘trisepta’ first appeared in the protologue of a saxicolous species the neotype chosen is an example of Hepp Flecht. Eur. 510 which was cited under, or in connection with, ‘trisepta’ in the treatments of Stizenberger, Bauch, Hedlund, and Vainio. Thallus effuse, sometimes partly endocuticular, endophloeodal or endoxylic, more usually developed on the surface of the substratum as rounded, shallow-convex, hemispherical, or + globose areolae. Areolae scattered or more usually + continguous, greenish white, or grey- white to blue-grey, occasionally becoming dark grey, often dark coloured on upper surface but pale and greenish white below, matt or slightly glossy, c. 40-200 um diam. Areolae in section, ecorticate but with an amorphous hyaline covering layer c. 2-5 wm thick; outermost hyphae often with grey-green to blue-green walls, K—, HNO3+ red. Phycobiont micareoid, cells 4-7 pm diam. Apothecia scattered or more usually crowded and often contiguous, adnate, plane to convex, sometimes becoming tuberculate; sometimes with an indistinct margin that is flush with the level of the disc; rarely (as in shaded corticolous forms) whitish or ivory-white, usually pale lead-grey to grey-black, sometimes grey-brown, often whitish or paler at the margin; (0-12—)0-14-0-4 (—0-6) mm diam, or to 1 mm when tuberculate. Disc finely roughened, matt, or slightly glossy (when black); margin (when evident) smooth and often more glossy than the disc. Hymenium 40-55 ym tall, usually olive-green or aeruginose (K—, HNO3+ red) in upper part and + hyaline below; in dark coloured apothecia the green pigment often occurs as vertical streaks through the hymenium. Asci clavate c. 40-55X12-17 um. Spores fusiform, clavate-fusiform, or oblong- fusiform, often slightly curved, (1—)3(—5)-septate, (11—)15—23(-—24) x3-5(-6) wm. Paraphyses numerous, branched, often anastomosing, 1-1-5 wm wide; apices often more richly branched and entangled, often slightly incrassate to c. 1-8 wm, or to 2:5 wm due to thickening by green pigment. Hypothecium c. 40-70 um tall, hyaline or dilute straw; hyphae interwoven, c. 1-1-7 wm wide; ascogenous hyphae with swollen cells c. 2-4 wm wide. Excipulum well developed, hyaline, or dilute straw in part, of richly branched and anastomosing, radiating hyphae c. 1-1-5 wm. Pycnidia frequently present (especially on bark or lignum), of two types: (a) immersed in areolae, white or greenish around the ostiole, 140-200 um diam, ostiole often widely gaping; conidia (macroconidia) markedly curved and often sigmoid, often faintly 1-S-septate, 21—-40(- 50) xX 1-1-5 um; (b) + sessile, white, 50-80 um diam, ostioles not, or only slightly, gaping; conidia (microconidia) narrowly fusiform-cylindrical, (5—)6—7(—7:7) X0-4-0-7 wm. Chemistry: Thallus and whitish apothecia K—, C+ red, PD-—; apothecia in section C+ orange-red; t.l.c.: gyrophoric acid. Observations: M. peliocarpa occurs in a wide range of habitats and is concurrently variable, especially regarding the colour of its apothecia and thallus. This variation is considered to be phenotypic, being due to the amount of green pigment produced in the hymenium and near the surface of the thallus in response to environmental factors, particularly light. When on the bark of old trees in sheltered woodland the apothecia are often very pale to blue-grey, more scattered than usual, and accompanied by numerous pycnidia; such forms are frequent in the ancient woodlands of the New Forest, Hampshire. There is some variation as to the production of areolae, these being often poorly developed or even absent when on cortex or lignum, and 172 BRIAN JOHN COPPINS usually abundant and well developed in more exposed situations on mossy rocks and peaty debris. M. peliocarpa is most closely related to M. alabastrites and M. cinerea (see under the former for further discussion). Apart from these two species, M. peliocarpa is most often confused with M. lignaria (q.v.) especially when occurring in exposed habitats where its apothecia may be black and markedly convex. Similarly, confusion could occur with the rare M. ternaria (q.v.), which like M. peliocarpa has rather flattened apothecia (in section) with a well developed excipulum. When fertile, M. leprosula has apothecia very like those of M. peliocarpa, but its thallus is composed of fragile areolae that readily dissolve into soredia and contains argopsin (PD+ red) as well as gyrophoric acid. In certain habitats, such as the stems of old shrubs (e.g. Calluna, Erica, and Ulex), M. peliocarpa could be mistaken for M. nitschkeana although the latter has an olivaceous, K+ violet pigment in the hymenium, pycnidial walls and thallus, and smaller spores. Specimens collected from Xanthorion communities and attributed to M. peliocarpa (or one of its synonyms) are mostly referable to Bacidia naegelii, which has simple paraphyses with markedly swollen apices, an excipulum of coherent hyphae (in K), and a very different thallus structure. Habitat and distribution: M. peliocarpa is rather catholic in its choice of habitat. As an epiphyte on bark (or on bryophytes thereon) it is mostly commonly found on mature Quercus, but has also been collected from Alnus, Betula, Fagus, Fraxinus, Ilex, Salix, Larix, Pinus, Juniperus, Calluna, Erica, and Ulex. It has a preference for trunks and main stems rather than small twigs and branches. Amongst the British collections associated lichens on bark include Arthonia spadicea, Biatorina atropurpurea, ‘Botrydina vulgaris’, Cladonia spp., Hypogymnia physodes, Lecanora pallida, L. symmicta agg., Lecidea icmalea, Leparia incana agg., Micarea alabastrites, Micarea cinerea, Normandina pulchella, Ochrolechia androgyna, Parmelia saxati- lis, Parmeliella jamesii, Pertusaria hymenea, Phyllopsora rosei, Platismatia glauca, Stenocybe septata and Trapelia sp. In the north and west of Britain it is frequently encountered on the lignum of fallen trunks and large branches (especially conifers), accompanied by such species as Lecidea granulosa agg., L. turgidula, Micarea denigrata, M. lignaria (including var. endoleuca), Mycoblastus sterilis, Ptychographa xylographoides, Xylographa abietina, and X. vitiligo. In the same areas it is often common on peaty soil, and on moribund bryophytes or peaty debris on old walls, boulders and rock faces, preferring rather more sheltered conditions than M. lignaria, with which it is easily confused in the field. Furthermore, it is not found at such high altitudes as attained by that species, and all British collections seem to have been made at below 500 m. Associated species in these habitats include Cladonia crispata, C. coccifera, C. squamosa, C. uncialis, Coelocaulon aculeatum s. lat., Hypogymnia physodes, Lecidea icmalea, Micarea leprosula, Ochrolechia androgyna, Parmelia saxatilis, and Platismatia glauca. It is less frequent- ly found growing directly on rock, and records from Britain indicate that it is able to do so only in rather dry situations. Such occurrences are mostly in the rather low rainfall districts of, for example, Durham, east Yorkshire and Sussex; accompanying species noted include Baeomyces rufus, Cystocoleus ebeneus, Lecidea granulosa agg., L. icmalea, Lecanora polytropa, Lepraria incana agg., Parmeliopsis ambigua, Trapelia coarctata, and T. involuta. M. peliocarpa is widely distributed in Britain but is most common in the west and in upland districts (but at low altitudes). This pattern is reflected in Europe as a whole, and it is apparently common in countries along the Atlantic seaboard and on the Atlantic islands including Iceland, the Azores, and Canary Islands. In central and eastern Europe it is found mainly in mountainous districts. Northwards it extends to just beyond the Arctic Circle, but I do not know of its occurrence in the high arctic. From outside Europe I have seen material from eastern Canada and north-eastern USA, and from New Zealand. Exsiccata: Anzi Lich. Sondr. 170A (UPS). Arnold Lich. Exs. 167A, B (BM ex K, M), 837, 1051 (BM ex K). Arnold Lich. Mon. 118 (BM ex K), 269, 357, 482 (BM ex K, MANCH). Hepp Flecht. Eur. 284 (E, M), 285 (E, L, M), 510 (E, M). Hepp Ziir. 206 (BERN). Korber Lich. Sel.Germ. 133A, B (M). Krypt. Exs. Vindob. 165 (BM). Larb. Lich. Herb. 347 (BM). Leighton Lich. Brit. 238 p.p. (BM, DBN, FRS). Lojka Lich. Hung. 134 (BM ex K). Malme Lich. Suec. 169 (S). Rasanen Lichenoth. Fenn. 343 (BM). Vézda Lich. Sel. 1342, 1380 (BM). Zwackh. Lich. Exs. 276, 897 (UPS). LICHEN GENUS MICAREA IN EUROPE iW = i () 1 2 2 ye 4 5 Pa Pho = s i = : 9 i AN e° Lf aren ed Wea’ | es re 78° f (ie j wi ’ Ve pg : eet ee ee ered 6 Wore 1 Oo. V 6 “op ° y rey : ° 4 aries pn Meee i cae i “el oa ee iy me 2 se Ps a ' ° e KS ie i ~t 7 AGT ade es ‘ lr e ° ae / al i r 3 Map 17 Micarea peliocarpa @ 1950 onwards O Before 1950 34. Micarea prasina Fr. (Figs 1D, 26-27, 49, 53-54; Maps 18-20) Syst. orb.: 256-7 (Dec. 1825). — Biatora prasina Fr., Stirp. agri femsion.: 36 (June 1825), nom. illeg. (Art. 63). — Biatora prasina (Fr.) Trevisan, Linnaea 28: 288 (1856), nec Tuck. & Mont., in Mont. (1857). — Catillaria prasina (Fr.) Th. Fr., Lich. Scand. 2: 572 (1874). See note (i) below and note on Sporacestra under ‘Excluded taxa’. Biatora micrococca K6rber, Parerg. lich.: 155 (1860). — Catillaria micrococca (K6rber) Th. Fr., Lich. Scand. 2: 571 (1874). — Micarea prasina f. micrococca (K6rber) Hed. in Bih. K. svenska VetenskAkad. Hand. II, 18 (3): 77, 87 (1892). — Micarea micrococca (K6rber) H. Gams, Kleine Kryptfl. 3: 67 (1967); comb. inval. (Art. 33.2). Type: Germany, Baden-Wiirttemberg, ‘in regno Wurtembergico’, on Pinus bark, 1862, K. A. Kemmler, Korber Lich. Sel. germ. 250 (L 910, 139-1361-neotype! [t.l.c.: ‘prasina- unknown A’]). See note (ii) below. Lecidea subviridescens Nyl. in Flora, Jena 51: 474 (1868). — Micarea subviridescens (Ny1.) Hedl. in Bih. K. svenska VetenskAkad. Hand. III, 18 (3): 77, 87 (1892). — Bacidia subviridescens (Nyl.) Zahlbr., Cat. lich. univ. 4: 154 (1926). Type: Channel Islands, Jersey, Boulay Bay, on coastal turf and soil, 1868, C. Larbalestier (H-NYL 19056 [as ‘subvirescens’ | —lectotype!; BM — isolectotype! [t.l.c.: ‘prasina unknown C’]; BM - topotypes!). See note (iii) below. Lecidea prasiniza Ny\. in Flora, Jena 57: 312 (1874). — Micarea prasina f. byssacea subf. prasiniza (Nyl.) Th. Fr. in Hedl. in Bih. K. svenska VetenskAkad. Handl. III, 18 (3): 77 (1892). Type: Finland, Tavastia australis, Padasjoki, Nyyst6la, on Alnus, 1872, E. A. Lang [Vainio] (H-NYL 21604 — lectotype!, H — isolectotype!). [material insufficient for t.1.c.]. Lecidea sordidescens Nyl. in Flora, Jena 57: 312 (1874). — Lecidea erysiboides f. sordidescens Nyl. in Norrlin in Not. Sdllsk. Fauna Fl. fenn. Férh. 11: 188 (1870); nom nudum (Art. 32). — Catillaria prasina 174 BRIAN JOHN COPPINS var. byssacea f. sordidescens (Nyl.) Blomb. & Forss., Enum. Pl. Scand.: 91 (1880). — Micarea prasina f. byssacea subf. sordidescens (Nyl.) Th. Fr. in Hedl. in Bih. K. svenska VetenskAkad. Handl. III, 18 (3): 77 (1892). — Lecidea byssacea var. sordidescens (Nyl.) Vainio in Természetr. Fuz. 22: 320 (1899). — Catillaria prasina var. sordidescens (Nyl.) Lettau in Hedwigia 52: 136 (1912). — Micarea prasina var. sordidescens (Nyl.) Brodo in Bull. N. Y. St. Mus. Sci. Serv. 410: 152 (1968). Type: Switzerland, Ziirich, ‘An faulen Fichten-Strunken Z. H.’, Hepp, Flecht. Eur. 278 (E — lectotype! [t.1.c.: ‘prasina-unknown B’]; isolectotypes: BM (boxed set)! [t.l.c.: ‘prasina-unknown B’], H-NYL 21632 [fragment]!, M!). See note (iv) below. Lecidea prasiniza var. prasinoleuca Nyl. in Flora, Jena 64: 7 (1881). Type: Germany, Baden-Wiirttemberg, Heidelberg, K6nigstuhle, on Picea abies, ix 1880, Zwackh, Lich. Exs. 593a (H-NYL 21601 — lectotype! [t.l.c.: ‘prasina-unknown A’]). See note (v) below. Catillaria micrococca f. glebosula Erichsen in Annls mycol. 36: 139 (1938). Type: Germany, Schleswig- Holstein, Eiderstedt, Ording, at base of Betula, 4 viii 1913, C. F. E. Erichsen (HBG — holotype! [t.1.c.: ‘prasina-unknown B’]. ; Catillaria micrococca var. discrepans Erichsen in Annls mycol. 36: 139 (1938). Type: Denmark, Jylland, Abenra (Apenrade), on smooth bark of Alnus in a valley near Elisenlund, 3 ix 1913, C. F. E. Erichsen (HBG - holotype!). [material insufficient for t.1.c.]. Catillaria prasina var. occulta Erichsen in Annls mycol. 36: 140 (1938). Type: Germany, Schleswig- Holstein, Ratzeburg, in the wood ‘Bak’, on + smooth bark of Alnus, 21 iti 1918, C. F. E. Erichsen (HBG —holotype!). [material insufficient for t.1.c.]. Lecidea declivitatum Erichsen in Mitt. Inst. allg. Bot. Hamb. 10: 408 (1939). Type: Germany, Schleswig- Holstein, ‘Krs Eutin: an der Boschung eines Waldwegs an der Nordwest-Seite des Ugleisees’, 5 vi 1914, C. F. E. Erichsen (HBG — holotype! [t.l.c.: ‘prasina-unknown C’}). Micarea polytrichi Poelt & D6ébb. in Bot. Jb. 96: 343 (1975). Type: Austria, Steiermark, Grazer Bergland, a little north of Maria Trost near the Wenisbucher Strasse, c. 460 m, on Polytrichum formosum on shady side of a narrow gorge, 30 iii 1974, J. Poelt (GZU — holotype!) [material insufficient for t.1.c.]. Catillaria prasina a laeta Th. Fr., Lich. Scand. 2: 573 (1874); nom. inval. (Art. 26). ?Biatora byssacea Zwackh in Flora, Jena 45: 510 (1862), non Hampe in Linnaea 25: 709 (1852). Type: Germany, Heidelberg, ‘K6nigsstuhle’, on bark of old Quercus, ? Zwackh (not traced). See note (vi) below. ?Catillaria melanobola f. biseptata B. de Lesd., Rech. Lich. Dunkerque: 198 (1910). Type: France, Nord, Dunkerque, St Pol, on a piece of leather in dunes, B. de Lesdain (not seen). ?Catillaria melanobola f. nigra B. de Lesd., Rech. Lich. Dunkerque: 199 (1910). Type: France, Nord, Dunkerque, lignum of hollow Salix, B. de Lesdain (not seen). Notes: (i) Biatora prasina Fr. (June 1825) is superfluous because the protologue includes the sentence: ‘Crusta est Byss. botryoides Linn!’. Byssus botryoides L. is a valid name (Sp. pl. 2: 1169 (1753)) but the material in the Linnaean herbarium (LINN) is a specimen (1278.16) labelled by Ehrhart; it is an alga and bears a determination label: ‘Protococcus viridis Ag. Determined by Francis Drouet iv. 1950’. In the protologue of Micarea and M. prasina, Fries (Dec. 1825) makes a less dogmatic statement, merely indicating that when sterile M. prasina is indistinguishable from B. botryoides; M. prasina is, thereby, not rendered superfluous. (ii) I have seen two specimens named by KGrber and collected from ‘Wiirtemberg’ by Kemmler: an undated specimen in WRSL and possibly part of the original gathering, and a later specimen issued in KO6rber’s exsiccata. The first specimen would have to be completely destroyed for t.l.c. analysis, whereas the exsiccate specimen is larger and was found to contain ‘prasina-unknown A’, and it is this specimen that I have chosen for typification. If future studies require this chemical race to be recognized at specific rank, then ‘micrococca’ is the earliest available epithet. (iii) L. subviridescens is the earliest name for the chemical race with ‘prasina-unknown C’. (iv) Although Nylander (1874) did not provide a description, he validated L. sordidescens by referring to it Hepp Flecht. Eur. 278; the printed label for this exsiccatum includes a short description and a drawing of spores. (v) If the chemical race containing ‘prasina-unknown A’ were to be awarded varietal rank, then this name provides the earliest available epithet, provided that ‘byssacea’ is not available; see also note (vi). (vi) I have not seen a holotype specimen or any material suitable for the lectotypification of Biatora byssacea Zwackh. A neotype has not been selected because this name is a later homonym of B. byssacea Hampe. Hampe’s name was given to a specimen collected by Charles Stuart in Tasmania, and the protologue ends with ‘an status degenerans’. Biatora byssacea Hampe is included by Zahlbruckner (Cat. lich. univ. 3: 897) in a list of excluded and dubious names and is not accorded a catalogue number; the type LICHEN GENUS MICAREA IN EUROPE 175 material has not been located in BM but it could be elsewhere (? G). Under the taxonomic arrangement given here Zwackh’s epithet is of no consequence. If the chemical races of M. prasina were to be awarded varietal status, then under the provisions of Article 72 the name ‘Catillaria prasina f. [var.| byssacea Th. Fr. (1874)’ could become important and require typification. If this name was typified with a specimen containing ‘prasina unknown A’ then it would be the earliest available name at that rank. Thallus effuse and often wide-spreading, very variable in colour, pale fawn (dry shaded situations), light green, pale to dark grey-green (often with glaucous hue), or olivaceous to Olive-black, often appearing + gelatinous when moist, composed of small + globose granules (goniocysts). Goniocysts c. 12-40(-60) xm diam, thinly scattered, or vertically proliferating to give the thallus a soft isidiose appearance, or densely aggregated to form a thick granular crust up to 200 ~m deep. Goniocysts ecorticate, often with shortly protruding hyphal filaments; outermost hyphae of superficial goniocysts sometimes surrounded by greenish, K+ violet pigment. Phycobiont micareoid, cells 4-7 wm diam. Apothecia usually numerous but sometimes few or absent, immarginate, sometimes adnate and shallow-convex, but more usually becoming markedly convex, subglobose or tuberculate; colour variable, whitish or pallid (’micrococca’ shade-forms), pale to dark grey, brownish grey, or grey-black, matt; 0-1—-0-4 mm diam, or to 0-6 mm when tuberculate. Hymenium 28—45(-S2) pm tall, hyaline or dilute dull straw, but more usually dull olivaceous, especially in the upper part (but coloured portion not sharply delimited) or in vertical streaks; pigment K+ violet, HNO3+ red, confined to gel-matrix. Asci clavate or cylindrical-clavate, 26—40(—50) x 8-12 um. Spores ovoid-ellipsoid, ovoid, ovoid-oblong or oblong, 0-1(—3)-septate; when 1-septate the lower cell is slightly longer and narrower than the upper, and there is often a slight constriction at the septum; variable in size, in range 8—14(—17) X2-3-4(-5) wm. Paraphyses rather numerous, branched and anastomozing, 0-5-1 wm wide; apical parts often wider, to 1:5 wm, but not thickened by pigment; epithecial pigment confined to surrounding gel matrix. Hypothecium c. 40-170 um tall, hyaline to dilute dull yellowish; hyphae interwoven, some becoming vertically orientated towards the hymenium, c. 0-7—1-3 wm wide; ascogenous hyphae with swollen cells, c. 2-4 wm wide. Excipulum poorly developed, sometimes evident as a narrow, hyaline or dilute olivaceous-straw (K+ violet), non-amyloid, reflexed zone; hyphae radiating, branched and anastomosing, c. 0-7—1 wm wide. Pycnidia often present but usually inconspicuous, white, or upper part around the ostiole grey (due to olivaceous, K+ violet pigment), immersed to sessile (never distinctly stalked), of two types: (a) c. 50-120 wm diam, emergent to sessile, ostiole sometimes gaping; conidia (mesoconi- dia) + cylindrical or narrowly obpyriform, often biguttulate and slightly constricted near the middle, (3-5—)4-6X1-2-1:7 wm; (b) c. 30-60(-100) wm, usually immersed in surrounding goniocysts, ostiole rarely gaping; conidia (microconidia) cylindrical or narrowly fusiform, (5—)5-5-8 X0-7-1 wm. Chemistry: All parts K—, C— (but olivaceous pigment C+ violet, never red), PD—-; t.l.c.: specimens have one of three unknowns (‘prasina unknown A’, ‘B’, or ‘C’; see p. 87); trace amounts of gyrophoric acid sometimes detected, possibly due to contamination by intimately associated species such as Lecidea icmalea and L. granulosa agg. Observations: Micarea prasina is the commonest and most variable member of the genus. However, it is one of the few European species whose thallus is composed of minute discrete granules (goniocysts). Two other species with a similar thallus are: M. hedlundii (q.v.) with a K+ red oily substance in the goniocysts and conspicuous tomentose, brown, stalked pycnidia; and M. melanobola which has a sharply delimited epithecium (pigment closely adhering to apices of the paraphyses) and shorter microconidia. Both of these species appear to be very rare. A comparison of the diagnostic features of M. melanobola, M. prasina and M. misella are given in Table 6. Diminutive forms of M. prasina with small pallid apothecia and a scanty thallus have sometimes been regarded as a distinct species (‘Catillaria micrococca’). Such forms have the same basic thallus structure, spores, paraphyses, anamorphs and chemistry (with either ‘prasina unknown A’ or ‘B’) as more typical forms (with coloured apothecia and more robust thallus), 176 BRIAN JOHN COPPINS and I have seen intermediates on many occasions. The ‘micrococca’ forms are found in heavily shaded situations (e.g. tree trunks in dense conifer plantations, and trunks of understory trees or shrubs) and I consider their diminutive stature and lack of pigmentation to be a phenotypic response to low light intensities. The specimens described as Micarea polytrichi are similarly diminished forms, in this case overgrowing mosses (mainly Polytrichum spp.) by woodland roads. European populations of M. prasina have been found to be represented by three chemical races, each with one of three distinctive, but as yet unidentified, substances (see p. 87). During the course of the present study I have been tempted to recognize the chemical races as formal varieties. However I defer from doing so at this time because: (a) the chemical structure and biogenetic relationships between the three substances is not yet known; (b) the three chemical races do not consistently correlate to any clear differences in morphology; (c) the correlations between chemistry and distribution and (or) ecology are as yet uncertain. In the British Isles (at least) the race containing the ‘unknown A’ is by far the commonest and is a ubiquitous coloniser of trees and shrubs in all types of woodland (including young conifer plantations), occurring on tree boles, bark and lignum of fallen trees and stumps, and fallen sticks; it also occurs on shaded mossy turf on the ground and amongst rocks (usually with an east- or north-facing aspect) in maritime situations. The race containing ‘unknown B’ has a more restricted ecology and is mainly found in old woodland situations (including the native pine-woods), growing on the rather dry lignum or old bark of large stumps or fallen trunks; it has not been found on coastal turf or rocks. Specimens of this race tend to have a thick, pale grey (often abrading to greenish SES, no nN Map 18 Micarea prasina sensu lato @ 1950 onwards O Before 1950 LICHEN GENUS MICAREA IN EUROPE sae et 3 @ 7 I p 7 3 ee icp ' =a a é re © 1 a Ree a c ©. ‘2 3 4 5 6 - JS Map 19 Micarea prasina substance A @ 1950 onwards O Before 1950 white) thallus, but equivalent morphological forms containing ‘unknown A’ are sometimes encountered. Specimens containing ‘unknown C’ are exceedingly rare and are known only from coastal turf in south-west England and the Channel Islands, and on argillaceous soil on a bank by a woodland path in south-eastern Schleswig-Holstein. The above ecological and phytogeo- graphical tendencies require to be more thoroughly tested throughout the range of M. prasina, and it is possible that a good case for the taxonomic recognition of these races as varieties, subspecies, or even species could be made. The type material of Lecidea subviridescens contains ‘unknown C’ and has spores that are at the high end of the overall size range for M. prasina s. ampl. (10-18 4-5-5 wm) and some with two or three septa. However the very large 2—3-septate spores are always old with secondarily thickened walls, and similar spores are sometimes encountered in other Micarea species that have predominantly 1-septate spores. Furthermore, similarly large, 2—3-septate spores have been found in some coastal specimens containing ‘unknown A’ and in some inland, lignicolous specimens containing ‘unknown B’. Although I regarded ‘subviridescens’ as a separate species in the recent British checklist (Hawksworth et al., 1980: 62), I now believe it should be subsumed under M. prasina pending further studies as intimated in the preceding paragraph. Habitat and distribution (see also ‘observations’): As an epiphyte on trees or shrubs M. prasina has been found on a wide range of phorophytes; in Britain these include Acer, Alnus, Betula, Castanea, Corylus, Fagus, Fraxinus, Metrosideros, Quercus, Salix, Sambucus, Sorbus, Ulmus, Calluna, Erica, Rhododendron, Vaccinium, Sarothamnus, Ulex, Abies, Larix, Picea, Pinus, Pseudotsuga, and Juniperus. It occurs on the bark or lignum (old dried wounds, etc.) of 178 BRIAN JOHN COPPINS 0 1 2 3 4 10) WT 0 aaa wae ae oe SS feel fk seen eceney: "| at ae ge” v4 acae me e as a 7 q € ‘ more 7 a lee ; ere 2 el ee | oe a pee ie = 3 4 5 e / 2 Map 20 Micarea prasina substance B @ 1950 onwards O Before 1950 substance C ®@ Before 1950 shaded trunks or main stems, but also on low (shaded) branches, fallen debris (especially in conifer plantations) and stumps. It is usually found in woodlands but sometimes occurs in sheltered niches in situations, e.g. bases of Ulex stems in more open, exposed hillside gorse-scrub. The phytosociological affinities of M. prasina are various and difficult to define. When on the bark of deciduous trees it sometimes occurs in shaded facies of communities belonging to the Lobarion pulmonariae, Parmelietum revolutae, and the Pseudevernietum furfuraceae. It is frequently found in small bark crevices amongst + smooth areas of bark which are colonized by communities of the Graphidetum scriptae, Pyrenuletum nitidae, and Lecanoretum subfuscae. When occurring in such situations as the shaded trunks of young conifers or shrubs M. prasina often exists as + pure stands; more detailed phytosociological studies may well lead to the formal syntaxonomic recognition of such communities. From amongst the British collections in E the associated species of M. prasina on the bark of deciduous trees include Arthonia spadicea, Bacidia vezdae, Chrysothrix candelaris, Cladonia coniocraea, Dimerella diluta, Evernia prunas- tri, Graphis elegans, Gyalideopsis anastomosans, Hypogymnia physodes, Lecidea icmalea, Lepraria incana, Ochrolechia androgyna, Opegrapha vulgata, Pachyphiale cornea, Parmelia glabratula, P. sulcata, Pertusaria multipuncta, Thelotrema lepadinum, Trapelia corticola, Dicra- noweissia cirrata, Frullania spp., Hypnum cupressiforme, Lejeunea ulicina, and Metzgeria furcata. When on lignum its affinities are similarly difficult to define but it is often found in more sheltered or shaded (humid) niches adjacent to communities of the Calicietum abietini, Cladonietum coniocraeae, Lecideetum ostreatae, and Parmeliopsidetum ambiguae. LICHEN GENUS MICAREA IN EUROPE 179 Species associated with M. prasina on the lignum of fallen trunks and large stumps in the British Isles include Bryoria fuscescens, Chaenotheca ferruginea, Chaenothecopsis spp., Clado- nia chlorophaea agg., C. macilenta, C. ochrochlora, Hypocenomyce scalaris, Hypogymnia physodes, Lecanactis abietina, Lecidea aeruginosa, L. icmalea, Micarea adnata, M. melaena, and Platismatia glauca. In coastal districts M. prasina is often found on plant debris, soil or moribund bryophytes in rock crevices or on ledges in sheltered gullies, and sometimes it occurs on the ground growing over plant debris or old Armeria tussocks, etc. Such habitats are usually sheltered and (or) witha north- to east-facing aspect. Associated lichens are few, but include Cladonia spp., Lecidea granulosa, L. icmalea, and Lepraria incana agg. Finds of M. prasina growing directly on rock are very rare, but I have seen a few collections made from sandstone rocks in woodlands (from east Sussex and north-east Yorkshire). M. prasina is still to be found close to the centres of large conurbations and cities (e.g. Bristol, Edinburgh, London, and the West Yorkshire conurbation), and its persistence is probably due more to its ability to avoid, rather than tolerate, the direct effects of air pollutants such as sulphur dioxide and its derivatives. It naturally favours substrata with a low pH and is able to grow in very sheltered and shaded situations where more light demanding pollution resistant species (especially Lecanora conizaeoides) are at a competitive disadvantage. M. prasina s. ampl. is widely distributed in the British Isles and much of Europe. In Scandinavia it occurs northwards to at least c. 67°N. It may be rare in districts adjoining the Mediterranean Sea, but those areas are mostly poorly known lichenologically, and I have seen one collection from Toscana (northern Italy). M. prasina is present in Macaronesia (Azores and Canary Islands) and I have seen collections of it from several states in the U.S.A. (Georgia, Maryland, Massachusetts, Michigan, and Wisconsin). In addition I have seen several collections from South America and Australisia which come close to M. prasina, but these require more critical study and will be treated in a later publication. Exsiccata. Containing ‘prasina unknown A’: Arnold Lich. Exs. 1122, 1472 (BM ex K, M). Ko6rber Lich. Sel. Germ. 250 (L). Lojka Lich. Univ. 29 (BM ex K, M), 30 (BM ex K). Malme Lich. Suec. 23 (M, S). Rabenh. Lich. Eur. 676 (BM, BM ex K, M, WRSL). Rasanen Lich. Fenn. 651 (BM, BM ex K, M), 652 (BM, BM ex K). Vézda Lich. Sel. 90 (BM, M), 1467 (BM). Zwackh. Lich. Exs. 593A (H-NYL 21601). Containing ‘prasina unknown B’: Arnold Lich. Exs. 230C (BM, GZU, M, WRSL). Cumm. Dec. N. Am. Lich. I. 355 (BM, WIS). Hepp Flecht. Eur. 278 (BM, E, M, WRSL). Lojka Lich. Univ. 31 (BM ex K, M). Chemistry not tested: Arnold Lich. Exs. 279 (WRSL), 280A (BM, BM ex K, M), 280B (BM ex K, M). Arnold Lich. Mon. 243 (UPS), 245 (BM ex K, M). Hepp Ziir. 224 (BERN). Kutak Lich. Bohem. 310 (O). Magnusson Lich. Sel. Scand. 134 (BM). Malme Lich. Suec. 24 (M, S). Rabenh. Lich. Eur. 733 (H). RAasanen Lich. Fenn. 653 (LD). Schaerer Lich. Helv. 196 p.p. (BM ex K, M). Vézda Lich. Sel. 1595 (BM, GZU, M). Zahlbr. Lich. Rar. 175 (BM). Zwackh. Lich. Exs. 416 (UPS), 591A (H-NYL 21598), 591B (H-NYL 21594, 21599), 592A (UPS), 592B (M, UPS), 592C (UPS), 592D (M, UPS), 592E (UPS), 593B (H-NYL 21600), 593C (H-NYL 21595), 656 (UPS). 35. Micarea pycnidiophora Coppins & P. James (Fig. 28: Map 21) in Lichenologist 11: 153 (1979). Type: England, South Hampshire, New Forest, near Cadnam, Shave Wood, 45 m, on bark of Fagus, 5 xi 1972, B. J. Coppins & F. Rose (E—holotype!; BM — isotype!). Thallus corticolous, often overgrowing moribund thalli of bryophytes and other lichens; effuse, often wide-spreading, thin, uneven, composed of scattered to confluent areolae arising from a thin varnish-like prothallus. Areolae flattened to convex-hemispherical, grey-green or dull green, c. 40-100 um diam; in section, without a distinct cortex or hyaline amorphous covering layer. Phycobiont micareoid, cells 4~7 ~m diam. Apothecia usually few or absent but sometimes abundant, immarginate, convex- hemispherical to globose, often tuberculate, ivory-white to pallid, translucent when moist, surface matt, 0-1-0-3 mm diam. Hymenium 35-50 wm, hyaline. Asci clavate, 30-35 x 10-12 um. Spores + straight and vertically aligned in the ascus (never tightly spiralled), shortly acicular, 180 BRIAN JOHN COPPINS upper end usually broader and more obtuse than the lower end, usually slightly curved, 3-7-septate, (14-)21-34 x 2-2-5(-2-7) um. Paraphyses numerous, branched and anastomosing, c. 1-1-5 wm, not swollen at apices. Hypothecium 20-80 um tall, hyaline; hyphae interwoven, but becoming vertically orientated towards the hymenium, c. 1-1-5 wm wide; ascogenous hyphae with swollen cells, c. 2-4 wm wide. Excipulum poorly developed (c. 10 wm wide); hyphae radiating, branched and anastomosing, c. 1 wm wide. Pycnidia always numerous and conspicuous, sessile or shortly stalked, whitish (concolorous with the apothecia), 100-300 um tall (including stalk) and 60-120 wm diam; stalks (pycni- diophores) simple but occasionally clustered and appearing as if branched at the base; stalk tissue composed of interwoven hyphae c. 1-1-5 wm wide that + separate in K. Conidiogenous cells + cylindrical, 5-10 1-1-5 wm. Conidia (mesoconidia) cylindrical, eguttulate, 3-8-6 x 1- 1-2(-1-5) wm. Chemistry: Apothecia, pycnidia and thallus (in section) K—, C+ red, PD—; t.l.c.: gyrophoric acid. Observations: Micarea pycnidiophora is characterized by its whitish apothecia and concol- orous, + stalked pycnidia, shortly acicular spores, and C+ red reactions (gyrophoric acid). It most closely resembles M. stipitata, which differs in having more elongate and often distinctly branched pycnidiophores, larger conidia, and C— reactions (gyrophoric acid absent). M. globulosella and M. synotheoides have similar acicular spores, but their apothecia are dark- coloured with an olivaceous (K+ violet) pigment, and their pycnidia are inconspicuous and usually immersed in the thallus. Confusion could arise with Scoliciosporum pruinosum (P. = od eee ieee D os ay og Fit pee “ ] 2m [hs ‘ > fer | arr RTT 4 = = BE = ct + Ae| | lar | le pris seks : 7 alee? a ) Se eee erica aa ee : AAs wee S| CHANNEL ISLANOS “3 <> h PLOTTED ON ¢ UTM GRID a’. C 1 2 4 € Map 21 Micarea pycnidiophora & + Micarea stipitata @ LICHEN GENUS MICAREA IN EUROPE 181 James) Vézda and S. schadeanum (Erichsen) Vézda, but those species have narrower spores which are spiralled in the ascus and non-micareoid phycobionts (? Trebouxia); moreover, S. pruinosum has a granular epithecium (granules dissolving in K). Furthermore, these species have inconspicuous (? unknown) pycnidia, and do not contain gyrophoric acid. For illustrations of conidia and conidiogenous cells, and for further discussions, see Coppins & James (1979). Habitat and distribution: M. pycnidiophora is a species of old woodlands, in damp, sheltered, and rather shaded situations. It is generally found on areas of + smooth-bark on the boles of old trees, especially Fagus and Ilex, but also Alnus, Quercus, and Rhododendron. Although characteristic of acid bark it has not yet been recorded on conifers. Its known centre of distribution is the New Forest of Hampshire in southern England, where it is locally frequent. Its other British localities are Sheffield Park in Sussex (on Rhododendron with Gyalideopsis anastomosans) and the Limb Valley, Yorkshire (its most northerly locality; at base of Fraxinus). The latter site is an ancient oak-wood just outside the large industrial city of Sheffield, but M. pycnidiophora occurs in an extremely sheltered situation, where it presumably avoids the effects of the high levels of air pollution prevalent in that area. M. pycnidiophora is little known outside of Britain, but is reported from Bretagne, France (Coppins, 1971, as ‘Bacidia sp. 2’) and the Canary Islands. 36. Micarea rhabdogena (Norman) Hed. (Fig. 29A) in Bih. K. svenska VetenskAkad. Handl. III, 18 (3): 75, 85 (1892). — Biatora (Lecidea) rhabdogena Norman in Ofvers. K. VetenskAkad. Férh. 27: 803 (1870). — Lecidea rhabdogena (Norman) Th. Fr., Lich. Scand. 2: 473 (1874). Type: Norway, Nordland, Maalselven, Skjeggenaes, J. M. Norman (O - lectotype!; isolectotypes: BM!, O!, S!). Lecidea glomerella f. ecrustacea Nyl. ex Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 457 (1934). — Lecidea glomerella f. ecrustacea Nyl. in Elfving in Medd. Soc. Fauna Fl. fenn. 2: 168 (1878); nom. nudum (Art. 32). Type: USSR, Karelskaya ASSR, Karelia olonetsensis, Gorki near Svir, F. Elfving (H-NYL 19120 - holotype!). Thallus inapparent, endoxylic; hyphal walls often olivaceous, K+ violet. Phycobiont prob- ably micareoid, cells c. 5-7 wm diam. Apothecia numerous, immarginate, convex-hemispherical or tuberculate, black, matt, 0-1- 0-3 mm diam, or up to 0-4 mm diam when tuberculate. Hymenium 23-30 um tall; upper part (epithecium) fuscous-brown, HNO3—, K— but pigment dissolving and fading into solution; remaining (lower) part dilute yellowish brown; hymenium sometimes tinged violet in K (evident after brown pigment has dissolved). Asci clavate, 19-28x7-9 um. Spores oblong-ellipsoid, ovoid-oblong or bacilliform, simple or rarely a few 1-septate, 6-9 1-5-2-3 um. Paraphyses numerous, hyaline throughout, c. 1 wm wide in mid-hymenium but often widening above to 1-5 fm; sparingly branched and sometimes anastomosing, but becoming richly branched above where their entangled apices, together with the brown pigment, form a + well delimited epithecium. Hypothecium c. 70-160 um tall, pale, dilute yellowish brown, K—, HNO;3-. Excipulum reflexed, dilute to dark fuscous-brown, K— but partly dissolving into solution; hyphae radiating, branched and anastomosing, c. 0-7—-0-1 ~m wide, sometimes widening to 1-5 jum towards the outer edge. Pycnidia of two types: (a) partly immersed to + sessile, black, matt, 40-80 um diam; wall olivaceous, K+ violet; conidia (mesoconidia) oblong-ellipsoid, oblong-ovate or obovate, c. 3-5—4-7X1-4-1-8 um; (b) immersed between surface wood fibres, black, c. 4 wm diam; wall olivaceous, K+ violet; conidia (microconidia) narrowly cylindrical, 4-4-5 x0-7-0-9 um. Chemistry: Sections of apothecia C—; material insufficient for analysis by t.l.c. Observations: M. rhabdogena resembles M. elachista in apothecial pigmentation and ana- tomy, but differs in its endoxylic thallus, smaller and mostly simple spores, and smaller, matt black pycnidia. M. rhabdogena is similar in appearance and spore characteristics to M. misella, 182 BRIAN JOHN COPPINS but the latter has an olivaceous, K+ violet hymenium (fuscous pigment lacking) and its mesoconidia are borne in stalked pycnidia. Diminutive, endoxylic forms of M. denigrata can be distinguished by the absence of a distinct (fuscous) epithecium, hymenium reacting K+ violet, C+ violet (due to olivaceous pigment) and, usually, C+ orange-red (gyrophoric acid). The mesoconidial states of M. denigrata and M. rhabdogena are very similar, but M. denigrata has longer (4-5-7-5 wm) microconidia, and sometimes has curved macroconidia; macroconidia are apparently not produced by M. rhabdogena or the related M. elachista. Habitat and distribution: M. rhabdogena appears to be a rare or much overlooked, exclusively lignicolous species, known so far only from north Norway, mid-Sweden and Karelia. All collections are on rather hard conifer lignum, and associated species include Calicium tra- binellum, Cetraria pinastri, Mycocalicium subtile, Parmeliopsis ambigua, P. hyperopta, Xylo- grapha abietina, and X. vitiligo. Exsiccata: Malme Lich. Suec. 20 (M, S). 37. Micarea stipitata Coppins & P. James (Fig. 29B; Map 21) in Lichenologist 11: 156 (1979). Type: Scotland, Argyll, Loch Creran, Glasdrum National Nature Reserve, on Betula, 27 v 1976, L. Tibell & Coppins 2357 (E — holotype!; isotypes: BM!, H!, UPS, hb Poelt!, hb Vézda!). Thallus and apothecia: more or less identical in appearance and internal anatomy to those of M. pycnidiophora (q.v.). Pycnidia always present numerous and conspicuous, borne on distinct whitish stalks (pycni- diophores) which may be simple, bifurcate or to S5-branched, mainly 400-800 ym tall and 60-100(—150) wm diam, often with small irregular clusters of + superficially encapsulated algae (small granular-areolae). Stalk tissue composed of interwoven hyphae, c. 1-1-5 wm wide, which + separate in K. Pycnidia innate in the apex or apices of pycnidiophores, + globose or doliiform, c. 60-90 xm diam. Conidiogenous cells + cylindrical, 7-11 1-4-2 wm. Conidia (mesoconidia) cylindrical or narrowly ellipsoid, eguttulate, 6-8 x 1-1-8 wm. Chemistry: All parts K—, C—, KC—, PD—; no substances detected by t.l.c. Observations: See under the related species M. pycnidiophora for the differences between them and for comparisons with other similar species. Habitat and distribution: M. stipitata is characteristic of acid, often leached, bark (or overgrowing bryophytes thereon) on trees in undisturbed woodlands in areas with a high rainfall. It occurs in communities referable (or closely akin) to the Parmelietum laevigatae on the trunks of Betula, Quercus, and, less frequently, Alnus, Pinus, Abies, and Pseudotsuga. It has once been found on mossy rocks. The British distribution of M. stipitata is a distinctly more northern and western, as well as more oceanic, than that of M. pycnidiophora (see Map 21). It has not yet been found in south-west England but presumably occurs there. Although known from the Azores and the Canary Islands it has not been found elsewhere in Europe, but should be looked for in the hyperoceanic woodlands of, for example, south-west Norway and France (Bretagne). Additional information: for additional illustrations and some further discussion see Coppins & James (1979). 38. Micarea subleprosula (Vézda) Vézda (Figs 30, 53-54; Map 25) in Vézda & V. Wirth in Folia geobot. phytotax, Praha 11: 101 (1976). — Bacidia subleprosula Vézda in Preslia 33: 366 (1961). Type: Czechoslovakia, Bohemia, Sudety, KrkonoSe, Mumlava valley near Harrachov, 900 m, over decaying mosses on granitic rocks, 1960, A. Vézda (E - isotype! [t.l.c.: alectorialic acid]). LICHEN GENUS MICAREA IN EUROPE 183 Thallus: As for M. leprosula (q.v.) but with a different chemistry (see below). Apothecia sometimes (? often) absent, immarginate, + globose and constricted at the base, sometimes irregularly aggregated or tuberculate, grey-brown, brownish black, or black and often with bluish tinge, 0-2-0-8 mm diam. Hymenium c. 65-90 ym tall, + hyaline but upper part olivaceous or dilute reddish brown, K—, HNO3+ reddish. Asci clavate, c. 60-70 19-22 wm. Spores fusiform, often slightly curved, 3—7(-9)-septate, (35—)40-50(-60) um. Paraphyses numerous, branched and often anastomosing, 1-1-5 wm wide; apices often more richly branched, not or only slightly incrassate (to 1-8 wm). Hypothecium c. 120-200 um tall, + hyaline or dull straw, but often mottled dilute fuscous-brown (K—) in the upper part; hyphae interwoven, but becoming outwardly orientated towards the hymenium and excipulum, c. 0-8—1-5 wm; ascogenous hyphae with swollen cells, c. 2-6 wm wide. Excipulum distinct in young apothecia, but reflexed and obscured in old apothecia, hyaline or dilute fuscous-brown in part; hyphae radiating, branched and anastomosing, c. 1-1-5 wm wide. Pycnidia not found. Chemistry: Thallus K—, C+ red, PD+ yellow; sections of apothecia C—; t.l.c.: alectorialic acid plus two unidentified accessory substances. Observations: Micarea subleprosula is closely allied to M. leprosula and sterile thalli of the two species can only be distinguished by their reactions with PD, or by t.l.c. See under M. leprosula for further discussion. For additional illustrations of M. subleprosula see Vézda (1961). Habitat and distribution: M. subleprosula grows over mosses on rocks in much the same habitats in which M. leprosula is mostly commonly found. It was originally described from the Sudety mountains of Czechoslovakia, where it occurred at an altitude of 900 m, and I have seen only two additional specimens: from Sweden (Varmland) at 200 m; and Wales (Snowdonia) at c. 760 m. It may have been overlooked for M. leprosula. However, I have tested numerous individual thalli personally gathered in the field (mainly in Scotland) during the last six years, and all of these were Pd+ red and referable to M. leprosula. 39. Micarea subnigrata (Nyl.) Coppins & Kilias (Figs. 31A, 50: Map 22) in Kilias in Herzogia 5: 391 (1980). — Lecidea subnigrata Nyl. in Flora, Jena 49: 370 (1866). — Lecidea denigrata* [subsp.] subnigrata (Nyl.) Crombie, Lich. Brit.: 70 (1870). — Catillaria subnigrata (Ny]l.) Herre in Proc. Wash. Acad. Sci. 12: 94 (1910). Type: Wales, Merioneth, Cader Idris, 1866, W. A. Leighton (H-NYL 19136 — lectotype!, sel. Kilias (loc. cit.); BM ex K — isolectotype!). Lecidea confusula Nyl. in Flora, Jena 55: 360 (1872). — Micarea confusula (Nyl.) Hedl. in Bih. K. svenska VetenskAkad. Handl. III, 18 (3): 76, 86 (1892). Type: Scotland, East Perthshire, Blair Atholl, Craig Tulloch, on mica-schist, 1871, Crombie (H-NYL 20191 — lectotype!; BM — isolectotype!). Thallus effuse, widespreading or as small patches amongst other lichens, of scattered to confluent or clustered, irregularly convex to subglobose areolae. Areolae 0-08—-0-45 mm diam (usually at their largest when adjoining apothecia), dark grey-brown, matt or slightly glossy; in section with a hyaline amorphous covering layer c. 3-10 wm thick, and outermost hyphae often pale brown (K—); a white medulla often developed in large areolae. Phycobiont micareoid, cells c. 4-7 wm diam. Apothecia usually numerous, scattered to crowded and confluent, immarginate, at first adnate and slightly convex, soon becoming convex-hemispherical and occasionally tuberculate, dark brown (reddish brown when wet) to brownish black, matt or slightly glossy (especially when young), 0-2—0-6 mm diam, or up to c. 1 mm diam when tuberculate. Hymenium 35-38 um tall, hyaline but upper c. 10-12 um (epithecium) fuscous brown, K—, HNO3-—. Asci clavate 33-35 10-14 um. Spores ellipsoid (0—)1-septate, 8-12x4-5 wm. Paraphyses numerous, branched and anastomosing, (1—)1-3-1-7 wm wide, sometimes, gradually widening to 2-5 wm towards apices; hyaline throughout (epithecial pigment confined to gel-matrix). Hypothecium 184 BRIAN JOHN COPPINS 45-160 um tall, hyaline; hyphae interwoven, some becoming vertically orientated towards the hymenium, c. 1-2—1-8 wm wide, intermixed with + inflated, short-celled ascogenous hyphae c. 2-5-5 wm wide. Excipulum well developed, 25-45 ym wide, dilute fuscous brown; hyphae radiating, branched and anastomosing, c. 1-3—1-7 wm wide. Pycnidia usually present, immersed within areolae with ostioles visible (50) as minute pores surrounded by a thin, often slightly raised, dark brown rim; in section with a simple circular or flask-shaped cavity, walls hyaline, or pale brown around the ostiole; of two types: (a) c. 80-200 jm diam; conidiogenous cells ampulliform to subcylindrical, some with 1-2 percurrent proli- ferations, 4-10 1-5—2-5 u~m, sometimes with swollen base to 4 wm wide; conida (macroconidia) helicoid with 2-3 spirals, often appearing 3—5-septate, overall size 7-105—6 um, filaments 1-8-2 um wide (b) c. 40-100 wm diam; conidiogenous cells + cylindrical or ampulliform, 4-10 1-1-4 wm, sometimes with swollen base to 3 wm wide; conidia (microconidia) cylindrical, 4-6-6 x0-8-1 wm. Chemistry: Thallus K—, C—, PD—; sections of apothecia C—; t.l.c.: no substances. Observations: Micarea subnigrata is characterized by its dark grey-brown, verrucose-areolate thallus, dark brown apothecia, fuscous brown epithecium and excipulum, colourless hypothe- cium, and (0-1-septate, ellipsoid spores. However, its most outstanding and characteristic feature is the helicoid shape of its macroconidia (Fig. 50A); such conidia are unique in the genus and apparently so in all pycnidial fungi, whether lichenized or not. In general appearance, habitat and shape and septation of spores M. subnigrata could be confused with M. intrusa, but g ‘ Oo 1 2 3 4 0 ot Te e | * | af ce cee \ aN ee > Ss 0 Fe 1 2 J 6 e Map 22 Micarea intrusa & 1950 onwards A Before 1950 + Micarea subnigrata @ 1950 onwards O Before 1950 LICHEN GENUS MICAREA IN EUROPE 185 that species has a green epithecium and a large-celled, non-micareoid phycobiont. The little known M. curvata may be closely related to M. subnigrata, but can be distinguished by the fabiform or strongly curved spores and C+ red reaction of apothecial sections. With its external and internal fuscous brown colourations M. subnigrata could be mistaken for a Fuscidea, but species of that genus have broader (c. 1-7-2:7 wm) mostly unbranched paraphyses (the apical cell of each being clavate or capitate and often provided with a dark brown ‘apical cap’), asci with a thick, strongly amyloid apical wall, and probably referable to the Teloschistes-type (Honegger, 1978), and a protococcoid phycobiont, with thick-walled cells, c. 10-14 wm diam. For additional illustrations of M. subnigrata see Kilias (1981: 391, 445). Habitat and distribution: M. subnigrata is confined to hard siliceous rocks in rather exposed situations. The communities in which it occurs are attributable to the Lecideion tumidae alliance, especially the Lecideetum lithophilae association. The collection (Coppins 8417) from Glentrool in Kirkcudbrightshire was made from the south-facing side of a doleritic boulder ona south-facing slope, at an altitude of 135 m; association species were: Cladonia subcervicornis, Huilia albocaerulescens, H. tuberculosa, Lecanora badia, Lecidea pycnocarpa, Lepraria neglec- ta, Rhizocarpon obscuratum, R. oederi, Stereocaulon vesuvianum, Trapelia involuta, T. aff. obtegens, and Andraea sp. From other British collections the following can be added: Candela- riella vitellina, Catillaria chalybeia, Lecanora intricata, L. polytropa, Lecidea fuliginosa, L. lactea, Lecidella scabra, and Parmelia verruculifera. For information on associated species in its Norwegian locality see M. intrusa (Table 5). M. subnigrata is known from scattered localities in upland districts in the Scottish highlands, Galloway, and Wales; recent field studies indicate that it may be far more common in these and similar areas (e.g. Lake District and Dartmoor) than present records (Map 22) suggest. Outside Britain it is known only from coastal Norway (Hordaland) and south-west Sweden (Halland). 40. Micarea subviolascens (Magnusson) Coppins, comb. nov. (Fig. 31B) Lecidea subviolascens Magnusson in Blyttia 7: 30 (1949). Type: Norway, Hordaland, Granvin, Stein- saethorgen, on easily weathered schist in an open windy place of a small hill, 780 m, ix 1944, J. J. Havaas, Lich. Exs. Norv. 694 (BG — lectotype!). Topotype material collected in 1949 distributed in Havaas, Lich. Exs. Norv. 710 (BG!) and Lich. Norv. Occid. 269 (BG!). Lecidea assimilata f. aberrans Th. Fr., Lich. Scand. 2: 523 (1874). Type: Norway, Troms, Tromsg¢, Figjfjellet, 1868, Th. M. Fries (UPS — holotype!). Lecidea assimilata var. hardangeriana Vainio in Acta Soc. Fauna. Fl. fenn. 57 (2): 374 (1934). — Lecidea assimilata var. hardangeriana Vainio in Havaas in Bergens Mus. Arb. 1909 (1): 29 (1909); nom. nudum (Art. 32). Type: Norway, Hordaland, Granvin, Sm¢reggen, 650 m, 22 viii 1902, J. J. Havaas, Lich. Exs. Norv. 139 (BG -— lectotype! [t.1.c.: no substances]; isolectotypes: BG!, H!). Thallus effuse, saxicolous, composed of + confluent, white to pale brown, convex, verrucose- areolae c. 0-1-0-5(-0-8) mm diam; with age the crust may thicken (to c. 1 mm) and become cracked and divided into ‘islands’ c. 1-2 wm wide. Areolae in section, sometimes with a hyaline amorphous covering layer c. 5-7 wm thick, upper c. 20 wm of areolae often tinged dilute olivaceous and Kf+ violet (pigment confined to the weak gel-matrix). Thallus hyphae 1-7—2-5 (-3) wm wide. Phycobiont micareoid, cells 4-7 wm diam. Cephalodia (?): spaces between areolae often filled by black, loose clusters of Stigonema. Apothecia numerous, immarginate, adnate, convex to subglobose, black, matt, 0-3—-0-6(-0-8) mm diam, often confluent, or forming tuberculate clusters up to 1:2 wm diam. Hymenium 40-55 wm tall, dark green (K+ violet, HNO3+ purple-red) above, and below in vertical streaks, otherwise dilute greenish or + hyaline. Asci clavate, 38-48x10-15 wm. Spores ellipsoid, ovoid-ellipsoid or oblong-ellipsoid, simple, 9-16(-17) x 4-5 wm. Paraphyses numerous, simple below, but in upper part often forked or with short lateral branches, 1—-1-8(-2) um wide, sometimes widening above to 3 um; apical walls hyaline although surrounded by densely pigmented gel-matrix. Hypothecium 150-300 um tall, dark purple-brown, K+ purple intensify- ing, or upperpart K+ dark green; all parts HNO3+ purple-red; hyphae interwoven, but + 186 BRIAN JOHN COPPINS vertically arranged in upper part, 1-5—2-5 wm wide, embedded in densely pigmented gel-matrix; ascogenous hyphae c. 2-5-5 wm wide. Excipulum + distinct in young apothecia, but soon reflexed, dark purple-brown (K+ dark green) within, changing to green (K+ green intensifying) towards the outer edge; hyphae radiating, branched, c. 1-5—2 um wide. Pycnidia: Not found. Chemistry: Thallus K—, C—, KC—, PD-; t.1.c.: no substances. Observations: In his protologue, Magnusson allied his new species with the Lecidea [Micarea] sylvicola group. However, Th. M. Fries, 75 years earlier, was probably nearer the truth when he described material of M. subviolascens as a form (‘aberrans’) of Lecidea [Micarea] assimilata. In my opinion M. subviolascens is very closely related to M. assimilata, and the reader is referred to the account of that species for further discussions. Habitat and distribution: M. subviolascens appears to have a very restricted distribution, being known only from the provinces of Troms and Hordaland in Norway. It is the only known saxicolous member of the M. assimilata group, and occurs on acidic rocks (schists and gneiss) in rather open situations. In the Hordaland localities it was collected at altitudes of 650 m and 795 m. The labels accompanying the specimens provide little extra ecological information, and the specimens have few associated species, although Pyrenopsis pulvinata and Rhizocarpon obscur- atum have been noted. Exsiccata: Havaas Lich. Exs. Norv. 139 (BG, H), 694 (BG), 710 (BG). Havaas Lich. Norv. Occid. 269 (BG). 41. Micarea sylvicola (Flotow) Vézda & V. Wirth (Figs 31C, 51A; Map 23) in Folia geobot. phytotax, Praha 11: 99 (1976). — Lecidea sylvicola Flotow, Lich. Schles. 171 (1829). Type: Poland, Schlesien, Flotow, Lich. Exs. 171A (UPS -— lectotype!, sel. Hertel (1975: 74); WRSL - isolectotype!). Lecidea aggerata Mudd, Man. Br. Lich.: 208 (1861). Type: England, Yorkshire, Battersby, Mudd, Lich. Brit. 175 (BM - lectotype!; isolectotypes: E!, H-NYL 14016!, M!, MANCH!). Lecidea incincta Nyl., Lich. Scand.: 231 (1861). Type: Finland, Satakunta, KAllfjard [Ahlainen: Kellahti], 1859, A. J. Malmgren (H — holotype!). Biatora smaragdina Arnold in Verh. zool. bot. Ges. Wien 19: 613 (1869). Type: Italy, Trentino-Alto Adige, ‘Melaphyr in Walde unterhalb Bad Razzes am Schlern [Monte Pez]’, vii 1867, Arnold (M — holotype!). Lecidea hellbomii Lahm in Flora, Jena 53: 177 (1870). Type: Sweden, Narke, Vredstorp, Urby, on granitic rock, 1869, P. J. Hellbom (M - lectotype!). Lecidea sylvicola f. sublivida Vainio in Medd. Soc. Fauna Fl. fenn. 10: 104 (1883). — Lecidea sylvicola var. sublivida (Vainio) Vainio in Acta Soc. Fauna FI. fenn. 57 (2): 298 (1934). Type: Finland, Ostrobottnia kajanensis, Kuhmo,Kylmala, 1877, E. A. Vainio (TUR-VAINIO 25226 — holotype!). Lecidea hypocyanea Vainio in Acta Soc. Fauna Fl.fenn. 57 (2): 300 (1934), non Stirton (1879). — Lecidea vainioi Magnusson in Blyttia 7: 31 (1949); nom. nov. Type: Finland, Regio aboénsis, Turku, Hirvensalo, 10 vii 1924, E. A. Vainio (TUR-VAINIO 33172 — lectotype!, sel. Hertel (1975: 74)). Lecidea sylvicola var. flotowii Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 297 (1934); nom. inval. (Art. 26). Thallus effuse, thin and + smooth or weakly rimose to rather thick, uneven and coarsely rimose, or sometimes with irregularly granular or verrucose areolae up to 0-3 mm diam, pale buff, pale to dark grey or bluish grey; in section without a cortex or hyaline covering layer, but walls of outermost hyphae sometimes greenish, K—, HNO3+ purple-red. Phycobiont not micareoid; cells thin-walled, + globose and c. S—12 wm diam, or ellipsoid and up to 15 10 wm. Apothecia numerous, convex-hemispherical from the start, often becoming + globose or tuberculate, black and often with bluish tinge, sometimes dark blue-grey (deep shade forms), 0-2-0-5 diam, or to 1:2 mm when tuberculate. Hymenium 40-60(—70) um tall, dilute bright or sordid aeruginose, but often darkish aeruginose in upper and lower parts, and in vertical streaks (due to presence of stout pigmented paraphyses), K— or + aeruginose intensifying, HNO3+ purple-red. Asci cylindrical-clavate, c. 35-45 8-12 wm. Spores ellipsoid or ovoid, simple (an occasional 1-septate spore seen in a few collections), (6-)7—10 x (2-5—)3—4-5 ym. Paraphyses LICHEN GENUS MICAREA IN EUROPE 187 rather scanty, of two types: p.p. evenly distributed, irregularly flexuose, simple or sparingly branched, often anastomosing, hyaline, thin, 0-7-1 wm wide, sometimes widening above to 1-8 um; p.p. fewer in number, as scattered individuals or in small fascicles, straight, simple or occasionally forked above, stout, coated + throughout (but most intensely so around the apices) by dark green pigment and appearing c. 2-3 wm wide, apices sometimes incrassate and up to 4 um wide. Hypothecium 90-200 um tall, dark aeruginose or olive-black, K— or + green intensifying; lower (‘core’) part, or rarely the entire hypothecium, dark purplish brown, K+ purple intensifying; all pigments HNO3+ purple-red; hyphae coated with dense dark green pigment and 2-3 wm wide (overall), interwoven but becoming vertically orientated towards the hymenium and some continuing into it as stout, pigmented paraphyses; ascogenous hyphae similarly pigmented, with swollen cells c. 3-5 xm wide. Excipulum not evident, even in sections of young apothecia. Pycnidia usually present and numerous, immersed, black, 40-200 um diam; in section with a single circular or ovate locule but often becoming internally convoluted with up to 6 locules, wall dark green K—, HNO3+ purple-red in upper part, turning paler below and dilute brownish or + hyaline at the base; conidiogenous cells irregularly subcylindrical, 5-10 x 1-2-1-5 wm, often with one or two percurrent proliferations; conidia (mesoconidia) + cylindrical or sometimes oblong- obovoid, sometimes biguttulate and often + constricted in the middle, 3-8-6(-6-6) x 1-1-7(-2) pm. Chemistry: Thallus K—, C—, KC—, PD-—; sections of apothecia C—; t.l.c.: no substances. Observations: Micarea sylvicola is characterized by its convex-globose, often tuberculate, black apothecia, green or blue-green hymenium, very dark green-black or brown-black hypothecium (appearing green, or rarely purplish, in K), and ellipsoid to ovoid, simple spores. The variation in hypothecial pigmentation is similar to that found in such species as M. assimilata, M. crassipes, and M. melaena. M. tuberculata is rather similar to M. sylvicola but has generally smaller apothecia, a more shallow hymenium, narrower, oblong-ovoid spores that are often 1-septate, and shorter conidia. Forms of M. bauschiana with greenish pigment in the hypothecium have been confused with M. sylvicola, however the pigment is always dilute and confined to the gel-matrix such that the hyphae appear hyaline in K (at X 1000). The thallus of M. sylvicola is often provided with a blue-grey tinge due to green-pigmented hyphae near the surface; such coloration has not been noticed with the thalli of M. bauschiana and M. tuberculata. Another species often confused with M. sylvicola is Lecidea erratica Kérber, which can be distinguished by its thinly marginate young apothecia, and distinct excipulum when observed in section. Habitat and distribution: M. sylvicola is mainly found on dry shaded rocks in the communities belonging to Micareetum sylvicolae, but it sometimes occurs in periodically wetter situations on rock faces, upper sides of boulders, and loose stones in woodlands where it may be associated with such species as Baeomyces rufus, Cystocoleus ebeneus, Fuscidea recensa, Huilia tuberculo- sa, Parmelia saxatilis, Rhizocarpon hochstetteri, R. obscuratum, R. oederi, Scoliciosporum umbrinum, Trapelia involuta, and in one instance (Coed Hafod, Denbigh) Bacidia vezdae. In addition, it is sometimes found on old fence posts in upland districts. In the British Isles M. sylvicola is more confined to upland districts and is less common than M. bauschiana. It seems to be particularly prevalent in Wales and south-east Scotland, and curiously rare in western Scotland and Ireland. Further field-work is required to establish whether or nor its apparent scarcity in these latter areas is real or merely an artefact resulting from uneven recording. However, my field observations in western Scotland suggest that its niche in the Micareetum sylvicolae is there occupied by M. lutulata. M. sylvicola is widely distributed in Scandinavia, but rarely found from north of about latitude 67°N. Elsewhere in Europe I have seen specimens from Germany (Hessen and Baden-Wirttemberg), eastern Austria, northern Italy, south-west Poland and the mountain regions of Czechoslovakia. From outside Europe I can confirm its presence in north-eastern North America (New York and Newfoundland). 188 BRIAN JOHN COPPINS le erp | ig Fag? 4 |. ars lech ye t ee Bird di a LaPeer I ) PA 7 a E + gat | te hor Po ) E Sade | 4 1 (CHANNEL (SLANOS o Ce aon \ PLOTTED ON UTM GRID a’, ns fe) i?) 4 ‘ / Map 25 Micarea subleprosula © 1950 onwards + Micarea cf. ternaria @ 1950 onwards O Before 1950 192 BRIAN JOHN COPPINS Lecidea milliaria f. ternaria is very fragmentary and my first impression was that it was a diminutive specimen of M. lignaria, even though I failed to obtain a positive reaction with PD. This opinion was changed by my subsequent examination of the Alaskan material which is identical to the holotype in all respects but larger and in better condition. Several problematical collections from the British Isles (Shetland Islands and western Ireland) are provisionally referred to M. ternaria. They were all collected in coastal districts on hard siliceous rocks and have a + obsolete thallus (PD— in microscopical preparations), rather flattened apothecia, and 0-3-septate spores. The specimen from Fair Isle has pycnidia, containing mesoconidia 5-2-6-8 x 1-2-1-8 wm (Fig. 52C). The discovery of an arctic species, such as M. ternaria, in coastal Britain is not without precedent. Examples are Lecanora straminea found in the Shetland Islands, Flannan Islands, and St Kilda, and Bacidia subfuscula found on North Rona, the Farne Islands, and as far south as the north coast of Norfolk and the Scilly Isles. A better understanding of the variability in habit and habitat of M. ternaria in the arctic should help to clarify the taxonomic relationships between the arctic and British populations. In addition, relevant material should be sought in coastal Norway. Habitat and distribution: On mosses and plant debris on the ground in coastal regions of arctic Europe and North America (northern Alaska). The Alaskan material (Thomson 9188) was growing with Micarea turfosa and Siphula ceratites. Southern populations on rocks may be represented in the British Isles (see ‘observations’ above). 44. Micarea tuberculata (Sommerf.) R. Anderson (Figs 32A, 34, 51B; Map 26) in Bryologist 77: 46 (1974). — Lecidea tuberculata Sommerf., Suppl. Fl. Lapp.: 160 (1826). Type: Norway, Nordland, Saltdalen, FiskevaagmGllen, iii 1822, S. C. Sommerfelt (O — lectotype!; UPS — isolectotype!). Lecidea latens Taylor in Mackay, Fl. Hib. 2: 259 (1836). Type: Ireland, Wicklow, The Dargle, T. Taylor (BM - lectotype!). Lecidea botryocarpa Nyl. in Flora, Jena 48: 603 (1865). Type: USSR, Karelskaya ASSR, Karelia onegensis, ’ad Onegam, Kapselka, 1863, 7. Simming (H-NYL 10766 — holotype!). Lecidea subinfidula Ny]. in Flora, Jena 52: 295 (1869). Type: Finland, Lapponia enontekiensis, Naimakka, 29 viii 1867, J. P. Norrlin 656 (H — lectotype!; UPS — isolectotype!). Lecidea tuberculata var. scandinavica Vainio in Acta Soc. Fauna Fl. fenn. 57 (2): 309 (1934). Type: Finland, Tavastia australis, Lammi, Evo, Lapinkallio, 1866, Norrlin (H-NYL 10767 — lectotype!). Thallus effuse, minutely scurfy-granular, or sometimes forming an irregularly rimose crust up to 0-2 mm thick, more rarely forming irregularly verrucose areolae c. 0-06—0-2(-0-3) mm diam; greenish buff, pale buff or greenish white; thin wefts of white prothalline hyphae sometimes visible. Phycobiont not micareoid; cells + globose, thin-walled, 5—10(-12) wm diam, or irregularly ellipsoid and up to 15X7 wm. Apothecia numerous, convex-hemispherical and immarginate from the start, often becoming + globose or tuberculate, black and often with bluish tinge, matt, 0-16-0-3(—0-4) mm diam, or 0:24-0:55 mm diam when tuberculate. Hymenium 30-35 pm tall, dilute green or aeruginose, K-— or + intensifying, HNO3+ purple-red, often with darker vertical streaks due to the presence of stout, pigmented paraphyses. Asci cylindrical-clavate, 25-30 7-9 um. Spores oblong-ovoid or oblong-ellipsoid, 0-1-septate, 5-5—8(—9) x 1-5—2-5 wm. Paraphyses rather scanty, of two types (Fig. 34): p.p. evenly distributed, irregularly flexuose, simple or sparingly branched, often anastomosing, thin, 1-1-5 wm wide, sometimes widening to 1-7 wm above, walls hyaline throughout and without adhering pigment; p.p. fewer in number, as scattered individuals or in small fascicles, straight, simple or occasionally forked above, stout, coated + throughout by dark greenish pigment and appearing c. 2-3 wm wide, apices sometimes + incrassate and up to 4 yum wide (including pigment). Hypothecium c. 80-115 wm tall, aeruginose- or olive-black, K+ aeruginose intensifying; hyphae coated with dense dark green pigment and 2-3 wm wide (overall), interwoven but becoming vertically orientated towards the hymenium and some LICHEN GENUS MICAREA IN EUROPE 193 continuing into it as stout, pigmented paraphyses; ascogenous hyphae similarly pigmented, with swollen cells c. 3-5 wm wide. Excipulum not evident, even in sections of young apothecia. Pycnidia often present, + immersed, black, 60-120 xm diam, ostioles often gaping; walls dark aeruginose, K—, HNO3+ red; conidiogenous cells slender, cylindrical, 5-10 1-1-5 wm, often with a swollen, sometimes pigmented, base up to 3-2 wm wide, and sometimes with one or two percurrent proliferations; conidia (mesoconidia) + cylindrical, sometimes faintly biguttulate, 3-4-3 1-1-4 um. Chemistry: Thallus K—, C—, KC—, PD~-; sections of apothecia C—; t.l.c.: no substances. Observations: Micarea tuberculata is characterized by its rather small, markedly convex, often tuberculate, blackish apothecia, blue-green hymenium, aeruginose-black hypothecium and small, ovoid-oblong, 0—1-septate spores. The relative proportion of simple to 1-septate spores can vary greatly between collections, and in some specimens no septate spores can be found. M. tuberculata has often been confused with forms of M. sylvicola with small, immature spores, but such spores are always ellipsoid or ovoid-ellipsoid, and quite distinct from the predominantly oblong-ovoid spores of M. tuberculata. If pycnidia are present the two species can be separated on the size of their conidia, which are significantly longer (c. 4-6 wm) in M. sylvicola. M. olivacea (q.v.) is superficially similar to M. tuberculata, but has slightly larger spores, shorter conidio- genous cells, and a micareoid phycobiont. Also of similar appearance is Psilolechia clavulifera (p. 201), but it has smaller spores, a pale hypothecium, numerous, stout, non-pigmented paraphyses, and a very different phycobiont. ey ie} 1 2 3 4 qi r a 76 “Ne [Sa de Sy | ee Se zo _ | Dee sae —— ERS, \ mete: os ig _ ts 4 =e a 1 | le C ~~ 3 i eres 3 Q = 2 Ps 2 ie : ioe ee aaa i SB 1 2 4 6 Map 26 Micarea tuberculata @ 1950 onwards O Before 1950 194 BRIAN JOHN COPPINS Habitat and distribution: M. tuberculata is found on rocks, stones, and tree roots, etc. in dry, sheltered underhangs, and is a faithful member of the Micareetum sylvicolae. In the British Isles it has scattered localities in the north and west; although not yet known from the mainland in England and Wales it is expected to occur in those areas in suitable terrain (e.g. the Lake District, Dartmoor and Snowdonia). It is undoubtedly much overlooked, but appears to be genuinely rarer than M. bauschiana and M. sylvicola, with which it often occurs. It appears to be widely distributed in Fennoscandia where it is known from as far north as Nordland in Norway and Lycksele Lappmark in Sweden. It seems to be rare elsewhere in Europe, but I have seen material from southern Germany, northern Italy, and the Tatra mountains of Czechoslovakia. Exsiccata: Arnold Lich. Exs. 1057 (BM ex K, M). Larb. Lich. Herb. 227 p.p. (BM). Rabenh. Lich. Eur. 648 p.p. (BM). Rasanen Lich. Fenn. 512 (BM, BM ex K), 672 p.p. (BM, LD-mixed with M. Jutulata, M-mixed with M. sylvicola). 45. Micarea turfosa (Massal.) Du Rietz (Fig. 33; Map 27) in Svensk bot. Tidskr. 17: 94 (1923). — Biatora turfosa Massal., Ric. Lich. Crost.: 128 (1852). Type: specimen unlocalized but probably from Sudety (Sudeten Mountains) in SW Poland, ‘Flotow Lich. exs. 130! hb. Flotow’ (VER - holotype!). Biatora turfosa* verrucula Norman in K. nor. Vidensk. Selsk. Skr. 5: 353 (1868). — Lecidea verrucula (Norman) Th. Fr., Lich. Scand. 2: 523 (1874). — Micarea verrucula (Norman) Hedl. in Bih. K. svenska VetenskAkad. Hand. III, 18 (3): 84, 95 (1892). Type: Norway, Nordland, ‘supra Heminghyt convallis Bejeren’, J. M. Norman (- lectotype!; isolectotypes: LD!, M!, O!). Thallus effuse but often forming rounded patches about 3-5 cm diam, thin, + uneven but not forming distinct areolae or goniocyst-like granules, blackish grey or brown-black but shaded portions sometimes dull grey-green, often appearing + gelatinous when moist. Thallus in section up to 70 wm thick, ecorticate and without an amorphous covering layer; outermost hyphae 2-4 wm wide with walls thickened by dark green pigment, K—, HNO3+ red; internal hyphae hyaline, c. 1-1-5 wm wide. Phycobiont micareoid, cells 4-7 wm diam. Apothecia numerous and often confluent, immarginate, convex to + globose, black, matt or slightly glossy, rarely brown (shade forms), 0-15—0-3(-0-4) mm diam. Hymenium 35-50 um tall; upper part (epithecium) sordid green, K—, HNO3+ red, or sometimes brownish in part; middle part dilute aeruginose or olivaceous; lower part dilute reddish brown and merging + impercep- tibly into the hypothecium. Asci clavate, 35-50 x 11-12 wm. Spores oblong-ellipsoid to fusiform, sometimes slightly curved, simple or 1—3-septate, (10—)12—21(-25) x (3-5—)4-5 wm. Paraphyses numerous, branched and sometimes anastomosing, c. 1-1-5 ~m wide in mid-hymenium but upper 5—15(—25) um thickened with dark greenish pigment and then 1-5-2-5(-3) wm wide. Hypothecium c. 70-140 pm tall, mottled reddish brown, K—, HNO3-— or turning orange-brown (never with purple tinge); hyphae interwoven but becoming vertically orientated towards the hymenium, hyaline or sometimes loosely coated with brown pigment, c. 1-5-1-7 wm wide; ascogenous hyphae with swollen cells up to 5 wm wide. Excipulum reflexed but distinct, reddish brown but darker than the hypothecium; hyphae radiating, branched and anastomosing, 1-1-5(—2) wm diam, hyaline (pigment confined to gel-matrix). Pycnidia often present but very inconspicuous, immersed, 35-40 zm diam, wall dark sordid green, K—, HNO3+ red. Conidia (microconidia) + cylindrical, c. 3-54-71 wm. Chemistry: Sections of thallus and apothecia K— , C— , PD—; no substances detected by t.l.c. Observations: M. turfosa is fairly constant in appearance, except that the thallus is less well developed when in boggy habitats. Microscopically there is much variation between collections with regard to spore septation: some having mostly simple spores (e.g. Vézda Lich. Sel. 538), others having numerous 1-septate and several 2- or 3-septate spores (e.g. Vézda Lich. Sel. 1135). In the type of Biatora turfosa the spores are mostly simple but a few with a single septum were found. In the type collections of Biatora turfosa* verrucula most spores are 1-septate. LICHEN GENUS MICAREA IN EUROPE 195 With the combination of blackish thallus and black, convex to + globose apothecia, M. turfosa is easily confused in the field with peat inhabiting forms of M. melaena, but the latter differs microscopically in having dark purple and/or green pigments in the hypothecium and thinner paraphyses. M. botryoides has similar apothecial pigmentation to M. turfosa, but its hypothecium is darker with nearly all the hyphae densely coated with pigment; in addition, M. botryoides has less numerous and (when hyaline) thinner paraphyses, a shallower hymenium, smaller spores, and its apothecia are always accompanied by numerous, stalked, black pycnidia. M. melaenida is unlikely to be mistaken for M. turfosa because of its whitish thallus and habitat of fine-grained, mineral soils; it can be further distinguished by its purple-brown epithecium, excipulum, and (usually) upper hypothecium. The little known M. osloensis differs from M. turfosa in several features but is most easily distinguished by its much smaller, non-septate spores. Habitat and distribution: M. turfosa seems to have an arctic-alpine distribution. In Britain it is known from summits in the Cairngorm, Grampian, and Breadalbane mountains of Scotland, where it grows on exposed turf (mostly over dead bryophytes) at altitudes of about 860-1245 m. In the Sudeten and Carpathian mountains of Czechoslovakia it is reported at altitudes of 1400 m and 1870 m respectively. It seems to be able to occur at lower altitudes in Sphagnum-bogs in Scandinavia, but no altitude data is given with any of the specimens seen. I have not seen any material from the Alps but it is likely to occur there. On the Cairngorm plateau M. turfosa occurs amongst Juncus trifidus. Associated lichens on the British gatherings include Cladonia spp., Lecidea caesioatra, and Lepraria neglecta. ~ ( Oo 1 2 3 4 0 wr 0 oa 14-38 Da ¢ : 0” se 4 e 4 Seal ae { 8 Sya? Pee alee ae & Kereta ws G7 L ) Ose Miles 3 bs e aaa ? Sy By, ye . i } > ¥ el De S.zan ae at a4 1 2 4 Map 27 Micarea turfosa @ 1950 onwards O Before 1950 196 BRIAN JOHN COPPINS Outside Europe M. turfosa is little known, but I have seen material from Greenland and northern Alaska. Exsiccata: Flotow Lich. 130 (VER). K6rber Lich. Sel. 12 (L). Magnusson Lich. Sel. Scand. 282 (GZU). Malme Lich. Suec. 865 (WIS). Norrlin & Nyl. Herb. Lich. Fenn. 321 (BM, M). Vézda Lich. Sel. 538 (BM), 1135 (BM). Excluded taxa Bacidia beckhausii Kérber, Parerga lich.: 134 (1860). — Micarea beckhausii (K6rber) Vézda in Poelt & Vézda, Bestimmungsschl. europ. Flechten, Erganzungsheft I: 162 (1977). Type: Germany, ‘West- phalen’, Beckhaus (L 910, 137 1363 — lectotype!). Bacidia miniuscula Anzi, Cat. lich. Sondr.: 70 (1860). — Micarea miniuscula (Anzi) Vézda, in Vézda & V. Wirth in Folia geobot. phytotax., Praha 11: 100 (1976). Type: not seen. Bacidia beckhausii must be excluded from Micarea, mainly on account of its excipulum structure (see p. 189 and 198). Bacidia nitschkeana var. perpusilloides Erichsen in Annals mycol. 41: 206 (1943). Type: West Germany, Schleswig-Holstein: Flensburg, Forst Clusries, near Wasserleben, on Picea twigs, 61923, C. F. E. Erichsen (HBG - lectotype!). Paratypes: Schleswig-Holstein: Flensburg, Jerrish6er Holz, 10 vi 1928, C. F. E. Erichsen (HBG!); Hamburg, Wohldorfer Wald, 12 vi 1905, C. F. E. Erichsen (HBG!); Hamburg, Volksdorfer Wald, 19 xii 1909, C. F. E. Erichsen (HBG!). All on young twigs of Picea. Thallus thin, green-grey. Apothecia minute, + pellucid, pale brown, immarginate, c. 0-1-0-15 mm diam. Hymenium, hyaline, c. 30-40 um tall. Hypothecium dark brown, K—, or + greenish in part, c. 40-45 wm tall. Excipulum thin, c. 12-15 wm wide, + pseudoparenchymatous with cells c. 3-5-8 2-5-3 wm (in K), hyaline or with greenish pigment in zone adjacent to hymenium. Asci clavate, numerous, 8-spored. Spores oblong-fusiform, straight or slightly curved, 3(—-5)-septate, 16-24(—28) x 3-4 wm (Fig. 56A). Pycnidia often numerous, blackish, c. 50 wm diam, wall greenish in K; conidia curved or sigmoid 12-30X0-8 wm. Phycobiont cells c. 5—10 wm diam. This variety agrees in all respects, except host substratum, with Bacidia myrtillicola Erichsen in Mitt. Inst. allg. Bot. Hamb. 10: 414 (1939); type: Schleswig-Holstein: Lauenburg, ‘im “Sachsenwald” bei Friedrichsruh an Vaccinium myrtillus am Rande des Reviers ‘‘Saupark’’.’, 2 xi 1924, C. F. E. Erichsen (HBG - holotype!). B. myrtillicola belongs to the mainly foliicolous groups of Bacidia s. lat. that includes B. fuscatula (Mill. Arg.) Zahlbr., B. rhapidophylli (Rehm) Zahlbr., and B. vezdae Coppins & P. James. Catillaria melanobola f. frullaniae B. de Lesd., Rech. Lich. Dunkerque, Suppl. 1: 119 (1914). Type: France, Nord, near Burgues, on Frullania on Ulmus, M. B. de Lesdain, Zahlbr. Lich. Rar. Exs. 152 (BM —isotype!). This is an apparently undescribed (at specific rank) species of Arthonia s. lat. The apothecia are very small, c. 0-1 mm diam, convex-globose and black. In section the epithecium and hypothecium are brownish (K+ olive), the asci clavate, c. 21X10 wm at maturity, with a minute internal amyloid ring [as described for Bryostigma leucodontis (Poelt & Débbeler, 1979) and Chrysothrix (Laundon, 1981), and noted by me in several Arthonia species, especially lichenicolous species and members of the subgenus Allarthonia], spores hyaline, 1-septate, 9-11 2-3-3 um, and paraphyses scanty, thin, c. 0-6-1 wm wide. This species is rather common in the British Isles where it is usually referred to as ‘Arthonia cf. exilis’. It is found in a variety of habitats, for example bark of shaded trunks of Acer, Fraxinus and Ulmus, branches and twigs of Corylus and Sambucus, over bryophytes (especially Frullania) on trunks of Quercus and Ulmus, tufts of Armeria on sea-cliffs, and possibly also on sandstone rocks. When on bark the apothecia may attain a larger size, to 0-2 mm diam. Corticolous material has recently been distributed as Vézda Lich. Sel. 1701. In E itis represented by material from the following British vice-counties: 1, 20, 29, 31, 62, 64, 67, 78, 83, 88, 90, 98, 99, 110, H18 and H20; and from Denmark and Germany (Schleswig-Holstein). There has been much confusion regarding the correct interpretation of Arthonia exilis (Flérke) Anzi. I have recently had the opportunity of examining type material of this species (Flérke Deutsch. Lich. 187 (WRSL - lectotype!)) and a brief description of it is as follows: Thallus lignicolous (probably on worked timber; accompanied by a few granular-areolae of Candelariel- la vitellina), thin, grey-white. Phycobiont not Trentepohlia, cells, 10-19 wm diam. Apothecia numerous, + regularly dispersed, black, 0-1-0-2 mm diam. Epithecium (including lower border with substratum) reddish brown, K—. Hymenium c. 30 um tall, I+ reddish, K/I+ blue. Asci clavate, with minute amyloid ring (‘Bryostigma — type’), 22-30 14-15 wm, 8-spored. Spores hyaline, 1-septate, 8-122-5—3-5(-4) um. Paraphyses + coherent (even in K), rather stout, c. 1-5-2 wm wide; apices + swollen with brown LICHEN GENUS MICAREA IN EUROPE 197 Fig. 56 A, spores of Bacidia myrtillicola (HBG -— lectotype of Bacidia nitschkeana var. perpusilloides). B-C, Catillaria bouteillei (H-NYL 1884 — holotype of Lecidea littorella); B, spores; C, conidiogenous cells and conidia. Scale = 10 wm. pigment-caps; closely adhering pigment sometimes continuing down to about mid-hymenium level. Hypothecium hyaline. The status of A. exilis s. str. as a British species requires confirmation. Catillaria rhodosphaera Th. Fr. & Hulting in Th. Fr. Lich. Scand. 2: 571 (1874). Type: Sweden, Dalsland, Haverud, 1870, J. Hulting (UPS — lectotype!; UPS — isolectotype!). This species was provisionally referred to Micarea by Kilias (1980: 392), but in fact belongs to Biatora Fr., aname which requires conservation against Biatora Ach. (= Stenhammarella Hertel). In my opinion C. rhodosphaera represents a saxicolous form of Catillaria sphaeroides (Dickson) Schuler. Lecidea arceutina var. hypnaea Nyl. in Flora, Jena 51: 165 (1968). — Bacidia arceutina f. hypnaea (Nyl.) A. L. Sm., Monogr. Brit. Lich. 2: 158 (1911). Type: Jersey, 1866, Larbalestier (H-NYL 17931 —holotype!). Based on examinations of supposed isotype material at BM, this was listed as a synonym of Bacidia lignaria by James (1965) and Micarea lignaria var. lignaria by Hawksworth et al. (1980). However, the holotype in H-NYL is a muscicolous form of the normally corticolous (occasionally saxicolous) Bacidia 198 BRIAN JOHN COPPINS arceutina (Ach.) Arnold. In his diagnosis Nylander gives the spore length as 45-70 wm and on the holotype packet he wrote ‘45-20 1 wm’; my examination of the holotype found the spores to be 38-51 1-5 wm. Lecidea clavulifera Ny|.—see Psilolechia clavulifera. Lecidea demarginata Nyl. in Flora, Jena 61: 245 (1878). Type: Finland, Tavastia australis, Evo, ‘supra saxum in sylva juniori, loco olim deusto’, 1873, J. P. Norrlin, Norrlin & Nyl. Herb. Lich. Fenn. 179 (H- lectotype!; isolectotypes (without exsiccate label): H-NYL 20162! & 20163!). In the protologue Nylander cited material of Norrlin’s from Finland and of Larbalestier’s from Ireland. The epithet ‘demarginata’ was first used in 1875 on the label of Herb. Lich. Fenn. 179, although without a diagnosis. An example of this exsiccate is here selected as lectotype, and is a specimen of Lecidea erratica Korber. The cited Larbalestier material is represented in BM and H and belongs to M. lutulata. Lecidea denigrata var. bacidiella Vainio in Medd. Soc. Fauna Fl. fenn. 10: 28 (1883). — Micarea bacidiella (Vainio) Vézda, in Vézda & V. Wirth in Folia geobot. phytotax., Praha 11: 100 (1976). Type: Finland, Lapponia kemensis, Sodankyla, Pyhatunturi, Kannolla, on lignum, 1878, E. A. Vainio (TUR-VAINIO 22505 — holotype!). As stated by Vainio (1934: 462) this is a synonym of Bacidia miniuscula Anzi (i.e. B. beckhausii K6rber). The following were notes made from the holotype of var. bacidiella: Thallus lignicolous, entirely endoxylic. Phycobiont cells c. 8-14 wm diam. Apothecia thinly marginate and + plane when young, soon becoming convex and immarginate, black, c. 0-2-0-3 mm diam. Hymenium c. 35 wm tall, with olivaceous vertical streaks, K+ violet, C+ violet (not red). Asci clavate, 30-35 x 10-12 ym, 8-spored. Spores rod-shaped or slightly curved, (1—)3(—7)-septate, 17—-26x1-7—2 um. Paraphyses simple or sparingly branched, 1-1-5 wm wide; apices somewhat incrassate, to 2 wm wide. Hypothecium hyaline. Excipulum dilute olivaceous (K+ violet), of radiating, branched hyphae c. 1-5-2 wm wide embedded in a dense gel; hyphae becoming more distinct in K but not separating. Pycnidia immersed in the substratum, black, c. 50-100 ~m diam; walls olivaceous, K+ violet. Conidia simple, hyaline, oblong- ellipsoid, often wider at proximal end, 2-8-3-5 x 1(-1-4) wm. B. beckhausii usually occurs on bark, although I have seen a few additional lignicolous specimens from Scotland. Micarea bacidiella sensu Vézda & Wirth (loc. cit.) is not B. beckhausii but the superficially similar Micarea globulosella (see p. 135). Lecidea erysiboides Ny}. in Not. Sallsk. Fauna Fl. fenn. 4: 232 (1859). — Catillaria erysiboides (Nyl.) Th. Fr. Lich. Scand. 2: 572 (1874). Type: Finland, Nylandia, Helsingfors [Helsinki], ‘Gumtackt’ on rotting decorticate pine trunk, 1858, W. Nylander (H — lectotype!; H-NYL p.m. 4514 —isolectotype!). This name has been misapplied by many lichenologists and the majority of specimens seen belong to Micarea prasina, or other species such as Catillaria sphaeroides, Lecania cyrtella, and Micarea denigrata. The type material of L. erysiboides is not a Micarea but a species of Catillaria s. lat., although I am uncertain of its affinities. It has small, plane to convex, reddish brown (testaceous) apothecia that are marginate when young, the excipulum is composed of much-branched, radiating hyphae which are distinct in K but still tightly bound by the gel matrix, and the spores are ovoid and often constricted at the septum, 1-septate with the upper cell usually enlarged and + globose, 8-9-5 x3—5 um. C. erysiboides has not been correctly reported from the British Isles but it should be looked for, especially in the eastern Scottish highlands. Additional specimens of C. erysiboides examined: NORWAY. Hordland: Granvin, on Betula lignum, 1904, Havaas, Lich. Exs. Norv. 292 (BG). FINLAND. Nylandia: Helsinki, 1859, W. Nylander (H-NYL 21650), and 1861 (BM). ITALY: Trentino (‘Sudtirol’): Paneveggio, on top of cut stump of Picea, 2 ix 1883, Arnold, Lich. Exs. 1002 (BM ex K). USSR. ‘Lapponia orientalis’, 1863, N. I. Fellman, Lich. Arct. 156 (BM ex K). Lecidea fuliginea Ach., Syn. Lich.: 35 (1814). — Micarea fuliginea (Ach.) Fr., Syst. orb.: 257 (1825). - Micarea fuliginea (Ach.) Fr., Stirp. agri femsion.: 37 (1825); comb. inval. (Arts 34.1, 43). I have not seen the type of this name but it is probably a synonym of Lecidea icmalea Ach. A later synonym of the latter is Pannularia perfurfurea Nyl. which is the type of the genus Placynthiella Gyelnik. Lecidea gelatinosa Flérke in Magazin Ges. naturf. Fr. Berl. 3: 201 (1809). — Micarea gelatinosa (Florke) Brodo in Bryologist 70: 216 (1967). I have not seen type material of this species, but if its general interpretation is correct then it is not a Micarea. In fact it belongs in the Lecidea granulosa group, which includes L. aeruginosa Borrer, L. aeneofusca Flérke ex Flotow, L. granulosa (Hoffm.) Ach. and L. viridescens (Schrader) Ach. These species are not congeneric with the type species of Lecidea Ach. (L. fuscoatra (L.) Ach.) and they await the formal transference to a genus in the Trapeliaceae. Further detailed studies are needed to see if they can be LICHEN GENUS MICAREA IN EUROPE 199 placed in Trapelia Choisy or Trapeliopsis Hertel & G. Schneider, or if a new genus is required to accommodate them. Lecidea littorella Nyl. in Flora, Jena 60: 229 (1877). — Catillaria littorella (Nyl.) Zahlbr., Cat. lich. univ. 4: 56 (1926). Type: Ireland, West Galway, ‘Hibernia occidentalis, Bord du Lough Inagh, mais extreme- ment rare’, 1876, C. Larbalestier (H-NYL 18884 — holotype!, isotypes: BM!, BM ex K!). The following notes were made from the holotype: Thallus saxicolous, not delimited but forming small patches amongst Hymenelia lacustris and Porina chlorotica, whitish, or dull ochraceous in part (? due to age), rimose, matt. Apothecia numerous, weakly marginate, plane to slightly convex, pallid to dull orange-red, 0-1—-0-3 mm diam; margin not exceeding the level of the disc, very thin, c. 0-1-0-3 mm diam; margin not exceeding the level of the disc, very thin, c. 0-01-0-02 mm wide, whitish with faintly pruinose appearance. Hymenium 40-47 um tall, hyaline. Asci cylindrical-clavate, ‘Lecanora-type’, 8-spored. Spores ovoid to ovoid-oblong, often ‘slipper-shaped’, 1-septate and often constricted at the septum, 9-5-14(—16) x4-6(-7) wm (Fig. 56B). Paraphyses numerous, thin, 0-8-1 wm wide, simple or sparingly branched below, becoming wider (to 1-7 ~m) and more frequently branched above. Hypothecium hyaline. Excipulum hyaline or dilute yellow-straw, inspersed with minute crystals in water mounts but + clearing in K, c. 20 um wide laterally, slightly widening below to 28 um, minutely paraplechtenchymatous with cells in the lower part measuring 4-62-54 wm. Pycnidia frequent, at first immersed, later becoming emergent, white (translucent when wet), c. 80-100 wm diam. Conidia (Fig. 56C) simple, ovoid to pyriform, 2°8X1-5-1-7 wm. L. littorella was provisionally referred to Micarea by Kilias (1981: 392). However, my subsequent examination of the holotype proved it to be a saxicolous form of the normally foliicolous Catillaria bouteillei (Desm.) Zahlbr.; previous reports of this species on rock are given by Degelius (1944) and Santesson (1952). C. bouteillei is apparently widely distributed in Ireland, especially on the leaves of Buxus (Knowles, 1929; Scannell, 1978). An additional British saxicolous specimen has been located: V.C.5, South Somerset, Broomfield, Ruborough Camp, 1914, W. Watson (BM ex K). Nylander’s choice of the epithet ‘littorella’ is rather misleading because the holotype was not collected on the sea-shore, but on the shores of an inland, freshwater lake some 10 km from the sea. British authors (e.g. James, 1970; Fletcher, 1975) have mistakenly applied the name Caztillaria littorella to a species that grows in crevices in siliceous rocks near the sea-shore; this species probably belongs to the perplexing Lecania erysibe complex. Lecidea milliaria a. [var.| terrestris Fr. Lich. Eur. 342 (1831). Type: not designated. This name is of unlikely application in Micarea. Fries cited his Lich. Suec. no. 213, the example of which in UPS is a member of the Lecidea limosa group. Lecidea ocelliformis Ny]. in Flora, Jena 48: 145 (1865). — Bilimbia ocelliformis (Ny1.) Branth. & Rostr. in Bot. Tidskr. 3: 226 (1869). — Lecidea atroviridis f. ocelliformis (Nyl.) Hedl. in Bih. K. svenska VetenskAkad Handl. III, 18 (3): 64 (1892). — Catillaria prasina f. ocelliformis (Ny1.) Erichsen in Annals. mycol. 41: 205 (1943). Type: Finland, Tavastia australis, Hollola, ad corticem Sorbi, 30 vi 1863, J. P. Norrlin 210 (H-NYL 20607! — labelled ‘Isotype’ by M. Inoue in 1979). This is not a Micarea. If the types of L. atroviridis (Arnold) Th. Fr. and L. ocelliformis are conspecific, as placed by Hedlund (op. cit.) and Vainio (1934: 218), then the latter epithet has priority. Lecidea pauxilla Krempelh. in Verh. zool.-bot. Ges. Wien 26: 455 (1876). Type: New Zealand, on bark [? Podocarpus], 18-, C. Knight (M — lectotype!; M — isolectotype!). L. pauxilla was given as a synonym of Catillaria synothea (i.e. Micarea denigrata) by Zahlbruckner (Cat. lich. univ. 4: 78, 1926). However, the type material belongs to Cliostomum griffithii (Sm.) Coppins. Lecidea recondita Erichsen in Annls mycol. 42: 25 (1944). Type: Germany, Schleswig-Holstein: ‘Apen- rade, an Blécken am Grunde einer Erosionsschluct im GehGlz Jiirgensgaard’ [pow Denmark, Jylland, near Abenral, 28 vii 1932, C. F. E. Erichsen (HBG - holotype!). In the protologue Erichsen compares this with Lecidea sylvicola (= Micarea sylvicola). My examinations of the holotype proved it not to be a Micarea but referable to Catillaria chalybeia (Borrer) Massal.; see Kilias (1980: 448). Lecidea sabuletorum f. simplicior Ny\., Lich. Scand.: 205 (1861). — Bilimbia trisepta f. simplicior (Nyl.) Vainio in Acta Soc. Fauna Fl. fenn. 53 (1): 258 (1922). Type: not designated. Nylander (loc. cit.) cited the following material ‘supra muscos ad Helsingforse [ipse] et in Sueciaé montibus [Thedenius] atque supra terram in Lapponia’. According to Fries (1874: 523) the ‘Helsingforse’ specimen is Lecidea verrucula (= Micarea turfosa); according to Vainio (1922: 140) the Swedish specimen from Funnesdalsberget in Harjedalen collected by 200 BRIAN JOHN COPPINS K. F. Thedenius is Lecidea dufourei (= Catillaria contristans (Nyl.) Zahlbr.). I am uncertain as to the identity of the specimen(s) referred to by Nylander as ‘in Lapponia’. This may be based on the specimen cited by Vainio (1922: 140, 258) as ‘supra Grimmias in Kipina Lapponiae a G. Selin collecta’, which Vainio refers to Bilimbia trisepta [sensu Vainio = Micarea peliocarpa| f. simplicior. However, material in Nylander’s herbarium (H-NYL 18841) labelled ‘Lapponia, G. Selin 1861’ is Catillaria contristans. Furthermore, another specimen of ‘Lecidea sabuletorum f. simplicior’ collected from Iceland by Isaac Carroll in 1861 (H-NYL 18862) is also C. contristans. It would appear that Nylander’s concept of ‘simplicior’ is best fitted by C. contristans and should be typified on the material collected by Thedenius or Selin. Lichen niger Hudson, Fl. angl., ed. 2, 2: 524 (1778). — Collema nigrum (Hudson) Hoffm., Deutschl. Fl.: 103 (1796). — Micarea nigra (Hudson) Fr., Syst. orb.: 257 (1825). — Micarea nigra (Hudson) Fr., Stirp. agri femsion.: 37 (1825); comb. inval. (Arts 34.1, 43). — Placynthium nigrum (Hudson) Gray, Nat. Arr. Br. Pi1>395;(1821). This is the type species of the genus Placynthium Gray. Lichen viridescens Schrader, in Gmelin Syst. nat. 2 (2): 1361 (1792). — Lecidea viridescens (Schrader) Ach. Meth. Lich.: 62 (1803). — Micarea viridescens (Schrader) Brodo in Bryologist 70: 216 (1967). I have not seen the type of this name but if its normal interpretation is correct then the species belongs to the Lecidea granulosa group; see notes above under Lecidea gelatinosa. Micarea chrysophthalma P. James in Lichenologist 5: 131 (1971). — Chrysothrix chrysophthalma (P. James) P. James & Laundon, in Laundon in Lichenologist 13: 104 (1981). This is a species of Chrysothrix and is discussed in detail by Laundon (1981). Micarea coccinea Fr., Syst. orb.: 257 (1825); nom. nudum (Art. 32.1). I do not know the identity of this species, and I have not seen this name in any other publication. On morphological grounds it is unlikely that Fries intended a new combination based on Lichen coccineus Dickson [= Haematomma ochroleucum (Necker) Laundon] or Lecidea coccinea Schw. [= Lecidea russula Ach. ]. Micarea cyanescens Poelt & Dobbeler in Bot. Jb 96: 339 (1975). Holotype: Germany, Bayern, Altbayer- ische Alpen, Chiemgauer Berge, 900-1000 m on Campylium halleri on north-facing calcareous rocks, 29 ix 1974, J. Poelt (GZU, not seen). Paratypes: Austria, Tirol, Lechtaler Alpen, ‘unteres Medriol-Tal uber Zams bei Landeck, an Dolomitblécken unter Griinerlen’, 1600-1700 m, on Campylium halleri, 1969, J. Poelt (hb Poelt 13295!); Austria, Steiermark, ‘iiber Kalk, Rote Wand bei Mixnitz, c. 1470 m, unweit des Gipfels’, 1974, J. Poelt (GZU!). This is a very curious species; its hymenium components are embedded in a dense gel which does not disperse in 50% of KOH. It is not a Micarea and a new genus is probably required to accommodate it. Micarea hylocomii Poelt & Dobbler in Bot. Jb. 96: 341 (1975). Type: Austria, Tirol, Samnaun Gruppe, on the way from Serfaus to Madatschen, c. 1500 m, on Hylocomium splendens, 15 ix 1972, J. Poelt (GZU - holotype!). I endorse the describing authors’ view that the position of this species in Micarea is doubtful, and I believe a new genus should be erected to accommodate it. Micarea leprosa P. James in Lichenologist 5: 133 (1971). — Vezdaea leprosa (P. James) Vézda, in Poelt & Dobbeler in Lichenologist 9: 170 (1977). This is a species of Vezdaea (Poelt & Débbeler, 1975). A large collection of it from Scotland has recently been distributed as Hertel Lecid. Exs. 60. Micarea minima Poelt & Débbeler in Bot. Jb. 96: 342 (1975). Holotype: Germany, Rheinland-Pfalz, ‘Boschung eines Waldweges bei Battweiler, Kreis Zweibriicken, c. 400 m’, on Polytrichum commune and P. formosum, 1974, J. Poelt (GZU, not seen). Paratype: Austria, Karnten, Nockgruppe, Afritzer Berge, ‘Bergwald und saure Weiden iiber Verditz, nérdlich Villach’, c. 1300 m, on Polytrichum formosum, 1974, J. Poelt (hb Poelt 13294!). I was unable to find any apothecia in the examined paratype and so I cannot confidently exclude this species from Micarea. Further studies are required to establish its generic disposition. Patellaria nigrata Mill. Arg. in Flora, Jena71: 540 (1888). — Bacidia nigrata (Mill. Arg.) Zahlbr., Cat. lich. univ. 4: 129 (1926). — Micarea nigrata (Mill. Arg.) Kalb, Lichenes Neotropici Fasc. 1: 8 (1981); comb. inval. (Art. 33.2). Type: Brazil, Apiahy, on argillaceous soil, 1883, Puiggari (G!). This species closely resembles a Micarea and even appears to have ‘micareoid’ algae. However, its LICHEN GENUS MICAREA IN EUROPE 201 excipular hyphae are conglutinated, broad (c. 2-2-5 wm in K), and pachydermatous. It is closely allied to Bacidia subrudecta (Vainio) Zahlbr. and may have affinities with Byssoloma, but I think it is best retained in Bacidia s. lat. pending further study. The material distributed by Kalb (Lich. Neotrop. Exs. 22) as ‘Micarea nigrata’ is M. lignaria var. lignaria. Psilolechia clavulifera (Ny|.) Coppins, comb. nov. — Lecidea clavulifera Nyl. in Flora, Jena 52: 294 (1869). — Micarea clavulifera (Nyl.) Coppins & P. James, in D. Hawksw., P. James & Coppins in Lichenologist 12: 107 (1980). Type: Finland, Lapponia kemensis, Muonionska [Muonio], Keimiéniemi, ad radicem abietis, 1867, J. P. Norrlin 627 (H — lectotype!; isolectotypes: H!, H-NYL 20855!). Lecidea clavulifera f. subviridicans Nyl. in Flora, Jena 60: 463 (1877). Type: Ireland, West Galway, Connemara, Kylemore, in a cave on the NW side of Doughraugh mountain, 1876, C. Larbalestier (BM —- lectotype! Isolectotypes: BM ex K!; also distributed as Larbal. Lich. Herb. 29: BM!, H!). Thallus effuse, whitish or greenish white, of dispersed to coalescing, irregular granular-areolae c. Fig. 57 Psilolechia clavulifera (Coppins 3614, E). A, spores. B, paraphyses. C, phycobiont cells (cell contents and mycobiont hyphae omitted). Scale = 10 um. 202 BRIAN JOHN COPPINS 0-1-0-2 mm diam. Areolae often disintegrating to form a scurfy-granular crust. Phycobiont not micareoid; cells in clusters and tightly bound by short-celled hyphae, but haustoria not observed; clusters intercon- nected by filamentous hyphae c. 1-5-2 wm wide. Phycobiont cells irregularly globose, broadly ellipsoid or oblong, c. S—12(—18) x3-8 ym, often arranged in pairs or in short chains of up to four cells (Figs. 57C). Apothecia convex-hemispherical and immarginate from the beginning, sometimes becoming subglo- bose, more rarely tuberculate, grey-black with bluish tinge, but shade forms sometimes whitish, blue-grey or grey-brown, 0-1—0-3(-0-4) mm diam; base of apothecia with a white rim (c. 50 4m wide) of outwardly radiating hyphae. Hymenium 28-35 um tall, dilute straw (shade forms), or dilute greenish or aeruginose (K+ green intensifying) especially in the upper part. Asci cylindrical-clavate, 25-30 5-7 um, 8-spored. Spores ovoid, oblong-ovoid or dacryoid, simple, 3-6 (1—)1-2-1-7(-2) wm (Fig. 57A). Paraphyses (Fig. 57B) numerous, usually branched, sometimes anastomosing, distinctly septate and appearing + articu- lated, stout, 1-3-2 wm wide; apices sometimes + clavate and up to 3 wm wide, walls not pigmented but often surrounded by deeply pigmented gel matrix. Hypothecium c. 50-70 um tall, hyaline, or dilute greenish or aeruginose (K—, HNO;+ red) but then never darker than the hymenium. Excipulum indistinct, of radiating hyphae that protrude as loose, hyaline hyphae c, 1-5-2 wm wide and up to 50 wm long. Pycnidia not found. Chemistry: All parts K—, KC—, C—, PD—; no substances detected by t.l.c. Preliminary studies of Lecidea clavulifera by Mr P. W. James and myself led us to transfer it to Micarea. However, critical studies have given me second thoughts on this placement. The distinct white rim of protruding excipular hyphae (superficially like those of Byssoloma spp.), stout and distinctly septate paraphyses, and the unusual phycobiont are all uncharacteristic of a Micarea. The consideration of Psilolechia Massal. as an alternative genus for L. clavulifera was prompted by observations of a lichenicolous member of the Caliciales, Microcalicium arenarium (Hampe ex Massal.) Tibell; M. arena- rium is usually found as a parasite of Psilolechia lucida (Ach.) M. Choisy (see Tibell, 1978), but at three localities in Scotland I have found it on L. clavulifera. At the Berwickshire locality the host and parasite occurred in abundance on roots, stones, and soil of two up-ended trees (Fraxinus); P. lucida was also present in quantity but was not parasitized by M. arenarium. With the exception of pigmentation, sections of the ascocarps of P. lucida and L. clavulifera show + identical anatomical features, e.g. nature of paraphyses, size and shape of asci and spores, and excipulum with numerous protruding hyphae. P. lucida differs from P. clavulifera in the yellow-green colour of its leprose-granular thallus and apothecia (due to presence of pulvinic acid derivatives), and a different phycobiont with + globose cells 5-14 wm diam. The thallus hyphae of both species are identical in appearance, and pycnidia are not known in either species. A recent account of P. lucida is given by James (in Poelt & Vézda, 1981). The phycobiont of P. clavulifera is very unusual; it is reminiscent of the Stichococcus phycobiont of species such as Chaenotheca stemonea and Coniocybe furfuracea, but its cells are much larger. P. clavulifera occurs in communities of the Micareetum sylvicolae, on roots, stones, and consolidated soil of underhangs on banks or the root systems of up-ended trees. Most British records are from Scotland, but it is also known from Cumbria, south Wales and western Ireland (Map 28). From outside Britain I have seen material of it from Finland, Germany, and Czechoslovakia (see below). Additional specimens of P. clavulifera examined: BritisH IsLes. Brecon (V.C.42). 22/94: Upper Dyfnant Valley, 1982, Woods (hb Woods). Cumberland (V.C.70). 35/54: Baron Wood, by River Eden, 60-75 m dry sandstone cliff, 1979, Coppins 4344 (E). Berwick (V.C.81). 36/76: W of Elba, S side of Whiteadder Water, exposed roots of up-ended Fraxinus, 1981, Coppins 8890 (E — Microcalicium arenarium folder). West Perth (V.C.87). 27/40: Aberfoyle, the Trossachs, on stone amongst roots of up-ended tree, 1978, Coppins 3661 (E). Mid Perth (V.C.88). 27/55: Black Wood of Rannoch, stone in bank by track, 1976, Coppins 4654 (E); 27/81: Crieff, Drummond Wood, roots and sandstone of up-ended tree, partly parasited by Microcalicium arenarium, 1978, Coppins 3164, 3625 (E). Angus (V.C.90). 37/33: Sidlaw Hills, Auchterhouse Hill, 396 m, underside of boulder, 1975, Coppins 846 (E — Microcalicium arenarium folder). South Ebudes (V.C.102). 16/49: Colonsay, Coille Mhé6r, roots and soil of up-ended Betula, 1981, Coppins 8878 (E). Mid Ebudes (V.C.103). Mull: 17/32: Bunessan, Ardfenaig Woods, 1970, James (BM); 17/54: Salen, 1968, James (BM); 17/55; Aros House, roots of up-ended tree, 1968, James (BM). West Galway (V.C.H.16). 84/72 (L/65): Connemara, near Clifden, 1878, Larbalestier (BM). FINLAND. Satakunta: Siikainen, Vuorijarvi, Vaasaneva, sandy soil amongst roots of Picea, 1936, Laurila (GZU, H). W. Germany. Baden-Wiirttemberg: Heidelberg, K6nigstuhle, on sandstone, 1883, Zwackh (H-NYL p.m. 4229). CzEcHosLovAKIA. Slovakia, Nizke Tatry, Liptovska Teplitka, Dzurova, 18 —, Lojka 4289 (BM). LICHEN GENUS MICAREA IN EUROPE 203 Ce) ° > =A aanl=: pees pa ae | Se ia DE) Ls : ee Gas a # e awl ae | ORS, 1 3 bp a o = Map 28 Psilolechia clavulifera @ 1950 onwards O Before 1950 Sporacestra Massal. in Atti R. Ist. veneto Sci. U1, 5: 264 (1860). Type species: Biatora prasina Tuck. & Mont. in Mont. (1857), non (Fr.) Trevisan (1856). Sporacestra was included as a synonym of Micarea by Vézda & Wirth (1976: 99) who confused the two quite separate applications of the name ‘Biatora prasina’. The name Biatora prasina Tuck. & Mont. was introduced for a corticolous species from Venezuela with long acicular spores, and no reference was made to the ‘prasina’ of Fries. Similarly, Massalongo made no reference to Fries’ name and his generic description (e.g. ‘. . . sporidii capillari aghiformi, lineari allungati . . .’) clearly refers to the stated type species, ‘Biatora prasina Mont., Tuck.’. The Venezuelan species was transferred to Bacidia de Not. by Zahlbruckner, but his use of the epithet ‘prasina’ is contrary to the present Code. From the descriptions of this species I believe it should be retained in Bacidia, pending further study. The nomenclature and required new combination for the species is as follows: Bacidia prasinata (Tuck. ) Coppins, comb. nov. — Biatora prasinata Tuck. in Syn. N. Am. Lich 2: 41 (1888); nom. nov. — Bacidia prasina Tuck & Mont. in Mont. in Annls Sci. nat. [V, 8: 296 (1857), non (Fr.) Trevisan (1856). — Bacidia prasina Zahlbr., Cat. lich. univ. 4: 253 (1926); nom. illeg. (Art. 11). Type: Venezuela, on bark, Fendler (not seen). Index to exsiccatae I have attempted to examine at least one example of all European and North American material referable to Micarea and distributed in recognized exsiccatae (Lynge 1915-22, 1939; Sayre, 1969). My preliminary list was compiled from Lynge (op. cit.), available schedae, major floras 204 BRIAN JOHN COPPINS (e.g. Fries, 1874; Kérber, 1855, 1859-65; Smith, 1926; Vainio, 1922, 1934) and numerous other papers dealing with species here included in Micarea. Most examples gleaned from these sources have been seen; the few not seen are listed after the main index. Herbarium locations for examined examples containing Micarea species can be found in the relevant taxonomic accounts; locations of examples not containing Micarea species are given in the index. The names on the exsiccate labels are given (in round brackets following the number) where they are taxonomically at variance with the species represented. In many cases, only one or two examples of a particular number have been seen by me, thus this index must be regarded as provisional. Some numbers listed as not containing a Micarea (e.g. Zwackh Lich. Exs. 778) may well in further examples (not seen by me) contain a Micarea. In addition, some Micarea species may have been misidentified and distributed under names not connected with Micarea, and consequently overlooked in the course of this study. It should be noted that many numbers in the exsiccatae of Harmand (Lich. Loth.), Mougeot & Nestler, and Schaerer are notoriously heterogeneous with regard to species compo- sition, substratum, and locality. Most numbers in other exsiccatae are reasonably uniform in these respects but heterogenicity sometimes arises in cases where species of similar outward appearance occur together in the field (e.g. M. bauschiana—M. lutulata—M. sylvicola—M. tuberculata, and M. denigrata—M. lignaria—M. peliocarpa). During the preparation of exsiccatae the need for the careful identification of individual samples is paramount before they are dis- tributed. Generic names are abbreviated as follows: Bacidia (B), Biatora (Bi.), Biatorina (Biat.), Bilimbia (Bil.), Catillaria (C.), Lecanora (Lec.), Lecidea (L.), Micarea (M.), Psilolechia (P.) and Scoliciosporum (S.). ANZI, Lich. Exs. Ital.: 256 M. denigrata; 259A (Bil. lignaria) M. melaena; 259B (Bil. hypnophila — given by Lynge as Bil. lignaria) B. sabuletorum (BM). ANZI, Lich. Lang.: 148 (Bil. syncomista) M. lignaria, one example in BM with a little M. peliocarpa. ANZI, Lich. Sondr.: 170A M. peliocarpa; 170B M. melaena. ARNOLD, Lich. exs.: 120 M. bauschiana; 167A (Bil. lignaria B saxigena Leighton) M. peliocarpa; 167B M. peliocarpa; 217 M. nitschkeana; 279, 230A-C M. prasina; 332A-—C M. melaena; 348A, B M. lignaria; 409A M. sylvicola; 409B (L. sylvicola) M. sylvicola, or M. lutulata; 503A—D M. nitschkeana; 548,549 M. cinerea; 556[A], B (L. assimilata) M. crassipes; 626, 627 M. misella; 836 M. lithinella; 837 M. peliocarpa; 1051 (Bil. ternaria) M. peliocarpa; 1057 M. tuberculata; 1121 M. crassipes; 1122 M. prasina; 1471 M. elachista; 1472 M. prasina. ARNOLD, Lich. Mon.: 46 M. denigrata; 47 M. cinerea; 48 M. nitschkeana; 49 M. melaena; 115, 116 M. cinerea; 118 (Bil. trisepta) M. peliocarpa; 172 M. misella; 241 M. misella; 243 M. prasina; 244 (Biat. prasiniza) M. adnata; 245 M. prasina; 246 M. elachista; 248, 249 M. melaena; 269 (Bil. trisepta) M. peliocarpa; 270 (Bil. trisepta) M. nitschkeana; 307 M. misella; 357 (Bil. trisepta) M. peliocarpa; 407 M. melaena; 482 (Bil. trisepta) M. peliocarpa. BOHLER, Lich. Brit.: 85 (L. viridescens) M. lignaria. BRITZELMAYR, Lich. exs.: 174 (Biat. prasiniza var. laeta) M. adnata; 175 (Biat. glomerella) L. viridescens (M); 208 M. misella; 310 (Biat. synothea) L. turgidula + Lec. sp. (H); 464 (Biat. synothea) M. denigrata + C. nigroclavata (H); 599 (Biat. synothea) Arthonia vinosa (H); 829 (Bil. milliaria f. nigrita) M. nitschkeana; 846 M. cinerea; 946 M. melaena; 961 (L. assimilata f. irrubata) L. hypnorum + L. limosa (H). CLAUDEL & HARMAND, Lich. Gall.: 43 M. lignaria; 89 M. nitschkeana; 445 M. misella. CROMBIE, Lich. Brit.: 174 (L. misella) M. misella +/or M. prasina. CUMMINGS, Decad. N Amer. Lich. Ed. I: 302 M. lignaria; 355 M. prasina. Ed. II: 232 M. lignaria. ELENKIN, Lich. Ross.: 189 M. misella. Erb. Critt. Ital., ser. I: 198 (C. synothea) C. nigroclavata (UPS). FELLMAN, Lich. Arct.: 159 M. melaena; 164 (L. assimilata f. alpestris) M. incrassata, + a little M. crassipes in example in H; 165 (L. assimilata) M. crassipes; 166 (L. limosa) M. incrassata. FLAGEY, Lich. Franche-Comté: 137 (Bi. synothea) Lec. sp. + Lecidella ? euphorea (UPS). Flora Hung. exs.: 714 M. melaenida. FLOTOW, Lich. exs.: 129A M. lignaria; 129B (L. milliaria b. major — according to Lynge) Mycoblastus affinis (UPS); 129C (L. milliaria var. lignaria — according to Lynge) M. melaena; 129D (L. milliaria) C. globulosa (UPS); 129E M. lignaria; 130 M. turfosa; 131 M. lignaria; 171A M. sylvicola. LICHEN GENUS MICAREA IN EUROPE 205 FRIES, Lich. Suec.: 29 M. lignaria, + some M. denigrata in example in UPS; 98 M. denigrata, + some M. lignaria in example in E; 212A, B M. melaena. HANSEN, Lich. Groenl.: 172 (L. assimilata) L. stenotera (BM); 247 (L. assimilata) L. limosa (BM). HARMAND, Guide Elém. Lich.: 91 M. lignaria. HARMAND, Lich. Loth.: 810 (Lecidea vernalis, and f. prasina) mixture from several localities, example in ANGUC has Bi. (? Moelleropsis) humida, C. sphaeroides, L. icmalea and L. vernalis, that in UPS has C. sphaeroides and L. vernalis; 838 M. denigrata; 852 M. lignaria; 853 M. nitschkeana. HAVAAS, Lich. Norv.: 139 M. subviolascens; 555 M. lignaria; 571 (L. sylvicola var.) M. lutulata; 694 M. subviolascens; 710 M. subviolascens. HAVAAS, Lich. Norv. Occid.: 269 M. subviolascens. HEPP, Flecht. Eur.: 14 M. denigrata; 20 (Bi. lignaria) M. nitschkeana; 21 (Bi. cinerea) M. cinerea, or M. nitschkeana; 278 M. prasina; 284 M. peliocarpa; 285 (Bi. lignaria var. milliaria) M. peliocarpa; 504 M. melaena; 510 M. peliocarpa; 524 (Bi. asserculorum) S. umbrinum (E). HEPP, Ziirich: 206 (L. cinerea) M. peliocarpa; 210 (L. synothea) Lecania cyrtella agg. (BERN); 224 M. prasina. HERTEL, Lecid. exs.: 34 M. lignaria; 54 M. ? bauschiana. JOHNSON, N. Engl. Lich. Herb.: 373 M. denigrata; 375 (L. turneri; listed under Bil. lignaria by Smith (1911, 1926)) Bacidia sabuletorum + Toninia lobulata (BM); 376 M. melaena; 434 L. sylvicola var. hellbomii) M. sylvicola, or M. bauschiana; 453 M. lignaria; 504 M. bauschiana. KALB, Lich. Neotropici: 22 M. lignaria (M. nigrata); 186 M. lignaria. KAVINA & HILITZER, Crypt. Cech.: 269 M. sylvicola. KORBER, Lich. Sel. Germ.: 12 M. turfosa; 75 M. sylvicola; 133A (Bil. lignaria) M. peliocarpa; 133B M. peliocarpa; 137 (Bi. denigrata) L. (Lecidella) alba (L); 250 M. prasina. Krypt. Exs. Vind.: 165 M. peliocarpa; 362 M. melaena; 658 M. lignaria; 1232 M. nitschkeana; 1532 M. misella; 2061 (C. prasiniza var. prasinoleuca) Dimerella diluta (BM, BM ex K, M); 2268 M. assimilata; 2561 (C. denigrata) C. globulosa (BM, BM ex K); 3153 M. denigrata; 3154 M. melaenida; 3651 (L. misella) M. denigrata; 4214 M. misella; 4858 M. denigrata. KUTAK, Lich. Bohem.: 205 (L. asserculorum) M. denigrata; 310 M. prasina; 417 M. lignaria, + alittle M. leprosula in example in O; 516 M. denigrata; 517 M. denigrata. LARBALESTIER, Lich. Caesar. Sarg.: 83 (L. arceutina var. hypnaea) M. lignaria; 84 M. sylvicola. LARBALESTIER, Lich. Herb.: 29 P. clavulifera; 68 M. bauschiana; 223 M. lutulata; 227 (L. polioides) M. tuberculata, but example in LIV is L. monticola; 272 M. lignaria; 304 M. sylvicola; 305 M. sylvicola or M. bauschiana; 347 M. peliocarpa. LEIGHTON, Lich. Brit.: 120 (Bi. uliginosa) M. melaena; 210 M. lignaria; 238 (L. milliaria var. terrestris) M. lignaria, or M. peliocarpa, example in MANCH has a little M. leprosula; 386 M. lignaria; 388 (L. sabuletorum var. milliaria) M. botryoides. LOJKA, Lich. Hung.: 60 M. cinerea; 61 M. lignaria; 134 M. peliocarpa. LOJKA, Lichenoth. Univ.: 29-31 M. prasina; 137 M. nitschkeana; 233 M. lithinella. MAGNUSSON, Lich. Sel. Scand.: 134 M. prasina; 208 M. leprosula; 282 M. turfosa; 340 M. nitschkeana. MALBRANCHE, Lich. Norm.: 287 (L. sphaeroides var. lignaria) M. nitschkeana; 387 (L. sphaeroides var. melaena) M. denigrata. MALME, Lich. Suec.: 20 M. rhabdogena; 21 M. elachista; 22 M. anterior; 23, 24 M. prasina, 25 M. nitschkeana; 26 M. eximia; 27 M. melaena; 28 M. contexta; 125 M. lithinella; 145 M. denigrata; 169 M. peliocarpa; 199 M. sylvicola; 216 M. assimilata; 288 M. lignaria; 362 M. crassipes; 365 (M. denigrata var. pyrenothizans) M. misella; 865 M. turfosa. MIGULA, Crypt. Germ., Aust., Helv.: 1 (Bil. milliaria) M. lignaria +/or M. leprosula, + M. nitschkeana; 132 M. denigrata; 226 M. lignaria. MOUGEOT & NESTLER, Stirpes Crypt. Vog.-Rehn.: 1329 (L. melaena) an example in E includes a little M. melaena, other collections in BM (2 sets), DBN and M (3 sets) contain a selection from the following, Lec. spp., is granulosa agg., L. turgidula, L. sp., M. denigrata; 1430 M. lignaria. MUDD, Lich. Brit.: 156-8 M. lignaria; 159 M. melaena; 164 (L. prasina) L. viridescens (E, BM, M); 175 M. sylvicola. NORRLIN & NYLANDER, Herb. Lich. Fenn.: 145 (L. sylvicola) L. conferanda (BM, H, M); 177 M. denigrata; 180 M. melaena; 182 (L. pelidna) M. intrusa [some examples may prove to be S. umbrinum]; 194A, B M. crassipes; 314 M. elachista, one example H is B. hegetschweileri; 319A, B (L. milliaria var. ternaria) M. lignaria; 321 M. turfosa; 724 M. elachista; 744 M. misella; 745 (L. asserculorum) M. denigrata; 763, 764 (L. tuberculata) M. sylvicola. OLIVIER, Lich. Orne: 237 M. melaena; 344 M. lignaria. PISUT, Lich. Slov.: 156 M. lignaria. 206 BRIAN JOHN COPPINS POELT, Lich. Alp.: 22 (L. misella) M. denigrata. RABENHORST, Lich. Eur.: 224 (Bi. turfosa) L. uliginosa (BM ex K); 322 M. lignaria, example in M has some M. peliocarpa; 582 (Bil. lignaria) M. nitschkeana; 583 M. nitschkeana, 603 (Bil. syncomista) M. lignaria; 626 M. denigrata; 648 (Bi. bauschiana) M. bauschiana, or M. tuberculata; 675 (L. sylvicola) L. erratica (BM, M); 676 M. prasina; 733 M. prasina. RASANEN, Lich. Fenn. exs.: 489 (Bil. nitschkeana) B. naegelii (LD); 512 M. tuberculata; 642 M. nitschkeana; 651, 652, 653 M. prasina; 672 (L. sylvicola) M. sylvicola, +/or M. tuberculata, +/or M. lutulata; 822 (C. elachista f. simplicata) L. turgidula (BM ex K); 963 M. melaena. RASANEN, Lichenoth. Fenn.: 137 (C. elachista f. simplicata) B. igniarii (BM ex K); 343 M. peliocarpa; 344 M. melaena; 426 (C. ilicis) M. globulosella. Reliq. Suza.: 42 M. misella. ROUMEGUERE, Lich. Gall.: 87 (L. vernalis var. milliaria) B. sabuletorum (BM, M); 193 (L. vernalis var. synothea) M. lignaria; 231 M. melaena, but example in BM is mixture of a L. sp. and Lec. spp.; 232 M. lignaria; 467 (L. melonida [melaenida]) Toninia aromatica (BM). SAMPAIO, Lich. Port.: 132 M. denigrata; 147 M. lignaria. SCHAERER, Lich. Helv.: 196 (L. sabuletorum var. lignaria) examples labelled ‘Ad ligna decorticata in alpibus’ are M. prasina, examples labelled ‘In m. Belpberg’ contain one, or a mixture, of the following, M. lignaria, M. melaena, M. peliocarpa and M. leprosula; 327 (Parmelia varia var. denigrata) L. (Lecidella) alba (BM ex K), or L. aff. cadubriae (E). SUZA, Lich. Bohem.: 131 M. sylvicola. THOMSON, Lich. Arct.: 12 (B. melaena) Toninia lobulata (LD). VAINIO, Lich. Bras.: 1420 M. misella; 1451 M. misella. VEZDA, Lich. Bohem.: 133 M. lignaria; 258 M. lignaria; 282 M. crassipes. VEZDA, Lich. Sel. exs.: 11 M. crassipes; 14 M. melaena; 90 M. prasina; 516 M. lignaria; 538 M. turfosa, 706 (M. violacea) B. naegelii (BM); 858 (M. ternaria) M. lignaria; 957 (L. tuberculata) M. sylvicola; 1036 M. lignaria; 1087 M. cinerea; 1088 (M. ternaria) M. lignaria; 1134 M. elachista; 1135 M. turfosa; 1341 (M. tuberculata) M. sylvicola; 1342 M. peliocarpa; 1380 M. peliocarpa; 1430 M. denigrata; 1467 M. prasina, 1595 (M. hemipoliella) M. prasina; 537 M. melaenida. WEBER, Lich. Exs.: 73 M. denigrata. ZAHLBRUCKNER, Lich. Rar. exs.: 110 M. nitschkeana; 152 (C. melanobola f. frullaniae =) Arthonia aff. exilis (BM); 174 (L. infidula) L. erratica; 175 M. prasina; 276 M. turfosa. ZWACKH, Lich. exs.: 121 M. lignaria; 122 (L. milliaria Fr. var. ?) M. elachista; 276 M. peliocarpa; 279A (Bi. gelatinosa var. minor!) M. bauschiana; 279B M. bauschiana; 394 M. denigrata; 416 M. prasina; 417 (Bil. nitschkeana) S. umbrinum + S. chlorococcum (UPS); 470 M. nitschkeana [two labels: ‘Bilimbia nitschkeana, bei Minster in Westfalen’; or ‘Lecidea nitschkeana, bei Dolbrueck in Kreise Paderborn .. .’, sometimes labelled ‘470 bis’, example in M. also has M. denigrata]; 534 M. nitschkeana; 535 (L. latens) M. sylvicola; 587 M. nitschkeana; 590 M. lithinella; 591A, B, 592A-E, 593A-C M. prasina; 594A, B, 595 M. bauschiana; 596 (L. latens) M. sylvicola; 597 M. sylvicola; 656 M. prasina; 657 M. melaena; 675 M. melaena; 778 (L. cinerea) L. symmicta agg. (UPS); 780 (L. trachona) M. sylvicola;, 897 M. peliocarpa; 898 M. cinerea; 900 M. turfosa; 919 M. sylvicola; 1085 M. misella. Examples of the following have not been examined by me; names taken from Lynge (op. cit.). BRITZELMAYR, Lich. exs.: 838 (Bil. lignaria); 890 (Bil. trisepta). FLOTOW, Lich. exs.: 112 (L. synothea ?). HARMAND, Lich. Loth.: 834 (L. denigrata). KORBER, Lich. Sel. Germ.: 343 (Bil. syncomista). LARBALESTIER, Lich. Herb.: 139 (L. micrococca). OLIVIER, Lich. Orne: 264 (L. nitschkeana). ROUMEGUERE, Gen. Lich. exs.: 36 (L. vernalis var. synothea); 51 (L. vernalis var. milliaria). No examples of this exsiccata have been traced; it is almost certain that it was never distributed (P. M. Jorgensen, in litt.). SCHADE, STOLLE & RIEHMER, Lich. Sax.: 256 (B. trisepta). ZWACKH, Lich. exs.: 131 (L. glomerella and L. confluens). Acknowledgements My sincerest thanks go to Mr P. W. James and Dr F. Rose for their encouragement, guidance and many kindnesses, not only during the present study, but also from the time of my first interests in lichenology in 1965. For further help and advice in those earlier years I here take the opportunity of extending my thanks to Dr D. L. Hawksworth, Mr J. R. Laundon, Dr M. R. D. Seaward, Dr T. D. V. Swinscow, and Mr A. E. Wade. I am much indebted to the curators and owners of the herbaria listed in ‘Materials’; especially as their searches for relevant.specimens and processing of loans have often been long and arduous procedures. Furthermore, many of these curators and collectors have provided me with much supplementary information on localities, habitats, and historical background. Special thanks are due to Miss F. J. Walker LICHEN GENUS MICAREA IN EUROPE 207 -of the British Museum (Natural History) for her efficient handling of my many ‘urgent’ requests for herbarium and bibliographical information. Recognition for their helpful comments and suggestions on various nomenclatural and taxonomic problems pertaining to Micarea and related groups goes to Mr B. L. Burtt, Prof. Dr H. Hertel, Dr H. Kilias, Prof. P. M. Jorgensen, Prof. Dr J. Poelt, Prof. R. Santesson, Dr V. Wirth, and Dr A. Vézda. Much advice and assistance has come from many of my colleagues at the Royal Botanic Garden, Edinburgh; in particular, I would like to thank Prof. D. M. Henderson, the Regius Keeper, and Mrs F. M. Bennell, Mr A. P. Bennell, Mrs N. M. Gregory, Dr R. Watling, Mrs J. M. Woods, Mrs H. Hoy, Mrs D. A. Morrison, Mr M. V. Mathew, Mr D. G. Long, and the staff of the typing section. All the people and organizations to whom I am grateful for assistance with field excursions are too numerous to mention individually. However, special thanks go to Dr L. Tibell for arranging and leading me on a tour of mid- and north Sweden in 1977, and to the Nordic Lichen Society (especially Mr U. Sgchting and Mr K. Ramkaer) for inviting me as guest visitor to their excursion in north Jylland in 1979. The following funds and institutions are thanked for financial support of field-work: Nature Conservancy Council, Natural Environment Research Council, Royal Botanic Garden, Edinburgh, and the World Wildlife Fund. 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[Regnum veg. 106.] Utrecht. Honegger, R. 1978. The ascus apex in lichenized fungi I. The Lecanora-, Peltigera- and Teloschistes-types. Lichenologist 10: 47-67. Hughes, S. J. 1972. New Zealand Fungi 17. Pleomorphism in Euantennariaceae and Metacapnodiaceae, two new families of sooty moulds. NZ. JI Bot. 10: 225-242. — 1976. Sooty moulds. Mycologia 68: 693-820. Huneck, S. & Follmann, G. 1972. Mitteilungen tiber Flechteninhaltsstoffe. XCIV. Zur vergleichenden Phytochemie der Gattung Bacidia (Lecideaceae). J. Hattori bot. Lab. 36: 54-56. —— & Lamb, I. M. 1975. 1’-chloropannarin, a new depsidone from Argopsis friesiana: notes on the structure of pannarin and on the chemistry of the lichen genus Argopsis. Phytochemistry 14: 1625-1628. Jahns, H. M. 1974 [‘1973’] Anatomy, morphology, and development. Jn V. Ahmadjian & M. E. Hale (Eds), The lichens: 3-58. New York. James, P. W. 1965a. A new check-list of British lichens. Lichenologist 3: 95-153. —— 1965p. Field meeting in Scotland, Lichenologist 3: 155-172. — 1970. The lichen flora of shaded acid rock crevices and overhangs in Britain. Lichenologist 4: 309-322. —, Hawksworth, D. L. & Rose, F. 1977. Lichen communities in the British Isles: a preliminary conspectus. Jn M. R. D. Seaward (Ed.), Lichen ecology: 295-413. London. — & Henssen, A. 1976. The morphological and taxonomic significance of cephalodia. Jn D. H. Brown, D. L. Hawksworth & R. H. Bailey (Eds), Lichenology: progress and problems: 27-77. London. Kilias, H. 1981. Revision gesteinsbewohnender Sippen der Flechtengattung Catillaria Massal. in Europa. Herzogia 5: 209-448. Knowles, M. C. 1929. The lichens of Ireland. Proc. R. Ir. Acad. B, 38 (12): 179-434. Korber, G. W. 1855. Systema lichenum Germaniae. Breslau. — 1859-65. Parerga lichenologica. Breslau. Korf, R. P. 1973. Discomycetes and Tuberales. Jn G. C. Ainsworth, F. K. Sparrow & A. S. Sussman (Eds), The fungi, an advanced treatise 4A: 249-319. Lamb, I. M. 1953. New rare or interesting lichens from the Southern Hemisphere. Lilloa 26: 401-438. — 1963. Index nominum lichenum. New York. Laundon, J. R. 1981. The species of Chrysothrix. Lichenologist 13: 101-121. Leighton, W. L. 1879. The lichen-flora of Great Britain, Ireland, and the Channel Islands. Ed. 3. Shrewsbury. *Hedlund, J. T. 1891. Om bilbildning genom pycnoconidier hos Catillaria denigrata (Fr.) och C. prasina (Fr.). Bot. Notiser 1891: 186-211. LICHEN GENUS MICAREA IN EUROPE 209 Lindsay, W. L. 1859. Memoir on the spermagones and pycnides of filamentous, fruticulose, and foliaceous lichens. Trans. R. Soc. Edinb. 22: 101-103. 1872. Memoir on the spermogones and pycnides of crustaceous lichens. Trans. Linn. Soc. Lond. 28: 189-318. Lynge, B. 1915-22. Index specierum et varietum lichenum quae collectionibus ‘Lichenes Exsiccati’ distributae sunt. Nyt Mag. Naturvid. 53-54, 57-60 [bound volume with separate pagination]. 1939. Index collectionum ‘Lichenes Exsiccati’. Suppl. I. Nytt Mag. Naturvid. 79. Magnusson, A. H. 1942. New or otherwise interesting Swedish lichens. XI. Bot. Notiser 1942: 1-18. Millbank, J. W. 1976. Aspects of nitrogen metabolism in lichens. Jn D. H. Brown, D. L. Hawksworth & R. H. Bailey (Eds), Lichenology: progress and problems: 441-455. London. Omar, M. B., Bolland, L. & Heather, W. A. 1979. A permanent mounting medium for fungi. Bull. Br. mycol. Soc. 13: 31-32. Ozenda, P. 1963. Lichens. In Handbuch der Pflanzenanatomie, Ed. 2 (W. Zimmerman & P. Ozenda, Eds), 6 (9): i-x, 1-199. Berlin. —— & Clauzade, G. 1970. Les Lichens. Etude biologique et flore illustrée. Paris. Peveling, E. 1974 [‘1973’]. Fine structure. Jn V. Ahmadjian & M. E. Hale (Eds), The lichens: 147-182. New York. Poelt, J. 1958. Uber parasitische Flechten. II. Planta 51: 288-307. — 1974a [‘1973’]. Systematic evaluation of morphological characters. Jn V. Ahmadjian & M. E. Hale (Eds), The lichens: 91-115. New York. —— 1974b. Classification. In V. Ahmadjian & M. E. Hale (Eds), The lichens: 599-632. New York. —— 1977. Die Flechte Micarea hemipoliella auf dem Schneidried Cladium mariscus. Herzogia 4: 409-413. —— & Dobbler, P. 1975. Uber moosparasitische Arten der Flechtengattungen Micarea und Vezdaea. Bot. Jb. 96: 328-352. —— & —— 1979. Bryostigma leucodontis nov. gen. et spec., eine neue Flechte mit fast unsichtbaren Fruchtk6rpern. Pl. Syst. Evol. 131: 211-216. —— & Vézda, A 1977. Bestimmungsschliissel europdischer Flechten. Erganzungsheft I. Vaduz. —— & —— 1981. Bestimmungsschlussel europdischer Flechten. Erganzungsheft II. Vaduz. Santesson, R. 1952. Foliicolous lichens 1. A revision of the obligately foliicolous lichenized fungi. Symb. bot. upsal. 12 (1): 1-590. Sayre, G. 1969. Cryptogamae exsiccatae — An annotated bibliography of published exsiccatae of Algae, Lichenes, Hepaticae and Musci. Mem. N.Y. Bot Gdn 19: 1-174. Scannell, M. J. P. 1978. Catillaria bouteillei (Desm.) Zahlbr., an epiphyll on leaves of Prunus lauro-cerasus from the collection of R. A. Phillips (1937). Jr. Nat. J. 19: 172. Schaerer, L. E. 1850. Enumeratio critica Lichenum Europaeorum. Bern. Smith, A. L. 1911. A monograph of British lichens 2. London. — 1926. A monograph of British lichens 2, 2nd ed. London. Stafleu, F. A. & Cowan, R. S. 1976-81. Taxonomic literature. A selective guide to botanical publications and collections with dates, commentaries and types. 2nd ed. 1 [Regnum veg. 94], 2 [Regnum veg. 98], 3 [Regnum veg. 105]. Utrecht. Steven, H. M. & Carlisle, A. 1959. The native pinewoods of Scotland. Edinburgh & London. Sutton, B. C. 1980. The Coelomycetes. Kew. Tibell, L. 1978. The genus Microcalicium. Bot. Notiser 131: 229-246. Vainio, E. A. 1922. Lichenographia fennica II. Baeomyceae et Lecideales. Acta Soc. Fauna Fl. fenn. 53 (1). 1934. Lichenographia fennica IV. Lecideales II. Acta Soc. Fauna Fl. fenn. 57 (2). Vézda, A. 1961. Lichenes novi vel rariores Sudetorum occidentalium. Preslia 33: 365-368. 1978. Neue oder wenig bekannte Flechten in der Tschechoslowakei. II. Folia geobot. phytotax., Praha 13: 397-420. 1979. Flechtensystematische Studien. XI. Beitrage zur Kenntnis der Familie Asterothyriaceae (Discolichens). Folia geobot. phytotax., Praha 14: 43-94. —— & Wirth, V. 1976. Zur Taxonomie der Flechtengattung Micarea Fr. em. Hedl. Folia geobot. phytotax., Praha 11: 93-102. Vobis, G. 1977. Studies on the germination of lichen conidia. Lichenologist 9: 131-136. 1980. Bau und Entwicklung der Flechten-Pycnidien und ihrer Conidien. Bibltheca lich., Lehre 14: i-v, 1-141. — & Hawksworth, D. L. 1981. Conidial lichen-forming fungi. In The biology of conidial fungi (G. T. Cole & W. B. Kendrick, Eds) 1: 245-273. New York, London & San Francisco. Walker, F. J. & James, P. W. 1980. A revised guide to microchemical techniques for the identification of lichen products. Bull. Brit. Lich. Soc. 46 (suppl.): 13-29. 210 BRIAN JOHN COPPINS Watson, W. 1930. The lichens of Somerset. Taunton. Wirth, V. 1973. Uber einige fiir Deutschland neue Flechten. Nova Hedwigia 24: 181-192. Zahlibruckner, A. 1921-40. Catalogus lichenum universalis. 10 vols. Berlin. Zahlbruckner, A. 1926. Lichenes. B. Spezieller Teil. In Die nattirlichen Pflanzenfamilien. Ed. 2. (A. Engler, Ed.) 8: 61-270. Leipzig. Index Accepted names are in roman and synonyms in italic. New names and principal references are in bold whilst an asterisk (*) denotes a figure or map. Some taxa mentioned incidentally (e.g. in lists of associated species) are not included. aberrans 185, 186 abietina (Lecanactis) 67 absistens (Bacidia) 88 adnata 24, 29, 33*, 61, 64, 65, 66*, 67, 68*, 89, 90, 93-95, 105, 108-110, 111*, 114, 204 aeneofusca (Lecidea) 198 aeolotera (Huilia) 26 aeruginosa 151 aeruginosa (Lecidea) 198 affinis (Mycoblastus) 204 aggerata 186 alabastrites 22, 23, 26*, 29, 32*, 34*, 64, 65, 85, 89-91, 93-95, 99, 103, 110-113*, 122, 172 alba (Lecidea, Lecidella) 205, 206 albella 121,122 albicans 170 albidolivens 170 alnicola 151 andesitica 128 aniptiza 127 anterior 24, 29, 32, 33*, 61, 64, 65, 69*, 90, 91, 95, 100, 105, 108-110, 112-114, 136, 205 antrophila 149 aphanoides 138, 140 apochroeella (Lecidea) 132 arceutina (Bacidia) 197, 198 arenarium (Microcalicium) 202 arnoldiana (Bacidia) 67 aromatica (Toninia) 206 Arthonia 31, 98 Arthoniaceae 31 Arthothelium 31 asserculorum (Lecidea) 158, 159 assimilata 22,25, 29, 32, 35*, 61, 62, 64, 67, 87-90, 95, 100, 104, 114-116*, 154, 186, 187, 205 atroviridis (Lecidea) 199 Bacidia 88, 97 bacidiella (Lecidea, Micarea) 135, 198 bauschiana 24, 25, 29, 30*, 31, 36*, 61, 64, 71*, 87, 89, 93, 95, 100, 105, 117-119*, 148, 149, 187, 194, 204—206 beckhausii (Bacidia, Micarea) 20, 30, 88, 102, 135, 189, 196, 198 berengeriana (Lecidea) 115 Biatora 97, 112, 197 Biatorina 97 biforme (Anisomeridium) 66 biseptata 174 Bispora 29 botryocarpa 192 botryoides 24, 29, 32, 36*, 61, 62, 64, 65*, 67, 69* , 84, 88-90, 95, 99, 106, 108, 114, 118-121*, 149, 156, 159, 163, 195, 205 botryoides (Byssus) 174 bouteillei (Catillaria) 197*, 199 brasiliana 158 Bryophagae 96, 196 Bryostigma 31 Buellia 31,61 byssacea 174,175 Byssoloma 98, 202 cadubriae (Lecidea) 206 caesioatra (Lecidea) 30, 116 calamophila 142 callicarpa 117 Caloplaca 31 carbonicola 132 carneoglauca (Bacidia) 67 Catillaria 61, 88, 97 catillarioides 151 Chaenothecopsis 64 chalybeia (Catillaria) 97, 199 chlorococcum (Scoliciosporum) 91, 97, 206 chrysophthalma (Chrysothrix, Micarea) 200 Chrysothrix 98, 196 cinerea 22-26, 29, 37*, 61, 63-66, 70*, 85, 87, 88-91, 93-95, 99, 104, 108, 109, 112, 121-123*, 124, 129, 141, 144, 166, 172, 204-206 clavulifera (Psilolechia, Lecidea, Micarea) 105, 107, 163, 193, 198, 201*, 202, 203*, 205 Cliostomum 97 coccinea (Micarea) 96, 200 Collemaceae 20,24 LICHEN GENUS MICAREA IN EUROPE conferanda (Lecidea) 205 confusula 183 conglomerata 169 contexta 24, 29, 31, 32, 38*, 61, 63, 64, 71*, 85, 87, 88, 90, 91, 95, 100, 107, 124, 125, 134, 164, 205 contristans (Catillaria) 98, 115, 200 contrusa 138 crassipes 20, 28*, 29, 38*, 62-64, 67, 72*, 87-89, 95, 96, 100, 106, 115, 116, 125, 126, 152, 187, 204-206 cupreola 170 curvata 29, 38*, 64, 85, 87-89, 92, 95, 105, 126, 127, 185 cyanescens (Micarea) 200 cyrtella (Lecania) 198, 205 cyrtellina (Lecania) 66 declivitatum 174 decrustata 158 delicatula 121, 122 demarginata (Lecidea) 149, 198 denigrata 22, 23*, 24, 26, 29, 30, 32, 39*, 61, 64, 66, 67, 73*, 84, 85, 87-90, 92-96, 99-102, 108, 109, 127-130*, 131, 133-136, 148, 166, 182, 198, 204-206 diluta 112 diluta (Dimerella) 205 dilutiuscula 117 discrepans 174 discretula | 127 dufourei (Lecidea) 200 ecrustacea Lamy 127 ecrustacea Nyl. ex Vainio 181 efflorescens (Lecidea) 85 elachista 22, 23*, 29, 40*, 64, 84, 88, 90, 91, 95, 99, 103, 105, 129, 131-133, 181, 204—206 elegantior (Huilia) 26 endocyanea 151 endoleuca 64, 84-86, 93-95, 104, 122, 146, 147* epiphaeotera 151 erratica (Lecidea) 67, 187, 198, 206 erysibe (Lecania) 199 erysiboides (Catillaria, Lecidea) 20, 114, 198 euphorea (Lecidella) 204 exigua 170 exilis (Arthonia) 196, 197, 206 eximia 24, 29, 40*, 61-64, 71*, 87, 90, 91, 95, 100, 107, 124, 134, 164, 168, 205 flotowii 186 fraterculans 170 friesiana 128 friesiana (Argopsis) 85 frullaniae (Catillaria) 196, 197, 206 fucatus (Mycoblastus) 88 fuliginea (Lecidea, Micarea) 96, 198 fusca, spododes var. 165 zit fusca, synotheaf. 128 fuscatula (Bacidia) 196 Fuscidea 22,97, 185 fuscoatra (Lecidea) 97, 198 fuscopurpurea (Arthonia) 31 gelatinosa (Lecidea, Micarea) 198 geomaea 142 geophana (Steinia) 98 glebosula 174 globifera 158 globularis 49*, 158 globulosa (Catillaria) 66, 204, 205 globulosella 22, 23, 41*, 64, 74*, 85, 88, 90, 95, 96, 99, 101, 134, 135, 166, 180, 189, 190*, 198, 206 glomerella 131 gomphillaceum 63, 96, 143, 144 graniforme (Cliostomum) 67 granulosa (Lecidea) 87, 98, 198, 200, 205 granvina 149 Graphidaceae 31 griffithii (Cliostomum) 67, 159, 199 gymnomitrii (Bryomyces) 143 hardangeriana 185 hedlundii 24, 31, 40*, 64, 75*, 85, 88, 90, 91, 95, 100, 102, 108, 114, 135-137, 140, 175 hegetschweileri (Bacidia) 205 hellbomii 186 Helocarpon 96, 126 hemipoliella 39*, 127, 131 hemipolioides 170 Herteliana 97 Huilia 26,97 humida (Biatora, ?Moelleropsis) 205 hylocomii (Micarea) 200 hymenea (Pertusaria) 88 hypnaea (Lecidea) 197, 198 hypnorum (Lecidea) 31, 88, 115-116, 204 hypocyanea_ 186 hypoleuca 122 hypopta (Lecidea) 132 icmalea (Lecidea) 87, 198, 205 igniarii (Bacidia) 206 ilyophora 151 incincta 186 incrassata 22, 23, 25, 28*, 29, 42*, 61, 64, 67, 72* , 88, 89, 95, 96, 100, 104, 115, 116*, 137, 138, 204 infidula 117 infuscata 115, 137 intrusa 20, 23, 25, 29, 32, 43*, 61, 64, 85, 87-89, 95, 105, 138*, 139, 140, 184*, 185, 205 irrubata 114 juistense (Anisomeridium) 66, 67 212 laeta 174 lapiseda 170 latens 192 laxula 148 Lecanora 61, 88 Lecanoraceae 98 Lecidea 88, 97 Lecideaceae 19, 31,71, 87,97, 98 Lecidella 22,31, 97 leprosa (Vezdaea, Micarea) 200 leprosula 22, 44*, 64, 85-87, 89, 90, 95, 99, 103, 107, 140-142*, 172, 183, 205 leucococca 170 leucodontis (Bryostigma) 31, 196 levicula (Lecidea) 87, 100 lignaria 20, 22, 23*, 24, 27*, 29, 31, 45*, 61, 63, 64, 66, 76*, 84-90, 92-96, 99, 104, 122, 126, 129, 141, 142-145*, 146, 166, 170-172, 191, 192, 197, 201, 204—206 limosa (Lecidea) 98, 115-116, 199, 204, 205 lithinella 24, 28*, 32, 46*, 61, 64, 85, 89, 92, 93, 95, 100, 105, 118, 147, 148, 150*, 204—206 littorella (Catillaria, Lecidea) 197*, 199 livescens 170 livida B. de Lesd. 165 livida Korber 170 lobariella (Dactylospora) 88 lobulata (Toninia) 145, 205, 206 longior 128 lucida (Psilolechia) 202 lutulata 24, 25, 29, 46*, 64, 65, 75*, 88, 89, 95, 100, 106, 118, 120, 148-150*, 161, 187, 198, 204-206 lynceola 117 major 128 malmeana 134 meizospora 143 melaena 22,29, 47*, 61-64, 77*, 85, 87-96, 100, 103, 107, 124, 150—153*, 154, 165, 171, 187, 195, 204—206 melaenida 22, 24, 29, 48*, 64, 88-90, 92, 95, 100, 106, 115, 154, 155, 195, 204-206 melaeniza 24, 29, 47*, 64, 65, 69*, 85, 90, 91, 95, 99, 106, 108, 114, 120, 155, 156, 159, 161, 164 melanobola 24, 31, 47, 61, 64, 77*, 85, 88, 90, 95, 100, 102, 156, 157, 175 melanochroza 158 Melanolecia 97 melaphana_ 43*, 138 meridionalis 154 Metacapnodiaceae 67 Micarea 20, 96-100 Micareaceae 98 microcarpa 165 micrococca 173-176 microspora 117 milliaria 84, 142 minima (Micarea) 200 BRIAN JOHN COPPINS miniuscula (Bacidia, Micarea) 196, 198 misella 24, 29, 31, 49*, 61, 64, 65, 67, 85, 88, 89, 90, 94-96, 100, 102, 107, 114, 120, 156, 157, 158-160* , 164, 175, 181, 182, 204-206 monticola (Lecidea) 205 moriformis 125 muhrii 24, 29, 49*, 61, 64, 88-91, 95, 106, 148, 149, 156, 160, 161 myriocarpa 24, 29,31, 32, 49*, 62, 64, 78*, 88-90, 95, 105, 106, 148, 149, 161, 162*, 163 myrtillicola (Bacidia) 196, 197* naegelii (Bacidia) 172,206 niger (Lichen) 200 nigra (Micarea) 96, 200 nigra, melanobolaf. 174 nigra, spododes var. 165 nigella 24, 29, 31, 50*, 61-64, 78*, 87, 90, 91, 94, 95, 100, 107, 108, 114, 120, 124, 134, 156, 159, 163, 164*, 165 nigrata Mill. Arg. (Bacidia, Micarea, Patellaria) 200, 201, 205 nigrataNyl. 143 nigroclavata (Catillaria) 204 nigrum (Collema, Placynthium) 200 nitschkeana 22, 23, 30, 50*, 64, 66, 67, 85, 87—96, 99-101, 129, 135-136, 165-167*, 171, 172, 189, 204-206 niveoatra (Opegrapha) 66, 67 Nostoc 25,96, 114, 155 obscurior 117 occulta 174 ocelliformis (Lecidea) 199 oculata (Pertusaria) 88 olivacea 24, 29, 31, 50*, 62—64, 78*, 84, 89, 90, 95, 100, 107, 124, 134, 164*, 165, 167, 168, 193 osloensis 29, 31, 54*, 61, 64, 88, 89, 95, 106, 169, 195 pallida (Lecidea) 26 panaeola (Huilia) 25, 140 parissima 127 paucula 149 pauxilla (Lecidea) 199 peliocarpa 22-25, 27*, 29, 51*, 61, 63-66, 79*, 85-87, 89-91, 93-96, 99, 103, 104, 108, 109, 111, 112, 122, 129, 141, 144, 149, 166, 169-173*, 191, 200, 204-206 perfurfurea (Pannularia) 198 perpusilloides 196, 197* phaeobaea (Arthonia) 67 Phyllopsora 22, 85 Placynthiella 198 poliococca 131 poliococcoides 131 poliodes 148 polytrichi 53*, 80*, 96, 174, 176 polytropoides 22, 45*, 143 LICHEN GENUS MICAREA IN EUROPE populina (Lecidea, Micarea) 31, 32 praeviridans 127 prasinaFr. 20, 23*, 24-26, 29, 30, 32, 52*, 53*, 61, 64, 66, 80*, 85—96, 100, 102, 108, 109, 129, 136, 148, 157, 173-176*, 177*, 178*, 179, 198, 204—206 prasina Tuck. & Mont. (Bacidia, Biatora) 203 prasinata (Bacidia, Biatora) 203 prasiniza 53*, 109, 173 prasinoleuca 174 pruinosum (Scoliciosporum) 30, 97, 180 pseudoglomerella 128 Psilolechia 98, 202 pulverula 125 pycnidiophora 23, 29, 54*, 64, 65, 67, 85, 89, 90, 93, 95, 99, 103, 108, 114, 179, 180*, 181, 182 pyrenothizans 127 quercicola(Lecanora) 66 radiata (Arthonia) 31 recondita (Lecidea) 199 relicta 154 rhabdogena 24, 29, 31, 55*, 64, 85, 88, 90, 91, 95, 99, 105, 133, 164, 181, 182, 205 rhapidophylli (Bacidia) 196 rhodosphaera (Catillaria) 197 Rhopalospora 97 rosei(Phyllopsora) 85, 146 rosella (Bacidia) 97 rusticella 117,118 sabuletorum (Bacidia) 145, 204-206 salevensis (Bacidia) 171 sanguineoatra (Lecidea) 115 saprophila 151 saxicola 142 saxigena Hepp 169, 170 saxigena Leighton 143 scandinavica 192 schadeanum (Scoliciosporum) 180 schumannii 48*, 154 Scoliciosporum 97, 98, 103, 140 semialbula 128 semipallens 117 simplicata 131 simplicior (Lecidea, Bacidia) 199, 200 smaragdina 186 sordidescens 173,174 sororians 131, 133 spadicea (Arthonia) 67 sphaeroides (Bacidia, Bilimbia, Catillaria, Lichen) 20, 112, 197, 198, 205 spodiza 127 spododes 165 Sporacestra 203 Steinia 98 Stenhammarella 197 stenotera (Lecidea) 98, 115-116, 205 213 Stereocauliscum 96 Stigonema 25, 96, 114, 115 stipitata 23, 29, 55*, 64, 65, 67, 89, 91, 93-95, 99, 103, 108, 114, 180*, 182 stizenbergeri 151 straminea (Lecanora) 192 Strangospora 98 subabietina (Lecanactis) 67 subassimilata 125 subdiscordans (Byssoloma) 98 subfuscula (Bacidia) 192 subinfidula 192 subleprosula 20, 24, 31, 32, 56*, 64, 85-87, 89, 94, 95, 99, 103, 107, 141, 182, 183, 191* sublivescens 127 sublivida 186 submisella 127 subnigrata 22, 23, 29, 32, 57*, 61, 64, 67, 81*, 88-90, 95, 105, 127, 139, 183, 184*, 185 subrudecta (Bacidia) 201 subviolascens 20, 22, 23, 25, 29, 57*, 64, 88, 89, 95, 100, 101, 115, 185, 186, 205 subviridescens 52*, 173,174,177 subviridicans 201 sylvicola 20, 23-25, 29, 31, 57*, 61, 62, 64, 66, 82* , 87-90, 93, 95, 96, 100, 107, 118, 149, 152, 186-—188*, 193, 194, 199, 204—206 symmicta (Lecanora) 30, 98, 206 synothea (Lecidea) 128, 199 synotheoides 24, 32,57, 58*, 64, 74*, 88, 90, 91, 93-95, 100, 102, 157, 180, 188-—190* tenebrosa (Schaereria) 88 tetnaria 22,23, 29, 59*,-63; 64; 83", 89)'95, 96, 99, 104, 126, 144, 171, 190, 191*, 192 terrestris (Lecidea) 199 tetramera (Bilimbia) 112 Toninia 22 trachona (Bacidia) 67, 206 Trapelia 199 Trapeliaceae 199 Trapeliopsis 199 Tremolecia 97 trisepta 170,171, 200 triseptataNyl. 143 triseptata Hepp (Bacidia) 143 triseptatula 170 triseptatuloides 170 tuberculata 24,25, 29, 31,59*, 61, 62*, 63, 64, 82*, 84, 87, 89, 90, 95, 100, 107, 163, 168, 187, 192, 193*, 194, 204-206 turfosa Massal. 24, 29, 32, 60*, 62, 64, 88, 89, 95, 96, 103, 106, 155, 194, 195*, 196, 199, 204—206 turfosa Fr. 47*, 150 turgidula (Lecidea) 30, 168, 204—206 Tylothallia 97 uliginosa (Lecidea) 169, 206 umbrinum (Scoliciosporum, Bacidia, 214 BRIAN JOHN COPPINS umbrinum — cont. vinosa (Arthonia) 204 Lecidea) 25, 92,97, 127, 128, 138, 140, 205, violacea 51*, 170 206 violacea (Bacidia) 170 umbrosa 149 viridescens (Lecidea, Micarea) 198, 200, 204, 205 viridis (Protococcus) 174 vainioi 186 vulgaris 128 vermicillifera (Opegrapha) 67 vulgata(Opegrapha) 67 vernalis (Lecidea, Biatora) 20, 205 verrucula 194 xylophila 143 vezdae (Bacidia) 196 Vezdaea 24, 30, 98 zsakii 22, 48*, 154 British Museum (Natural History) Seaweeds of the British Isles The synthesis of many years’ research carried out by the British Museum (Natural History) and the British Phycological Society, this is the second in a series of books that will be published under this title covering all the British and the majority of northern Atlantic seaweeds. Volume 1 Rhodophyta Part 2A Cryptonemiales (sensu stricto), Palmariales, Rhodyme- niales. Linda M. Irvine Following the style and format laid down in Part 1, this book deals with the species attributed to the orders Cryptonemiales, Palmariales and Rhodymeniales. Each species description incorporates notes on ecology and distribution and is supported by one or more line illustrations. Keys to aid identification are also included. When complete, this title will be the standard work of reference in its field; it will provide students and researchers with a digest of the most up-to-date and compre- hensive information available on the marine algae of Great Britain and Ireland which will be indispensable throughout the north Atlantic region and beyond. ISBN 0 565 00871 4 128 pagesillustrated £13.00. Titles to be published in Volume 11 The algae of Lightfoot’s Flora scotica. By Peter S. Dixon. A taxonomic study of the lichen genus Micarea in Europe. By B. J. Coppins. The hepatics of Sierra Leone and Ghana. By E. W. Jones and A. J. Harrington. Studies in the Corallinaceae with special reference to Fosliella and Pneophyllum in the British Isles. By Y. M. Chamberlain. Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, Dorchester Bulletin of the British Museum (Natural Hite) The hepatics of Sierra Leone and Ghana EE. W. Jones & A. J. Harrington Botany series Vol 11 No 3 24 November 1983 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), 1983 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 11 No 3 pp 215-289 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 24 November 1983 The hepatics of Sierra Leone and Ghana | E. W. Jones The Green, Kirtlington, Oxford OXS 3HJ A. J. Harrington a Department of Botany, British Museum (Natural History), Cromwell Road, London SW7 SBD AM Contents BEV ERS RE Re parce nnodraccr os ta outs Sewers ese enn wcsetaaa te ono at dee enn t ceamnneetennecsr ons oa: POLE La a LT AL (01) BARE eae OEE Re ANE PEERS trl cy gc Ae gr eins 7S EE ROL ESA Se eG UREN AAT Bn Rene BOS ae OCIS a ce recy ae ee cn eae cb nC wo me oan) Spee te BR COCR ST rss ee hoo aaa s eS aE MG Ceara inh Sule wi Ga bnew Coes tunes DMS csr fealty ose ena Ho ee nse Contant Ait cn crmaveae suvuawura anne ete ree ae PUCRINIWIOURCIIOIS occ 22 feo oe cee ares chs Se acaben tee penta dubanguadesdacasatceceveeeuretts’ MORO COS oe oie ee a Ek Lasaeadn cud ce hen aed Adee aah push iaered coreaeca ves otiNendints ben Synopsis The vegetation and hepatic flora of Sierra Leone and Ghana are briefly described. Detailed information on the distribution of 178 taxa is presented, together with notes on their ecology and taxonomy. Of these taxa, 95 have been collected in both countries, 44 are known from Sierra Leone but not Ghana, and 39 have been found in Ghana but not Sierra Leone. Many of the taxa recorded only in Sierra Leone are montane in distribution and therefore unlikely to occur in Ghana. On the other hand it is probable that most of those at present recorded only from Ghana will eventually be found in Sierra Leone. Three new species are described and illustrated: Cololejeunea calcarata E. Jones and Drepanolejeunea ankasica E. Jones from Ghana, and Lejeunea lomana E. Jones from Sierra Leone. The following are placed in synonymy: Ceratolejeunea saxbyi Pearson (= C. calabariensis Stephani); Lejeunea eplicata Stephani (= L. ramosissima Stephani); Lejeunea triquetra Mitten and Mastigolejeunea tarkwana Pearson (= Thysananthus spathulistipus (Reinw., Blume & Nees) Lindenb.); Prionolejeunea aberrans Stephani (= Lejeunea papilionacea (Stephani) E. Jones); Taxilejeunea nicholsonii Pearson (= Lejeunea grossecristata (Stephani) E. Jones), and Riccia undulata S. Arnell (= R. moenkemeyeri Stephani). Introduction During the nineteenth century various botanists who collected bryophytes elsewhere in Africa made brief visits to Sierra Leone (mainly Freetown) or, less frequently, the Ghana coast, but in general they seem to have collected mosses rather than hepatics; they have left very few specimens of hepatics from either country in our large herbaria, and published records are equally scanty. Since 1910 a number of botanists, agriculturalists, etc. (see index of collectors), some resident, some as visitors, but most with no special knowledge of bryophytes, have made small collections of hepatics, the majority of which we have identified in recent years. Some of these consist of only one-three specimens; those of Hossain, Irvine, and Richards are the most considerable, apart from Jenik’s, which we have not seen in its entirety. One of us (A.J.H.) worked in the Botany Department of Fourah Bay College, Freetown, from March to December 1966. He was engaged primarily in studying the ecology of epiphyllous hepatics, but also collected other bryophytes, to a total of about 1000 specimens. E.W.J. collected bryophytes in Ghana from 23 January to 25 February 1971 (collecting, numbers 1214-1418). Working in the Botany Department of the University of Ghana at Legon, near Accra, and with the facilities provided by the Department, he was able to visit a Bull. Br. Mus. nat. Hist. (Bot.) 11 (3): 215-289 Issued 24 November 1983 216 E. W. JONES & A. J. HARRINGTON representative selection of bryophyte-rich sites ranging over the whole of southern Ghana. He collected in Sierra Leone between 28 February and 1 April 1971 (collecting numbers 1419- 1579). Fourah Bay College also served as his base, but he paid brief visits to the Loma Mountains, the Kambui Hills, and the Gola Hills in the east of the country. In both countries the period February—March falls in the latter part of the dry season — an advantage in so far as the weather is favourable for travelling, but a disadvantage in that some genera of terrestrial and ephemeral species such as Fossombronia and Riccia cannot be found, and sporogonia of many other species have too often disintegrated. The early dry season (October-November) is amuch more favourable time for collecting bryophytes in these climates. Our own collections form the most substantial contribution to our knowledge of the hepatic flora of the two countries. A few specimens are still unidentified and some taxonomic problems remain to be solved, but further progress is likely to be very slow, and the residue which remains to be studied is small. E.W.J. has cited some of his own specimens, together with a few from the small collections mentioned above, in his recent papers in the series ‘African hepatics’ (J. Bryol. 7-12, 1972-82). No comprehensive account of the hepatic flora of any West African territory exists, and the only publication dealing exclusively with the hepatics of either Sierra Leone or Ghana is a post- humously published paper by Pearson (1931) describing a small collection made by H. H. Saxby near Tarkwa, Ghana, in 1910-11. Therefore it has seemed worth preparing a list of the hepatics of the two countries based primarily on our own collections, but incorporating information from the other sources that we have indicated; this provides a picture of the hepatic flora of two representative, but in certain respects contrasting, West African countries. Sierra Leone Sierra Leone lies between 6°55’ and 10°N and between 10°16’ and 13°18’W. Most of the central part of the country has a mean annual rainfall of 2500-3000 mm (100-120 ins). This increases markedly towards the coast and shows a contrasting, though less pronounced, decrease towards the north. Even the driest areas of the country receive 2000 mm — more than all except the wettest parts of Ghana and Nigeria — but the dry season is longer and more severe than in these two countries. Thus Kabala, in the relatively dry north, with a mean annual rainfall of 2285 mm, is shown by Walter & Lieth (1967) as having a ‘dry’ period* of four months, and Bo, representative of much of the central lowlands, with a mean annual rainfall of 2937 mm, has a ‘dry’ period of three months. Much of the country is situated ‘within the climatic limits of closed forest’ (Cole, 1968) but, except in the Freetown Peninsula and some remote hill country in the east, no mature forest remains. Shifting cultivation prevails everywhere, but in the southern half of the country fallow farmland tends to revert to forest, whereas in the drier northern half it tends to revert to savanna. We have very little information on the bryophyte flora of this large area of farmland and its derivative vegetation; a few collections (none made by bryologists) from Njala and Bo come from the moister part of the country with ‘forest’ tendency, and some from Musaia, Kabala, and the Lake Sonfon area come from the drier part with ‘savanna’ tendency. In his vegetation map, Cole (op. cit.) shows a belt of savanna woodland some 20-50 km wide along the northern and north-eastern borders of the country. The hepatic flora of this woodland is unknown but is likely to be very poor. The Freetown Peninsula, bryologically the best known part of the country, is also the wettest. Freetown itself has an average of about 3340 mm of rain a year, but the mountain ridge which rises sharply to altitudes of 600-800 m receives more than 5000 mm. The Peninsula, however, also has the longest and most severe dry season of any part of the country, and even Number * Walter & Lieth plot rainfall and temperature so that 10°C on the temperature scale corresponds to 20 mm rain on the rainfall scale; a ‘dry’ period is one in which the precipitation curve lies below the temperature curve, thus indicating the likelihood of vegetation suffering a moisture deficit. The severity of the dry period is indicated both by the length of such periods and by the distance below the temperature curve to which the rainfall curve descends. Where only monthly mean rainfall figures, but no temperature figures are available, a crude indication of the length of dry season is given by the number of consecutive months with less than some convenient arbitrary amount of rain. Hall & Swaine (1976) use 100 mm, but a lower figure, e.g. 50 mm (approximately 2 ins) is more sensitive. HEPATICS OF SIERRA LEONE & GHANA ley Two River, with a mean annual rainfall of 5490 mm has five consecutive months each with 50 mm or less rain (Gregory, 1965). The mountains are still clothed with forest in which bryophytes grow with great luxuriance, but the number of species is small. The almost complete absence of Frullania spp. is a remarkable feature of the flora which presumably has a climatic cause. Other species which might be expected but have not been recorded are Bazzania decrescens subsp. molleri, Mastigolejeunea auriculata, M. nigra, Porella subdentata, and corticolous species of Radula. Despite the length of the dry season epiphyllous hepatics are abundant in deep valleys where perennial streams maintain a high local humidity, as in the Fourah Bay College Botany Re- serve. Towards the interior of Sierra Leone the general level of the land rises; much of the eastern half of the country lies above 300 m, and there are some groups of hills where the most important of the remaining areas of forest are situated. Hepatics have been collected in the Kambui Hills Forest Reserve to the north of Kenema, and in the Gola Hills to the south-east, close to the frontier with Liberia. Though the hepatic flora is probably richer than that of the Freetown Peninsula it seems to show some of the same curious limitations; thus after a day spent in the Gola North Forest Reserve E.W.J. noted ‘No Mastigolejeunea or Ptychocoleus were seen, and no Frullania’, and similarly in the Kambui Hills E.W.J. noted that holostipean Lejeuneaceae, other than Lopholejeunea and Archilejeunea, were ‘almost absent’, and Frullania ‘not seen’. In eastern Sierra Leone, near the border with Guinea, rise the Loma Mountains — the highest mountains in Africa west of Cameroon — and, smaller in extent but only about a hundred metres lower, the Tingi Hills. Both consist of undulating plateaux at 1220-1520 m, with peaks rising to about 1900 m. Bintimani (1948 m), the highest peak of the Loma Mountains, has a small summit plateau surrounded by imposing dolerite cliffs up to 80 m high. Hepatics found on the plateau and cliffs include Fossombronia husnotii, Gongylanthus richardsii, which is otherwise known only from basic volcanic soils in Cameroon and Tanzania, and Plagiochasma eximium. Some records from Bintimani are included in a short paper by Potier de la Varde (1948) on the mosses of the Loma Mountains. The main plateau of the Lomas is covered by grassland (the ‘sub-montane shrub savanna’ of Cole (1968)) which is frequently burned and therefore devoid of bryophytes, except sometimes Anthoceros sp., Fossombronia sp., Gongylanthus ericetorum, and various small mosses between grass tussocks. However, ‘gallery’ forest extends up the valleys to about 1650 m and there are also small trees and bushes on rock outcrops. The edges of the forest are punctuated in many places by very large trees of Parinari excelsa which evidently create ‘fire-breaks’ by reducing the amount of grass that can grow beneath them. The sides of these large rough-barked boles which face the grassland are well illuminated and bear a remarkable bryophyte flora which includes some species not seen in other habitats; particularly noteworthy are Chandonanthus hirtellus and Conoscyphus trapezioides, tropical montane species with wide but very disjunct distribu- tions. The understorey of the forest has a rich flora of shade-tolerant bryophytes, amongst which Odontolejeunea tortuosa is conspicuous. Exposed branches of small trees and bushes growing amongst rocks are clothed with Usnea spp. and other lichens, and with hepatics that include Frullania arecae, a pantropical montane species, F. depressa, first described from Mont Cameroun (Cameroon Mountain), but known from many other African mountains ranging from Ethiopia to Madagascar and South Africa, and Lejeunea ramosissima. Brachiolejeunea tristis, Dicranolejeunea madagascariensis var. madagascariensis, Marchesi- nia moelleriana, Ptychanthus striatus, and Strepsilejeunea brevifissa are other characteristic members of the African montane flora which have been recorded; it is remarkable that Lejeunea acuta Mitten has not been found. The Tingi Hills have not been visited by bryologists but it is to be expected that their flora will resemble that of the Loma Mountains; in both areas Polytrichum commune Hedwig has been found growing on the fringes of grass mats overlying gently sloping slabs of rock. Sphagnum davidii Warnstorf was discovered in the Tingi Hills by J. K. Morton and D. Gledhill in 1965. 218 E. W. JONES & A. J. HARRINGTON Bryophytes have never been collected in the extensive forest reserves on the lower ground to the west and south of these two massifs. Ghana Ghana extends from 4°45’ to 11°10’N and from 1°12’E to 3°15’W. North of about 6°30’N in the east and about 7°45’N in the west the country is covered mainly by savanna woodland (Taylor, 1960). From this area we have only three records of hepatics —- Fossombronia sp. (probably F. occidento-africana) , Mastigolejeunea auriculata, and Targionia hypophylla. Though the flora is unlikely to be rich, it will doubtless resemble that of the corresponding savanna in Nigeria, characterized by the more drought-resistant Lejeuneaceae such as Acrolejeunea emergens and Mastigolejeunea auriculata on trees, and on the ground by ephemeral species that perennate either by spores or tubers (e.g. Riccia spp.). To the south is the forest region, to which almost all collecting of bryophytes has been confined. Hall & Swaine (1976) recognize seven main divisions within the forest on the basis of an objective analysis of its phanerogam flora; these divisions correspond to some extent with rainfall regimes. In the south-west is a lowland area of ‘Wet Evergreen’ forest (‘Guineo- Congolian Rain forest, wetter types’, UNESCO—AETFAT vegetation map of Africa (White, in press)) which extends westwards into the Ivory Coast, with a mean annual rainfall exceeding 1750 mm. Hall & Swaine point out that it has a ‘wetter rainy season’ than the rest of Ghana, with the mean rainfall for some months exceeding 500 mm. Going eastwards from Ghana no other lowland region with as high a rainfall is encountered until the extreme south-east of Nigeria and Cameroon is reached. The Ankasa River Forest Reserve and Aiyinasi Agricultural Research Station are in this zone. They have a rich hepatic flora which includes Cololejeunea cornuta and Trachylejeunea serrulata, two species that have hitherto been known only from the Calabar (S.E. Nigeria)-Cameroon region. East and north-east of the Wet Evergreen forest, and still mainly lowland, is a zone of ‘Moist Evergreen’ forest with 1500-1750 mm mean annual rainfall. Saxby’s collections from Tarkwa, and E.W.J.’s collections from the Subri and Pra-Suhien Forest Reserves, come from this zone, while the Ochi Headwaters Forest Reserve is on its northern boundary. Still further to the north and east is a broad zone of ‘Moist Semi-deciduous’ forest, the most extensive forest region of Ghana, covering the broad belt of high ground which crosses southern Ghana from south-east to north-west. Much of this zone is between 150 and 300 m above sea-level; to the north-east it is bounded by a scarp which, over much of its length, controls the boundary between forest and savanna. Hall & Swaine divide this Moist Semi-deciduous forest into a ‘South-east sub-type’, with 1500-1750 mm mean annual rainfall (thus not appreciably different from that of the Moist Evergreen zone) and a ‘dry’ period (see footnote on p. 216) of usually four consecutive months each with a mean rainfall of less than 100 mm, and a distinctly drier ‘North-west sub-type’, with 1250-1500 mm mean annual rainfall and a ‘dry’ period of usually five months. The hepatic flora of the South-east sub-type is exemplified by Aburi, Kade Agricultural Research Station, and the neighbourhood of the Tafo Cocoa Research Institute, including the Southern Scarp Forest Reserve; that of the North-west sub-type by the Bia North Forest Reserve and the Krokosua Hills. The hepatic flora of the South-east sub-type of the Moist Semi-deciduous forest closely resembles that of the Moist Evergreen forest; a few species have been recorded from the Moist Evergreen forest but not from the Moist Semi-deciduous; probably mostly the result of insufficient collecting, but the absence of Arachniopsis diacantha, Cephalozia fissa, Plagiochila africana, and Thysananthus spathulistipus is likely to be due to climate. The hepatic flora of the drier North-west sub-type seems to be very much poorer. The contrast was particularly striking between the corticolous flora of the crowns of emergent trees in the Aiyaola Forest Reserve near Kade (12 species) and in the Bia North Forest Reserve, where only four species — Acrolejeunea emergens, Cheilolejeunea intertexta, Frullania ericoides, and Masti- golejeunea auriculata, all drought-tolerant — were found. Here also the boles of cocoa trees were almost devoid of bryophytes, whereas near Tafo at least six species of hepatic could be found ina cocoa plantation. HEPATICS OF SIERRA LEONE & GHANA 219 In general the flora of these two zones of Moist Evergreen and Moist Semi-deciduous forest closely resembles that of southern Nigeria, as seen for example in the forest reserves around Akure and Benin. The abundant species are the same, and all but seven (i.e. about 90 per cent) of the species that have been recorded from the two Ghana zones are known from Nigeria west of the Cross River; of the other seven, five (Cheilolejeunea trifaria, Cololejeunea dentata Prionolejeunea grata, Radula holstiana, and Taxilejeunea pulchriflora) are known from east of the Cross River, either in Nigeria or Cameroon, and are likely to occur to the west. Hall & Swaine find that within the general area of the South-east sub-type of the Moist Semi-deciduous forest certain high plateaux have a floristically distinct forest which they name ‘Upland Evergreen’ forest. The Atewa Hills (c. 730 m) is the best known of these botanically, and the only one from which bryophytes have been collected. The hepatic flora includes some montane species — Plagiochila pectinata, Radula boryana, and R. stenocalyx — which are not known from the surrounding country. One small patch of swamp forest is said to be the only site of its kind in the Atewa Hills. Between dominant Lophira alata and Gilbertiodendron limba water stands 20-30 cm deep. Tree boles and the stems of the abundant undershrubs are thickly clothed with bryophytes, amongst which ramify many small and delicate Lejeuneaceae; epiphyllous hepatics are luxuriant. At some time trial pits have been dug here to explore deposits of kaolin; hence it is referred to as the ‘Kaolin Swamp’ to distinguish it from other less remarkable areas of wet forest. A thunder-storm and failing light greatly curtailed E.W.J.’s collecting when he visited it; it would certainly be worth much closer examination. The Krokosua Hills form a plateau at 450-600 m in the drier North-west sub-type of the Moist Semi-deciduous forest; although they do not carry Upland Evergreen forest, the bryophyte flora seems much richer than that of the surrounding lower ground. In eastern Ghana a narrow line of hills rises abruptly to 760-880 m, forming the frontier with Togo. One of these hills, Amedzofe (760 m), with its spectacular waterfall, has been frequently visited by botanists, and is presumably in the Moist Semi-deciduous forest zone; although no forest now remains, the bryophyte flora of the cocoa plantations of the district closely resembles that seen near Kade and Tafo. The summit of the hill bears grassland on the flank exposed to the south and south-west, with a small patch of savanna woodland in the lee of a rock outcrop. Other higher hills further north are covered entirely by savanna and have not been visited by collectors of bryophytes. We include in this paper a few records from collections made in these hills near Klouto in Togo, only a few kilometres from the Ghana frontier. We know of no bryophyte collections from Hall & Swaine’s remaining forest-types — the ‘Dry Semi-deciduous’, ‘Southern Marginal’, and ‘South-east Outlier’ — but their hepatic flora is probably an impoverished version of the Moist Semi-deciduous forest flora. The coastal belt of eastern Ghana, in which lie Accra, Legon University, and also Achimota — the seat of botanical studies in Ghana before Legon was opened — receives an average of only 630-890 mm of rain a year; it is the driest part of Ghana, though the dry season is shorter and less severe than that of the northern savanna. Little attention has been paid to the hepatics in this area, and only Anthoceros buettneri, Notothylas spp., Riccia rhodesiae, and Ricciocarpos natans have been recorded. Though we should expect more species to be found, they are likely to be mainly terricolous. A botanist who has worked in the forests of other parts of tropical Africa is likely to be impressed in Ghana both by the luxuriance of the forest and the richness of the bryophyte flora, especially in the Moist Semi-deciduous forest. The total annual rainfall is relatively low, but probably exercises less influence on the bryophyte flora than the frequency, severity, and length of dry periods, and in Ghana the dry periods appear to be relatively short. Thus, to take figures from Walter & Lieth (1967), Tafo, in the Moist Semi-deciduous forest, has a mean annual rainfall of 1658 mm, and a ‘dry’ period of less than one month, and even Kumasi, near the border between the Moist and the Dry Semi-deciduous forest, with a mean annual rainfall of 1481 mm, has a ‘dry’ period of only two months. We may contrast these figures with Calabar, 3028 mm mean annual rainfall and a ‘dry’ period of two months, or Benin with a mean annual rainfall of 1975 mm and a ‘dry’ period of three months — wetter than Kumasi, but with a longer dry season. A comparison of monthly rainfalls in the forest regions of Ghana and Nigeria shows that the 220 E. W. JONES & A. J. HARRINGTON two-peak rainfall regime is more pronounced in Ghana, thus tending to spread a given amount of rain over a longer period. The savanna zone of Ghana, where the rainfall is of the single peak type, also has more moderate dry seasons than comparable Nigerian savanna: e.g. Tamale, 1100 mm mean annual rainfall, four consecutive months with less than 25 mm; Minna, on the same latitude in Nigeria, 1380 mm mean annual rainfall, five consecutive months with less than 25 mm. Nomenclature and the citation of literature Author abbreviations follow the Draft index of author abbreviations compiled at the Herbarium, Royal Botanic Gardens, Kew (1980). We have given additional bibliographic references for many of the taxa included in the catalogue; these mostly refer to descriptions more detailed than those provided in the original publications. Apart from new synonyms, we have cited only the principal synonyms that have been used in connection with the African flora. Nomenclature of flowering plants follows Hutchinson & Dalziel (1954-72). Identification and the citation of specimens Except where otherwise indicated E.W.J. has named his own collections from both countries, all other collections from Ghana, and specimens collected in Sierra Leone by Arnell, Brenan, Revell, and Richards. A.J.H. has identified his own collections, unless otherwise indicated, and the small collections from Sierra Leone housed at the British Museum (Natural History). In general, localities are cited from west to east and from north to south; we have not cited them by administrative districts as these tend to be impermanent and have indeed undergone considerable changes in both countries during the period over which records have been accumulating. Records from hilly districts have been arranged in order of altitude, from the lowest to the highest. Where there are numerous records of a given taxon from one locality, we have sometimes cited only a selection. The index of collectors will usually indicate sufficiently closely the date of collection and the herbarium in which the specimen is preserved. E.W.J.’s collections are in his private herbarium except where the collecting number is followed by a herbarium abbreviation (see below); in such instances the collection is not represented in his private herbarium. ! indicates a field record unsubstantiated by a herbarium specimen; such records have been cited only when there is no possible doubt as to identity. The herbarium abbreviations used are those adopted by Holm- gren, Keuken & Schofield (1981). Abbreviations A.R.S. Agricultural Research Station BR. Botany Reserve Ek. Forest Reserve p-p. pro parte — in part (where a specimen includes more than one taxon) s.n sine numero — without a number Index of localities The index gives the latitude, longitude, and, in most cases, altitude of all localities mentioned more than once in the catalogue. Not all of the figures are as precise as we would wish; e.g. some of the forest reserves are of considerable area and the exact point within them at which collections were made is not recorded. Sierra Leone Co-ordinates are taken from 1:50,000 maps published by the Directorate of Overseas Surveys for the Sierra Leone Government, Series G742 (D.O.S. 419), 2nd ed. 1964-73. HEPATICS OF SIERRA LEONE & GHANA 221 Bagru River Bambawo — Forest School and collecting area Bo Freetown Peninsula Bathurst Charlotte Falls Fourah Bay College Botany Reserve Guma Valley Havelock Plateau Hill Station Kongo Dam Leicester Leicester Peak Lumley Cove Mount Aureol (Fourah Bay College) Mountain Torrent Picket Hill Regent Sugar Loaf Gap Sugar Loaf Mountain Toke Toke-valley three km to the north-east York York Pass Giema Gola Hills — see Lalehun Gola North F.R. Jawo Kabala Kasewe F.R. (= Kasewe Hills F.R.) Koinadugu Kurubonla Lake Sonfon Lalehun Loma Mountains and surroundings Bintimani (Bintumane, Loma Mansa) Camp 1 Camp 2 Dawule (Da-Oulen) Kongbundu Valley Seradu Sokurela Masimo Musaia Musaia Agricultural Station Musaia-Gbentu road one mile north of river Mongo Njala Ghana 7 43N 12 31W; 0-15 m (0—S0 ft) 8 O1N 11 09W; 300-460 m (1000-1500 ft) 7 58N 11 45W; 90-105 m (300-350 ft) 8 26N 13 12W; 225-255 m (750-850 ft) 8 25 30N 13 12W; 210 m (700 ft) 8 28 30N 13 13 30W; 180-225 m (600-750 ft) 8 22N 13 12W; 270-300 m (900-1000 ft) 8 28N 13 13 15W; 340-370 m (1100-1200 ft) 8 27 30N 13 15W; 210-270 m (700-900 ft) 8 25N 13 12 15W; 385-400 m (1250-1300 ft) 8 27 30N 13 13 15W; 355-415 m (1150-1350 ft) 8 27N 13 13 30W; 595 m (1950 ft) 8 27N 13 17W; 0-15 m (0-50 ft) 8 28 30N 13 13 15W; 240 m (800 ft) 8 28N 13 12 45W; 150-340 m (500-1100 ft) 8 17N 13 07W; 888 m (2913 ft) 8 26N 13 13W; 270-300 m (900-1000 ft) 8 25N 13 13 30W; c. 550 m (1800 ft) 825 15N 13 13 45W; 761 m (2497 ft) 8 18 N 13 11 30W; 0-15 m (0-50 ft) 8 20N 13 10 30W; 45-120 m (150-400 ft) 8 17N 13 11W; 15-60 m (50-200 ft) 8 18N 13 08W; 150-270 m (500-900 ft) 8 02N 11 04W; 180m (600 ft) 731 to 7 45 N 10 44 to 11 04W; 150-370 m (500-1200 ft) 752. N 11 08W; 120-135 m (400-450 ft) 9 35N 11 33W; 430-490m (1400-1600 ft) 8 19 to 8 22N 12 10 to 12 13W; 75-210 m (250-700 ft) 9 32N 11 22W; 520-535 m (1700-1750 ft) 9 12N 10 57W; 430 m (1400 ft) 9 15N 11 31W; 550-565 m (1800-1850 ft) 7 41N 10 58W; 225-240 m (750-800 ft) 9 13 30N 11 07W; 1948 m (6390 ft) 9 13N 11 06W; 1220 m (4000 ft) 9 13 15N 11 07W; 1650 m (5400 ft) 9 10N 11 06W; 1565 m (5140 ft) 9 13 30N 11 07 15W at 1520 m (5000 ft) 9 13N 11 03W; 595-610 m (1950-2000 ft) 9 12N 11 04W; 625 m (2050 ft) 8 43N 11 39W; 180-195m (600-650 ft) 9 45N 11 34W; 340-370 m (1100-1200 ft) 9 44N 11 35W; 370 m (1200 ft) 9 47N 11 35W; 370 m (1200 ft) 8 O6N 12 0SW; 45-60 m (150-200 ft) Co-ordinates are taken mainly from Hall (1980). Aburi Achimota 5 51N 0 11W; c. 430 m (1400 ft) 5 37N 0 14W; c. 60 m (200 ft) 22. E. W. JONES & A. J. HARRINGTON Afram Headwaters F.R. Agogo Aiyaola F.R. Aiyinasi A.R.S. Amedzofe (= Amedzope) Ankasa River F.R. (now Ankasa Game Production Reserve) Asanta Asenanyo F.R. (= Asenanyo River F.R.) Atewa Hills F.R. (= Atewa Range F.R.) Bame Begoro Bia North F.R. (= Bia Tributaries North F.R.) Bosuso Bunso (= Bunsu) Esiama Esukawkaw F.R. Kade Kade A.R.S. Kibi Kpedze Krokosua Hills F.R. Kumasi Legon Mole Game Reserve (now Mole National Park) Mpraeso Nsuta Ochi Headwaters F.R. (= Ochi Headwaters Block 1 F.R.) Oda Osenasi Pampramasi Pra-Suhien F.R. Southern Scarp F.R. Subri F.R. (= Subri River F.R.) Tafo Cocoa Research Institute Tarkwa Todome Vane 708 to 7 15N 132 to 1 48W; 300-370 m (1000-1200 ft) 6 48N 1 05W; 300-610 m (1000-2000 ft) 6 06 to 6 12N 054 to 059 W 5 02N 2 28W; 30-60 m (100-200 ft) 6 51N 0 26E; 610-760 m (2000-2500 ft) 5 10 to 5 25N 2 28 to 2 44W;; c. 60m (200 ft) 4 54N 2 17W; 0-15 m (0-50 ft) 6 18 to 6 35N 2 04 to 2 17W; c. 150 m (500 ft) 5 58 to 6 20N 031 to 0 41W; 610-730 m (2000-2400 ft) 6 40N 0 21E 6 23N 0 23W; c. 610 m (2000 ft) 6 37 to 6 57N 257 to 3 06W 6 18N 025W 617N028W 4 56N 2 21W; 0-15 m (0-50 ft) 6 18 to 6 26N 0 43 to 0 52W; 180-240 m (600-800 ft) 6 05N 0 50W; c. 150 m (500 ft) 6 08N 0.55W; c. 150 m (500 ft) 6 10N 0 33W; c. 300 m (1000 ft) 6 50N 0 30E; c. 150 m (500 ft) 6 19 to 6 38N 2 40 to 2 56W; 300-610 m (1000-2000 ft) 6 41N 137W; c. 225 m (750 ft) 5 39N 0 11W; c. 90 m (300 ft) 9 12 to 10 11N 1 33 to 2 13W; c. 180 m (600 ft) 6 35N 044W 5 16N 159W; c. 30 m (100 ft) 5 43 to 5 44N 1 16 to 1 19W; 150-180 m (500-600 ft) 5 55N 0 59W; c. 120 m (400 ft) 5 57N 045W 6 37N 257W; c. 150 m (500 ft) 5 14 to 5 22N 1 22 to 1 36W; 90-120 m (300-400 ft) 6 12 to 6 45N 0 17 to 0 59W; 300-610 m (1000-2000 ft) 5.08 to 5 29N 1 36 to 1 54W; 90-120 m (300-400 ft) 6 13N 0 22W; c. 150 m (500 ft) 5 18N 1 59W; 90-120 m (300-400 ft) 6 42N 0 22E 6 49N 0 25E; c. 460 m (1500 ft) Index of collectors The index contains the names of all collectors who are mentioned in the catalogue more than once, together with the country(ies) and year(s) in which they collected, and the herbaria where their specimens are preserved. We have given some additional information on those people who are not included in the biographical index of collectors compiled by Hepper & Neate (1971). EW/J denotes the private herbarium of E. W. Jones. Arnell, S. W. Sierra Leone, 1951. S, UPS. HEPATICS OF SIERRA LEONE & GHANA 223 Barter, C. Sierra Leone, 1857. NY (Herb. Mitten), BM. Boughey, A. S. Ghana, 1953. EWJ. Brenan, J. P. M. Sierra Leone, 1948. EWJ. Cummins, H. A. Ghana, 1896. BM, G (Herb. Stephani). Deighton, F. C. Donald, L. H. Foote, Miss V. J. Gardner, Mrs T. S. Gledhill, D. Hall, J. B. Harrington, A. J. Sierra Leone, 1934-54. BM. American postgraduate student of linguistics. Sierra Leone, 1966. BM. Ghana, 1937-39. BM. Sierra Leone, 1958. BM. Sierra Leone, 1966. BM. Ghana, 1959-79 (some with Jones). GC, EWJ. Sierra Leone, 1966. BM, FBC (a few specimens only). Haswell, D. R. Sierra Leone, 1963. BM. H¢geg, O. A. Norwegian lichenologist. Sierra Leone 1929. BM. Hossain, M. Ghana, 1967-69 (some with Richards). GC, EWJ. Irvine, F.R. Ghana, 1935-40; Sierra Leone, 1939. BM. Jaeger, P. Sierra Leone, 1944-45. PC. Jenik, J. Ghana, 1964-7. GC, EWJ, OP. Jones, E. W. Ghana, Sierra Leone, 1971. EWJ, BM, GC, FBC. Jones, T. S. Sierra Leone, 1951. BM. Mann, G. Sierra Leone, 1861. NY (Herb. Mitten), BM. Marmo, V. Sierra Leone, 1955. BM. Marshall, C. British student, member of Durham University Expedition to the Kambui Hills. Sierra Leone, 1966. BM. Morton, J. K. Sierra Leone, 1965-66. BM, FBC. Revell, R. J. British school teacher. Sierra Leone, 1968. EWJ. Richards, P. W. Ghana, 1967-69 (some with Hossain); Sierra Leone, 1971. NMW, GC, FBC, EWJ. Saxby, H. H. British businessman. Ghana, 1910-11. MANCH, CGE, BM. Sellar, P. W. Sierra Leone, 1963. BM. Thomas, N. W. Sierra Leone, 1914-15. BM. Tindall, H. D. Sierra Leone, 1954. BM. West-Skinn, R. Ghana, c. 1957. BM. Species I. ACROLEJEUNEA (Spruce) Schiffner 1. Acrolejeunea emergens (Mitten) Stephani in Engl., Pflanzenw. Ost-Afrikas C: 65 (1895); Gradst. in Bryophyt. Biblthca 4: 71 (1975), var. emergens Ptychocoleus emergens (Mitten) Stephani, Sp. hepat. 5: 24 (1912); E. Jones in Trans. Br. bryol. Soc. 2: 406 (1954). SIERRA LEONE. Freetown Peninsula: Mount Aureol [‘Oriel’], Arnell 2304, 2305, 2308, 2434, 2506 (UPS, fide Gradstein, op. cit.); on mango near Hill Station, Hgeg s.n.; Grafton Valley (8°24’N, 13°10’W), on small tree, Harrington 639. Musaia Agricultural Station, on Uapaca togoensis, Sellar B3. Tekaw, Veterinary Department HQ (8°51'N, 12°01'W), on large tree near the Rest House, Harrington 113, 115. Kabala, near the District Officer’s bungalow, Harrington 136, Jones 1524a. About four km south of Kabala, on small tree in swampy valley, Harrington 118 p.p. Five km south of Kabala, in secondary forest, Harrington 125 p.p. Njala, upper part of trunk of Nauclea diderrichii, Deighton 5313 p.p.; on north side of trunk, especially buttresses, of Amphimas pterocarpoides, Deighton 5603. GHANA. Bia North F.R., near Pampramasi, in crown of large Entandophragma, Jones 1411. Asenanyo F.R., in crown of Terminalia, Jones 1413a. Kumasi, in garden, Irvine 18. Pra-Suhien F.R., small tree by the Rest House, Jones 1348. Esukawkaw F.R., Jenik (Herb. Jones). Kade, bole of fallen tree in forest, Jones s.n. Aburi, Jones s.n. A lowland species, widely distributed in tropical Africa, with a preference for isolated trees, open woodland, and the crowns of trees, and climates with a strong dry season. The absence of records from the wettest parts of Sierra Leone and Ghana is noteworthy. It is also remarkable 224 E. W. JONES & A. J. HARRINGTON that neither of us gathered it near Mount Aureol (Fourah Bay College), where Arnell found it frequently in 1951. var. confertissima (Stephani) Gradst. in Bryophyt. Biblthca 4: 76 (1975). Ptychocoleus confertissimus (Stephani) Stephani, Sp. hepat. 5: 23 (1912); E. Jones in Trans. Br. bryol. Sac. 2: 405 (1954). SIERRA LEONE. Freetown Peninsula, Leicester Peak, c. 520 m (1700 ft), Jones 1457f. Distinguishable from var. emergens only by the perianth, which has four main deep plicae, sometimes with one to two subsidiary plicae, whereas the perianths of var. emergens have six to nine shallow subequal keels. Var. confertissima has hitherto been recorded only from west Cameroon and southern Nigeria, where it is locally frequent. 2. Acrolejeunea pycnoclada (Taylor) Schiffner in Engl. & Prantl, Nat. Pflanzenfam. 1 (3): 128 (1893); Gradst. in Bryophyt. Biblthca 4: 108 (1975). GHANA. Aiyaola F.R. near Kade, in crown of large Piptadeniastrum, Jones 1232, det. Gradstein. A species which is widely distributed in the Indo-Malayan region and Oceania, and apparently also in Madagascar, but known from continental Africa only from this collection and a single collection at Stanleyville (now Kisangani), Zaire. Gradstein (op. cit.) states that ‘the species grows preferably in anthropogenic habitats’; the occurrence in the Aiyaola F.R. does not show this preference, but suggests rather that Acrolejeunea pycnoclada is a component of the ‘tree crown flora’. II. ANEURA Dumortt. 1. Aneura pinguis (L.) Dumort., Sy//. Jungerm. Europ.: 86 (1831). Riccardia pinguis (L.) Gray, Nat. arr. Brit. pl. 1: 684 (1821); E. Jones in Trans. Br. bryol. Soc. 3: 84 (1956). SIERRA LEONE. Kambui Hills, Bambawo, on damp silt in floor of old mine workings, Jones 1542. GHANA. Pra-Suhien F.R., on rotting logs, Jones! III. ANTHOCEROS L. 1. Anthoceros buettneri Stephani, Sp. hepat. 5: 997 (1916). GHANA. Legon Botanical Garden, Hall (GC 47098). Agrees well with the isotype from Bismarkburg, Togo (FH); the holotype (G) is too fragmentary to be useful. It is a small plant with the habit of Anthoceros crispulus auct., but differing in the spores, which are black, 40—-45um diam., and densely clothed on all faces with spines 1-2 wm long. 2. Anthoceros mandonii Stephani, Sp. hepat. 5: 997 (1916); Vanden Berghen in Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8(1): 195 (1972); Bull. Jard. bot. natn. Belg. 48: 371 (1978). SIERRA LEONE. Loma Mountains, Bintimani, 1830-1940 m (6000-6360 ft), on soil over rocks, west-facing cliffs. Harrington 212 p.p. and summit plateau, Morton s.n. Anthoceros spp. are certainly more widespread than the above records indicate; we have seen plants which cannot be named specifically from Amedzofe and elsewhere in Ghana. IV. ARACHNIOPSIS Spruce 1. Arachniopsis diacantha (Montagne) M. Howe in Bull. Torrey bot. Club 29: 288 (1902). SIERRA LEONE. Loma Mountains: on bole of large tree in forest near ‘Camp 2’, 1650 m (5400 ft), HEPATICS OF SIERRA LEONE & GHANA 225 Jones 1482; on moist bank of streamlet in deep shade, c. 1520 m (5000 ft), Jones 1496 p.p. Kambui Hills, Bambawog, in forest on plateau, on charred stump, Jones 1546 p.p. Gola North F.R., rotten log by stream in forest, Jones 1566b. GHANA. Ankasa River F.R., on rotting log, Jones 1366 p.p.; on rotting bark in swampy forest one mile south of the Ankasa River, Richards R7061 p.p. Aiyinasi A.R.S., on rotting log in forest, Jones 1361a, 1362. Subri F.R., Jones ! Pra-Suhien F.R., on rotting log, Jones 1335. Amedzofe, near the waterfall, /rvine 411. Widely distributed throughout the forest of the wettest parts of W. Africa, usually on rotting logs, but occasionally on earth or tree boles, and associated (as in all the above records except Jones 1482) with Lophocolea martiana subsp. newtonii (Stephani) R. M. Schuster; Cephalozia fissa Stephani is also a frequent associate. Fulford (1968) followed earlier authors in recognizing two allied species in S. America: Arachniopsis diacantha with stiff leaf segments 5-8 cells long, the cells mostly 130 wm long, and A. coactilis Spruce with flaccid leaf segments 6-8-(10) cells long, the cells 78-90 wm long. African plants are intermediate in one respect or another between these two, though they tend to be nearer to A. diacantha. We follow Arnell (1963) in considering the two species to be synonymous. V. ARCHILEJEUNEA (Spruce) Schiffner 1. Archilejeunea abbreviata (Mitten) Vanden Berghen in Revue bryol. lichén. 20: 117 (1951). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., on bole of isolated tree near Heddle’s Farm, Jones s.n.; Leicester Peak, on mango, Jones 1464a; near Toke, on shaded boulder by stream, Jones 1448, and on branches of isolated shrub nearby, Jones s.n. Lake Sonfon, on low tree in forest surrounding the ‘lake’, Harrington 40 p.p. Bagru [‘Bagroo’] River, Mann s.n. (Lejeunea abbreviata Mitten —NY, holotype; BM, isotype). GHANA. Krokosua Hills, 460 m (1500 ft), on liane, Jones 1389. Asenanyo F.R., on buttress of large tree and on pole-size tree boles by the river, Jones 1413b, c. Subri F.R., on bole of large tree in Raphia swamp, Jones 1358. Bunso, on cocoa, Jones 1317a p.p. Bunso—Begoro road, on liane by stream, Jones 1302a. Southern Scarp F.R., near Begoro, on small pole, Jones 1306. Kade, on small tree in deep shade, Jones s.n. Amedzofe, in old coffee farm, Jones 1249; on log near the waterfall, Jones 1282. See note following Archilejeunea autoica. 2. Archilejeunea africana Stephani, Sp. hepat. 4: 705 (1911); Vanden Berghen in Revue bryol. lichén. 20: 114 (1951). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., epiphyllous, Jones s.n.; Regent, tree in farm bush, Harrington 409 p.p.; York Pass, on twigs of sapling in forest understorey, Harrington 631, 637a, 638. Masimo, Marmo 161 p.p. Kasewe F.R., on twig in rocky stream bed in forest, c. 90 m (300 ft), Richards R7165b, and epiphyllous, Richards R7170 p.p., both det. Jones. Njala, upper part of trunk of Nauclea diderrichii, Deighton 5313 p.p. Kambui Hills, Bambawo, on small trees in forest near the Rest House, Jones 1527, 1532b. Giema, epiphyllous in secondary forest near stream, Harrington 528 p.p.. 529 p.p. Gola Hills: Gola North F.R., in primary forest, epiphyllous, Jones 1568 p.p., and on twigs of understorey ‘shrublets’, Jones 1569 p.p.; near Lalehun, epiphyllous, Marshall s.n. GHANA. Pampramasi, on small pole in damp forest, Jones 1408c. Aiyinasi, in forest, on buttress of Uapaca, Jones 1361d. Nsuta, on liane in secondary forest on bank of Pra river, Jones 1351b. Southern Scarp F.R., near Begoro, on small pole, Jones 1306b. See note following Archilejeunea autoica. 3. Archilejeunea autoica Vanden Berghen in Revue bryol. lichén. 20: 119 (1951). SIERRA LEONE. Kambui Hills, forest on plateau above. Bambawo, Jones 1544. GHANA. Pampramasi, rotting log by stream in forest, Jones 1410b, on rocks in the stream bed, 1410c, and tree root in the bank, 1410d. Krokosua Hills F.R., on small poles and lianes in understorey, c. 460 m (1500 ft), Jones 1391, 1392b. Pra-Suhien F.R.., side of tree root, Jones 1343b. Kade, base of liane in forest in deep shade, Jones 1219. 226 E. W. JONES & A. J. HARRINGTON Archilejeunea abbreviata, A. africana, and A. autoica are closely allied; indeed Vanden Berghen (1972: 90) has suggested that the first two are synonymous. Nevertheless I have rarely had difficulty in separating them. A. africana often seems distinct in its habit, with closely-spaced long-beaked perianths ranked along one side of the branches of a cyme. In contrast A. abbreviata is more irregularly branched, with the subgynoecial innovations not again quickly fertile, and the perianths usually less cordate and with short beaks. Moreover, the two species often grow in different places, with A. abbreviata showing a preference for less-sheltered habitats. In Jones 1306 the two species were noted in the field as growing close together on the same branches, but quite distinct, A. africana being green when fresh, and A. abbreviata being brown. [E.W.J.] On the other hand I find great difficulty in separating Archilejeunea autoica from A. abbreviata. Intermediates are frequent (some of the specimens listed above are intermediate), and A. autoica is almost always associated with A. abbreviata (e.g. in the Krokosua Hills, Jones 1389 and 1391, and at Kade, Jones s.n. and 1219) under conditions that suggest that A. autoica is merely a modification of A. abbreviata induced by deep shade or moisture. [E.W.J.] The frequency with which A. africana (but not A. abbreviata) occurs as an epiphyllous hepatic in Sierra Leone is remarkable; I have not recorded it as such elsewhere. [E.W.J.] 4. Archilejeunea linguaefolia Stephani, Sp. hepat. 4: 708 (1911); Vanden Berghen in Revue bryol. lichén. 20: 113 (1951). SIERRA LEONE. Musaia Agricultural Station, pumping station, epiphyte, Haswell B1 p.p. Kenema, on the banks of the Moa river at Jawo, on tree boles, roots and branches, and on banks of sand in the flood zone, ‘usually the only bryophyte present, and in great quantity from near present water level throughout the flood zone’ (field note), Jones 1537. Gola Hills, Lalehun, on boulders in dry stream bed in forest, Jones 1593: VI. ASTERELLA P. Beauv. 1. Asterella abyssinica (Gottsche) Grolle ex Vanden Berghen in Résult. scient. Explor. hydro- biol. Bassin Lac Bangweolo & Luapula 8 (1): 170 (1972). Fimbriaria abyssinica Gottsche, Syn. hepat.: 569 (1846). SIERRA LEONE. Loma Mountains, Bintimani, cliffs near the summit, in crevices, Jones 1486, det. Grolle. Asterella abyssinica is frequent on Mont Cameroun (Cameroon Mountain) at similar altitudes (Jones 330, 433, 434). VII. BAZZANIA Gray 1. Bazzania decrescens (Lehm. & Lindenb.) Trev. St. Léon in Memorie Ist. lomb. Sci. Lett. 13: 414 (1877), subsp. molleri (Stephani) E. Jones in J. Bryol. 8: 303 (1975). SIERRA LEONE. Loma Mountains: 1220-1680 m (4000-5500 ft), frequent on the boles of large Parinari etc. on the margins of the forest, Jones 1478b, 1489, 1507; Harrington 334 p.p., 336 p.p. GHANA. Ankasa River F.R., on tree boles near the ground, Jones 1372a, b. Atewa Hills F.R., in swamp forest on boles of large Lophira, Jones 1328 p.p. 1331 p.p.; on bole of Gilbertiodendron limba, 790 m (2600 ft), Richards & Hossain (GC36670). For a discussion of the variability of this plant see Jones (op. cit). It is widely distributed but local in Africa, and is usually abundant where it occurs. It is usually montane; its occurrence in the Ankasa River F.R. is remarkable, and the fact that it was growing close to the river, some of it (Jones 1372b) in a site where it may possibly be irrigated in periods of high water, may indicate that high rainfall is usually a factor that controls its distribution. VIII. BRACHIOLEJEUNEA (Spruce) Schiffner 1. Brachiolejeunea tristis Stephani, Sp. hepat. 5: 112 (1912); Vanden Berghen in Bull. Soc. r. Bot. Belg. 92: 115 (1960). HEPATICS OF SIERRA LEONE & GHANA PRA: Brachiolejeunea camerunensis E. Jones & Vanden Berghen in Bull. Jard. bot. Etat Brux. 21: 88 (1951). SIERRA LEONE. Loma Mountains, on branches of isolated trees in rocky ground etc. near ‘Camp 2’, 1680 m (5500 ft), mixed with Marchesinia moelleriana, Lopholejeunea jonesii, Frullania spp. etc., Jones 1480 p.p., 1495 p.p. This species is widely distributed in the African mountains but is unlikely to occur elsewhere in Sierra Leone unless in the Tingi Hills, or in Ghana. IX. CALYPOGEIA Raddi 1. Calypogeia fissa (L.) Raddi, Jungermanniografia Etrusca: 33 (1818); Bischler in Revue bryol. lichén. 37: 97 (1970). SIERRA LEONE. Loma Mountains, moist bank of streamlet in deep shade, c. 1520 m (5000 ft), Jones 1496 p.p. Oil bodies have not been seen, but the plant when fresh lacked any blue tinge, and it agrees well with African plants accepted as Calypogeia fissa by Bischler (op. cit.), and with plants from Tanzania (Jones 1959, 1964) referred to by Jones (1976: 44). 2. Calypogeia longifolia Stephani, Sp. hepat. 6: 449 (1924); Bischler in Revue bryol. lichén. 37: 106 (1970). SIERRA LEONE. Loma Mountains, head of the Kongbundu Valley, north-west of Bintimani, c. 1520 m (5000 ft), on vertical banks of moist humic earth in grassland, Jones 1502 p.p. Bischler (op. cit.) gives the distribution of this species as South Africa (Cape Province, Natal, Transvaal), Lesotho, Swaziland, Southern Rhodesia (= Zimbabwe), and central Madagascar. X. CAUDALEJEUNEA (Stephani) Schiffner 1. Caudalejeunea africana (Stephani) Schiffner in Engl. & Prantl, Nat. Pflanzenfam. 1 (3): 129 (1893); Vanden Berghen in Revue bryol. lichén. 17: 96 (1948). GHANA. Ankasa River F.R., epiphyllous in crown of Glutea, Jones 1378c; on palm in swamp one mile south of the Ankasa bridge, Jones 1382 p.p. Subri F.R., on fronds of Raphia in swamp forest, Jones 1355. A very local species, which is probably confined to the lowland rain forest of West Africa and the Congo Basin; usually, as in the three Ghana records, epiphyllous, though by no means exclusively so. 2. Caudalejeunea dusenii Stephani, Sp. hepat. 5: 11 (1912); E. Jones in Trans. Br. bryol. Soc. 2: 165 (1953). GHANA. Aiyinasi A.R.S., boughs of a small tree by the Rest House, Jones 1384a. Aiyaola F.R., near Kade A.R.S., on twigs and small branches in the crown of a Piptadeniastrum, Jones 1236 p.p., and of Khaya, Jones 1237b. Amedzofe, on cocoa trees, Jones 1272c. Caudalejeunea dusenii usually grows in rather open well-illuminated places on small branches and twigs, thus in the crowns of large forest trees, but also on bushes and small trees of farms and gardens. It has been recorded from Nigeria (Jones, op. cit.), Cameroon (Jones, op. cit.; Augier, 1974) and, more recently, from the Uluguru Mountains in Tanzania (Bizot & Pécs, 1979). 3. Caudalejeunea hanningtonii (Mitten) Schiffner in Engl. & Prantl, Nat. Pflanzenfam. 1 (3): 129 (1893); Vanden Berghen in Revue bryol. lichén. 17: 98 (1948). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., frequent both as an epiphyll and on twigs, Harrington 78 p.p., 80, 82 p.p., Jones 1440b and s.n.; Leicester Peak, on old mango, Jones 1464 p.p. Kafogo [‘Kafoko’] (9°24’N, 11°44’W), 240 m (800 ft), Thomas 2126a. Loma Mountains, forest above Seradu, 1040 m (3400 ft), epiphyllous, Jones s.n. Kasewe F.R., on twig in rocky stream bed in forest, c. 90 m (300 ft), Richards R7165a, and epiphyllous, Richards R7170 p.p. Giema, on low vegetation by stream in secondary forest, Harrington 529 p.p., 532 p.p. 228 E. W. JONES & A. J. HARRINGTON GHANA. Krokosua Hills, on cocoa, Jones 1388. Aiyinasi, on Uapaca bole in forest near the A.R.S., Jones 1361e; epiphyllous near the river Fiakpole, Jones 1364b. Pra-Suhien F.R., epiphyllous, Jones 1345b. Bunso, on cocoa, Jones 1317b; near the Bosuso—Begoro road, on liane, Jones 1302c. Forest near Kade A.R.S., epiphyllous, Jones s.n. Osenasi (16 km south-east of Kade), on cocoa, Jones s.n. Aburi Botanical Gardens, Jones s.n. (GC). Kpedze, on cocoa, Jones s.n. Much the most widespread and commonest species of Caudalejeunea in tropical Africa. It is generally distributed throughout southern Ghana, especially on small trees such as cocoa in farms, and on bushes and small trees in secondary forest, usually on tree branches, though occasionally epiphyllous; this agrees with Jones’ experience of it in most other parts of Africa. It is curious that in Sierra Leone almost all the records show it as being epiphyllous. It is not clear whether this indicates a genuine difference in habitat-preference, which might be due to the climatic peculiarities of the country, or whether it is merely due to the lack of collecting in the farmland and secondary forest of central and southern Sierra Leone. 4. Caudalejeunea tricarinata E. Jones in Trans. Br. bryol. Soc. 2: 169 (1953). SIERRA LEONE. Freetown Peninsula: three km north-east of Toke, on branch of a small isolated shrub growing amongst rock slabs, Jones 1448c; York Pass, epiphyllous, Harrington 626 p.p. Kasewe F.R.., on savanna tree, Richards s.n. (Herb. Jones). Njala, upper part of trunk of 66 foot-tall Nauclea diderrichii, Deighton 5313 p.p. Hitherto known only from Nigeria (Jones op. cit., 1968b) and Cameroon (Augier, 1978). Schuster (1980) finds ‘no reliable differences’ to separate Caudalejeunea tricarinata from the polymorphic American species, C. lehmanniana (Gottsche) A. W. Evans. Accordingly he reduces the name to a synonym of C. /ehmanniana. XI. CEPHALOZIA (Dumort.) Dumort. 1. Cephalozia fissa Stephani in Hedwigia 30: 204 (1891). GHANA. Ankasa River F.R., on rotting log, Jones 1366. Aiyinasi A.R.S., in forest, Jones 1361a. Tarkwa, Saxby s.n. A lowland species, widely distributed in rain forest in tropical Africa on rotting wood, where it is often associated with Arachniopsis. It is likely to be present in south-eastern Sierra Leone. XII. CEPHALOZIELLA (Spruce) Schiffner 1. Cephaloziella ?vaginans Stephani in Wiss. Ergebn. dt. ZentAfr. Exped. 2: 119 (1911). SIERRA LEONE. Loma Mountains, valley west of Bintimani, c. 1650 m (5400 ft), on bole of large tree on forest margin, Jones 1493, det. Vana. Hitherto known from Ruwenzori (G, type), Kilimanjaro (Jones 2246, det. Varia, BM and Herb. Jones) and, as var. camerunensis S. Arnell (Arnell, 1958), from Mont Cameroun (Cameroon Mountain). XIII. CERATOLEJEUNEA (Spruce) Schiffner 1. Ceratolejeunea beninensis E. Jones & Vanden Berghen in Bull. Jard. bot. Etat Brux. 21: 63 (1951). GHANA. Ankasa River F.R., on leaves of low shrubs in small gaps in forest by the river, mixed with Ceratolejeunea diversicornua, Cololejeunea cornuta, Trachylejeunea serrulata etc., Jones 1375 p.p. Ceratolejeunea beninensis was previously known only from Benin, Nigeria, and Cameroon (Banga F.R., Jones 284). Perianths are uncommon. 2. Ceratolejeunea calabariensis Stephani in Hedwigia 34: 234 (1895); Vanden Berghen in Bull. Jard. bot. Etat Brux. 21: 71 (1951); Revue bryol. lichén. 39: 381 (1973). HEPATICS OF SIERRA LEONE & GHANA 229 Ceratolejeunea saxbyi Pearson in Mem. Proc. Manchr lit. phil. Soc. 65 (1): 4 (1921). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., on trees, Harrington 579, 580, 666 p.p., Jones 1420; Sugar Loaf, 460 m (1500 ft), 7. S. Jones 245 p.p. Bagru [‘Bagroo’] River, mixed with Thysananthus spathulistipus, Mann s.n. Kambui Hills, Bambawo, on large roadside trees, Harrington 569 p.p., 570 p.p.; Jones 1550a; on smooth bole of small tree in forest, Jones 1530; on earth bank by road in forest, Jones 1529. Gola Hills: Lalehun, upper part of bole of large tree, Jones 1554b p.p.; buttress of Piptadeniastrum, Jones 1555b; Gola North F.R., on boughs in crown of large tree, mixed with Marchesinia excavata, Jones 1562. GHANA. Tarkwa, Saxby (MANCH, holotype of Ceratolejeunea saxbyi; BM, isotype), and mixed with Taxilejeunea nicholsonii Pearson, Saxby (CGE). Pra-Suhien F.R., on mango by the Rest House, Jones 1349. Atewa Hills F.R., on trunk of Cassipourea, Richards & Hossain (GC 36668 p.p.); boughs in crowns of Cassipourea, Jones 1297, 1322 p.p. Kade A.R.S., mixed with Cheilolejeunea trifaria on main bough of Parinari glabra in forest, Hossain (GC 36696). The problems concerning the delimitation and taxonomy of Ceratolejeunea calabariensis are discussed under C. zenkeri. Of the plants recorded above that have been examined by Jones, 1554b, 1349 and C. saxbyi have perianth horns 100-150 um long, and would thus be C. calabariensis as defined by Vanden Berghen in 1951, while the remainder have longer horns, mostly 200-300 um long. The plants from the Atewa Hills have perianth horns up to 350 wm or exceptionally 400 um long, and might be considered intermediate to C. zenkeri. 3. Ceratolejeunea diversicornua Stephani, Sp. hepat. 5: 410 (1913); Vanden Berghen in Bull. Jard. bot. Etat Brux. 21: 68 (1951); Bull. Jard. bot. natn. Belg. 47: 206 (1977). GHANA. Ankasa River F.R., on leaves of shrubs with Ceratolejeunea beninensis, Jones 1375 p.p., det. Vanden Berghen; Jenik s.n. (OP); on palm leaves in swamp one mile south of the Ankasa bridge, Jones 1382 p.p. Ceratolejeunea diversicornua is a rare species otherwise known only from Cameroon and Kivu. In many respects the Ankasa River plants are intermediate between C. diversicornua and C. cornutissima Stephani as described by Vanden Berghen (1951), thus supporting his later (1977) suggestion that the two species may be synonymous. The sterile shoots differ from those of C. beninensis, when the two species are mixed, in bearing utricles and in having leaves which are less falcate with margins distantly serrulate (as illustrated by Vanden Berghen (1951: fig. 23F) for C. cornutissima) due to slightly projecting radial walls. The perianth horns are only c. 200 wm long; shorter than described by Vanden Berghen (1951). 4. Ceratolejeunea zenkeri Stephani, Sp. hepat. 5: 449 (1913), sensu Vanden Berghen in Bull. Jard. bot. Etat Brux. 21: 78 (1951). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., on stones, Harrington 605, on Dacryodes klaineana, Harrington 586, on large tree, Harrington 666 p.p., epiphyllous, Jones, s.n.; Leicester Peak, on tree bole, Brenan (Jones 461, det. Vanden Berghen), on bough of Combretum, Jones 1454 p.p. Giema, on sapling, Harrington 554 p.p. GHANA. Krokosua Hills F.R., tree boles on sheltered woodland margin, 550 m (1800 ft), Jones 1406. Ankasa River F.R., in crown of Glutea, Jones 1378a. Pra-Suhien F.R., bough in crown of tree, Jones 1346. Vanden Berghen (1951) recognized three closely allied species (and also other more distinct species) in Africa (excluding the Mascarenes), for which he used the names Ceratolejeunea calabariensis Stephani, C. jungneri Stephani, and C. zenkeri Stephani. They were said to differ chiefly in the lengths of the perianth horns, which were 100-150 wm, 200-280 (exceptionally 360) um, and 330 (exceptionally as short as 280)—530 um respectively. In addition C. zenkeri differed from the other two in having ocelli which, in many leaves, formed a conspicuous vitta extending to near the middle of the leaf. Vanden Berghen pointed out that the vegetative differences between C. calabariensis and C. jungneri were slight, and in 1973, influenced by the variability in length of horn that he observed in certain collections, he made them synonymous under the name C. calabariensis. Specimens with horns about 300 um long, thus in the range of overlap between ‘C. jungneri’ and ‘C. zenkeri’ are common (12 of the 40 specimens in Herb. Jones under 230 E. W. JONES & A. J. HARRINGTON the preceding three names), and these may or may not have a vitta. The vitta thus becomes the chief distinguishing feature of C. zenkeri. A vitta is, however, often present in some leaves but not in others; in Jones 1346, for example, it is present in branch leaves but not in stem leaves. The different forms often occur in association with each other, and sometimes indeed intimately mixed, but it is not clear whether such mixtures consist of different genotypes or are the result of intraclonal variation. Thus there are strong reasons for merging C. zenkeri with C. calabariensis. Nevertheless the variation within the taxon that would be so formed is so great that some division seems desirable; moreover a reasonably large proportion of collections are homogeneous. We have therefore tentatively continued to recognize two taxa, C. calabariensis and C. zenkeri, and retained the names used by Vanden Berghen, even though these names are almost certainly not correct. Ceratolejeunea calabariensis is probably identical with the earlier C. belangeriana (Gottsche) Stephani from Mauritius. C. belangeriana itself is, however, almost certainly identical with one or more of the still earlier species from the West Indies and Central and South America — C. cornuta (Lindenb.) Schiffner, C. maritima (Spruce) Stephani, and C. variabilis (Lindenb.) Schiffner. Examination of the literature and of herbarium specimens shows that there is great confusion between these plants, and it will not be possible to decide the correct name for C. belangeriana until the American members of the complex have been more closely studied. We therefore retain a late name, C. calabariensis, the application of which is certain, rather than adopt an earlier name, the application and correctness of which are doubtful. Vanden Berghen (1951) used the name Ceratolejeunea zenkeri for plants with a vitta and long-horned perianths with considerable hesitation, his justification being that part of the collection Zenker 3935g preserved in the BM possesses these features. The only specimen labelled C. zenkeri in Stephani’s own herbarium is a portion of Zenker 3935g, but is C. calabariensis. Moreover Stephani’s ‘Icones’ show C. zenkeri as a short-horned plant, while his original descriptions are too lax to differentiate between C. zenkeri sensu Vanden Berghen and C. calabariensis. Zenker 3935g was a large collection, as the portion in the BM shows, and it doubtless included both taxa. XIV. CHANDONANTHUS Mitten 1. Chandonanthus hirtellus (Fried. Weber) Mitten in J. Linn. Soc. (Bot.) 22: 321 (1886); Vanden Berghen in Bull. Soc. r. Bot. Belg. 98: 137 (1965). SIERRA LEONE. Loma Mountains, 1650 m (5400 ft), on the boles of large Parinari on the forest margin, Jones 1478a, 1493. A species with a wide but disjunct distribution, chiefly in the mountains of Africa and the Old World tropics. On the Loma Mountains it occurs in a wide range of forms, from robust shoots with four-lobed leaves down to slender Lophozia-like plants with bilobed leaves, creeping amongst Bazzania decrescens subsp. molleri, Conoscyphus trapezioides, and various Le- jeuneaceae. XV. CHEILOLEJEUNEA (Spruce) Schiffner 1. Cheilolejeunea decursiva (Sande Lacoste) R. M. Schuster in Beih. nov. Hedwigia 9: 112 (1963); Grolle in J. Bryol. 9: 531 (1977). Cheilolejeunea tisserantii Vanden Berghen & Jovet-Ast in Revue bryol. lichén. 20: 105 (1951). SIERRA LEONE. Freetown Peninsula: by Mountain Torrent below Havelock Plateau, on base of small tree, mixed very sparsely with other small Lejeuneaceae, Jones 1441b, and on Vitex, Jones 1443; three km north-east of Toke, c. 90 m (300 ft), on bole of small tree growing in rock pavement, Jones 1448b. Loma Mountains: epiphyllous in gallery forest c. one km west of ‘Camp 2’, 1520 m (5000 ft), Morton (Harrington 347 p.p.); 1680 m (5500 ft), Jones 1483a p.p. There are very few records of this species from Africa, though it is evidently widely distributed, as it is known from Réunion, Madagascar, Tanzania, Zaire, the Central African Republic, and HEPATICS OF SIERRA LEONE & GHANA 251 Guinea. It would be difficult to detect if it were to be present, as it is in Jones 1441b, in small amounts mixed with common species, such as Lejeunea caespitosa and L. eckloniana, so that it has probably been overlooked. 2. Cheilolejeunea exinnovata E. Jones in J. Bryol. 12: 37 (1982). SIERRA LEONE. Gola North F.R., in unexploited forest, epiphyllous, Jones 1568 p.p. (BM, holotype). GHANA. Ankasa River F.R., epiphyllous on shrublets in small gap near the river, Jones 1375 p.p. The species is also known from Zimbabwe, and is present in Spruce’s L419 from Sao Gabriel, Brazil (MANCH, sub Cheilolejeunea aneogyna). 3. Cheilolejeunea intertexta (Lindenb.) Stephani in Bull. Herb. Boissier 5: 79 (1897); Grolle in J. Hattori bot. Lab. 46: 344 (1979). Cheilolejeunea inflata Stephani, Sp. hepat. 5: 645 (1914); E. Jones in Trans. Br. bryol. Soc. 2: 385 (1954). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., epiphyllous, Harrington 78 p.p., 82 p.p., 84, 87 p.p., and on bark, Harrington 586 p.p., 596 p.p.; Mountain Torrent, below Havelock Plateau, on Vitex, Jones 1443 p.p.; Leicester Peak, 550 m (1800 ft), Jones s.n.; valley three km north-east of Toke, on isolated shrub, Jones s.n.; Black Johnson Beach near York, epiphyllous on low shrub in thicket behind the beach, Harrington s.n. Kambui Hills, Bambawo, on tree boles in forest, Jones 1527 p.p., 1532d, 1533. Giema, on sapling by road, Harrington 554 p.p. GHANA. Bia North F.R., in crown of Entandophragma, Jones 1411c. Ankasa River F.R., Richards (GC 36679). Aiyinasi A.R.S., on Citrus, Jones 1383c. In scrub near Esiama, Jones 1365. Pra-Suhien F.R., trees by the Rest House, Jones 1350a. Bunso, on cocoa, Jones 1317a. Between Bame and Todome, on cocoa, Jones s.n. See notes under Cheilolejeunea serpentina. 4. Cheilolejeunea newtonii Stephani ex Schiffner in Engl. & Prantl, Nat. Pflanzenfam. 1 (3): 124 (1893); E. Jones in Trans. Br. bryol. Soc. 2: 388 (1954). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., on tree boles, Jones 1419a, b, Harrington 582, on rock, Jones 1422; Havelock Plateau, Jones s.n.; Leicester Peak, on tree bole, Jones 1430a, on rock, Jones 1452a; Regent, Irvine 204b, 205, 231; Bathurst, Kongo Dam, on boulder, Jones 1435; Dighton Dam. No. 1, valley below Sugar Loaf, on boulder, Harrington 661 p.p.; three km north-east of Toke, on shaded boulder, Jones 1447. Kambui Hills, Bambawo, shady earth bank by road, Jones 1529. Gola North F.R., on buttress of tree, Jones s.n. Banks of the Moa river at Jawo, on tree bole at top of flood zone, Jones 1538b. GHANA. Ankasa River F.R., on bole of Parinari, Richards (GC36679 p.p.), epiphyllous, Jones 1375 p.p. Tarkwa, Saxby (CGE). Esukawkaw F.R., Jenik. Kade A.R.S., on old lianes in forest in deep shade, Jones 1217 p.p. (GC). Amedzofe, 760 m (2500 ft), on decaying tussocks of Catagyna (= Afrotrilepis) , Jones 1253, and on shaded rocks, Jones 1255. Cheilolejeunea newtonii is known only from West Africa; it seems to be exceptionally abundant on the Freetown Peninsula, where it is probably the commonest species of Cheilolejeunea. It resembles C. intertexta, with which it often grows, in being monoecious. Well-developed plants differ in being somewhat larger (strong shoots of C. intertexta c. 0-9 mm wide, of C. newtonii 1-0-1-2 mm wide), with the keels of the lobule making a more distinct narrower sinus with the postical margin of the lobe (in C. intertexta the keel is almost in line with the postical margin of lobe), but weak shoots of C. newtonii may be indistinguishable from C. intertexta vegetatively. Thus mixtures of the two species, which are not infrequent, are very confusing, and it was such mixtures that led Jones (1973) to conclude that the two species were synonymous. Grolle (1979), however, pointed out that in C. intertexta the first leaf to be formed on a subgynoecial innovation is an underleaf, while in C. newtonii the first leaf is a lateral leaf. This order is not absolutely constant, however; though there is a great preponderance of the one type of innovation in any plant, as many as one in ten innovations may be of the other kind. Occasionally when twin 232 E. W. JONES & A. J. HARRINGTON subgynoecial innovations have been formed, one member of the pair has an underleaf first, while its companion has a lateral leaf first. Cheilolejeunea newtonii is probably more restricted to areas of high rainfall than either C. intertexta or C. serpentina. 5. Cheilolejeunea serpentina (Mitten) Mizutani in J. Hattori bot. Lab. 26: 171 (1963); Grolle in J. Hattori bot. lab. 46: 346 (1979). Cheilolejeunea principensis Stephani ex Paris in Revue bryol. 33: 38 (1906); E. Jones in Trans. Br. bryol. Soc. 2: 383 (1954). SIERRA LEONE. Freetown Peninsula: Mount Aureol [‘Oriel’], Arnel! 2507; Fourah Bay College B.R., on tree boles, Jones 1420 p.p., on liane, Harrington 596 p.p.; Leicester Peak, Brenan (Jones 464 p.p., 476 p.p.); Lumley Cove, tree bole on the beach, Richards (Jones 467); Regent, small tree in farm bush, Harrington 408 p.p.; Sugar Loaf Mtn, Tindall 62 p.p.; valley three km north-east of Toke, Jones s.n.; Toke, on small trees in ‘coastal savanna’ behind the beach, Jones 1444c, 1446b. Musaia Agricultural Station, pumping station, Haswell Bl p.p. Lake Sonfon, on Erythrophleum on edge of forest surrounding the ‘lake’, Harrington 53 p.p. Between Sokurela and Kurubonla, on Uapaca, Jones 1522d. Njala, upper part of trunk of Nauclea diderrichii, Deighton 5313 p.p. Kambui Hills, Bambawo, on mango, Harrington 427 p.p., 437 p.p., on roadside tree, Harrington 569 p.p., 570 p.p. Giema, on fallen tree in secondary forest, Harrington 536 p.p. Lalehun, upper part of bole of large tree, Jones 1554 p.p. Gola North F.R., in crown of large tree, Jones 1560 p.p. GHANA. Ankasa River F.R., on Pentadesma bole, Richards (GC 36674, 36678), epiphyllous, Jones 1375 and Jenik s.n. Aiyinasi A.R.S., on Citrus, Jones 1383a. Pra-Suhien F.R., bough of tree by the Rest House, Jones 1350a p.p. Aiyaola F.R. near Kade, in crown of Piptadeniastrum, Jones 1232b. Osenasi, on cocoa, Jones s.n. Aburi Botanical Gardens, /rvine s.n. Amedzofe, Irvine 412; on Terminalia and Lophira near top of hill, Jones 1257e. Cheilolejeunea serpentina differs from C. intertexta and C. newtonii in being dioecious and in having a rounder leaf with a more strongly arched keel; the first leaf on a subgynoecial innovation is a lateral leaf (as in C. newtonii). C. serpentina and C. intertexta are the two commonest species of the genus throughout much of West Africa, where indeed they are amongst the most abundant schizostipulean Lejeuneaceae. They also occur in East Africa where, however, they are very local, as they are essentially lowland species and confined to limited areas of high humidity, chiefly near the coast. The two species often grow mixed, and appear to be more tolerant of drought and exposure than most other schizostipulean Le- jeuneaceae; thus they often occur in the crowns of large trees, and on the small branches and twigs of isolated bushes. Usually they grow on bark or on rock, but in districts with very high rainfall, such as the Freetown Peninsula and the Ankasa River F.R., they are epiphyllous. C. serpentina is often abundant on the seacoast, where it may occur (as at Toke) very close to the high-water-mark, and clearly must tolerate some salt. C. newtonii is certainly much more restricted in range, probably by a lower tolerance of drought, and in most districts is much less abundant, though it grows in similar habitats, and is often mixed with one or both of the other two species. Like them, it seems to occur as an epiphyll only in the wettest sites. Schuster (1980) has provisionally placed Cheilolejeunea principensis into the synonymy of C. rigidula (Montagne) R. M. Schuster. If his treatment is confirmed by further investigation the epithet rigidula will have priority over serpentina. 6. Cheilolejeunea surrepens (Mitten) E. Jones in J. Bryol. 9: 49 (1976). Cheilolejeunea africana (Stephani) E. Jones in Trans. Br. bryol. Soc. 2: 388 (1954). Cheilolejeunea silvestris (Gottsche) E. Jones in J. Bryol. 7: 548 (1973). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., epiphyllous, Jones s.n.; Toke, epiphyllous, Jones s.n. (these two specimens are sterile and named with slight doubt, though they are both the same species). Giema, forest near road, epiphyllous, Harrington 552b p.p., 552c p.p., 552d p.p. GHANA. Pampramasi, on bole of small tree in forest, Jones 1408b. Aiyinasi A.R.S., on Citrus, Jones 1383 p.p.; on small tree by the Rest House, Jones 1363d. Pra-Suhien F.R., bough of small tree by the Kest House, Jones 1347b. Bunso, on cocoa, Jones 1317a p.p. Aiyaola F.R. near Kade, branches in crown of HEPATICS OF SIERRA LEONE & GHANA 233 large Piptadeniastrum, Jones 1236 p.p.; branchlets in crown of Khaya, Jones 1237a; on bole of middle- storey tree, Jones 1239b. Kpedze, on cocoa, Jones s.n. Cheilolejeunea surrepens is a variable species which is widely distributed in both West and East Africa, but usually forms very small colonies. It is often mixed with C. intertexta or C. serpentina, and slender branches may be difficult to distinguish from C. intertexta, though on strong shoots the much longer lobules are distinctive. It appears to be one of those species which are characteristically inhabitants of the smaller branches and twigs in the forest canopy, but which can also occur near the ground on the twigs of bushes and small trees in open places. Thus it tends to be collected around forest rest houses and in plantations. The paucity of records from Sierra Leone may be due to the lack of collections from suitable sites. It is very rarely epiphyllous, so that it is all the more remarkable that it has been recorded from Sierra Leone only as an epiphyll. 7. Cheilolejeunea trifaria (Reinw., Blume & Nees) Mizutani in J. Hattori bot. Lab. 27: 132 (1964); Grolle in Wiss. Z. Friedrich-Schiller -Univ. Jena 27: 9 (1978). Euosmolejeunea brachytoma (Gottsche) Stephani, Sp. hepat. 5: 577 (1914). SIERRA LEONE. Freetown Peninsula: Leicester Peak, Brenan (Jones 476 p.p.); Sugar Loaf, 460 m (1500 ft), 7. S. Jones 245 p.p. Lake Sonfon, on trunk of Erythrophleum at edge of forest, Harrington 53 p.p. Loma Mountains, Seradu, tree boles in valley forest by the village, Jones 1515b, 1516b. Kambui Hills, Bambawo, shaded earth bank, Harrington 508 p.p., Jones 1529 p.p.; tree bole by road, Harrington 570 p.p., Jones 1550 p.p. GHANA. Ankasa River F.R., on bole of Pentadesma, Richards (GC 36674 p.p.). Aiyinasi A.R.S., on Citrus boles, Jones 1383d, e p.p. Ochi Headwaters F.R., West-Skinn 155, 183. Atewa Hills F.R., boughs in crown of Lophira, Jones 1319b. Aburi Botanical Gardens, on buttress of tree, Jones s.n. Between Vane and Amedzofe, rocks in steep roadside bank, Jones 1266a. Cheilolejeunea trifaria has a wide but disjunct distribution in tropical Africa, which is difficult to understand. Where it is present it is often abundant. It tends to be montane — thus the records from the Freetown Peninsula are from the higher ground — but its occurrence nearly at sea-level in western Ghana suggests that this tendency may be determined by some factor such as the frequency of mists. XVI. CHILOSCYPHUS Corda 1. Chiloscyphus dubius Gottsche in Abh. naturw. Ver. Bremen 7: 346 (1882); E. Jones in Trans. Br. bryol. Soc. 2: 200 (1953). SIERRA LEONE. Freetown Peninsula, York Pass, on boulder on small island in stream, Harrington s.n. Gola North F.R., on rotten log by stream in forest, Jones 1566d (FBC). GHANA. Ankasa River F.R., Jones s.n. (GC). Tarkwa, Saxby. Subri F.R., Jones ! Ochi Headwaters F.R., West-Skinn 184. Pra-Suhien F.R., on exposed tree root, Jones 1343 p.p. Esukawkaw F.R., Jenik (Herb. Jones). Begoro, below the waterfall, Jones ! Southern Scarp F.R., near Begoro, Jones ! Atewa Hills F.R., Jones 1331 p.p. (GC). Kade A.R.S., on stump in forest, Hossain (GC 36681). Amedzofe, Irvine 411, Jones s.n. (GC). Chiloscyphus dubius is widely distributed and usually abundant in the lowland rain forests of West Africa, often associated with Lophocolea martiana subsp. newtonii. It also extends through East Africa to Natal, Madagascar, and the Mascarenes, but in these areas it is much more local, doubtless because of the very restricted occurrence of lowland forest with a sufficiently high rainfall. It usually grows on rotting wood in deep shade, but frequently also near the bases of trees and exposed tree roots. It is evidently frequent throughout the forests of southern Ghana. The paucity of records from Sierra Leone is remarkable; it may be explained at least in part by the fact that C. dubius is a lowland species (though in Ghana it is frequent up to 700 m) and is strictly confined to forest habitats. Little lowland forest survives in Sierra Leone, and we have few collections of hepatics from that which is left. In the Freetown Peninsula the rarity of C. dubius may be due to the long dry season, though the lowland forest here is mostly 234 E. W. JONES & A. J. HARRINGTON farm bush, which does not provide the decaying logs and the sheltered conditions that the species seems to need. XVII. COLOLEJEUNEA (Spruce) Schiffner 1. Cololejeunea africana (Stephani) R. M. Schuster in Beih. nov. Hedwigia 9: 173 (1963). Leptocolea africana (Stephani) E. Jones in Trans. Br. bryol. Soc. 2: 148 (1953). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., Morton s.n. (Herb. Jones); Toke, Jones s.n. Gola Hills, near Lalehun, on low vegetation, Marshall s.n. Apparently a rare epiphyllous species, previously known only from a few gatherings in Cameroon and Sao Tomé, and from the mountains of south-eastern Kenya and eastern Tanzania (Bizot & Pécs, 1974, 1979). It is present in only very small amounts in the Sierra Leone collections. 2. Cololejeunea androphylla E. Jones in Trans. Br. bryol. Soc. 2: 432 (1954). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., on rough bole of a large tree 12 cm from ground, Jones 1424. Kambui Hills, Bambawo, in forest on plateau, 370 m (1200 ft), on charred stump, Jones 1546, and on rotten log, Jones 1548. Gola Hills: Lalehun, log in forest, Jones s.n.; Gola North F.R., rotting log by stream in forest, Jones 1566c. GHANA. Kade A.R.S., large rotten log in forest, Jones 1215a. Aiyaola F.R. near Kade, on rotten log, Jones 1230a. Previously known from Nigeria and Cameroon (Jones, op. cit.), the Congo Republic (Pécs, 1980) and the Seychelles (Grolle, 1978). 3. Cololejeunea appressa (A. W. Evans) Benedix in Beih. Repert. nov. Spec. Regni veg. 134: 31 (1953); E. Jones in Trans. Br. bryol. Soc. 5: 571 (1968). SIERRA LEONE. Giema, forest near road, epiphyllous, Harrington 552a p.p. In continental Africa, hitherto known only from Uganda, Kenya (Bizot & Pécs, 1974), and Tanzania (Bizot & Pécs, 1979). 4. Cololejeunea bolombensis (Stephani) Vanden Berghen in Bull. Jard. bot. natn. Belg. 42: 475 (1972). Leptocolea bolombensis (Stephani) E. Jones in Trans. Br. bryol. Soc. 2: 155 (1953). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., on leaves of Carapa procera, Harrington s.n.; York Pass, low vegetation in forest, Harrington 633 p.p.; Picket Hill, in forest near summit, c. 850 m (2800 ft), Gledhill s.n. About five km south of Kabala, in secondary forest by road, Harrington 127 p.p. Lake Sonfon, in forest surrounding the ‘lake’, Harrington 33 p.p., 34, 35 p.p., 70e. Loma Mountains: forest above Sokurela, 1040 m (3400 ft), Jones 1514b; valley forest at Seradu, 610 m (2000 ft), Jones 1518. Kambui Hills, Bambawo, Harrington 467 p.p., 468 p.p., 483 p.p., Jones 1526, 1543a. Giema, in secondary forest, Harrington 548 p.p., 549a p.p. Gola Hills, near Lalehun, low vegetation in primary (?) forest, Marshall s.n. In all the Sierra Leone collections Cololejeunea bolombensis was epiphyllous, on low vegetation in forest. The species is widely distributed, though perhaps local, in Africa from Guinea eastwards to Kenya, and southwards through the Congo Basin to the Uluguru Mountains in Tanzania, and, according to Pécs (Bizot & Pécs, 1979), the Umtali district of Rhodesia (= Zimbabwe). The absence of records from Ghana is, therefore, remarkable. In West Africa and the Congo Basin C. bolombensis is a lowland species, but the East African records come from altitudes of 900-2000 m, and thus indicate a montane taxon. Moreover, the closely allied Cololejeunea adhaesiva (Mitten) R. M. Schuster grows in the East African mountains. Cololejeunea bolombensis is very variable, and it is likely that Cololejeunea auriculata (E. Jones) R. M. Schuster will prove to be an extreme form, with narrow lobules and faintly HEPATICS OF SIERRA LEONE & GHANA 235 punctate cuticle. In many examples of C. bolombensis, including the type, the punctate cuticle is conspicuous but, as in many Lejeuneaceae with this form of cuticle (e.g. Lejeunea flava (Sw.) Nees), it varies greatly in distinctness, and the ‘points’ are sometimes so faint that careful observation with critical illumination is required to see them. The type of C. auriculata (Jones 73A) from Benin has a faintly punctate cuticle. The other differences from C. bolombensis are the width of the lobule, the insertion of the lobule, the extent of the hyaline border, and the general shape of the leaf (oval in C. auriculata, obovate in C. bolombensis). 5. Cololejeunea calcarata E. Jones, sp. nov. Figs 1 & 2. Tenella; surculi cum foliis 0-6-1-0 mm lati, repentes. Folia 0-35-0-6 x 0-3-0-4 mm, contigua, antice secunda, semicirculares basi cuneata, margine postico stricto, margine antico arcuato, grosse 6-7 dentato dentibus triangulariis; dens distalis semper conspicuus, calcariforme retrocurvatus. Cellulae lobi laeves, medianae 15-20 x 20-30 um, trigonis mediocris. Lobulus quam lobo 1/2 brevior, oblongo-ovatus, proximale leviter inflatus, distale planus, carina stricta vel leviter arcuata, dens apicalis ex angulo lobuli oriens, vulgo bicellularis, (nonnunquam unicellularis), cellulo apicale sphaerico incurvo; papilla hyalina sphaerica 10-12 wm diametro in facie interno celluli basalis inserta, dens alter in margine libera subnullus. Monoica. Bracteae masculae 3-4 jugae, laxe imbricatae, monandriae. Bracteae femineae lobis falcatis, apice acutis, interdum obscure dentatis; lobuli longitudine 2/3—3/4 loborum aequantes, anguste oblongi, apice acuti aut obtusi, carina substricta. Perianthium late pyriforme, 0-42 x 0-50 mm, dorso planum, ventre leviter inflatum, laterale obtuse carinatum, rostro brevissimo aut nullo. Typus: Ghana, Jones 1355b, in Herb. Mus. Brit. conservatus. Small and delicate, much branched, creeping on fronds of Raphia palm. Shoots 0-6-1-0 mm broad. Leaves approximate, arching upwards and forwards, 0-35-0-6 x 0-3-0-4 mm, nearly semicircular with cuneate base, broadly arched apex and antical margin, and straight or slightly concave postical margin. Apex and antical margin dentate with 4-7 distant large triangular teeth, very variable in size, but with the distal tooth at the junction of apex and postical margin always large and directed backwards, at right angles to the postical margin. Cells smooth, trigones medium, straight-sided or convex, intermediate thickenings occasional on some of the longer walls, chiefly in the mid and proximal part of the leaf; median cells 15-20 x 20-30 um, cells of third and fourth rows from antical margin 14-15 x 15-22 um. Lobule large, with the slightly arched keel extending half the distance from insertion to apex of lobe, c. 250-300 x 125-150 um, oblong-oval, moderately inflated proximally, plane distally, the free margin mostly visible in situ for about half its length, the apex usually truncate with a tooth at the angle, but very variable, and the angle sometimes extended to form a triangular lobe. The apical tooth mostly of two cells, the apical cell spherical and strongly incurved when mature, the hyaline papilla spherical, 10-12 wm diam., on the proximal half of the inner face of the basal cell of the tooth. An obscure blunt tooth sometimes present on the free margin, separated from the apical tooth by 3-4 cells. Stylus not found. Stem 45-50 wm diam., the cortical cells 20 x 40-50 zm; in transverse section five cortical cells and one medullary cell, all uniformly rather thick walled. Monoecious. Male bracts in 3-4 pairs, making a slender lax spike, the bracts monandrous; antheridia 50-55 xm diam. Female bracts resembling the leaves but narrower (c. 450 x 200 um), and with longer lobules, falcate, irregularly and variably dentate, often entire except for the acute outwardly directed apex; lobule c. 2/3—-3/4 the length of the lobe, the keel 1/2—2/3 its length, nearly straight, the ligulate apex truncate, rounded or acute. Perianth broadly pyriform, 0-5 x 0-4 mm, subcompressed, plane dorsally, ventrally inflated, ecarinate or the margin bluntly keeled around the apex and distally. GHANA. Subri F.R., in swamp forest, on dead fronds of Raphia, with Caudalejeunea africana, Colura digitalis, Cololejeunea crenatiflora, C. nigerica, Diplasiolejeunea cornuta, Cheilolejeunea surrepens, etc., Jones & Hall (Jones 1355b; BM, holotype). Cololejeunea calcarata is very distinct from C. dentata, the only other Cololejeunea with coarsely dentate leaves known from West tropical Africa; the two plants were indeed growing together. 236 E. W. JONES & A. J. HARRINGTON Fig. 1 Cololejeunea calcarata E. Jones. (a) shoot with perianth, x60; (b) female bracts, x 100; (c) tooth ‘w’ and (d) tooth ‘x’ from leaf (a) in Fig. 2, x500; (e) apical tooth of young lobule; (f) apical tooth of mature lobule, both 500. Drawn from Jones 1355b (holotype). Though they resemble each other in the general shape of the leaf, they differ in the pattern of dentition and in the tendency shown by C. dentata (in common with the allied C. malanjae Stephani from East Africa, C. cuneifolia, and others) to produce cells in the leaves, and even more in the perianths, which are conically protuberant and capped by lenticular thickenings of the wall. Thus in C. dentata the leaf margin between the major teeth is irregularly (though only slightly) crenulate or serrulate with protuberant cells, and slight lenticular thickenings of the wall can often be detected above the lumina of some of the other cells of the lobe. C. calcarata shows no trace of this tendency on either leaves or perianth. The postical margin of the leaf of C. calcarata is straight or slightly concave, and completely without teeth except for the large backwardly directed spur at its apex, whereas the postical margin in C. dentata tends to be convex with some minor denticulation. C. malanjae differs from C. calcarata even more in the shape of the leaf. Judging from descriptions and drawings Cololejeunea calcarata resembles C. plagiochiliana P. Tixier from Madagascar in the shape of leaf, but has a very different perianth. HEPATICS OF SIERRA LEONE & GHANA 539 Fig. 2 Cololejeunea calcarata E. Jones. (a)-(e) leaves, X60-‘w’ and ‘x’, teeth illustrated in Fig. 1 (c) and (d); (f), (g) female bracts, x60; (h) cells from ‘y’ and (i) the tooth ‘z’ on leaf (c), 500; (j) median cells from another leaf, 500. Drawn from Jones 1355b (holotype). 6. Cololejeunea cardiocarpa (Montagne) R. M. Schuster in Bryologist 62: 54 (1959); Vanden Berghen in Revue bryol. lichén. 44: 448 (1978). Leptocolea cardiocarpa (Montagne) A. W. Evans in Bull. Torrey bot. Club 38: 268 (1911); E. Jones in Trans. Br. bryol. Soc. 3: 200 (1957). SIERRA LEONE. Freetown Peninsula, on branch of small bush by Mountain Torrent, Jones s.n. A pantropical species, usually epiphyllous, which has not hitherto been recorded from West Africa, though it is widely distributed but apparently local from Cameroon (Tixier, 1975) and Zaire eastwards to Kenya and Tanzania, and south to Zimbabwe. As Vanden Berghen (op. cit.) notes, it tolerates considerable desiccation, and sometimes grows where no other epiphyllous species is present. Like many other epiphyllous species, it tends to be corticolous where dry conditions prevent the growth of epiphyllous communities. Its occurrence in the Freetown Peninsula is doubtless related to the severe dry season that the Peninsula experiences. 7. Cololejeunea cornuta E. Jones in Trans. Br. bryol. Soc. 2: 436 (1954). SIERRA LEONE. Gola North F.R., epiphyllous on low vegetation, Jones 1567 p.p., 1568 p.p. GHANA. Ankasa River F.R., on low shrubs near the river, Jones 1375 p.p. Previously known only from the Kwa Falls, Calabar, Nigeria. 238 E. W. JONES & A. J. HARRINGTON 8. Cololejeunea crenatiflora (Stephani) Vanden Berghen in Bull. Jard. bot. natn. Belg. 42: 487 (1972). Leptocolea crenatiflora Stephani, Sp. hepat. 5: 842 (1916); E. Jones in Trans. Br. bryol. Soc. 2: 411 (1954). SIERRA LEONE. Freetown Peninsula, Leicester Peak, in forest c. 60 m below the summit, Jones 1428 p.p. Kambui Hills, Bambawo, low vegetation by pool in forest, Harrington 468 p.p., 469 p.p., 474 p.p., 483 p.p., 484 p.p. GHANA. Krokosua Hills F.R., leaves of low shrubs in forest, 490-550 m (1600-1800 ft), Jones 1396 p.p. Ankasa River F.R., low shrubs near the river, Jones 1375 p.p.; on palm leaves in swamp forest one mile south of the Ankasa bridge, Jones 1382. Subri F.R., on Raphia fronds in swamp forest, Jones 1355 p.p. Pra-Suhien F.R., Jones 1345b. Kade A.R.S., the dominant epiphyllous hepatic on herbs in wet places in forest, Jones 1221. Atewa Hills F.R., abundant in swamp forest, Jones 1294. Amedzofe, abundant on aroids beneath dripping rocks, Jones s.n. Cololejeunea crenatiflora is a widely distributed epiphyllous species in tropical Africa, from Sierra Leone eastwards to Tanzania and southwards to north-western Zambia (Pécs, 1975) and the Uluguru Mountains in Tanzania. In West Africa it is predominantly a lowland species, though it has been recorded as high as 1430 m on Mont Cameroun (Cameroon Mountain). It is especially abundant in wet places in forest, where it is sometimes one of the dominant epiphyllous species. The paucity of records from Sierra Leone is probably attributable to the lack of collections from suitable sites. 9. Cololejeunea cuneifolia Stephani ex A. W. Evans in Bull. Torrey bot. Club 38: 253 (1911). Leptocolea cuneifolia (Stephani ex A. W. Evans) A. W. Evans in Bull. Torrey bot. Club 38: 280 (1911); Stephani, Sp. hepat. 5: 843 (1916); E. Jones in Trans. Br. bryol. Soc. 2: 414 (1954). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., Harrington 86 p.p., 91 p.p., 103 p.p., Jones 1423; Leicester Peak, Jones 1428 p.p.; Bathurst, Kongo Dam, Jones s.n.; Guma Valley, mature forest, Harrington 101 p.p., 112 p.p.; Toke, Jones s.n.; Black Johnson Beach near York, thicket behind the beach, Harrington 15; York Pass, forest understorey, Harrington 625 p.p., 630 p.p.; Picket Hill, forest near summit, c. 850 m (2800 ft), Gledhill s.n. Lake Sonfon, on herbs in forest surrounding the ‘lake’, Harrington 70 p.p. Masimo, Marmo 160 p.p., 161 p.p. Kambui Hills, Bambawo, Harrington 411, 484 p.p., Jones 1526 p.p., 1543 p.p. Giema, secondary forest, Harrington 528 p.p., 549a p.p. Gola Hills, near Lalehun, Marshall s.n., by stream in forest, Jones 1567. GHANA. Krokosua Hills F.R., frequent on low shrubs in small gaps in forest, 490-550 m (1600-1800 ft), Jones 1396. Ankasa River F.R., low herbs and small shrubs near the river, Jones 1374b, 1375 p.p. Pra-Suhien F.R., frequent on herbs, Jones 1345c p.p. Cololejeunea cuneifolia often accompanies C. crenatiflora and is likewise often one of the most abundant epiphyllous hepatics in moist, sheltered places on herbs and low shrubs in the forest. The relative abundance of C. cuneifolia in Sierra Leone and its scarcity in Ghana is remarkable, and may perhaps indicate a preference for a somewhat different rainfall regime from that which favours C. crenatiflora. Cololejeunea cuneifolia is very close to C. duvignaudii E. Jones and also to C. zenkeri (Stephani) E. Jones. Jones (op. cit.) followed Evans (op. cit.) and Stephani (op. cit.) in placing C. cuneifolia in Leptocolea —i.e. as having a compressed perianth — whereas he placed the other two species in Cololejeunea, though their perianths are plane dorsally and therefore subcom- pressed, and thus intermediate between the compressed perianths of Leptocolea and the fully terete perianths of Cololejeunea sensu stricto. C. zenkeri is sufficiently distinct from the other two in its small, spherical hyaline papilla situated at the base of the apical tooth; moreover it is a montane species. Vegetatively C. cuneifolia and C. duvignaudii resemble each other in having a pyriform hyaline papilla on the side, or near the apex, of the apical tooth. Jones described C. cuneifolia as having a lobule often greatly reduced, but always less than 1/3 the length of the lobe, measured from insertion to apex, whereas C. duvignaudii had lobules 1/4-2/5 the length of the lobe. The greater range of material now available shows that C. cuneifolia is considerably more variable than was implied in 1954. The length of the lobule ranges from 1/9 to 1/3-5 the HEPATICS OF SIERRA LEONE & GHANA 239 length of the lobe, though in 80 per cent of the West African specimens it is 1/4 or less. The minority with lobules 1/3-5 the length of the lobe come from open sites on the Freetown Peninsula (Harrington 630 p.p., Jones 1428 p.p., Toke, Jones s.n.) and might be considered to be small-lobuled forms of C. duvignaudii, though they are clearly the same species as plants with smaller lobules from more sheltered sites on the Peninsula. The perianths of these large-lobuled Sierra Leone plants agree with C. cuneifolia. The lobules of the type collection of C. duvignaudii (Duvignaud 998c) vary from 1/3-7—1/2-4 of the length of the lobe. A review of all the available material suggests that C. cuneifolia is chiefly West African and north of the equator, while C. duvignaudii is exclusively south of the equator; the few records of C. cuneifolia from south of the equator may well be C. duvignaudii with reduced lobules. The leaves of C. cuneifolia frequently have papillae over one or more marginal rows of cells — a single papilla over the lumen of each cell. These papillae vary greatly in development; they may be detectable over the whole distal half of the leaf. 10. Cololejeunea dentata (E. Jones) R. M. Schuster in Beih. nov. Hedwigia 9: 175 (1963). Leptocolea dentata E. Jones in Trans. Br. bryol. Soc. 2: 161 (1953). SIERRA LEONE. Kambui Hills, Bambawo, low vegetation by pool in forest, Harrington 471 p.p. GHANA. Subri F.R., epiphyll in Raphia swamp, Jones 1355a. Apparently a rare species, known also from Mont Cameroun (Cameroon Mountain) and from Zaire (Kindu, Jones 798). 11. Cololejeunea elegans (Stephani) E. Jones in Trans. Br. bryol. Soc. 2: 424 (1954). Physocolea elegans Stephani, Sp. hepat. 5: 870 (1916). GHANA. Atewa Hills F.R., on tree boles in swamp forest (the ‘Kaolin Swamp’), scattered in small amounts through mats of other bryophytes, Jones 1328 p.p., 1331 p.p. A rare species, known otherwise only from the type collection in Cameroon (Bomana, Dusen 133) and from a recent collection on Kasigau Mtn, S.E. Kenya, by Faden (det. Pécs, Bizot & Pécs, 1974). 12. Cololejeunea floccosa (Lehm. & Lindenb.) Schiffner, Consp. hepat. archip. ind.:243 (1898); E. Jones in Trans. Br. bryol. Soc. 5: 571 (1968). SIERRA LEONE. Gola Hills, near Lalehun, epiphyllous on low vegetation in primary (?) forest, Marshall s.n. Probably the only record of this species from Africa. 13. Cololejeunea hyalino-marginata (Nees ex Montagne) Grolle in J. Hattori bot. Lab. 49: 85 (1981). Leptocolea leloutrei E. Jones in Trans. Br. bryol. Soc. 2: 146 (1953). Cololejeunea leloutrei (E. Jones) R. M. Schuster in Beih. nov. Hedwigia 9: 173 (1963). SIERRA LEONE. Gola North F.R., epiphyllous on herbs in unexploited primary forest, Jones 1568 p-p. GHANA. Krokosua Hills F.R., 490-550 m (1600-1800 ft), epiphyllous on low shrubs in small gaps in forest, Jones 1396 p.p. A variable epiphyllous species, hitherto known from Cameroon (Tixier, 1975), Sao Tomé, Kenya, and Tanzania where it is frequent on some of the mountains. It is also frequent in the Seychelles, the Comoro Islands, Madagascar, Réunion, and Mauritius (Grolle, op. cit): 14. Cololejeunea lanceolata E. Jones in Trans. Br. bryol. Soc. 2: 428 (1954). SIERRA LEONE. Kambui Hills, Bambawo, Marshall (Harrington 475 p.p.); on very low vegetation in forest near pool, Harrington 558a. 240 E. W. JONES & A. J. HARRINGTON GHANA. Ankasa River F.R., on low herbs in moist place in deep shade, Jones 1374a. Pra-Suhien F.R., local, Jones 1345a. Atewa Hills F.R., epiphyllous in swamp forest (the ‘Kaolin Swamp’), Jones 1331b p.p. Previously known only from Nigeria and Cameroon. 15. Cololejeunea myriocarpa (Nees & Montagne) A. W. Evans in Bull. Torrey bot. Club 38: 256 (1911). Cololejeunea minutissima subsp. myriocarpa (Nees & Montagne) R. M. Schuster in J. Elisha Mitchell scient. Soc. 71: 232 (1955). GHANA. Kade A.R.S., on branches of Bauhinia by the Rest House, Jones 1243a. Only a single colony of this plant was seen; it seems to be without gemmae, but in other respects it closely matches Cololejeunea myriocarpa, a species known from the West Indies, Mexico and southern Florida. Plants which are closely allied, if not identical, are widely distributed, but apparently rare, in tropical and South Africa, and have given rise to considerable differences of opinion as to their taxonomic status. Amongst these is C. myriantha (Herzog) S. Arnell from South Africa. Arnell (1963) records it from many localities. The most important difference from C. myriocarpa noted by Herzog (1952) is its larger cells. C. dissita E. Jones (Jones 19546) from Benin is a form which is, exceptionally, epiphyllous, and in which all the leaves have fully developed saccate lobules. C. minutissima subsp. utriculifera Vanden Berghen (Vanden Ber- ghen, 1961) is distinguished chiefly by the presence of some leaves which take the form of ‘utricles’, the lobules being as large as, or larger than, the lobes; it has been recorded only from Rwanda at 2000-2400 m, and may perhaps be a montane form. Schuster (op. cit.) considered Cololejeunea myriocarpa to be a subspecies of C. minutissima (Smith) Schiffner, but Tixier (1979) made them fully synonymous. I cannot accept Tixier’s opinion. C. minutissima was described from Britain, where it is not a variable species, and appears to differ consistently from C. myriocarpa and its allies from tropical Africa in its larger lobules, which are almost always saccate (only occasionally does the basal leaf of a branch or the pair of leaves immediately below a female inflorescence have explanate lobules), its larger gemmae, and female bracts with a distinct sinus between lobe and lobule, and with a blunt keel. Schuster noted these differences in American material of the two species, though he found some overlapping in the range of number of cells in the gemmae. [E.W.J.] The Ghana plant (and others that I have seen from Nigeria) cannot be ascribed to either Cololejeunea myriantha or to C. minutissima subsp. utriculifera. It seems to me that C. myriocarpa and C. minutissima are sufficiently distinct to make it a matter of no great importance whether the one is regarded as a subspecies of the other or not, and to maintain C. myriocarpa as a species makes nomenclature less cumbersome. The whole complex clearly needs critical revision on a world scale. [E.W.J.] 16. Cololejeunea nigerica (E. Jones) R. M. Schuster in Beih. nov. Hedwigia 9: 177 (1963). Leptocolea nigerica E. Jones in Trans. Br. bryol. Soc. 2: 150 (1953). SIERRA LEONE. Freetown Peninsula, York Pass, epiphyllous on low vegetation, Harrington 636 p.p. Three km south of Kabala, by the road to Makeni, in secondary forest, Harrington 126 p.p., 127 p.p. Bo, near the school, Revell (Herb. Jones). Kambui Hills, Bambawo, on leaves of mango, Jones 1541 p.p. (lobules atypical). GHANA. Asenanyo F.R., on branches in crown of Triplochiton, Jones 1412 p.p., and in crown of Terminalia superba, Jones 1413a p.p. (form in which some leaves lack hyaline cells). Pra-Suhien F.R., on bough of tree by the Rest House, Jones 1347a p.p. Kade A.R.S., on twigs of Citrus, Jones 1226. Aburi Botanical Gardens, on stone, Jones s.n. (GC) and on Bougainvillea twigs, Jones s.n. Amedzofe, epiphyllous on herbs in cocoa farms above the waterfall, Jones 1273 p.p. Kpedze Rest House, on twigs of cocoa, Jones s.n. Cololejeunea nigerica appears to be a West African lowland species. As the above records show it is frequent on the bark of twigs and small trees in open places, where it is probably often HEPATICS OF SIERRA LEONE & GHANA 241 overlooked. Although it was first described as an epiphyllous species, it seems to be less frequent in this habitat. 17. Cololejeunea peraffinis (Schiffner) Schiffner, Consp. hepat. archip. ind.: 245 (1898), var. elegans Benedix in Beih. Repert. nov. Spec. Regni veg. 134: 35 (1953); E. Jonesin Trans. Br. bryol. Soc. 5: 571 (1968). SIERRA LEONE. Lake Sonfon, by seasonal stream in forest surrounding the ‘lake’, epiphyllous, Harrington 70b p.p. 18. Cololejeunea pusilla Stephani in Hedwigia 34: 251 (1895); E. Jones in Trans. Br. bryol. Soc. 2: 426 (1954), var. pusilla GHANA. Southern Scarp F.R., near Begoro, on small barkless smooth log in moderate shade, Jones 1305. var. obtusifolia E. Jones in Trans. Br. bryol. Soc. 2: 427 (1954). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., Harrington 103 p.p.; Jones 1423; Guma Valley. mature forest, Harrington 112 p.p.; Black Johnson Beach near York, in thicket behind the beach, Harrington s.n.; York Pass, on low vegetation, Harrington 636 p.p.; Picket Hill, in forest near summit, 850 m (2800 ft), Gledhill s.n. About five km south of Kabala, secondary forest, Harrington 126 p.p., 127 p.p. Lake Sonfon, on herbs and small shrubs in forest surrounding the ‘lake’, Harrington 35 p.p., 44, 45, 70 p.p., 70c, 70d, 70e. Loma Mountains: between Kurubonla and Seradu, in forest by stream, Harrington 141; Seradu, in forest in valley by the village, c. 610 m (2000 ft), Jones 1518 p.p.; forest above Sokurela, Harrington 385 p.p., and at 1040 m (3400 ft), Jones 1514b p.p.; near ‘Camp 1’, c. 1220 m (4000 ft), on Marattia by stream, Harrington 292d p.p. Kambui Hills, forest above Bambawo, Harrington 421 p.p., 485 p.p., Jones 1526 p.p., 1543 p.p. Giema, secondary forest, on low vegetation, Harrington 528 p.p. Jawo, bank of the river Moa, on branches of tree at top of, or just above, the flood level, Jones 1536. Gola Hills: Lalehun, in forest by stream, Jones s.n.; near Lalehun, on low vegetation in primary (?) forest, Marshall s.n. GHANA. Krokosua Hills F.R., 490-550 m (1600-1800 ft), abundant on leaves of low shrubs in small gaps in forest, Jones 1396 p.p. Ankasa River F.R., on low shrubs in small gap in forest, Jones 1375 p.p. Begoro, on rocks in dry stream bed below the waterfall, in light-canopied forest, Jones 1307c. Atewa Hills F.R., on low herbs, Jones 1294 p.p. The relationship between Cololejeunea pusilla var. pusilla and var. obtusifolia requires further study; the two sometimes grow together, and intermediates sometimes occur, but var. obtusifo- lia is apparently always much the more abundant. Both are almost always epiphyllous, though occasionally occurring on other substrata. Var. obtusifolia is one of the most widely distributed epiphyllous hepatics in tropical Africa, often being the only epiphyllous species present in districts that are climatically marginal for their occurrence. It is, however, mainly a species of lowland forest; thus in the Loma Mountains it is frequent at 1000-1200 m, but has not been recorded from the higher forests of the plateau. Unlike some relatively drought-tolerant epiphyllous Lejeuneaceae which can grow in the crowns of forest trees, C. pusilla seems to be restricted to leaves of herbs and low shrubs. Like many other epiphyllous hepatics, it often grows near water. Perianths are relatively uncommon. 19. Cololejeunea zenkeri (Stephani) E. Jones in Trans. Br. bryol. Soc. 2: 420 (1954). SIERRA LEONE. Loma Mountains: forest near ‘Camp 1’, 1220 m (4000 ft), Harrington 289 p.p., and on Marattia by stream, Harrington 292d p.p.; valley forest near ‘Camp 2’, 1520-1680 m (5000-5500 ft), Morton (Harrington 364 p.p.), Jones 1483a p.p., Jones 1488. Cololejeunea zenkeri is closely related to C. cuneifolia; both have leaves of the same general shape and with the same type of marginal dentition and areolation. C. zenkeri is, however, quite distinct in having a spherical (not pyriform) hyaline papilla on the inner face of the base of the apical tooth (not on the side of the tooth). It appears to be strictly a montane species, occurring at higher altitudes than C. cuneifolia, probably in all the main groups of tropical African mountains eastwards to Tanzania. 242 E. W. JONES & A. J. HARRINGTON XVIII. COLURA (Dumort.) Dumort. 1. Colura digitalis (Mitten) Stephani, Sp. hepat 5: 931 (1916); E. Jones in J. Bryol. 10: 387 (1979). SIERRA LEONE. Freetown Peninsula, three km north-east of Toke, epiphyllous, Jones s.n. Loma Mountains: gallery forest c. one km west of ‘Camp 2’, 1520 m (5000 ft), Morton (Harrington 347 p.p., 354); bushes by spring by ‘Camp 2’, 1680 m (5500 ft), Jones 1495a. Kambui Hills, Bambawo, low vegetation by pool in forest, Harrington 472 p.p. GHANA. Ankasa River F.R., on twigs in crown of Samanea by the river, rare, Jones 1377b p.p. Subri F.R., on leaves of Raphia in swamp forest, Jones 1355 p.p. Atewa Hills F.R., epiphyllous in swamp forest (the ‘Kaolin Swamp’), Jones 1331b p.p. 2. Colura dusenii (Stephani) Stephani, Sp. hepat. 5: 931 (1916); E. Jones in J. Bryol. 10: 387 (1979). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., on Eugenia jambos, Harrington 750 p.p., in forest in valley bottom, Jones s.n.; York Pass, in forest by stream, Harrington 612 p.p., 614 p.p. GHANA. Ankasa River F.R., abundant on twigs in crown of Samanea by the river, Jones 1377 b p.p.; on leaves of undergrowth in moist gully, Richards R7054 p.p. (Herb. Jones). Subri F.R., on Raphia fronds in swamp forest, Jones 1355 p.p. Colura digitalis and C. dusenii sometimes grow together, as in Jones 1355 and 1377, but whereas C. digitalis is generally distributed in suitable localities throughout tropical Africa, and extends to the Comoro Islands, C. dusenii is very much more local. Both species may be epiphyllous or grow on twigs and branchlets, and both occur through a wide altitudinal range. 3. Colura tenuicornis (A. W. Evans) Stephani, Sp. hepat. 5: 942 (1916). Colura calyptrifolia subsp. tenuicornis (A. W. Evans) Vanden Berghen in Bull. Soc. r. Bot. Belg. 42: 463 (1972). SIERRA LEONE. Kambui Hills, Bambawo, low vegetation by pool in forest, Harrington 472 p.p.; on mango leaves near the ground, Jones 1541 p.p. XIX. CONOSCYPHUS Mitten 1. Conoscyphus trapezioides (Sande Lacoste) Schiffner in Engl. & Prantl, Nat. Pflanzenfam. 1 (3): 92 (1893). Lophocolea devexa Mitten in J. Proc. Linn. Soc. 7: 165 (1863 [‘1864’]). SIERRA LEONE. Loma Mountains, on boles of large Parinari excelsa on the margin of the forest, 1520-1680 m (5000-5500 ft), locally common, but sometimes as extremely slender shoots creeping amongst other bryophytes, Jones 1477a, 1490. Previously known in Africa only from Mont Cameroun (Cameroon Mountain), Sao Tomé, and more recently from the Uluguru Mountains in Tanzania (Bizot & Pécs, 1974), but with a disjunct distribution in the mountains of the Mascarenes, the Indo-Malayan region, and Samoa. XX. CYATHODIUM Kunze 1. Cyathodium africanum Mitten in J. Linn. Soc. (Bot.) 22: 327 (1886); E. Jones in Trans. Br. bryol. Soc. 2: 58 (1952). SIERRA LEONE. Freetown Peninsula: Mount Aureol, path below Kortright, in very sheltered rock crevice, Harrington 664; Leicester, vertical bank of ferruginous stone by road, 400 m (1300 ft), Jones 1450; Regent, beside stream, /rvine 222; by path from Regent to the Sugar Loaf, on brick supports of pipeline in forest, Harrington 645. Musaia Agricultural Station, on bare ground near the Rest House, Haswell B17 (sterile). Near road-bridge over tributary of the Rokel River, c. 32 km (20 miles) south-west of Kabala, on earth bank, Harrington 402 (sterile). Kambui Hills, Bambawo, on stonework below bridge, Harrington 562 (sterile). Sunkori, 19 km south of Kurubonla (9°02'N, 11°01’30"W), by stream, W. N. Woodhead (Herb. Richards). HEPATICS OF SIERRA LEONE & GHANA 243 GHANA. Mpraeso Scarp, Hall 2644. Amedzofe Mission House, /rvine 403. Doubtless far more widely distributed, especially in Ghana, than the records indicate. It appears to need a seasonally wet and nutrient-rich substratum, and often grows in deep shade, when it is rendered conspicuous by the brilliant luminous emerald green of the young thalli. It dies in the dry season. It frequently grows in man-made habitats, e.g. drains in villages. It is probably conspecific with the South American Cyathodium cavernarum Kunze. XXI. CYLINDROCOLEA R. M. Schuster 1. Cylindrocolea atroviridis (T. Sim) Vana in Lejeunia II, 98: 7 (1979). Cephaloziella atroviridis (T. Sim) S. Arnell in Bot. Notiser 108: 309 (1955); E. Jones in Trans. Br. bryol. Soc. 3: 431 (1958). SIERRA LEONE. About five km south of Kabala, in forest by the Makeni road, on fallen partly burned tree, Harrington 124. GHANA. Afram Headwaters F.R., on rotting logs in plantation, Jones 1416. Widely distributed in tropical Africa, but chiefly in climates with a strong dry season, and especially, as near Kabala, on wood that has been charred. Thus it is likely to be frequent in the drier parts of both Ghana and Sierra Leone where very few bryophytes have been collected. 2. Cylindrocolea chevalieri (Stephani) R. M. Schuster in Bull. natn. Sci. Mus. Tokyo 12: 666 (1969); E. Jones in J. Bryol. 9: 46 (1976). Alobiella chevalieri Stephani, Sp. hepat. 3: 351 (1908). Cephaloziella retusa E. Jones in Trans. Br. bryol. Soc. 3: 438 (1958). GHANA. Begoro, on wet rocks and on rotting log at edge of stream close to water level just above the waterfall, Jones 131la & b. Amedzofe, on wet rocks, sheltered but well lit, in stream just above the waterfall, Jones 1278e; also sterile, depauperate and scanty at the foot of cliff just below summit of the hill, Jones ! Previously known only from single records from Brazzaville on the Congo, the Falls of the Mougoungoulou in Gabon, and Idanre in western Nigeria. See Jones (op. cit.) for details. 3. Cylindrocolea ?ugandica (E. Jones) R. M. Schuster in Nova Hedwigia 22: 171 (1972 [‘1971’]). Cephaloziella ugandica E. Jones in Trans. Br. bryol. Soc. 3: 433 (1958). SIERRA LEONE. Loma Mountains, on bole of large tree on forest margin close to ‘Camp 2’, 1680 m (5500 ft), probably occasionally scorched, Jones 1476a p.p. This plant resembles Cylindrocolea ugandica in perianth, but differs in having a more shallowly lobed leaf with a broader insertion. In this respect it resembles an un-named sterile plant from Uganda in Herb. Jones (Murchison Falls, Gittins 19). These two plants may represent an undescribed species. It is, however, clear that this complex, which includes also C. gittinsii (E. Jones) R. M. Schuster, C. madagascariensis (Stephani) R. M. Schuster, and Cephaloziella abyssinica Gola, requires critical revision with the aid of more copious collections than are at present available. XXII. DICRANOLEJEUNEA (Spruce) Schiffner 1. Dicranolejeunea madagascariensis (Gottsche) Stephani, Sp. hepat. 5: 158 (1912); E. Jones in Trans. Br. bryol. Soc. 6: 73 (1970), var. madagascariensis SIERRA LEONE. Loma Mountains: forest near ‘Camp 1’, c. 1220 m (4000 ft), on tree bole, Harrington 264 p.p.; valley north-west of Bintimani, on stem of liane in deep shade, Jones 1498c; west-facing cliffs of Bintimani, c. 1830 m (6000 ft), on rock face, Harrington 213. Previously known from Cameroon, East Africa, and Madagascar. 244 E. W. JONES & A. J. HARRINGTON XXIII. DIPLASIOLEJEUNEA (Spruce) Schiffner 1. Diplasiolejeunea aulae E. Jones in J. Bryol. 7: 552 (1973). GHANA. Atewa Hills F.R., boughs in crown of Lophira, Jones 1319c (BM, type); upper part of bole of Antiaris africana, Jones 1320 p.p. 2. Diplasiolejeunea cavifolia Stephani in Bot. Jb. 20: 318 (1895); Vanden Berghen in Bull. Jard. bot. natn. Belg. 47: 220 (1977). Diplasiolejeunea brachyclada A. W. Evans in Bull. Torrey bot. Club 39: 216 (1912); Grolle in Feddes Reprium 73: 86 (1966). GHANA. Aiyaola F.R. near Kade, on branches and twigs in crown of Piptadeniastrum, Jones 1236 p.p., and on branches and twigs of Khaya, Jones 1237 p.p. 3. Diplasiolejeunea cornuta Stephani, Sp. hepat. 5: 918 (1916); Vanden Berghen in Bull. Jard. bot. natn. Belg. 47: 221 (1977). Fig. 3. GHANA. Aiyinasi A.R.S., on Citrus leaves, Jones 1385a. Subri F.R., on Raphia leaves in swamp forest, Jones 1355 p.p. Diplasiolejeunea cornuta is widely distributed but apparently local in tropical Africa. It is a variable plant; the type, illustrated by Vanden Berghen (1960a), has entire leaves with a rounded apex, (‘marginibus repandis vel erosulis’ according to Stephani (op. cit.)) but forms are frequent in which the leaves are more or less dentate, as illustrated by Jones (1973) and Vanden Berghen (1977). Jones 1385a differs in several respects from other dentate-leaved forms of the species that I have seen from Africa, and may prove to be a distinct species, although Grolle considers (in litt.) that it falls within the range of variation of D. cornuta. Usually in dentate forms of D. cornuta the postical margin of the lobe is arched and makes a well marked angle with Fig. 3 Diplasiolejeunea cornuta Stephani forma. (a), (b) leaves, X60; (c) tooth at distal end of postical margin of leaf, x300. Drawn from Jones 1385a. the keel, and it bears one or two blunt teeth at its proximal base, quite close to the keel; the teeth on the antical margin are very irregular, and many are formed by strongly protuberant single cells. In Jones 1385a the postical margin is straight, almost in line with the keel, and entire except for a strong triangular spur-like backwardly directed tooth at its apex. The antical margin bears three or four similar evenly-spaced triangular teeth. The group of ocelli at the base of the leaf consists of up to ten cells (usually two to five in D. cornuta), and the ocelli are brown. [E.W.J.] 4. Diplasiolejeunea harpaphylla Stephani, Sp. hepat. 5: 919 (1916). Diplasiolejeunea incurvata Jovet-Ast & P. Tixier in Revue bryol. lichén. 31: 29 (1962); fide Grolle in litt. SIERRA LEONE. Kasewe F.R., on savanna tree amongst Calymperes sp., Richards s.n. (Herb. Jones). GHANA. Ankasa River F.R., on twigs in crown of Samanea by the river, Jones 1377b. Aiyinasi A.R.S., on bough of small tree by the Rest House, Jones 1384b. Asenanyo F.R., twigs in crown of Terminalia superba, Jones 1413a p.p. Pra-Suhien F.R., on bough of Acacia by the Rest House, Jones 1347a. Aiyaola F.R. near Kade, twigs and branchlets in crown of Piptadeniastrum, Jones 1236 p.p., and in crown of Cylicodiscus, Jones 1240. HEPATICS OF SIERRA LEONE & GHANA 245 XXIV. DREPANOLEJEUNEA (Spruce) Schiffner 1. Drepanolejeunea ankasica E. Jones, sp. nov. Figs 4 & 5. Monoica. Folia aliquantum falcata, apice late acuto, decurvo, margine subintegro aut irregulariter obtuseque dentata, basi antica proximale stricta, distale arcuata, caulem excedente. Cellulae medianae 24-36 X 15-18 wm, parietibus vix incrassatis; ocelli tres uniseriati adsunt, ocellis duobus basalibus contiguis, tertio superiore sejuncto. Lobulus fusiformis margine libero incurvo, ex quatuor cellulis constructo, dente apicali longe libero, curvato. Amphigastria obtrapeziformia, quam caule 2-2'2-plo latiora lobis latis, acutis, patulis, sinu latissimo, basi cuneata. Perianthium circa 0-5 x 0-35 mm, obconicum, quinquecarinatum, carinis altis, distale crenulatodentatis. Typus: Ghana, Ankasa River Forest Reserve, Jones 1377 pro parte, in Herb. Mus. Brit. conservatus. Green, creeping over twigs. Stem 45-60 «m diam. with cortical cells c. 18 X 35 wm. Leaves approximate to imbricate, 0-23—-0-26 mm long x 0-2-0-3 mm wide, antically secund, more or less falcate with decurved acute but rather widely pointed apex; antical base proximally straight and parallel to the stem, then arching across the stem and passing into the arched antical margin, the postical margin concave, the apex ending in a single short cell, rather widely or even bluntly pointed; the margin nearly entire or the antical margin irregularly dentate with up to 6-8 low, obtuse, irregularly spaced teeth, each formed by a conically protuberant cell. Lobule fusiform, about two fifths the length of the lobe, the free margin incurved, not visible in situ, of four cells; the apical tooth long (c. 30 x 8um), curved, acute, free for most of its length with the large hyaline papilla near its base on the proximal side, lying against the margin of the lobule, the tip of the apical tooth engaging with protuberant cells on the inner face of the lobe. Fig.4 Drepanolejeunea ankasica E. Jones. (a) part of shoot; (b) leaves; (c) female involucre; (d) perianth with bracteole; (e) perianths. All x56. (f) underleaves, x280; (g) base of leaf, antical view, x280-keel of lobule and its junction with the lobe to the left, the basal ocelli partly overlapped by adjacent cells; (h) apical tooth and free margin of lobule, x 470; (i) apical tooth of lobule, x 470. All drawn from Jones 1377 (holotype). 246 E. W. JONES & A. J. HARRINGTON oO weal Fig. 5 Drepanolejeunea ankasica E. Jones. (a) leaf, antical view; (b) leaf, postical view; (c) part of male inflorescence showing two bracts with their underleaves. All x280. Drawn from Jones 1377 (holotype) . Cells of the antical margin of the lobe c. 15 x 15 wm, of the third and fourth rows in from the antical margin 20-25 x 15-22 wm, subapical cells 15-30 x 15-22 um, median cells 24-36 x 15-18 uum, the walls thin with trigones small or absent, and weak intermediate thickenings; usually with three ocelli in an interrupted line, the two basal ocelli contiguous, just above the margin of the lobule, the upper ocellus isolated. Underleaves two to two and a half times the width of the stem, obtrapeziform with widely diverging, broadly triangular lobes ending in a single short cell, separated by a very shallow broad sinus, the base cuneate. Monoecious. Androecia on long or short shoots, often close to a gynoecium (sometimes on a subgynoecial innovation), of four or five pairs of bracts with large imbricate underleaves throughout, the bracts monandrous, antheridia 55—60 wm diam. Gynoecia often on short lateral shoots, sometimes on long shoots, with one or two subgyroecial innovations. Bracts and bracteole irregularly and bluntly dentate and containing ocelli. Perianth 0-33-0-35 x 0-4 -0-5 mm, obconic with five long deep equal keels and a strong rostrum; the keels more or less crenulate-dentate distally, especially on the angle, with conically protuberant cells. HEPATICS OF SIERRA LEONE & GHANA 247 GHANA. Ankasa River F.R., on twigs in the crown of a Samanea near the river, with Colura spp., Diplasiolejeunea harpaphylla, Cheilolejeunea surrepens etc., Jones 1377 p.p. (BM, holotype). Drepanolejeunea ankasica belongs to the subgenus Drepanolejeunea Grolle (Grolle, 1976). In general appearance it resembles D. claviformis Stephani and D. friesii Vanden Berghen. It differs from both of these in being monoecious, in the slight development of trigones in the cells of the leaf, and in the apical tooth protruding far beyond its proximal neighbour; in D. claviformis and D. friesii the apical tooth is engaged for most of its length and free only near the apex. Drepanolejeunea cultrella (Mitten) Stephani is monoecious (though it has sometimes been described as dioecious), but has very different horned perianths and underleaves with narrow setiform lobes of elongate cells. D. claviformis also differs in its perianths (which are, however, usually rare) with their inflated obtuse horns. 2. Drepanolejeunea cultrella (Mitten) Stephani, Sp. hepat. 5: 324 (1913); E. Jones in Trans. Br. bryol. Soc. 5: 568 (1968). SIERRA LEONE. Freetown Peninsula, three km north-east of Toke, epiphyllous, Jones s.n., de- pauperate and sterile, the determination therefore uncertain. 3. Drepanolejeunea ?friesii Vanden Berghen in Svensk bot. Tidskr. 45: 366 (1951). Fig. 6. STERRA LEONE. Loma Mountains: on sheltered tree boles on margin of the forest near ‘Camp 2’, 1680 m (5500 ft), Jones 1474 p.p., 1475 p.p., 1477 p.p.; west of Bintimani, 1650 m (5400 ft), Jones 1491. GHANA. Atewa Hills F.R., boughs in crown of Cassipourea, Jones 1299d; boughs in crown of Lophira, Jones 1319a; upper part of bole of Antiaris, Jones 1320b; mixed with other bryophytes on boles of trees in swamp forest, Jones 1331a p.p. Fig.6 Drepanolejeunea ?friesii Vanden Berghen. (a) leaves; (b) perianth; (c) perianth with involucre. All x80. (d) apical tooth of lobule, x666. Drawn from Jones 1320b. 248 E. W. JONES & A. J. HARRINGTON Female inflorescences, but no perianths, are present in the collections from the Loma Moun- tains, but they appear to be the same as those from the Atewa Hills. Perianths (hitherto undescribed for Drepanolejeunea friesii) are present in Jones 1319a and, more copiously, in Jones 1320b. Pearson (1931) recorded Drepanolejeunea clavicornis Stephani (= D. claviformis Stephani) amongst Saxby’s collections from Tarkwa, Ghana, but it has not been possible to trace the specimen at either Manchester or Cambridge; it may have been the same species as that from the Atewa Hills. D. friesti is very similar to D. claviformis in vegetative characters; Vanden Berghen (1961) described them as differing chiefly in the absence of any basal ocellus from the leaves of D. friesii — an unusual feature in the genus. D. claviformis has been recorded only from West Africa, whereas D. friesii has been recorded from many localities in the East African mountains. The plants from the Loma Mountains and Atewa Hills, tentatively identified as D. friesii, have a line of two to three basal ocelli, sometimes interrupted, in some leaves, above the lobulus, but they are apparently not always present. The perianths are different from those described for D. claviformis, and the plants also seem to differ from D. claviformis and resemble D. friesii in the female bracts and bracteole which are almost entire, with only a few short coarse teeth — not laciniately dentate as in D. claviformis. The perianths of Jones 1320b are 0-6-0-7 mm long and 0-36-0-4 mm in diameter, inflated and terete proximally, contracted distally, with a strong rostrum and with five compressed equal keels in the upper half, the keels tuberculate with prominent cells. XXV. DUMORTIERA Nees | 1. Dumortiera hirsuta (Sw.) Nees in Reinw., Blume & Nees in Nova Acta physico-med. 12: 410 (1824); Vanden Berghen in Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8 (1): 182 (1972). SIERRA LEONE. Loma Mountains, forest near ‘Camp 1’ above Sokurela, 1220 m (4000 ft), on stones and boulders partly submerged in stream, Harrington 269. XXVI. FOSSOMBRONIA Raddi 1.Fossombronia husnotii Corbiére in Mém. Soc. natn. Sci. nat. Math. Cherbourg 26: 353 (1889); Vanden Berghen in Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8 (1): 147 (1972). SIERRA LEONE. Loma Mountains: Bintimani, 1890 m (6200 ft), on vertical rock face, Jones s.n.; also on west-facing cliffs of summit, on earth, Harrington 212 (without spores). Fossombronia husnotii is widely distributed in tropical Africa, but is probably montane; it is, for example, known from the Uluguru Mountains in Tanzania (Pécs 6004/S, Herb Jones), and Jones collected a sterile plant which is probably this species on Mont Cameroun (Cameroon Mountain) at 2500 m (Jones 371). Its colourless rhizoids and the constant production of descending stolons and tubers form useful diagnostic characters. It is variable in size of spore and in the pattern formed by the lamellae on the face of the spore. Areolae may or may not be present in the centre of the face, but the lamellae are high, projecting as spines 5—10 um tall around the periphery. In the plant from Bintimani the spores are 50-60 wm diam., without areolae, and with 20—24 spines. 2. Fossombronia occidento-africana S. Arnell in Bot. Notiser 105: 317 (1952). SIERRA LEONE. Freetown Peninsula: Mount Aureol [‘Oriel’], Arnell 2413, 2448, 2517 (UPS), 2231, 2249, 2409 (S), 2497 (S, lectotype); Fourah Bay College, moist roadside bank, Richards R7086; Fourah Bay College B.R., on boulder by path to Heddle’s Farm, Harrington 599; Charlotte Falls, shaded earth bank by road above the falls, Harrington 640 p.p. Musaia—Gbentu road, one mile north of the river Mongo, on bare earth in dried-up river bed, Sellar B35 p.p. Kambui Hills, Bambawo, shaded earth bank, Harrington 504. Giema, earth bank in roadside hollow, Harrington 555. HEPATICS OF SIERRA LEONE & GHANA 249 GHANA. Amedzofe, earth bank near the Training College, Hall (GC 47071); eroded earth below a land drain, Hall (GC 47226 & Herb. Jones). Konkori Scarp, Mole Game Reserve, abundant on sandstone rocks under trees, Hall s.n. — probably this species but without sporogonia. Arnell described Fossombronia occidento-africana from his collections made on ‘Mount Oriel’ in 1951, but did not specify a type; 2497 is his most copious collection, with numerous mature sporangia, and it is one from which he made a microscope preparation; we therefore select it as the lectotype. The original description is inaccurate in certain respects. The following descrip- tion is based on Arnel! 2497 and on other collections from Sierra Leone and Ghana: Monoecious, but probably protandrous. Shoots to c. 8 mm long, unbranched or occasionally dichotomous, not forming tubers. Stem in the fertile part of a shoot approximately semicircular in transverse section, 0-4—-0-5 mm wide X 0-25-0-3 mm thick, in the older parts of the shoot narrower than thick, c. 0-2 x 0-25 mm; medullary cells with endophytic fungus. Rhizoids purple. Leaves more or less oblong or rounded-trapezoidal, slightly narrowed to the insertion, the apex truncate and retuse or rounded and very shallowly lobed, often undulate, 0-7—1-5 mm long x 0-75-1-5 mm wide. Cells of leaf near the apex 30 X30—40 X 40 or 30 X 50 um, in the middle of the leaf 25-45 x 35-60 um, at the base up to 60 x 100 um, the walls thin, without trigones, cuticle smooth. Involucres obconic, c. 1-4 mm tall, the mouth irregularly lobed. Spores 36-50 ym diam. (40-45-50 xm in Arnell 2497), the distal face irregularly areolate, the areolae 4-6 wm diam., 8-10 across the face of the spore, the walls raised at the angles to form processes 2-3 um tall which appear on the periphery of the spore as 30—40 or more ‘spines’; the proximal face with only a few irregular thickenings. Elaters mostly 3-spiral, but many 2-spiral at one or both ends, anda few entirely 2-spiral. Cells of sporangium wall with massive radial thickened bands, very few of which extend across the tangential walls (thus not making ‘semicircular’ bands), 3-4 bands on each radial wall, spaced 10-15 wm apart in the undehisced sporangium, but more crowded after dehiscence. . Arnell described the cuticle as having ‘1—2 long striae per cell’, and cells with ‘sometimes small trigones’. We have not been able to find either striae or trigones, and they are certainly normally absent. Fossombronia occidento-africana is the commonest species of Fossombronia in lowland West Africa; it is known from several localities in Nigeria (e.g. Jones 854 from Enugu) and is the plant that Augier (to whom we are indebted for lending us specimens) found at Yaoundé in Cameroon and which he assigned ‘a titre provisoire’ to F. crozalsii Corbiére (Augier, 1972). We agree with Augier that this West African Fossombronia may indeed prove to be F. crozalsii, which is, however, imperfectly known. We have had to rely on descriptions and comparison with a single small specimen of F. crozalsii collected by Jones in Tubney Wood, Berkshire, England, in 1938. The chief differences seem to be: (1) Spores of Fossombronia crozalsii are usually said to be not more than c. 40 wm diam. (e.g. Paton (1973) finds that they are 34-41 ym diam.), whereas spores of F. occidento-africana are often c. 45 wm or larger. But some collections have smaller spores; thus the plants from Amedzofe have spores 33-40 wm diam., and Richards R7086 from Fourah Bay College has spores 36—42 wm diam. (2) The elaters are described as bispiral in F. crozalsii, though the Tubney Wood plant has some 3-spiral elaters. (3) The Tubney Wood plant has much thinner and sparser bands of thickening in the sporangium wall. Of these differences the last two seem to be the most important. We prefer to keep the two species separate until the range of variation of F. crozalsii is adequately known. XXVIII. FRULLANIA Raddi 1. Frullania africana Stephani in Hedwigia 30: 269 (1891); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 151 (1976). GHANA. Southern Scarp F.R., near Begoro, on Celtis, Hall s.n. (GC). Amedzofe, in coffee planta- 250 E. W. JONES & A. J. HARRINGTON tions, Jones s.n. (GC). Unlocalised and undated, Akpabla 588 (BM), the collecting number indicates that this was collected c. August 1936 in the Achimota district, possibly at Aburi (J. B. Hall, in Jitt.). Not hitherto recorded from Ghana. Known only from a few scattered localities near the west coast of Africa, from the mouth of the Congo to Guinea. It appears to be characteristic of hill country, but occurs at lower altitudes than Frullania arecae which it resembles, except in its paroecious inflorescence and its 4- or 5-keeled perianth. 2. Frullania apicalis Mitten in Phil. Trans. R. Soc. 5: 168 (1879); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 47 (1976), var. apicalis SIERRA LEONE. Freetown Peninsula: Leicester Peak, on Combretum in scrub amongst rocks, 520 m (1700 ft), Jones 1453b; Sugar Loaf Mountain, Richards R7129, det. Vanden Berghen; three km north-east of Toke, on bole of small tree in thicket on rocky slope, Jones 1579, det. Vanden Berghen; Toke, on bole of large Chrysobalanops in thicket at the back of the beach, Jones 1445, det. Vanden Berghen; Picket Hill, on tree in forest just below the summit, c. 850 m (2800 ft), Gledhill s.n. Loma Mountains: between ‘Camps 1 and 2’, c. 1370 m (4500 ft), on rocks sheltered by trees, Jones 1472, det. Vanden Berghen; near ‘Camp 2’ (below Bintimani), 1520-1680 m (5000-5500 ft), on branch of isolated small tree, Jones 1480a, det. Vanden Berghen, and on adjacent rock, Jones 1480c, and on branches of Craterispermum laurinum on forest margin, Harrington 147, det. Vanden Berghen; c. three km south of ‘Camp 2’ between Bintimani and Dawule, 1370 m (4500 ft), on Dissotis leonensis growing amongst boulders, Harrington 229, det. Vanden Berghen. Gola Hills, Lalehun, on upper part of bole of large tree, Jones 1554a, det. Vanden Berghen. GHANA. Ankasa River F.R., twigs in crown of Samanea by the Ankasa River rapids, Jones 1377 p.p. Atewa Hills F.R., c. 760 m (2500 ft), boughs in crown of Braeya, Jones 1286a, of Lophira, Jones 1319a, of Antiaris, Jones 1320, all det. Vanden Berghen; in crown of Cassipourea, Richards & Hossain (GC 36667). Frullania apicalis has a wide, but disjunct, distribution in East Africa, where it is locally abundant from Uganda southwards to Mozambique, and in Madagascar and the Mascarenes. The distribution in West Africa is similarly disjunct, from the Bight of Biafra westwards to Sierra Leone. Usually its habitats seem to be montane, despite its occurrence at sea-level near Freetown. Thus, in Ghana it is abundant in the crowns of trees in the Atewa Hills, but was not found in the crowns of trees in the lowland forest reserve of Aiyaola. Its occurrence near sea-level in the Ankasa River Forest Reserve in western Ghana is paralleled by the presence of other montane species — e.g. Radula boryana and Bazzania decrescens. In the Freetown Peninsula F. apicalis is the only species of Frullania that has been recorded, and it appears to be rare. In the Loma Mountains it is abundant above c. 1400 m, together with lichens (e.g. Usnea sp., Parmelia spp., Anaptychia sp.) clothing the branchlets of isolated small trees and bushes that grow on rock outcrops. Frullania apicalis grows on well-illuminated branches in the open crowns of large trees, or on isolated bushes and small trees. It appears to be restricted to natural communities. 3. Frullania apiculata (Reinw., Blume & Nees) Dumort., Recueil observ. Jungerm.: 13 (1835); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 102 (1976). GHANA. Ankasa River F.R., twig in crown of Samanea, Jones 1377 p.p. Aiyinasi A.R.S., on branches of Citrus, Jones 1383, det. Vanden Berghen. Atewa Hills F.R., in crown of Braeya, Jones 1286c, det. Vanden Berghen; in crown of Antiaris, Jones 1320b. Apparently a local species in Africa, in similar habitats to Frullania apicalis, with which it was mixed in Jones 1377, 1286, and 1320. 4. Frullania arecae (Sprengel) Gottsche in K. dansk. Vidensk. Selsk. Skr. 6: 322 (1867); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 154 (1976). Frullania ecklonii (Sprengel) Gottsche & Lindenb., Syn. hepat.: 413 (1845). SIERRA LEONE. Lake Sonfon, on bole of mango at edge of forest, Harrington 26. Loma Mountains: Jaeger 1376 (PC), det. Vanden Berghen; near ‘Camp 2’, 1520 m (5000 ft), on branches of Craterispermum laurinum on edge of forest, Harrington 145 p.p., 146, det. Vanden Berghen. GHANA. Aburi Botanical Gardens, on Cupressus, Jones s.n. (GC). HEPATICS OF SIERRA LEONE & GHANA 250 5. Frullania depressa Mitten in J. Proc. Linn. Soc. 7: 168 (1863 [‘1864’]); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 206 (1976). SIERRA LEONE. Loma Mountains: near ‘Camp 2’, 1520-1680 m (5000-5500 ft), on branches of small isolated trees, Jones 1480b, 1495 p.p., 1497b; Bintimani, on dolerite rocks at foot of west-facing cliffs, c. 1830 m (6000 ft), Harrington 221, det. Vanden Berghen. On the Loma Mountains usually associated with Frullania apicalis. A local montane species known elsewhere in Africa from Cameroon and in widely scattered localities in East Africa, from Ethiopia southwards to Zimbabwe, and in Madagascar. 6. Frullania diptera (Lehm. & Lindenb.) Gottsche, Lindenb. & Nees, Syn. hepat.: 420 (1845); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 194 (1976). GHANA. Bunso, on cocoa, Jones s.n. Kpedze Rest House, on cocoa, Jones s.n. Between Bame and Todome, on cocoa, Jones s.n. A local and apparently rather rare species, which apparently always occurs in small quantities, not hitherto recorded from Ghana. The records cited by Vanden Berghen (op. cit.) show three main areas where it occurs: (1) the lower hills in south-eastern Kenya and north-eastern Tanzania, from the Shimba Hills to the Southern Highlands, (2) central Madagascar, (3) the coastal region of South Africa, from Capetown to Durban. For West Africa he gives only a single record, from northern Nigeria. It may, however, be much overlooked, as it is not conspicuous and rarely, if ever, occurs abundantly in pure patches. It is pale, often yellowish green in colour, with flat complanate leaves closely appressed to the substratum. Although it occurs as high as 1750 m in Tanzania, it seems to be chiefly a lowland species. 7. Frullania eplicata Stephani, Sp. hepat. 4: 679 (1911); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 83 (1976). GHANA. Aiyaola F.R., on boughs of Piptadeniastrum, Jones 1233, det. Vanden Berghen. Amedzofe, in savanna woodland just below the summit of the hill, on the upper part of the boles of old but small Lophira and Terminalia, Jones 1258, det. Vanden Berghen. 8. Frullania ericoides (Nees) Montagne in Annls Sci. nat. (Bot.) II, 12: 51 (1839); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 161 (1976). Frullania squarrosa (Reinw., Blume & Nees) Dumort., Recueil observ. Jungerm.: 13 (1835). SIERRA LEONE. Musaia town, on mango, Donald s.n. Kabala, on tree trunk in open woodland below the District Officer’s bungalow, Harrington 137 p.p. About four km south of Kabala, on isolated small tree in swampy valley, Harrington 118 p.p. About five km south of Kabala, on small tree in secondary forest, Harrington 125 p.p. Lake Sonfon, on tree trunk at edge of forest surrounding the ‘lake’, Harrington 27. Loma Mountains, Seradu, on tree boles in forest by the village, in small amounts mixed with Mastigole- Jeunea auriculata etc., Jones 1515, 1517. GHANA. Bia North F.R., boughs in crown of Entandophragma etc., rare, Jones ! Asenanyo F.R., in crown of Triplochiton, Jones 1412 p.p., det. Vanden Bergen. Kumasi, in garden, /rvine 17. Oda, on cocoa, Irvine 10. Southern Scarp F.R., near Begoro, Jones ! Aburi Botanical Gardens, Foote 3, Irvine 19a, Jones s.n. Amedzofe, abundant in farms, Jones ! Between Bame and Todome, on cocoa, Jones s.n. Pantropical and polymorphic, Frullania ericoides is probably the most abundant species of its genus, predominantly lowland (though ascending to 2000 m in East Africa), and extending through a wide range of rainfall regimes. It is particularly frequent in farmland, secondary forest, and similar disturbed sites. The above records do not adequately indicate its abundance in southern Ghana, as, despite it polymorphism, it is easily recognized in the field and therefore need not be collected. On the other hand the indications of its rarity or absence from southern and western Sierra Leone are probably genuine. It is remarkable that it has not been recorded from around Freetown, where apparently suitable disturbed habitats abound, nor from Njala. 252 E. W. JONES & A. J. HARRINGTON 9. Frullania nodulosa (Reinw., Blume & Nees) Nees, Syn. hepat.: 433 (1845); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 132 (1976). SIERRA LEONE. Gola North F.R., on fallen bough, Jones 1571 p.p. GHANA. Subri F.R., on fallen bough, Jones 1353. Oda, on cocoa, Irvine 9a. Kade A.R.S., crown of Parinari, Hossain (GC 36700). Aiyaola F.R. near Kade, crown of Triplochiton, Jones 1227b (GC). Also unlocalised specimens — Burton & Cameron s.n. (BM) and Cummins s.n. (BM). Apparently confined to the crowns of trees in old natural forest, and therefore most frequently seen on fallen boughs. 10. Frullania obscurifolia Mitten in Phil. Trans. R. Soc. 5: 168 (1879); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 143 (1976). Frullania borgenii Pearson in Forh. VidenskSelsk. Krist. 1890 (2): 8 (1891). SIERRA LEONE. Kabala, on tree-bole in open woodland below the District Officer’s bungalow, Harrington 136 p.p., det. Vanden Berghen. GHANA. Aburi Botanical Gardens, on palm, Jones 1577, det. Vanden Berghen. 11. Frullania purpurea Stephani, Sp. hepat. 4: 626 (1911); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 77 (1976). SIERRA LEONE. Kasabere Hills, on small savanna tree, Richards (Jones 1580), det. Vanden Berghen. Kasewe F.R., on small savanna tree, Richards s.n. (Herb. Jones). The two localities are very close together. A rare species, known elsewhere only from Benin (Jones 148), Calabar (Jones 211 p.p.), and, according to Vanden Berghen (op. cit.), Sao Tomé (Moller 20), Madagascar (five records), and Mauritius (two records). 12. Frullania spongiosa Stephani in Hedwigia 33: 147 (1894); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 167 (1976). GHANA. Bia North F.R., boughs in crown of Entandophragma, Jones 1411 p.p., det. Vanden Berghen. Krokosua Hills, on cocoa in farms near Bia bridge, Jones 1388a, det. Vanden Berghen. Asenanyo F.R., bough in crown of Triplochiton, Jones 1412, and of Terminalia superba, Jones 1413, det. Vanden Berghen. Oda, /rvine 7, and on cocoa, Irvine 9b, 10. Bunso, on base of oil palm, Jones 1301, det. Vanden Berghen. Kade A.R.S., upper branches of Aningeria, Hossain (GC 39321, 39325, 39335). Osenasi, on cocoa, Jones s.n. Aburi Botanical Gardens, on palm trunks, /rvine 22; on Cupressus, Jones s.n., det. Vanden Berghen; Jones 1576, det. Vanden Berghen. Amedzofe, 730 m (2400 ft), abundant on cocoa in farms above the waterfall, Jones 1270, det. Vanden Berghen. Vane, on rocks in roadside banks, Jones 1266b, det. Vanden Berghen. Perhaps the most abundant species of Frullania throughout much of southern Ghana, and like F. ericoides to which it is allied, especially frequent in farms on planted cocoa, etc., but also found in the crowns of large trees in natural forest. It is similarly abundant in parts of southern Nigeria and Cameroon, and like several other species of Frullania seems to occur in a disjunct area in East Africa. 13. Frullania trinervis (Lehm. & Lindenb.) Gottsche, Lindenb. & Nees, Syn. hepat.: 427 (1845); Vanden Berghen in Bull. Jard. bot. natn. Belg. 46: 198 (1976). SIERRA LEONE. Kabala, on tree trunk in open woodland below the District Officer’s bungalow, Harrington 137. GHANA. Between Bame and Todome, on cocoa, Jones s.n. Widely distributed throughout tropical and South Africa. From analogy with Nigeria it may be expected to have its greatest frequency in northern Ghana and Sierra Leone; Vanden Berghen (op. cit.) cites records from that part of Guinea between Sierra Leone and the headwaters of the Senegal River. HEPATICS OF SIERRA LEONE & GHANA 253 XXVIII. GONGYLANTHUS Nees 1. Gongylanthus ericetorum (Raddi) Nees, Naturgesch. europ. Leberm. 2: 407 (1836); E. Jones in Trans. Br. bryol. Soc. 4: 649 (1964); Vanden Berghen in Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8 (1): 45 (1972). SIERRA LEONE. Loma Mountains, frequent on earth amongst grass tussocks from c. 1220 m (4000 ft) upwards: south of ‘Camp 2’, Harrington 296 p.p.; near ‘Camp 2’, 1520 m (5000 ft), Harrington 144; lower slopes of Bintimani Peak, on peaty soil, 1680 m (5500 ft), Harrington 178. 2. Gongylanthus richardsii E. Jones in Trans. Br. bryol. Soc. 4: 650 (1964). SIERRA LEONE. Loma Mountains, frequent on Bintimani, but generally at higher levels than Gongylanthus ericetorum, not descending below c. 1830 m (6000 ft): summit plateau, beneath overhanging rock, c. 1940 m (6360 ft) Harrington 208; on earth around tussocks of Trilepis, Hypoxis etc., 1860-1900 m, Jones 1487. Known elsewhere only from the volcanic soils of Mont Cameroun (Cameroon Mountain) and, in Tanzania, of Kilimanjaro (Bizot & Pécs, 1974) and Rungwe. XXIX. JUNGERMANNIA L. 1. Jungermannia ?borgenii Gottsche ex Pearson in Forh. VidenskSelsk. Krist. 1892 (14): 11 (1893); Vania in Folia geobot. phytotax. 9: 289 (1974). SIERRA LEONE. Musaia—Gbentu road, one mile north of the river Mongo, on bare earth in dried-up river bed, Sellar B35 p.p. (no perianths seen). Musaia Agricultural Station, on bare ground near the Rest House, Sellar B12 p.p. (no perianths seen). Jungermannia borgenii is widely distributed in tropical Africa and extends to South Africa and the Mascarenes. I am sceptical as to whether Jungermannia borgenii can be safely separated from J. dusenii (Stephani) Stephani in the absence of perianths. [E.W.J.] XXX. LEJEUNEA Libert 1. Lejeunea autoica R. M. Schuster in J. Hattori bot. Lab. 25: 6 (1962); E. Jones in J. Bryol. 10: 389 (1979). GHANA. Atewa Hills F.R., c. 730 m, on boughs in crown of Braeya, Jones 1286 p.p.; on bole of middle-storey tree, Jones 1291; on stem of Vernonia, Jones s.n. Originally described from Florida, and recorded elsewhere only from the Atewa Hills and Mont Cameroun (Cameroon Mountain), this imperfectly known species is likely to prove widely distributed. It could readily be passed over as the much more abundant dioecious species, Lejeunea confusa. 2. Lejeunea brenanii E. Jones in J. Bryol. 10: 391 (1979). SIERRA LEONE. Freetown Peninsula: Leicester Peak, in low forest just below the summit, Brenan (Jones 463, 464); on twigs and stems of shrublets amongst rock outcrops, 520 m (1700 ft), Jones 1455; on twigs of Phyllanthus in open scrub, 520 m, Jones 1457b. Gola North F.R., on upper part of bole of large tree, Jones 1554b. GHANA. Krokosua Hills F.R., abundant on boughs in the crowns of trees, 520-550 m (1700-1800 ft), Jones 1398; on exposed stems of shrubs on edge of forest, c. 550 m, Jones 1401a; epiphyllous on Culcasia, Jones 1399 p.p. Atewa Hills F.R., in crown of Cassipourea in small amounts on the sides of boughs and in crooks, where few other plants are present, Jones 1298. Aburi Botanical Gardens, /rvine s.n., Foote 4, Jones s.n. Amedzofe, near the waterfall, /rvine 412. Lejeunea brenanii is one of a group of critical taxa allied to L. flava, distinguished from the other members by its large size, large cordate underleaves, dioeciousness, long androecia with underleaves throughout, narrowly but longly keeled perianths, etc. It is evidently locally 254 E. W. JONES & A. J. HARRINGTON abundant and somewhat montane, often clothing the well-illuminated branchlets of trees and bushes with a mantle of long pendent shoots. When sterile it would be indistinguishable from L. ramosissima. J. Augier (in litt.) has recently detected it amongst his collections from Cameroon. 3. Lejeunea caespitosa Lindenb., Syn. hepat.: 382 (1845); E. Jones in J. Bryol. 7: 37 (1972). SIERRA LEONE. Freetown Peninsula: Mount Aureol [‘Oriel’], Arnell 2334b (Herb. Jones); boles of small trees by Mountain Torrent, Jones 1443, also s.n. Bo, near the school, epiphyllous, Revell (Herb. Jones). Loma Mountains, above Seradu, epiphyllous, 1040 m (3400 ft), Jones s.n. Kambui Hills, Bambawo, on bole of Musanga, Jones 1534; on mango leaves near the ground, Jones 1541b. Gola North F.R., epiphyllous by stream in forest, Jones 1567 p.p. GHANA. Aiyinasi A.R.S., on Citrus, Jones 1363b. Asanta, abundant on coconut palms on the beach, Jones 1359c. Asenanyo F.R., on boughs in crown of Triplochiton etc., Jones 1412; buttress of Terminalia, Jones s.n. Pra-Suhien F.R., rotting log in deep shade, Jones 1344a; epiphyllous, Jones 1345c p.p. Bobiri F.R. (6°39’ to 6°42'N, 1°16’ to 1°23’W), rotting log in deep shade, Jones 1414. Ochi Headwaters F.R.., West-Skinn 147, 153. Kade A.R.S., bole of Parinari, Hossain (GC 36686); bole of Diospyros, Jones 1223. Aiyaola F.R. near Kade, in crown of Triplochiton, Jones 1227c, 1229; on liane, Jones 1241. Atewa Hills F.R., on bole of Cassipourea, Richards & Hossain (GC 36685); epiphyllous on low herbs at edge of small gap in forest, Jones 1294 p.p. Begoro, rocks in dry stream bed below the waterfall, Jones 1307b p.p. Begoro—Bunso road, on buttress of Celtis, Jones 1303. Bunso, on boles of cocoa, Jones 1317a. Aburi Botanical Gardens, on bough of Cupressus, Jones s.n. Amedzofe, on shade trees in coffee farm, Jones 1247; on bole of old Combretum in savanna woodland just below summit of the hill, Jones 1259b; on rocks by stream just above the waterfall, Jones 1277; epiphyllous by the stream in cocoa farms, Jones 1273. Kpedze Rest House, on cocoa, Jones s.n. One of the commonest species of Lejeunea in lowland habitats throughout the wetter parts of tropical Africa, and possibly pantropical, having been recorded from other countries under other names; polymorphic. It is most commonly corticolous or lignicolous in shady, sheltered places. As the above records show, it sometimes grows in the crowns of large forest trees, but is much rarer here than on tree boles near the ground. In many districts it does not occur as an epiphyll, or does so only exceptionally, but in some localities in Ghana and Sierra Leone it was abundant on leaves. The epiphyllous forms tend to have underleaves which are conspicuously lunate, being deeply divided into narrow lobes which are often only 2-3 cells wide at the base, and the cells 2-3 times as long as wide, thus resembling the American Lejeunea longifissa Stephani (regarded as conspecific by Schuster (1980) ). Such plants may look distinctive, but the underleaves vary greatly, and often undergo transitions within a single shoot to the broader-lobed, short-celled form which is commonest in other habitats. 4. Lejeunea camerunensis (Stephani) E. Jones in J. Bryol. 7: 33 (August 1972) [non L. kamerunensis (Stephani) Vanden Berghen in Bull. Jard. bot. natn. Belg. 42: 446 (Decem- ber 1972)]. Eulejeunea camerunensis Stephani, Sp. hepat. 6: 417 (1923) (see Art. 68.1). Lejeunea tuberculiflora E. Jones ex Pécs in Acta bot. hung. 25: 231 (1979), nom. illeg. (Art. 63.1). ? Eulejeunea microclada Pearson in Annls Cryptog. exot. 4: 66 (1931). SIERRA LEONE. Freetown Peninsula: Mount Aureol, c. per., Arnell2334a (S, as Lejeunea saccatiloba Stephani); Fourah Bay College B.R., on bole of tree near Heddle’s Farm, c. per., Jones 1421; Leicester Peak, on old mango, sterile, Jones 1464 p.p.; Toke, on old Terminalia scutifera on the beach, male, Jones 1444a. Kabala, on trunk of mango near the District Officer’s bungalow, c. per., Harrington 140 p.p. Between Sokurela and Kurubonla, c. 500 m, on bole of Uapaca, c. per., Jones 1522c. Loma Mountains: forest above Sokurela, 910-1220 m (3000-4000 ft), on tree bole, mixed with Porella subdentata, Plagiochila spp. & Lejeunea eckloniana, c. per., Jones 1467b; Bintimani Peak, c. 1830 m (6000 ft), amongst dolerite boulders, c. per., Harrington 203 p.p. Kambui Hills, Bambawo, on boles of small trees in moderate shade, in forest above the old mine, c. per., Jones 1531, 1535; on old mango bole by the staff bungalows, c. per., Jones 1549. Jawo, near Kenema, by the river Moa, on tree boles at or just above the top of the flood zone, mixed with Lejeunea setacea, c. per., Jones 1536b. Gola Hills, Lalehun, shaded bole of tree, c. per., Jones 1556. HEPATICS OF SIERRA LEONE & GHANA 255 GHANA. ? Ankasa River F.R., in crown of Glutea, sterile, Jones 1378 p.p. ? Tafo Cocoa Research Inst., on trees in garden, sterile, Jones s.n. ? Bunso, on cocoa, female but without perianths, Jones 1317a. Kade A.R.S., on upper branches of Aningeria robusta, male, Hossain (GC 39330); female, but without perianths, Hossain (GC 39331); on bole of Parinari excelsa, male, Hossain (GC 36689). Amedzofe, on bole of old shade-tree in coffee farm, c. per., Jones 1245; in savanna woodland just below summit of the hill, on bole of Combretum, c. per., Jones 1256, and on Terminalia, male, Jones 1257c. Lejeunea camerunensis is widely distributed and often abundant in the wetter parts of tropical Africa, extending eastwards to Mauritius. In the absence of the characteristic perianths, however, it cannot be identified with certainty; thus records based on male or sterile plants must be regarded as doubtful. Jones (op. cit.) described the apical tooth of the lobule as ‘up to twice as long as wide’, and the perianth as ‘ecarinate, terete or bluntly angled at the sides and ventrally in the upper half, dorsally plane’. Some of the plants from eastern Sierra Leone (Jones 1531, 1556) show considerable variation in the perianths, some of which are weakly carinate near the apex, and may have a weak dorsal keel, though they have the characteristic tuberculate cells. Jones 1535 has completely ecarinate perianths, whereas Jones 1531 from the same site has perianths that are distinctly carinate. The apical teeth of the lobules of these plants are also somewhat longer than previously described — up to 30 x 10 wm. The type of Eulejeunea microclada Pearson has not been found; Pearson described only male inflorescences. What is presumably the same species exists mixed with Saxby’s specimen of Cheilolejeunea newtonii at CGE, but is sterile. The synonymy of EF. microclada with Lejeunea camerunensis thus remains uncertain. 5. Lejeunea confusa E. Jones in J. Bryol. 7: 24 (1972). Microlejeunea jungneri Stephani, Sp. hepat. 5: 812 (1915), non Lejeunea jungneri Stephani (1896). SIERRA LEONE. Freetown Peninsula, on stems of bushes by Mountain Torrent, mixed with Lejeunea caespitosa, Cheilolejeunea serpentina et al., c. per., Jones 1443b. Lake Sonfon, on small tree in grassland near the ‘lake’, Harrington 31 p.p. Loma Mountains, Bintimani Peak, 1830 m (6000 ft), amongst dolerite boulders, Harrington 203 p.p. Kambui Hills, Bambawo, epiphyllous on low vegetation near pool in forest, Harrington s.n.; epiphyllous on low vegetation near stream, Harrington 423 p.p. Gola North F.R., on bole of Vitex mixed with Calymperes sp., c. per., Jones 1558. GHANA. Ankasa River F.R., on bole of Vitex, Jones 1369, and in the crown, Jones 1368b. Widely distributed and frequently abundant in tropical Africa, extending eastwards to Tanza- nia, but difficult to identify when sterile. Perianths are usually rare and difficult to find. It may readily be confused with small forms of Lejeunea caespitosa or with L. autoica, both of which differ in being monoecious and in the structure of the well-developed lobulus; L. caespitosa also has larger cells and a more translucent texture. L. confusa is usually corticolous; its exceptional occurrence as an epiphyllous species in Sierra Leone parallels the exceptional occurrence there of L. caespitosa and Cheilolejeunea serpentina as epiphylls. 6. Lejeunea eckloniana Lindenb., Syn. hepat.: 381 (1845); E. Jones in J. Bryol. 8: 78 (1974). SIERRA LEONE. Freetown Peninsula: Leicester Peak, 520 m (1700 ft), on rock, sheltered by deciduous bushes, Jones 1456; valley above Kongo Dam, Bathurst, 490 m (1600 ft), on barkless rotten log in forest, Jones 1434b p.p.; Guma Valley, on trunk of small tree in forest, Harrington 109. Loma Mountains: Sokurela, 610 m (2000 ft), on bole of oil palm in swamp forest, Jones 1465a; between Sokurela and ‘Camp 1’, Harrington 394, 395, Jones 1467c; forest above Sokurela, 1220 m (4000 ft), on boulder, Jones 1470 p.p. Kambui Hills, Bambawo, epiphyllous on low vegetation by stream in forest, Harrington 423 p.p.; on boles of small trees in forest, 340 m (1100 ft), Jones 1527 p.p., 400 m (1300 ft), Jones 1547b. GHANA. Krokosua Hills, 550 m (1800 ft), on well-illuminated but sheltered stems of shrubs on edge of forest (sterile and elobulate), Jones 1401a p.p. Afram Headwaters F.R., on log in fragment of natural forest, Jones s.n. Subri F.R., on decaying Raphia palm in swamp forest, Jones 1352. Begoro, in dry stream bed below the waterfall, Jones 1307b; on flat rocks just above water level by the stream above the falls, Jones 1311 p.p. Atewa Hills F.R., on stem of Vernonia, Jones s.n. Kade A.R.S., on boles of small trees in forest, Jones 1216b, 1223. Aiyaola F.R. near Kade, bole of small tree, Jones 1231; on liane, Jones 1241. Aburi Botanical Gardens, frequent, Jones s.n. Amedzofe, on old Combretum in savanna woodland just below summit of the hill, Jones 1259; on wet rocks in stream just above the waterfall, Jones 1278b; 256 E. W. JONES & A. J. HARRINGTON epiphyllous, abundant, mixed with Taxilejeunea conformis, on tall herbs beneath dripping cliffs in ravine above the waterfall, Jones 1285b. Kpedze Rest House, on cocoa, Jones s.n. Between Todome and Bame, on cocoa, Jones s.n. Widely distributed, variable, and often abundant, Lejeunea eckloniana is difficult to separate in the field from other medium-sized species of Lejeunea, but microscopic examination of fresh living plants will show the distinctive simple oil-bodies. The variation and difficulties of determination are discussed by Jones (op. cit.); it is possible that more than one taxon is included. Jones 1456 from Leicester Peak is a remarkable robust form, in which all the perianths are borne on short lateral branches which lack subgynoecial innovations; a similar plant (Jones 1470 p.p.) was gathered in the Loma Mountains. If these plants were examined alone they would be ascribed to a different species or even (if the absence of subgynoecial innovations was considered to be a generic character) to a different genus. Intermediate conditions exist, however, and L. eckloniana shows a strong tendency to produce some gynoecia without, or with reduced, innovations. 7. Lejeunea flava (Sw.) Nees, Naturgesch. europ. Leberm. 3: 277 (1838); E. Jones in Trans. Br. bryol. Soc. 5: 548 (1968), subsp. flava SIERRA LEONE. Freetown Peninsula: Havelock Plateau, bole of small tree in scrub, Jones s.n.; Leicester Peak, 490-595 m (1600-1950 ft), abundant on boles of small trees and on twigs of bushes both in deep shade and in open scrub in rocky ground, Brenan (Jones 462), Jones 1429, 1430, 1457, 1462d; forest in valley above Bathurst, c. 430 m (1400 ft), on tree bole, Jones 1437. Loma Mountains, valley-head west of Bintimani, 1680 m (5500 ft), under bushes in deep shade, with Taxilejeunea conformis, Jones 1498b. GHANA. Kade A.R.S., on smooth well-lit tree bole in natural forest, Jones 1220. Aiyaola F.R. near Kade, side of main bough in crown of Khaya, Jones 1238. See the discussion on the taxonomic treatment of Lejeunea flava following L. grossecristata. 8. Lejeunea grossecristata (Stephani) E. Jones in Trans. Br. bryol. Soc. 5: 556 (1968). Taxilejeunea nicholsonii Pearson in Annls Cryptog. exot. 4: 64 (1931). SIERRA LEONE. Lake Sonfon, on edge of forest surrounding the ‘lake’, on mango, Harrington 24 p.p., 25 p.p., on small tree, Harrington 41. Bo, on dead tree stump, Gardner 130 p.p. GHANA. Krokosua Hills F.R., 550 m (1800 ft), on exposed stems of pioneer shrubs on margin of forest, Jones 1401b. Atewa Hills F.R., bough in crown of Cassipourea, Jones 1322 p.p. Bunso, on cocoa, Jones 1317a. Tafo, on bole of isolated tree in garden, Jones 1333. Aburi Botanical Gardens, on roots of epiphyte on palm, and on tree bole, Jones s.n. Amedzofe, in savanna woodland just below summit of the hill, on boles of old Lophira and Terminalia, Jones 1257a, b, c, e; on cocoa in farms just above the waterfall, Jones 1272b. The complex of autoecious plants included under, or closely allied with, Lejeunea flava is a taxonomically difficult group. Jones (op. cit.) tentatively recognized in Africa L. flava subsp. flava, L. flava subsp. tabularis (Sprengel) S. Arnell, L. longirostris (Stephani) E. Jones, and L. grossecristata, though he expressed doubts as to the distinctness of the last three, and in 1979 he accepted L. longirostris as synonymous with L. grossecristata. While collecting in 1971 he paid particular attention to this group. The plants from Sierra Leone and Ghana seem to fall into two groups: (1) Lax, often elongate plants with leaves distant or approximate, the underleaves small (0-26-0-36 mm wide), a quarter to a third the width of the shoot, distant to approximate, ovate or cordate-ovate, the perianths small, up to c. 0-56 mm long, keeled in the upper third or half, usually with one subgynoecial innovation. (2) Shorter plants, often closely applied to bark, with leaves approximate to imbricate, underleaves large (0-3-0-45 mm wide), a third to a half the width of the shoot, rounder than those of (1), thus circular rather than ovate, and more strongly cordate at the base, imbricate at least below female inflorescences, perianths larger, 0-6-0-7 mm long, the keels longer and deeper than in (1), extending below the middle of the perianth, often with two subgynoecial innovations. HEPATICS OF SIERRA LEONE & GHANA 257. Weak shoots of group (2), and the basal portions of branches, often have small distant underleaves. It is arguable that the two groups represent hygromorphic and xeromorphic, or perhaps shade- and light-forms, of a single taxon, but on the whole the field evidence suggests that they are different. The fact that the two groups seem to have different geographical distributions supports this belief. Group (1) corresponds with Lejeunea flava subsp. flava; the other group can be assigned either to subsp. tabularis or to L. grossecristata, if indeed these two are different taxa. In their tendency to be closely applied to bark, and to have imbricate leaves accrescent to the female inflorescence, they differ from subsp. tabularis as it occurs in South Africa, and are clearly the same as Jones 212 from Calabar, Nigeria, which was considered to be L. grossecristata by Jones (1968a). A satisfactory taxonomic treatment of this complex can probably only be attained after a much fuller study of the variation within and between populations than has yet been undertaken, combined with experimental work. It has not been possible to trace the holotype specimen of Taxilejeunea nicholsonii Pearson at MANCH,;; it was collected at Tarkwa, Ghana, by Saxby. Pearson (op. cit.) described it as autoecious, with perianths ‘narrowly obovate, 5-angled, slightly winged, rostellate’, and androecia ‘sessile . . .; 2-4 pairs of bracts’; perianths were said to be abundant. A packet at CGE contains copious material but is nearly sterile, with only a few poor androecia, suggesting that the plant might be dioecious. Assuming that the CGE packet represents the same plant as the type, and that Pearson’s observations are correct (as, indeed, they are likely to be) T. nicholsonii should be regarded as a synonym of Lejeunea grossecristata. A ‘Taxilejeuneoid’ female inflorescence would conform with this identification. 9. Lejeunea ibadana A. Harrington & E. Jones in J. Bryol. 12: 40 (1982). ? Lejeunea corbieri Stephani in Bull. Mus. natn. Hist. nat. Paris 18: 119 (1912). SIERRA LEONE. Kambui Hills, Bambawo, 370 m (1200 ft), on rotting log in forest, Jones 1548. GHANA. Pra-Suhien F.R., on rotting log in deep shade, Jones 1344c. Kade A.R.S., on base of large liane in deep shade in forest, Jones 1219 p.p. Amedzofe, on bole of cocoa in farms above the waterfall, Jones 1272a. Lejeunea ibadana is widely distributed and locally abundant in West Africa, especially on decaying vegetable matter such as rotting logs and bases of oil palms. It is present in Stephani’s herbarium (G 1173 from Bouroukrou, Ivory Coast) as ? L. corbieri. The type of L. corbieri, from Bangui, Central African Republic, consists mostly of L. setacea Stephani, but includes a small admixture of a Lejeunea which may be L. ibadana, but which is too scanty and imperfect for a safe determination. Unfortunately, Stephani’s original description is not adequate to indicate whether his L. corbieri is the plant that we have described as L. ibadana. 10. Lejeunea lomana E. Jones, sp. nov. Fig. 7. Surculi cum foliis c. 1 mm lati, caule 90 wm diam., appressi, irregulariter ramosi. Folia complanata, approximata vel leviter imbricata, c. 0-5 x 0-4mm, ovalia vel ovata, apice rotundata; lobulus magnus, lobo triplo brevior, oblongus, basin et carinam versus inflatus, apicem versus compressus; carina leviter arcuata, cum margine lobi postico sinus latum et non profundum delimitans; margo lobuli liber ex 6 cellulis constructus; dens apicalis substrictus, c. 40 wm longus X 13 wm latus, apice solum liber, praeterea ad marginem lobuli apicalem conjunctus; papilla hyalina ex apice dentis lateraliter oriens et marginem librum contingens. Amphigastria orbicularia, ad medium bifida, quam caule 1%2~2-plo latiora. Cellulae lobi medianae 18-24 x 24-35 um, leptodermatae, trigonis parvis aut mediocris sed manifestis, incrassationibus intermediis nullis. Dioica. Androecia vulgo 6-7 jugae, in ramis longis intercalata. Perianthia 0-5—0-8 x 0-33-0-5 mm, teretia, pyriformia, circum apicem 5-carinata, carinis non altis, consimilibus. Bracteae femineae perian- thio paulo breviores, obovatae, lobulis lobo 1!2-plo brevioribus, carinis aut bractearum superiorum aut bractearum ambarum leviter alatis. Typus: E. W. Jones sub no. 1503 in Monte Loma, Sierra Leone, lect., in Herb. Mus. Brit. conservatus. Green, making intricate patches closely applied to leaves or to the bark of shrubs. Strong shoots 0-8-1-0 mm broad, the stem 90 wm diam., with underleaves c. twice as wide as the stem. Leaves approximate or somewhat imbricate, complanate, spreading at an angle of 60—90° to the stem, 258 E. W. JONES & A. J. HARRINGTON Fig. 7 Lejeunea lomana E. Jones. (a) male inflorescence; (b) part of shoot; (c) shoot with perianth. All x40. (d) leaf, x48; (e) female bracts and bracteole, x48; (f) apex and (g) apex with free margin of lobule, compressed, 240; (h) underleaf, X240; (i) cells from ‘x’ on (d), X240; (j) transverse section of stem, 400. All drawn from Jones 1503 (holotype). gently convex, ovate or oval, 0:46-0:53 x 0-34-0-45 mm, the base truncate, antical margin arched, apex broadly rounded, postical margin gently arched and making a wide shallow sinus with the gently arched keel. Well developed lobule about half the length of the lobe (measured from stem to apex), oblong-oval, inflated along the keel and proximally but the apical part flat and lying against the lobe, with most of the free margin readily visible in situ; the free margin of six cells (in addition to the cell in the notch proximal to the apical tooth); the apical tooth long, c. 13 x 40 um, slightly curved, in line with the margin of the apex and free only at its rounded tip; hyaline papilla inserted on the side of the tip of the tooth and lying along the free margin. Subapical cells of leaf 17-20 x 19-23 um, averaging 18 X 21-6 wm, median cells 18-24 x 24-35 pm, averaging 22 X 26 um, cells of antical margin 15-18 x 16-20 wm, and 3-4 rows in from the antical margin 15-22 x 20-24 um, walls thin, trigones small or medium but distinct, intermedi- ate thickenings absent, cuticle finely punctate; oil bodies 2-6 per cell, compound, subspherical, oval or fusiform. Underleaves 1!/—2 times as wide as the stem, orbicular, 150-180 wm wide and long, bilobed to the middle, the lobes broadly triangular, 5—6 cells broad at the base, with V-shaped sinus. Stem in transverse section with seven cortical cells 15—20 um wide, their outer walls 2-3 wm thick, their radial walls thin, over six medullary cells, 10-15 x 10-20 um, thin-walled with slight thickening at wall junctions. Dioecious. Male plants profusely branched, androecia large, of (2)-6-7 pairs of closely imbricate bracts with two basal underleaves only, intercalary on long shoots. Gynoecia on long shoots, with one innovation, the first leaf of which is a lateral leaf. Female bracts obovate, 0-5—0-6 x 0-24-0-33 wm, the apex broadly rounded, lobules about 2/3 as long, with keels 1/2—2/3 the length of the lobule, the keels straight or weakly inarched, that of the upper bract, and sometimes both keels, slightly winged with one row of projecting cells; the apices of the lobule of HEPATICS OF SIERRA LEONE & GHANA 259 the upper bract, or of both lobules, acute. Bracteole oblong, about 2/3 the length of the bracts, bilobed to 1/3 its length. Perianth 0-5—0-8 x 0-33-0-5 mm, slightly exserted from the bracts, terete, pyriform, with five equal, rather low keels around the apex and upper half; the rostrum very variable, even in consecutive perianths, 20-80 wm long. The preceding description relates to well-developed shoots; there are frequent abrupt transitions to lengths of shoot bearing smaller leaves with greatly reduced lobules, and such reduced leaves are normally present at the bases of branches and succeeding a male infloresc- ence. SIERRA LEONE. Loma Mountains, 1520-1650 m (5000-5400 ft): gallery forest c. one km west of ‘Camp 2’, on leaves of Memecylon, Morton (Harrington 350 p.p., 351 p.p.), epiphyllous, Jones 1483 p.p., 1488 p.p., on small branches of shrubs, Jones 1479c, d & e, on dead branch, Jones 1495 p.p., 1497 p.p.; on margin of scrub in valley-head west of Bintimani, on bole of Vernonia, Jones 1503 (BM, holotype). In all these collections, except Jones 1503, Lejeunea lomana was present in only small amounts, and mixed with other Lejeuneaceae. All collections were made in an area of about two km radius, but this circumscription is presumably due to the absence of any collecting beyond this area on the Loma plateau. Although the collections are small they are sufficient to show that within this area L. lomana is a well-defined taxon, and distinct from several other closely allied, and apparently as yet undescribed, species that occur elsewhere in the African mountains. One of these, which I collected in very small amount on Mont Cameroun (Cameroon Mountain) at 1220 m (4000 ft) in 1948 (Jones 310 p.p.), differs chiefly in being monoecious and having large underleaves two to three times as wide as the stem. Two other apparently different species were collected by Dr D. Steel in the Bale Mountains of Ethiopia, and I have other plants from East Africa awaiting study that may belong to the same complex. All have a lobule of closely similar structure, with a free margin of six to nine cells, and similar perianths and female involucres. They should probably be separated from the genus Lejeunea, but do not seem to fit well into any other existing genus. [E.W.J.] 11. Lejeunea papilionacea (Stephani) E. Jones in J. Bryol. 7: 42 (1972). Prionolejeunea aberrans Stephani, Sp. hepat. 6: 387 (1923). SIERRA LEONE. Masimo, epiphyllous, Marmo 160 p.p. Kambui Hills, Bambawo, epiphyllous on low vegetation near stream, Harrington 415 p.p., 417 p.p., 440 p.p. GHANA. Ankasa River F.R., epiphyllous, abundant, Jenik; on palm leaves in swamp forest one mile south of the Ankasa bridge, Jones 1382 p.p. A local species allied to Lejeunea caespitosa, usually though not exclusively epiphyllous, and known elsewhere only from Cameroon and (as Prionolejeunea aberrans) Madagascar. Schuster (1980) has recently transferred it to Cardiolejeunea R. M. Schuster & Kachroo. 12. Lejeunea ramosissima Stephani in Bot. Jb. 8: 88 (1886 [‘1887’]); E. Jones in Trans. Br. bryol. Soc. 5: 300 (1967). Lejeunea eplicata Stephani, Sp. hepat. 5: 711 (1915); E. Jones in Trans. Br. bryol. Soc. 5: 303 (1967). SIERRA LEONE. Loma Mountains, 1370-1830 m (4500-6000 ft): ‘sur rameaux d’Eugenia, forét relique d’altitude vers 1850 [sic]m’, Jaeger 1188 p.p. (PC); on tree boles on margin of forest near ‘Camp 2’, Harrington 159 p.p. Jones 1472b, 1478; on tree boughs in gallery forest west of Bintimani, Harrington 327; epiphyllous in forest, Harrington 299, Jones 1483a p.p., 1488; in rock crevices, Harrington 227, 323 p.p. A montane species which is abundant in the forest on Mont Cameroun (Cameroon Mountain) from c. 1320 m upwards, and also recorded from the Congo Republic (Pécs, 1980), Sao Tomé, Pagalu, and from the Nimba Mountains (Schnell 3525, PC) and the Macenta Massif (Lisowski, OP and Herb Jones) in Guinea. Grolle (1978) notes that it is frequent at altitudes of 320-790 m in the Seychelles. Its occurrence in the East African mountains is doubtful. A dioecious species in which perianths are often rare or absent, it cannot be separated vegetatively from Lejeunea brenanii. Male inflorescences of L. ramosissima have underleaves only at the base, whereas 260 E. W. JONES & A. J. HARRINGTON those of L. brenanii have underleaves throughout their length. Jones 1488 has perianths which are bluntly angled near the apex, and thus resembles L. eplicata Stephani, and helps to demonstrate the identity of the two taxa. 13. Lejeunea setacea Stephani in Bull. Mus. natn. Hist. nat. Paris 18: 120 (1912); E. Jones in Trans. Br. bryol. Soc. 5: 784 (1969). SIERRA LEONE. Freetown Peninsula, Mount Aureol, earth bank of path, Jones s.n. Musaia Agricultural Station, pumping station, on tree bole, Haswell B1 p.p. Njala, in crown of Nauclea diderrichii, Deighton 5313 p.p. Bo, Gardner 128. Jawo near Kenema, by the Moa river, abundant on tree branches and boles at top of and just above the flood zone, Jones 1536. GHANA. Krokosua Hills, on cocoa in farm between the Forest Reserve and Bia bridge, Jones 1388 p.p. Kwahu-Tafo (6°39'N, 0°40’W), on cocoa, Jones s.n. Brukru Rock near Kwahu-Tafo, on shaly sandstone, Jones 1417. Aburi Botanical Gardens, on root of Hura, Jones s.n. Kpedze Rest House, on cocoa, Jones s.n. A lowland species, widely distributed throughout the wetter parts of the Guinea region of West Africa and the Congo Basin. The leaves are flat and the shoots strongly complanate — a feature that helps with recognition in the field. Outside this area it has been recorded only with doubt from south-eastern Kenya by Pécs (Bizot & Pécs, 1974), though it is closely allied to, and possibly conspecific with, the South American Lejeunea pililoba Spruce. Schuster (1980) has recently transferred both taxa to Rectolejeunea. XXXI. LEPTOLEJEUNEA (Spruce) Schiffner 1. Leptolejeunea astroidea (Mitten) Stephani, Sp. hepat. 5: 363 (1913); Vanden Berghen in Bull. Jard. bot. Etat Brux. 23: 68 (1953). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., Harrington 85, 89a p.p., 90, 99 p.p. etc., Jones s.n.; Leicester Peak, c. 550 m (1800 ft), in forest near summit, Jones 1428; Sugar Loaf Gap, on low vegetation in forest, Harrington 652 p.p.; Bathurst, Kongo Dam, Jones s.n.; Guma Valley, in mature forest, Harrington 101 p.p., 102 p.p.; Black Johnson Beach near York, on low shrub in thicket behind the beach, Harrington 16; York Pass, forest understorey, Harrington 625 p.p., 630 p.p.; Picket Hill, forest near summit, c. 850 m (2800 ft), Gledhill s.n. Masimo, Marmo 160 p.p., 161 p.p. Bo, near the school, Revell (Herb. Jones). Kambui Hills, Bambawo, in forest on plateau, c. 370 m (1200 ft), Harrington 411 p.p., 412 p.p., 421 p.p., 483 p.p., 484 p.p., 485 p.p., Jones 1543b. Giema, in secondary forest, Harrington 512 p.p., 549 p.p. Gola Hills, near Lalehun, Marshall s.n.; by river, Jones s.n. GHANA. Krokosua Hills F.R., 490-550 m (1600-1800 ft), abundant on low shrubs in small gaps, Jones 1396. Ankasa River F.R., leaves in crown of Glutea, Jones 1378c. Subri F.R., on Raphia fronds in swamp forest, Jones 1355 p.p. A lowland species, rarely ascending above 800 m, though it has been recorded as high as 1000 m in the Macenta region of Guinea (Duda & Vanden Berghen, 1967); widely distributed throughout the ‘rain forest’ districts of West Africa and the Congo. It appears to need shelter and high atmospheric humidity, and is almost invariably epiphyllous on herbs and low understorey shrubs. It is exceptional for it to occur as an epiphyll in the crown of a large tree, as in Jones 1378c, which, it should be noticed, was close to a river in a district of very high rainfall. It is often associated with the other two species of Leptolejeunea, and is perhaps the most abundant of the three. Like many other epiphyllous hepatics it requires more light than reaches the floor of the unbroken dense forest, and tends to occur beneath slight gaps in the canopy or in thin-canopied forest. 2. Leptolejeunea maculata (Mitten) Schiffner, Consp. hepat. archip. ind.: 275 (1898); Vanden Berghen in Bull. Jard. bot. natn. Belg. 47: 213 (1977). Leptolejeunea thomeensis (Stephani) Stephani, Sp. hepat. 5: 366 (1913); Vanden Boies in Revue bryol. lichén. 32: 50 (1963). Leptolejeunea truncatiloba Stephani, Sp. hepat. 5: 368 (1913); Vanden Berghen in Bull. Jard. bot. Etat Brux. 23: 65 (1953). SIERRA LEONE. Gola Hills, near Lalehun, on low vegetation in forest, Marshall s.n. HEPATICS OF SIERRA LEONE & GHANA 261 GHANA. Krokosua Hills F.R., 490-550 m (1600-1800 ft), abundant with Leptolejeunea astroidea, Jones 1396 p.p. Ankasa River F.R., on leaves in crown of Glutea, Jones 1378c p.p. Aiyinasi A.R.S., on Citrus leaves, Jones 1385 p.p. Pra-Suhien F.R., widely distributed on herbs in regenerated forest, Jones 1345b & c. Leptolejeunea maculata closely resembles L. astroidea in its general ecological requirements and, in W. Africa, the two species are usually associated. Unlike L. astroidea it occurs in the lowland rain forest of East Africa (e.g. at Amani in Tanzania), and extends to the Mascarenes. 3. Leptolejeunea quintasii Stephani, Sp. hepat. 5: 365 (1913); Vanden Berghen in Bull. Jard. bot. Etat Brux. 23: 70 (1953); Revue bryol. lichén. 32: 51 (1963). SIERRA LEONE. Giema, in secondary forest, Harrington 528 p.p., 548 p.p.,549a p.p. Gola Hills: near Lalehun, Marshall s.n., abundant by stream in forest, Jones 1553b; Gola North F.R., abundant in unexploited ‘moist forest’, Jones 1568 p.p. GHANA. Subri F.R., on Raphia fronds in swamp forest, Jones 1355 p.p. Kade A.R.S., very abundant in swamp forest on herbs, and on the forest margin, Jones 1222. Opposite Kade, in swamp on the Kadeva rivulet, on Raphia hookeri, Jenik (Herb. Jones & OP). Like Leptolejeunea maculata, L. quintasii extends eastwards through the Congo Basin to the lowland rain forest of East Africa, but has not been recorded from western Sierra Leone. Two or more of the three species of Leptolejeunea often grow together. When the leaves on which they have been growing are dried without undue pressure, the three species can be seen to differ in the positions assumed by their own leaves when dry. The leaves of L. quintasii remain flat and appressed to the substratum; those of L. astroidea become channelled by the up-curving of antical and postical margins, and remain spreading or obliquely ascending, while those of L. maculata roll longitudinally and stand up vertically, perpendicular to the substratum. Harring- ton found that plants are intensely fragrant when fresh, and their presence on a leaf can readily be detected by the nose. Presumably the fragrance emanates from the oil bodies in the ocelli, which differ somewhat in appearance; those of L. astroidea are finely granular and very highly refractive, appearing almost homogeneous, whereas those of L. maculata are coarsely granular and opaque. XXXII. LOPHOCOLEA (Dumort.) Dumort. 1. Lophocolea concreta Montagne in Annis Sci. nat. (Bot.) II, 4: 350 (1845); Grolle in Trans. Br. bryol. Soc. 3: 595 (1959). Lophocolea subrotunda Mitten in Phil. Trans. R. Soc. 168: 396 (1879); E. Jones in Trans. Br. bryol. Soc. 2: 175 (1953). SIERRA LEONE. Loma Mountains: in valley forest between Seradu and Kurubonla, 610 m (2000 ft), on rotten log, Jones 1519; in forest between ‘Camp 1’ and Sokurela, on rotten bough, Harrington 383; in forest above Sokurela, 1070 m (3500 ft), on rotten log, Jones 1511, and at 940 m (3100 ft), Jones 1512 p.p. GHANA. Subri F.R., on rotten log, Jones ! Amedzofe, shaded rock near summit of hill, Jones 1255a (GC). Widely distributed in tropical Africa; perhaps somewhat montane, and mostly in climates with a strong dry season. Thus it is likely that it is much more abundant in the drier parts of Sierra Leone and Ghana than the above records suggest. 2. Lophocolea difformis Nees, Syn. hepat.: 166 (1845); Grolle in Trans. Br. bryol. Soc. 3: 596 (1959). Lophocolea molleri Stephani in Bot. Jb. 8: 83 (1886 [‘1887’]); E. Jones in Trans. Br. bryol. Soc. 2: 181 (1953). SIERRA LEONE. Loma Mountains, forest in centre of plateau, on fallen bough, Morton (Harrington 366, 370). GHANA. Rocks in steep roadside bank between Vane and Amedzofe, c. 610 m (2000 ft), Jones 1264. 262 E. W. JONES & A. J. HARRINGTON Widespread in tropical and warm temperate Africa, but, like Lophocolea concreta, chiefly in country with a moderate rainfall and strong dry season, and thus likely to be more abundant than the above records suggest. 3. Lophocolea martiana Nees, Syn. hepat.: 152 (1845), subsp. newtonii (Stephani) R. M. Schuster, Hepat. Anthocerot. N. Amer. 4: 237 (1980). Lophocolea congoana Stephani in Bull. Soc. r. bot. Belg. 41: 119 (1904); Grolle in Trans. Br. bryol. Soc. 3: 597 (1959). Lophocolea newtonii Stephani, Sp. hepat. 3: 170 (1907); E. Jones in Trans. Br. bryol. Soc. 2: 188 (1953). SIERRA LEONE. Kambui Hills, Bambawo, steep earth bank by path, Harrington 443, 445 p.p.; rotten logs in forest, Jones 1528, 1544, 1545. Gola North F.R., on rotten log, Jones 1566a. GHANA. Bia North F.R., on rotting logs in forest, Jones ! Krokosua Hills F.R., 550 m (1800 ft), on grass tussock in open ‘glade’, Jones 1400. Tarkwa, Saxby (CGE). Ochi Headwaters F.R., West-Skinn 159. Esukawkaw F.R., Jenik (Herb. Jones). Begoro, Jones ! Southern Scarp F.R., Jones ! Atewa Hills F.R., on bole of large tree in swamp forest, Jones 1331 (GC). Amedzofe, on steep cliff near the waterfall, /rvine 411. A common species throughout the lowland rain forest of West Africa and the Congo Basin, extending also into East Africa, where it is much more local. The lack of records from western Sierra Leone is noteworthy; presumably it is a species for which the dry season of the Freetown Peninsula is too severe, nor does it occur freely in young secondary forest. It is commonest on rotting logs, but is also frequent on the bases of tree boles, and is very often associated with Chiloscyphus dubius. On rotting wood Arachniopsis diacantha and Cephalozia fissa are common associates. It is less frequent on earth and rock. XXXII. LOPHOLEJEUNEA (Spruce) Schiffner 1. Lopholejeunea abortiva (Mitten) Stephani, Sp. hepat. 5: 70 (1912); Vanden Berghen in Revue bryol. lichén. 39: 376 (1973). SIERRA LEONE. Freetown Peninsula: Dighton Dam No. 1, valley below Sugar Loaf, on boulder above the dam, Harrington 661 p.p.; York Pass, by stream, Harrington 628 p.p. Giema, on partly submerged boulder in stream, Harrington 538, 539. GHANA. Southern Scarp F.R., near Bunso, on bole of small tree, Jones s.n. Amedzofe, on wet rocks in stream just above the waterfall, Jones 1278a. Specific distinctions in the genus Lopholejeunea are frequently indistinct; L. abortiva is closely allied to the much more abundant L. fragilis, of which it may prove to be ahygromorphic state. It frequently grows, like Jones 1278a, in wet places close to streams, though it may also grow on tree boles and on fallen logs. Apart from being a somewhat larger plant than L. fragilis it is said to differ in the absence of a dorsal keel to the perianth, in the female bracteole being oboval (not suborbicular), and in the lobules of the female bracts being small (often more than half the length of the bract in L. fragilis), but there is much variation in all these features. A low dorsal keel may be present or absent, even in the same colony, but the keel is not winged, as it usually is in L. fragilis. 2. Lopholejeunea fragilis Stephani, Sp. hepat. 5: 65 (1912); Vanden Berghen in Bull. Jard. bot. Etat Brux. 20: 176 (1950). SIERRA LEONE. Freetown Peninsula, Leicester Peak, on bole of old mango, 440 m (1450 ft), Jones 1464; on boulder in open forest near the summit, Brenan (Jones 466); on tree bole in low forest near the summit, Brenan (Jones 476 p.p.); on rocks, 520 m, Jones 1452b. Kabala, on mango near the District Officer’s bungalow, Harrington 140 p.p. Lake Sonfon, epiphyte in forest surrounding the ‘lake’, Harring- ton 28 p.p. Loma Mountains: forest between Seradu and Kurubonla, 610 m (2000 ft), on tree boles, Jones 1515 p.p., 1520b, 1521 p.p., on Uapaca, Jones 1522b; Seradu, in forest by the village, 610 m (2000 ft), Jones 1517a; above Sokurela, 910-1220 m (3000-4000 ft), on tree boles in forest, Jones 1468a. Masimo, Marmo 160 p.p. Kambui Hills, Bambawo, on mango on edge of fores:, Harrington 434, 437 p.p.; on trees near the Rest House, Harrington 569 p.p., Jones 1527 p.p. HEPATICS OF SIERRA LEONE & GHANA 263 GHANA. Ankasa River F.R., on twigs of Samanea by the river, Jones 1377a. Pra-Suhien F.R., side of tree root, Jones 1343c. Widely distributed throughout West Africa and the Congo Basin, and extending more locally to Mozambique, Natal, and Transvaal; usually on bark, rarely on rock. 3. Lopholejeunea jonesii Vanden Berghen in Bull. Jard. bot. Etat Brux. 20: 178 (1950). SIERRA LEONE. Loma Mountains, frequent on tree boles in the forest from c. 1520 m (5000 ft) upwards: near ‘Camp 2’, on tree boles, Jones 1483b, on lianes, Jones 1476b, 1495b; between Dawule and Bintimani, Harrington 239 p.p.; forest west of Bintimani, Harrington 325 p.p., 326 p.p., Jones 1497, 1500. Previously known only from c. 2100 m on Mont Cameroun (Cameroon Mountain). Lophole- jeunea jonesii differs from L. subfusca in its acute leaves and the large lobules of its female bracts, which resemble those of L. fragilis. 4. Lopholejeunea subfusca (Nees) Schiffner in Bot. Jb. 23: 593 (1897); Vanden Berghen in Bull. Jard. bot. Etat Brux. 20: 173 (1950); Revue bryol. lichén. 39: 381 (1973). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., on lianes in forest, Harrington 92, 596 p.p., bole of tree near Heddle’s Farm, Jones s.n.; Havelock Plateau, on small tree in scrub, with Schiffneriolejeunea polycarpa, Jones s.n.; Regent, trees in farm bush, Harrington 408 p.p., 409 p.p.; three km north-east of Toke, on small tree in thicket, Jones 1448b. Kasewe F.R., on lateritic boulder in bed of forest stream, Richards R7171 (no perianths seen). Kambui Hills, Bambawo, on mango on edge of forest, Harrington 427 p.p., 437 p.p.; on small Drypetes in young-growth forest, Jones 1532a. Giema, on sapling, Harrington 554 p.p. GHANA. Krokosua Hills F.R., on exposed stems of pioneer shrubs on forest margin, Jones 1401a. Ankasa River F.R., on trunk of Parinari glabra, Richards (GC 36676). Aiyinasi A.R.S., on small tree by Rest House, Jones 1363a; on 17-18 year-old Citrus, Jones 1383. In low secondary bush near Esiama, Jones 1365. Asanta, on coconut palms on the beach, Jones 1359 p.p. Bunso, on cocoa, Jones 1317b. Tafo Cocoa Research Inst., in garden, Jones 1333. Pra-Suhien F.R., bole of small tree, Jones 1337b p.p. Kade A.R.S., upper branches of Parinari excelsa, Hossain (GC 36700); upper branches of Aningeria, Hossain (GC 39324). Amedzofe, on Combretum in savanna woodland just below summit of the hill, Jones 1256 p.p. (GC). A lowland species, widely distributed and often abundant in the moister districts throughout West Africa and the Congo Basin, and extending to the coastal districts of Kenya and Tanzania, and to Madagascar. It is also known from tropical America, Malaysia, Indonesia, etc., and is doubtless pantropical. It is probably photophilous, and commonly occurs on isolated trees, on the boles of trees in plantations, young farm bush, etc., and in the crowns of forest trees, rather than in the forest understorey. It is very variable, especially in the shape of the female bracteole, which may vary from obovate to almost semicircular, and the size and form of the perianth wings. The female bracts are obovate with small lobules, whereas in Lopholejeunea fragilis they are oval and more or less acute, with large lobules. The mature perianth is almost immersed in the bracts, whereas in L. fragilis the perianth emerges and is readily visible. The two species also differ in colour, L. subfusca typically being chestnut brown, while L. fragilis is blackish brown. XXXIV. MARCHANTIA L. 1. Marchantia planiloba Stephani in Bolm Soc. broteriana 4: 181 (1886); Vanden Berghen in Revue bryol. lichén 29: 51 (1960). GHANA. Marchaniia planiloba is in cultivation at the Department of Botany, Reading University; the records indicate that the original stock was collected in Ghana by Professor T. M. Harris; the locality is unknown. Marchantia planiloba and the closely allied M. parviloba Stephani (which Vanden Berghen (op. cit., 1965) considers may be phenotypes of a single species) appear to be somewhat montane in their distribution; they might be expected to occur, for example, in the Atewa Hills. M. parviloba has been recorded chiefly from East Africa, but also from S40 Tomé and Mont 264 E. W. JONES & A. J. HARRINGTON Cameroun (Cameroon Mountain), whereas M. planiloba has been recorded from S40 Tomé and Burundi. 2. Marchantia wilmsii Stephani in Hedwigia 31: 126 (1892); Vanden Berghen in Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8 (1): 178 (1972). Marchantia chevalieri Stephani ex Bonner in Candollea 14: 103 (1953). SIERRA LEONE. Nijala, abundant on the bank of the river on mudstones between highest and lowest river level, Deighton 2870; on clay at the river bank, inundated in the wet season, Deighton 6032. Loma Mountains, Sokurela, stream bank near the village, Harrington 142. GHANA. Aiyinasi, banks of river Fiakpole, Jones 1360. Kibi [‘Kibbi’], on damp shady bank in evergreen forest, /rvine 26. Road cutting between Kibi and Bunsu [‘Bunso’], ten miles from Kibi, (perhaps the same locality as Irvine 26), Boughey (Herb. Jones), det. Vanden Berghen. Buem-Krache District, waterfall at Shiare (c. 8°20'N, 0°40’E), Hall 1534 (Herb. Jones). The commonest species of Marchantia in tropical Africa, extending southwards to Transvaal and Natal, and from sea-level to 1250 m or more. In the field it is readily distinguished from M. parviloba and M. planiloba by the presence of a dark median line on the dorsal surface of the thallus. XXXV. MARCHESINIA Gray 1. Marchesinia excavata (Mitten) Stephani, Sp. hepat. 5: 145 (1912); E. Jones in Trans. Br. bryol. Soc. 6: 78 (1970). Archilejeunea apiculata Pearson in Annls Cryptog. exot. 4: 61 (1931). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., on bole of tree near Heddle’s Farm, Jones 1420; Leicester Peak, 520 m (1700 ft), on boughs of Combretum in scrub, Jones 1453a; forest above Bathurst, 430 m (1400 ft), on bole of large Dialium, Jones 1436; Sugar Loaf Mountain, 700 m (2300 ft), on bark, Tindall 57 p.p.; Sugar Loaf Mountain, north side, on tree, T. S. Jones 333 p.p., 340 p.p.; Toke, on large Chrysobalanops at back of beach, Jones 1466a, and also in valley three km to north-east, Jones s.n. Lake Sonfon, on trunk of Erythrophleum in forest by ‘lake’, Harrington 54, 55. Bumban (9°07'N, 11°54’W), 200 m (650 ft), Thomas 2006. Mesima near Kalu [‘Messima near Kailu’] (7°12'N, 11°46’W), on tree in coastal savanna, Richards (Herb. Jones). Gola North F.R., on main bole of large Mimusops, Jones 1559a; on upper bole and covering main boughs in crown of large tree, Jones 1562. GHANA. Krokosua Hills F.R., 550 m (1800 ft), on bole of tree on forest margin, Jones 1404. Afram Headwaters F.R., in relic of natural forest, Jones s.n. Ankasa River F.R., on bole of Samanea by the river, Jones 1376b. Tarkwa, Saxby (CGE, sub Archilejeunea apiculata). Pra-Suhien F.R., on upper part of bole of tree, Jones 1338. Atewa Hills F.R., near top of bole of large Lophira, Jones 1318; on large branch of Cassipourea, Hossain (GC 36666). Aiyaola F.R., on bole of middle-storey tree, Jones 1239a. A West African species, extending from Guinea to Zaire, and locally very abundant, especially in Sierra Leone. It usually grows on large trees, and has a preference for the upper parts of boles and boughs in the crowns, or the boles of isolated but sheltered trees; in such situations it often forms extensive sheets. It is essentially a lowland species. 2. Marchesinia moelleriana Pearson in Ark. Bot. 19 (5): 10 (1925); E. Jones in Trans. Br. bryol. Soc. 6: 80 (1970); J. Bryol. 10: 394 (1979). SIERRA LEONE. Loma Mountains: gallery forest in centre of plateau, on large branch, Morton (Harrington 374 p.p.); near ‘Camp 2’, 1520-1680 m (5000-5500 ft), on lianes in forest, Jones 1479, 1495 p.p.; valley-head west of Bintimani, 1680 m, on Vernonia on margin of forest, Jones 1497a. Marchesinia moelleriana is a montane species which appears to replace M. excavata at altitudes above 1000 m. It is known elsewhere from East Africa, where it extends from Mount Elgon to Rhodesia (now Zimbabwe); it has not been recorded from Cameroon, though it is to be expected there. HEPATICS OF SIERRA LEONE & GHANA 265 XXXVI. MASTIGOLEJEUNEA (Spruce) Schiffner 1. Mastigolejeunea auriculata (Wilson) Schiffner in Engl. & Prantl, Nat. Pflanzenfam. 1 (3): 129 (1893); Gradst. & Inoue in Bull. natn. Sci. Mus. Tokyo B, 6: 25 (1980). Mastigolejeunea carinata (Mitten) Stephani, Sp. hepat. 4: 759 (1912); Vanden Berghen in Bull. Jard. bot. Etat Brux. 19: 378 (1949). SIERRA LEONE. Musaia Agricultural Station, on Uapaca togoensis, Sellar B4. Kabala, on tree near the District Officer’s bungalow, Harrington 131; on boulder, 135. Secondary forest five km south of Kabala, Harrington 119a, 125 p.p. Between Seradu and Kurubonla, in valley forest, Harrington 400, Jones 1520. Bo, on tree stump, Gardner 130 p.p. GHANA. Bia North F.R. north of Pampramasi, in crown of Mansonia, Jones 1411 p.p. Asenanyo F.R., in crowns of Triplochiton et al., Jones 1412 (GC), 1413a. Oda, on cocoa, Irvine 8a. Esukawkaw F.R.., Jenik. Kade, abundant, Jones 1227a (GC). Tafo, trees in garden, Jones 1333. Aiyaola F.R., in crown of Triplochiton, Jones 1227a. Aburi Botanical Gardens, /rvine s.n., Jones s.n. (GC). South of Kpandu (7°N, 0°18’E), in savanna woodland, on Bauhinia, L. E. Newton 596. Amedzofe, in savanna woodland just below summit of the hill, Jones 1259e (GC). Mastigolejeunea auriculata is widely distributed and polymorphic in tropical America and Africa. In West tropical Africa it extends from rain forest into savanna country with less than 1000 mm average annual rainfall. It is widely distributed and abundant in southern Ghana, especially on the boles of isolated trees and on branches in the crowns. In Sierra Leone it seems to be much more local, occurring chiefly in the drier districts in the north-east; the lack of records from the Freetown Peninsula and the Kenema district is remarkable. 2. Mastigolejeunea florea (Mitten) Paris in Revue bryol. 33: 42 (1906); Vanden Berghen in Bull. Jard. bot. Etat Brux. 19: 373 (1949). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., on tree bole, Harrington 581 p.p.; Havelock Plateau, on tree bole in scrub, Jones s.n.; Leicester Peak, on boulder in rather deep shade, Brenan (Jones 477); Regent, on small tree in farm bush, Harrington 408 p.p. Near road-bridge c. 32 km south-west of Kabala, on boulder, Harrington 404. Lake Sonfon, in forest surrounding the ‘lake’, Harrington 31, 47. Between Seradu and Kurubonla, in valley forest, 610 m (2000 ft), Jones 1520c. Loma Mountains, on large branch of tree in valley forest in centre of plateau, c. 1520 m (5000 ft), Morton (Harrington 374 p.p.). Kambui Hills, Bambawo, base of large tree, Jones 1551. GHANA. Pampramasi, on tree root in stream bank, Jones 1410. Krokosua Hills: on boles of cocoa on western slopes, c. 300 m (1000 ft), Jones 1388 p.p.; on liane in forest, 490 m (1600 ft), Jones 1392a (GC); on trees on the margin of a ‘glade’, 550 m (1800 ft), Jones 1403b. Begoro, Southern Scarp F.R., Jones ! Tafo, on cocoa, Thorold TG.122 (BM & Herb. Jones); in garden, Jones ! Kade, Hossain (GC 39327). Aburi Botanical Gardens, Foote s.n., Irvine 23, Jones 1214a. Also Cummins s.n., without locality. Widely distributed in West tropical Africa, but more local than Mastigolejeunea auriculata, and much less tolerant of dry conditions and exposed situations; it prefers some shade and often grows near water. 3. Mastigolejeunea nigra Stephani in Bot. Jb. 20: 319 (1895). Brachiolejeunea nigra (Stephani) Stephani, Sp. hepat. 5: 115 (1912); Vanden Berghen in Bull. Jard. bot. Etat Brux. 21: 90 (1951). SIERRA LEONE. Musaia Agricultural Station, pumping station, epiphyte, Haswell Bl p.p. Loma Mountains, frequent between Sokurela and the upper margin of the forest from c. 910-1220 m (3000-4000 ft), on boles of large trees and on rocks, Harrington 382, on boulders, Jones 1470, on tree boles on the forest margin, Jones 1505. Kambui Hills, Bambawo, on mango on edge of forest, Harrington 437 p.p. Giema, on fallen tree in secondary forest, Harrington 536 p.p. Gola North F.R., boughs in crown of large tree, Jones 1572a. GHANA. Amokwan Suezo, two miles south of the Ankasa River bridge, on the roots of an epiphytic orchid which was collected for GC (Herb. Jones). As widely distributed in tropical Africa as Mastigolejeunea auriculata, but more local. Although it may occur nearly at sea-level it tends to be montane, being more abundant at higher altitudes 266 E. W. JONES & A. J. HARRINGTON than M. auriculata, as can be seen in the Loma Mountains. It is also more restricted to exposed, well-illuminated habitats. In lowland rain forest it often seems to grow chiefly in the crowns of trees; the absence of records from this habitat in the Bia North, Aiyaola, Asenanyo, and Atewa Hills forest reserves in Ghana in remarkable. The absence of records from the Freetown Peninsula is also worthy of note. 4. Mastigolejeunea turgida Stephani, Sp. hepat. 4: 762 (1912); Vanden Berghen in Bull. Jard. bot. Etat Brux. 19: 376 (1949). GHANA. Nsuta, by the Pra river, in the crown of a small Scottellia, Jones s.n. A species which seems to be almost restricted to the crowns of trees, and therefore rarely gathered, but which is quite local. It was known hitherto only from a few collections in southern Nigeria, Cameroon, Bioko, and Principe. XXXVII. METZGERIA Raddi 1. Metzgeria saxbyi Pearson in Annis Cryptog. exot. 4: 70 (1931); Vanden Berghen in Bull. Jard. bot. Etat Brux. 19: 200 (1948). GHANA. Tarkwa, Saxby. Atewa Hills F.R., branch in crown of Cassipourea, Richards & Hossain (GC 36663); on lianes brought down by a fallen Cassipourea, Jones 1326 (confirmed by Y. Kuwahara); on small stems of shrubs close to ground in deep shade, in swamp forest, Jones 1329. Evidently a locally abundant species in Ghana, Metzgeria saxbyi is one of the few monoecious members of the genus. It has also been recorded from Cameroon (Vanden Berghen, op. cit.). It is closely allied to the monoecious M. conjugata Lindb., from which it differs in having shorter marginal hairs which are mostly solitary, smaller cells in the wings of the thallus (40-45 wm wide x 55-65 um long in M. conjugata, 25—35(—40) wm wide x 40-50 wm long in M. saxbyi), and more numerous hairs on the ventral surface of the wings; ventral hairs are, however, not present everywhere. M. saxbyi sometimes forms gemmae on the margins of the fronds. It has not been possible to trace a type at MANCH, nor is Metzgeria saxbyi represented at CGE. 2. Metzgeria thomeensis Stephani in Hedwigia 30: 271 (1891); Vanden Berghen in Revue bryol. lichén. 29: 53 (1960); Kuwahara in J. Hattori bot. Lab. 40: 287 (1976). Metzgeria camerunensis Stephani, Sp. hepat. 1: 293 (1899); Vanden Berghen in Bull. Jard. bot. Etat Brux. 19: 192 (1948). SIERRA LEONE. Freetown Peninsula: Leicester Peak, base of pole in dense scrub, c. 550 m (1800 ft), Jones 1427; Sugar Loaf Gap, intermingled with pendent mosses, Harrington 662 p.p. Loma Mountains: near ‘Camp 1’ in forest above Sokurela, 1220 m (4000 ft), on small bough, Harrington 253 p.p., same locality, on boulder, Jones 1473a, on bole of large tree, Jones 1509; forest near ‘Camp 2’, 1520-1680 m (5000-5500 ft), on tree trunk, Harrington 156, abundant on undershrubs, Jones 1484a. GHANA. ? Subri F.R., on stems of shrublets in Raphia swamp, Jones s.n. ? Amedzofe, wet sheltered but well-lit rocks by stream just above the waterfall, Jones s.n. Metzgeria thomeensis is widely distributed in West and East Africa. According to Kuwahara (op. cit.) it also occurs in Central and South America, Java, Sabah, and Papua New Guinea, and is thus pantropical. 3. Metzgeria sp. SIERRA LEONE. Loma Mountains, 1370-1680 m (4500-5500 ft): Jones 1473b; in forest near ‘Camp 2’, Harrington 156, Jones 1479b; ibid., draping twigs of undershrubs, Jones 1484b; on Vernonia in scrub in valley-head north-west of Bintimani, Jones 1503c; on small tree, valley forest c. one km west of ‘Camp 2’, Harrington 331; valley forest in centre of plateau, Morton (Harrington 371). The above collections represent a dioecious propaguliferous Metzgeria which becomes blue on HEPATICS OF SIERRA LEONE & GHANA 267 drying, and is often mixed with M. thomeensis. Gemmiferous fronds are not markedly attenuate, and the gemmae, which are ecostate, are borne on the margins of the frond. Ventral hairs are lacking, except on the costa, and marginal hairs mostly solitary, 60-100 wm long. The costa varies considerably; most sections show two dorsal and four ventral cells, but in at least some of the specimens there are short lengths of costa with three or four dorsal cells, the number varying over a short distance, while the best developed of the collections (Morton (Harrington 371)) consistently has four dorsal cells. Medullary cells in transverse section of costa 14-16. Cells of wing 32-41 x 43-49 um, often with minute trigones; cuticle in Morton (Harrington 371) and Jones 1503c finely and distantly papillose, in Jones 1473b, 1479b and 1484b smooth. Male involucres naked, female involucres with hairs 150 wm long. The plants seem to differ in one detail or another from all the species that have been described from Africa, but the specimens are inadequate for a satisfactory determination, nor is it certain that they all belong to the same taxon, though they probably do so. Species of Metzgeria are far more widely distributed in the forest regions of both Sierra Leone and Ghana than the above records would imply, and additional species are probably present. The plants often grow as isolated fronds thinly scattered amongst other bryophytes, and they are most often sterile and possibly depauperate; well developed specimens with both sexes adequately represented can rarely be gathered. The fact that two or more species are sometimes mixed adds to the difficulty of determination. The African species have been studied mainly by Vanden Berghen (1948, 19605). More recently Kuwahara (1978) has studied the genus extensively, but he has examined few recent collections from Africa. His taxonomic treatment frequently conflicts with Vanden Berghen’s, and clearly a much closer study of the variation within populations, based on careful field work, is needed before a satisfactory delimitation of the African species can be achieved. XXXVI. MICROLEJEUNEA Stephani 1. Microlejeunea africana Stephani in Hedwigia 27: 61 (1888); E. Jones in Trans. Br. bryol. Soc. 5: 777 (1969). Lejeunea ulicina subsp. africana (Stephani) Vanden Berghen in Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8 (1): 130 (1972). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., on trunk of Dacryodes klaineana, Harrington 586 p.p.; on the road to Hill Station, Hgeg s.n.; Leicester Peak, Jones 1464 p.p.; Lumley Cove, Jones 468; York Pass, epiphyllous, Harrington 626 p.p. Musaia town, on mango, Donald s.n. Lake Sonfon, on edge of forest surrounding the ‘lake’, on mango, Harrington 24 p.p. Loma Mountains: Seradu, 610 m (2000 ft), in forest by the village, Jones 1515 p.p., 1517 p.p.; gallery forest c. one km west of ‘Camp 2’, 1520 m (5000 ft), on leaves of Memecylon, Morton (Harrington 348 p.p., 350 p.p., 351 p.p.). Njala, in crown of Nauclea diderrichii, Deighton 5313 p.p. Bo, near the school, epiphyllous, Revell (Herb. Jones). Kambui Hills, Bambawo, on low vegetation near pool in forest, Harrington 557 p.p. Giema, epiphyllous, Harrington 552b p.p. Gola Hills, Lalehun, low vegetation in forest, Marshall s.n. GHANA. Krokosua Hills F.R., 490-550 m (1600-1800 ft), on leaves of low shrubs in forest, Jones 1396 p.p. Asanta, on coconut palms on the beach, Jones 1359 p.p. Aburi Botanical Gardens, Jones s.n. (GC). Amedzofe, on tree trunks amongst Cheilolejeunea serpentina, Irvine 412 p.p. A widely distributed and common species, which is certainly far more abundant than the above records indicate; the inexperienced collector gathers it only by accident and the experienced collector avoids gathering it because it is too common. It usually grows on branches and twigs in the crowns of trees or shrubs, or on the boles of more or less isolated trees — not in the dense shade of the forest. It is less frequent though by no means rare as an epiphyll. It is closely allied to Microlejeunea ulicina (Taylor) A. W. Evans but as Jones (op. cit.) has shown there are sufficient differences to justify keeping the two taxa distinct. It is perhaps not a matter of great importance whether M. africana is regarded as a distinct species or merely as a subspecies, but the former procedure is more convenient in that it simplifies nomenclature. There has been an increasing tendency in recent years to reunite the genus Microlejeunea with Lejeunea; this arises, perhaps, from stressing superficial features, rather than structural 268 E. W. JONES & A. J. HARRINGTON features, in the definition of the genera. If defined in terms of the characters enumerated by Jones (op. cit.)*, Microlejeunea forms a taxon which is better defined than some of the Lejeuneacean taxa which are generally accepted as genera. 2. Microlejeunea ankasica E. Jones in J. Bryol. 10: 394 (1979). ? SIERRA LEONE. Freetown Peninsula, flaky bark of Vitex on bank of Mountain Torrent, Jones 1443d. GHANA. Ankasa River F.R., on twigs in the crown of a Vitex by the river, Jones 1368 (BM, holotype). ?Kade, forest near the Agricultural Research Station, on smooth well-lit bole of Maesobotrya, chiefly around inflorescence-cushions, Jones 1218. Microlejeunea ankasica closely resembles M. kamerunensis, but was described as differing in being autoecious, and in being completely devoid of innovations beneath the gynoecial bracts. Jones 1443d and Jones 1218 have innovations beneath the gynoecial bracts, but in other respects they agree completely with M. ankasica. This provides additional evidence for thinking that in the schizostipulean Lejeuneaceae the subgynoecial innovations do not have the taxonomic significance that they have in the holostipean Lejeuneaceae, where they are usually considered as providing generic characters (cf. Lejeunea eckloniana). 3. Microlejeunea kamerunensis Stephani, Sp. hepat. 5: 812 (1915); E. Jones in Trans. Br. bryol. Soc. 5: 782 (1969). Lejeunea kamerunensis (Stephani) Vanden Berghen in Bull. Jard. bot. natn. Belg. 42: 446 (December 1972), non L. camerunensis (Stephani) E. Jones (August 1972) (Art. 64.1). SIERRA LEONE. Loma Mountains: c. 1680 m (5500 ft), on twigs of shrubs near ‘Camp 2’, Jones 1495 p.p.; epiphyllous in forest near ‘Camp 2’, Jones 1483a p.p. Previously known from Cameroon and the East African mountains; most records are from moderately high altitudes, but some of those from Cameroon are from sufficiently low down to make it likely that the species will occur in Ghana. XXXIX. NOTOTHYLAS Sulliv. 1. Notothylas decurva (Mitten) Stephani, Sp. hepat. 5: 1020 (1917); Hassel de Menendez in J. Hattori bot. Lab. 41: 23 (1976). Notothylas angolensis Stephani, Sp. hepat. 5: 1020 (1917). SIERRA LEONE. Musaia, bare ground at base of Ceiba pentandra, Sellar B53. GHANA. Mpraeso Scarp, in ditch near road, Hall 2631. Legon University Botanical Gardens, Hall (GC 47099 p.p.). Apparently the commonest species of Notothylas in West Africa, and doubtless more frequent than the above records suggest. 2. Notothylas indica Kashyap in Proc. Lahore philos. Soc. 4: 54 (1925). GHANA. Legon University Botanical Gardens, mixed with Notothylas decurva, Hall (GC 47099 p.p.). Notothylas indica differs from N. decurva in having black (not yellow) spores, a somewhat longer sporangium with a well developed persistent columella, and long narrow exothecial cells. The plant from Legon matches specimens from Lucknow, kindly provided by Professor Ram Udar. A plant from Sierra Leone (one mile south of Musaia town, on road bank under trees, Donald s.n.) agrees with N. indica in the sporangium wall, but has yellow spores. * Usually dioecious, the keels of the female bracts usually more or less winged, stem in transverse section with only three medullary cells, lobule inflated with round mouth and long curved apical tooth, cells of lobe small. HEPATICS OF SIERRA LEONE & GHANA 269 XL. ODONTOLEJEUNEA (Spruce) Schiffner 1. Odontolejeunea tortuosa (Lehm. & Lindenb.) Stephani, Sp. hepat. 5: 173'(1912); Vanden Berghen in Revue bryol. lichén. 32: 52 (1963). SIERRA LEONE. Loma Mountains: forest near ‘Camp 1’, 1220 m (4000 ft), Harrington 293; gallery forest c. one km west of ‘Camp 2’, on leaves of Memecylon, Morton (Harrington 359, 362, 365 et al.); c. 1650 m (5400 ft), epiphyllous in forest by stream west of Bintimani (perhaps the same locality as Morton’s) Jones 1488a. Locally abundant on the leaves of shrubs in the forest of the Loma plateau. A montane species with a wide, but disjunct, distribution in tropical Africa, usually at altitudes of 1500-1700 m, though a few records are from much lower altitudes, and the first African collection, by Palisot de Beauvois from ‘Oware’ (Warri), Nigeria must have been from near sea-level. It is thus possible that it occurs in Ghana, e.g. in the Ankasa River area or the Atewa Hills. Where it is found it usually seems to be abundant. XLI. PHAEOCEROS Proskauer 1. Phaeoceros laevis (L.) Proskauer in Bull. Torrey bot. Club 78: 347 (1951), subsp. carolinianus (Michaux) Proskauer, Rapps Communs VIII Congr. int. Bot. Paris, Sect. 14-16: 69 (1954); S. Arnell, Hepat. S. Afr.: 400 (1963). SIERRA LEONE. Roadside one mile south of Musaia town, on rock under trees, Donald s.n. Widely distributed in tropical and South Africa, and doubtless more frequent than the single record suggests. It is likely to occur in wet sites in drier parts of Sierra Leone and Ghana, where hepatics have not been collected. XLII. PLAGIOCHASMA Lehm. & Lindenb. 1. Plagiochasma eximium (Schiffner) Stephani, Sp. hepat. 1: 78 (1898); Bischler in Revue bryol. lichén. 44: 248 (1978). SIERRA LEONE. Loma Mountains, Bintimani, frequent on earth in crevices in the dolerite cliffs below the summit, 1830 m (6000 ft) upwards: R. R. Glanville 479, Harrington 181, 183, Jones 1485, det. H. Bischler. Plagiochasma eximium is a montane species with a very wide but disjunct distribution; known also from Guinea, Cameroon, Zaire (Shaba), East Africa, Natal, southern Arabia, etc. XLUI. PLAGIOCHILA (Dumort.) Dumort. 1. Plagiochila africana Stephani, Sp. hepat. 2: 263 (1902); E. Jones in Trans. Br. bryol. Soc. 4: 293 (1962). SIERRA LEONE. Kambui Hills, Bambawo, in forest on twig of small tree, Harrington 414; epiphyl- lous, Harrington 441a. GHANA. Ankasa River F.R., on branchlet of small tree by the river, Jones s.n.; epiphyllous on small shrub, Jones 1375 p.p. Pra-Suhien F.R., Jones 1337. A lowland species, apparently restricted to the wetter parts of the West African forest from Cameroon, where it is frequent, westwards. Usually on small branches, lianes etc.; only exceptionally epiphyllous. 2. Plagiochila barteri Mitten inJ. Linn. Soc. (Bot.) 22: 320 (1886); E. Jones in Trans. Br. bryol. Soc. 4: 276 (1962). SIERRA LEONE. Presumably from the Freetown Peninsula, where, however, it has not been collected recently, Barter (NY, type). Loma Mountains: near ‘Camp 1’, 1220 m (4000 ft), on tree on margin of forest, Harrington 245; near ‘Camp 2’, 1520 m (5000 ft), on partly sheltered rock face in plateau grassland, Harrington 152. A montane species, abundant in some of the East African mountains — e.g. on Kilimanjaro 270 E. W. JONES & A. J. HARRINGTON between c. 1850 and 2800 m — and also present, though apparently not abundant, on Mont Cameroun (Cameroon Mountain) and the islands of the Gulf of Guinea. On Principe it has been gathered from as low as 460 m, so Barter may have found it in 1857 in the hills close to Freetown at similar altitudes, where the forest is now secondary. Plagiochila barteri seems to be uncommon on the Loma plateau. 3. Plagiochila fusifera Taylor in J. Bot., Lond. 5: 268 (1846); E. Jones in Trans. Br. bryol. Soc. 4: 311 (1962). Fig. 8. SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., side of partly buried boulder by bridge, Harrington 589; Guma Valley, on trees by road, Harrington 111. Lake Sonfon, on rotten bough in forest fringing stream, Harrington 36 (a depauperate form). Loma Mountains: near ‘Camp 2’, c. 1520 m (5000 ft), on trees in forest, Harrington 155, 157, 159, 297, 298, 318; sheltered rock in grassland, c. 1520 m, Harrington 152, Jones 1481; on Vernonia in scrub north-west of Bintimani, 1680 m (5500 ft), Jones 1497d p.p.; Bintimani summit plateau, c. 1940 m (6360 ft), in very sheltered crevices between boulders, Harrington 200 (leaves large enough for Plagiochila divergens but habit of P. fusifera); c. 1370 m (4500 ft), on sheltered boulder face in broken woodland, Harrington 231; branch in crown of tree on edge of forest, 1370 m, Jones 1504. Jawo near Kenema, branch of tree on bank of the river Moa, at the top of the flood zone, Jones 1540 (determination doubtful). GHANA. Pra-Suhien F.R., bole of small tree in moderate shade, Jones 1336a. Atewa Hills F.R., on Vernonia, Jones 1325a. Amedzofe, on trees in old coffee farm, Jones 1246. Rocks in roadside bank above Vane, Jones 1267. The plants grouped under the polymorphic Plagiochila fusifera form a taxonomically difficult complex. Jones (1962) tentatively distinguished three taxa in the group, Plagiochila divergens Stephani var. divergens, P. divergens var. myriocarpa (Pearson) E. Jones, and P. fusifera, but found it difficult to define clear boundaries between them; moreover poorly developed or atypical forms of P. fusifera might be difficult to separate from forms of P. moenkemeyeri, P. praemorsa, and P. strictifolia. The examination of many more specimens, especially from Tanzania and from West Africa, has neither reduced the difficulties nor suggested any better treatment. Typical Plagiochila divergens var. divergens differs from typical P. fusifera in habit, size, and dentition of leaf: P. divergens is richly and regularly pinnately or bipinnately branched, forming large oblong or triangular fronds, the shoots S—6 mm broad, with leaves 2:0-3-0 x 1-4-2-:0 mm, the teeth long and sharp, ending in 1-3 uniseriate long cells, the apical cell about 30 um long, twice as long as broad. P. fusifera is short, little branched, with an unbranched basal segment 10-14 mm long and 2-0-4-0 mm broad, and a head of a few subpinnate branches; leaves 1-4-1-7-(2-0) x 0-95-1-4 mm, the teeth short, broadly triangular, ending in 1-2 cells, the apical cell less than twice as long as broad, usually about as long as broad. P. divergens var. myriocarpa is intermediate in size (leaves 1-9-2:0 x 1-25-1-4 mm), but with the habit and dentition of P. divergens. Some of the plants from the Loma Mountains (especially Harrington 157, 297, Jones 1504) have leaves which are in the size-range of var. myriocarpa and also resemble it in dentition, with the apical cells of the teeth up to 30 wm X 12-15 um, but they resemble Plagiochila fusifera in habit, and they seem inseparable from more typical P. fusifera growing near by. Yet other plants from Loma, notably Harrington 200, have leaves which are large enough for var. divergens, though with short teeth and P. fusifera habit. The plants show very great variation in the degree to which the postical leaf-bases are ampliate, thus forming a ventral crest, and also in the development of trigones. Jones (op. cit.) commented that ‘P. fusifera appears to be characteristic of the country with a sufficiently high rainfall but a strong dry season’; the records for Sierra Leone and Ghana agree with this statement. Thus in Ghana the species seems to be uncommon in the forest districts, and was not found in the wet west. 4. Plagiochila integerrima Stephani in Bolm Soc. broteriana 4: 173 (1886); E. Jones in Trans. Br. bryol. Soc. 4: 267 (1962). HEPATICS OF SIERRA LEONE & GHANA 211 SIERRA LEONE. Loma Mountains: near ‘Camp 1’, 1220 m (4000 ft), on boulder by stream in forest, Harrington 270a, b; near ‘Camp 2’, 1520 m (5000 ft), on stone in stream in forest, Harrington 162. Kambui Hills, Bambawo, in forest on small tree root, partly submerged in stream, Harrington 465. Giema, on ‘boulders in and by stream (periodically submerged) in secondary forest, Harrington 520. GHANA. Pampramasi, by stream in forest, Jones! Ochi Headwaters F.R., West-Skinn 181, 186. Assin Cocoa Station (?5°42’N, 1°17’W), West-Skinn 129. Asuansi (5°18'N, 1°14’W), at side of river Kakum, H. E. Box 2080. Agogo, Foote 16. Begoro, on rocks below the waterfall, Jones ! Southern Scarp F.R., by stream, Jones ! Atewa Hills F.R., on tree boles in swamp forest (the ‘Kaolin Swamp’), Jones 1331 (GC); on stones in stream, Jones ! Bunso, Foote s.n. Perhaps the most generally distributed Plagiochila of tropical Africa, frequent throughout a wide range of rainfall regimes and altitudes, but always in wet places around springs, streams etc., often in deep shade. The absence of records from the Freetown Peninsula is remarkable. P. integerrima probably needs sites that are permanently moist, and the Freetown streams may be too seasonal in their flow and too torrential in the rains. 5. Plagiochila moenkemeyeri Stephani, Sp. hepat. 2: 412 (1904); E. Jones in Trans. Br. bryol. Soc. 4: 321 (1962). Fig. 8. SIERRA LEONE. Freetown Peninsula: Leicester Peak, abundant from c. 520 m (1700 ft) upwards, on boles of small trees and on twigs of shrubs, both in deep shade and in open rocky ground, Brenan (Jones 476), Jones 1430, 1431, 1459, 1460, 1461b, 1462a, and less frequently on rocks, Brenan (Jones 477), Jones 1458a; Regent, Irvine 218; Sugar Loaf Gap, on understorey shrubs and small trees, Harrington 648; York Pass, on small tree, Harrington 611a. Loma Mountains: forest between Sokurela and ‘Camp 1’, 910-1220 m (3000-4000 ft), Jones ?1471b; forest in valley near ‘Camp 2’, c. 1650 m (5400 ft), intimately mixed with Plagiochila fusifera, Jones ?1481a; near ‘Camp 1’, on small tree in forest, 1220 m (4000 ft), Harrington 255, 263. GHANA. Krokosua Hills F.R., 520-550 m (1700-1800 ft), on stems of small shrubs in forest, mixed with Plagiochila strictifolia, Jones 1397b. Aburi Botanical Gardens, Jones s.n. Amedzofe, on trees in cocoa farms above the waterfall, frequent, Jones 1271a. Fig. 8 (a), (b) leaves of Plagiochila fusifera Taylor. (a) Harrington 589 from Fourah Bay College B.R.; (b) Harrington 297 from the Loma Mountains — like some other specimens from here more coarsely and irregularly dentate than in the usual lowland forms of P. fusifera. (c)-(f) leaves of Plagiochila moenkemeyeri Stephani. (c) Jones 1430 from Leicester Peak, base of tree in small gap, open situation in rocky ground; (d) Jones 1462a from Leicester Peak, vertical bole of tree in deep shade; (e) Jones 1271a from Amedzofe; (f) Jones 1471b from the Loma Mountains — form with postical bases of leaves making a well-marked crest and antical bases more shortly decurrent than in typical P. moenkemeyeri, though more longly decurrent than in P. fusifera. All x20. 212 E. W. JONES & A. J. HARRINGTON Plagiochila moenkemeyeri is predominantly, if not exclusively, a West African species which is abundant in the region of the Bight of Biafra and is widely distributed through the forest region to the west, though perhaps local. It tends to be montane, although there are records from near sea-level in Nigeria. It is very variable, and forms occur which are difficult to separate from P. fusifera, though typically the two species are quite distinct. P. moenkemeyeri is more rigid, with more open, apparently dichotomous branching, and the dorsal leaf bases are more longly decurrent and much longer relative to the ventral base (Fig. 8). The trigones tend to be larger than in P. fusifera, and often somewhat nodular, though they are sometimes small. The longly decurrent dorsal leaf bases and the tendency of the ventral bases to form a crest may also give rise to difficulty in distinguishing P. moenkemeyeri from depauperate forms of P. pinniflora, though well developed forms of the two species are very different. The queried numbers from the Loma Mountains are determined with doubt. Jones 1471b and 1481a have leaves which are more shortly decurrent dorsally than in typical Plagiochila moenkemeyeri, with a tendency for the dorsal wing to be abruptly contracted at its base, and the trigones are quite small; the ventral bases are more strongly ampliate than in typical P. moenkemeyeri, and form a strong ventral crest. These plants may possibly represent a different taxon. 6. Plagiochila neckeroidea Mitten in Trans. Linn. Soc. Lond. 23: 57 (1860); E. Jones in Trans. Br. bryol. Soc. 4: 291 (1962). SIERRA LEONE. Freetown Peninsula, Fourah Bay College B.R., on bole of tree in valley, Jones s.n. Gola North F.R., fallen from tree in unexploited forest, Jones 1570b. GHANA. Krokosua Hills F.R., 550 m (1800 ft), on bole of tree on margin of forest, Jones 1404. Pampramasi, on bole of pole-tree in small gap in forest, Jones 1408. Ankasa River F.R., abundant on stems of small shrubs near the river, Jones 1375; occasionally epiphyllous, Jones 1375 p.p., Jenik (Herb. Jones & OP). Subri F.R., Jones ! Pra-Suhien F.R., on boles of small trees, Jones 1337a. Ochi Headwaters F.R., West-Skinn 144, 189. Kade A.R.S., bole of small tree in swamp forest, Jones 1224b. Plagiochila neckeroidea is frequent throughout the lowland rain forest of West Africa, and extends into the Congo Basin. It seems to require high atmospheric humidity, and is thus frequent in swamp forest, by rivers, and around very sheltered gaps in the forest, though never where liable to be submerged. It is a very distinct species which is likely to be confused only with P. salvadorica. P. neckeroidea occasionally produces deciduous plantlets on the leaves (e.g. in Jones 1408), though more rarely than does P. salvadorica. 7. Plagiochila pectinata (Willd.) Lindenb., Sp. hepat.: 14 (1839); E. Jones in Trans. Br. bryol. Soc. 4: 265 (1962). SIERRA LEONE. Presumably from near Freetown, Barter, two un-named collections in Herb. Mitten (NY), det. Jones. GHANA. Atewa Hills F.R., on base of a Gilbertiodendron limba, Richards & Hossain (GC 36671); on boles of trees in swamp forest (the ‘Kaolin Swamp’), Jones 1330a. Plagiochila pectinata is a montane species with a very disjunct distribution. It is abundant in the Mascarenes, in Tanzania in the Uluguru Mountains, and on the islands of the Gulf of Guinea; there are no records from Cameroon. The occurrence of outliers further west is therefore remarkable. In the Atewa Hills, P. pectinata is certainly very local, and it is probable that both records come from the same small swamp, where the plant is very abundant. A specimen in the British Museum collected c. 1823 by George Don, labelled ‘Africa tropica occidentalis’ probably came from near Freetown, but as Don also visited Sao Tomé he may have collected it there. It is likely that, as with P. barteri, the destruction of the primary forests has exterminated P. pectinata. 8. Plagiochila pinniflora Stephani in Hedwigia 30: 212 (1891); E. Jones in Trans. Br. bryol. Soc. 4: 281 (1962). SIERRA LEONE. Loma Mountains, Sokurela, on branch of tree in swamp forest, Jones 1465b. HEPATICS OF SIERRA LEONE & GHANA 213 Jagbahun (8°08'N, 11°56’W), hanging from branches of kola trees, Deighton 3083. Gama [‘Ngaama’] (7°44'N, 11°13'W), upper branches of tree, Deighton 5882. Gola North F.R., on branch of tree in unexploited forest, Jones 1571. GHANA. Oda, on cocoa, Irvine 9. Bunso, on cocoa, Jones 1317 p.p. Kade A.R.S., in forest, dominant on upper branches of Aningeria, Hossain (GC 39316); on branches of Entandophragma, Richards 6318. Amedzofe, 730 m (2400 ft), on cocoa, Jones 1271. Plagiochila pinniflora is characteristic of the lowland rain forest of West Africa, and extends into the Congo Basin and western Uganda. It is often abundant in the crowns of trees, and, like other species of this habitat, is rarely collected unless from felled trees or fallen branches. It does, however, also occur freely in plantations, especially of cocoa. In Sierra Leone it may, perhaps, be present chiefly in the south-east. It is a distinctive species which never gives rise to difficulties of identification unless it is very depauperate. 9. Plagiochila praemorsa Stephani in Bot. Jb. 8: 92 (1886 [‘1887’]); E. Jones in Trans. Br. bryol. Soc. 4: 300 (1962). SIERRA LEONE. Freetown Peninsula: Leicester Peak, on rock, Brenan (Jones 478) Sugar Loaf Gap, on understorey shrubs, Harrington 648 p.p. Gola Hills: near Lalehun, Marshall s.n.; Gola North F.R., on fallen branch in unexploited forest, Jones 1570a, and on stem of small undershrub, Jones s.n. GHANA. Krokosua Hills F.R., c. 550 m (1800 ft), festooning twigs and boles of trees in seasonally wet forest around a ‘glade’, slender forms, Jones 1404 p.p., 1405. Ankasa River F.R., epiphyllous, Jenik (Herb. Jones & OP). ‘Ashanti’, Cummins (G 5165). Tarkwa, Saxby (CGE, sub Plagiochila strictifolia). Subri F.R., on Mitragyne in swamp forest, Jones 1356, and on Raphia in swamp, Jones 1354b. Pra-Suhien F.R., on exposed tree root in bank, Jones 1342a, and on bole 1-3 feet from ground, Jones 1342b. Ochi Headwaters F.R., West-Skinn 187, 193. Begoro, on flat rocks by stream above the waterfall, Jones 131la p.p.; on horizontal bough in open place at foot of waterfall, Jones ?1313 (perhaps Plagiochila strictifolia forma). Atewa Hills F.R., on small shrubs by stream in deep shade, Jones 1324b; clothing the boles of trees in swamp forest (the ‘Kaolin Swamp’), slender forms, Jones 1330b. Kade A.R.S., on small poles in swamp forest, Jones s.n. Like Plagiochila neckeroidea, P. praemorsa is widely distributed in the lowland rain forest of West Africa, and extends into the Congo Basin, usually in sites of high atmospheric humidity. It is extremely variable, and forms occur which are difficult to separate from forms of P. strictifolia and P. fusifera. It sometimes grows in extremely slender states that would be impossible to name without the evidence of associated more robust shoots. Such forms, mingled with many other bryophytes, clothe the tree boles with a thick mantle in the ‘Kaolin Swamp’ of the Atewa Hills F.R. (Jones 1330b; also s.n.), and they also festoon the twigs of small trees in a piece of wet woodland on the Krokosua plateau. 10. Plagiochila salvadorica Stephani in Hedwigia 30: 272 (1891); E. Jones in Trans. Br. bryol. Soc. 4: 293 (1962). SIERRA LEONE. Freetown Peninsula: Guma Valley, epiphyte in unexploited forest, Harrington 100; Picket Hill, in forest just below the summit, c. 850 m (2800 ft), Gledhill s.n. Gola North F.R., abundant on the boles of trees, Jones 1557, 1559b, 1564, 1569. GHANA. Ankasa River F.R., on stems of understorey shrubs near the river, Jones 1375c. Atewa Hills F.R., very abundant on the boles of large Lophira etc., in swamp forest (the ‘Kaolin Swamp’), Jones 1328; dominant on the trunk of Gilbertiodendron limba, Richards & Hossain (GC 36669) (perhaps the same locality as Jones 1328). Plagiochila salvadorica is closely allied to P. neckeroidea, and this is the only species with which it is likely to be confused. The two species have a generally similar geographical range, and indeed they sometimes grow together, though in Zaire and in Nigeria (if indeed it occurs) P. salvadorica seems to be rare, and P. neckeroidea frequent. In the western part of the range P. salvadorica is at least as abundant as P. neckeroidea. The essential difference between the two species lies in the shape of the postical leaf base — diverging away from the stem close to the insertion in P. neckeroidea, but prolonged forwards parallel to the stem for an appreciable distance before bending outwards into the postical margin in P. salvadorica. P. salvadorica 274 E. W. JONES & A. J. HARRINGTON often, though by no means always, forms deciduous plantlets on the leaves, while P. neckeroidea rarely does so; this character cannot be relied on to separate the species. 11. Plagiochila strictifolia Stephani in Hedwigia 30: 210 (1891); E. Jones in Trans. Br. bryol. Soc. 4: 297 (1962). SIERRA LEONE. Freetown Peninsula: Leicester Peak, on tree trunk in rather open forest, Brenan (Jones 479), on stems of small shrubs, Jones 1431 p.p., on base of mango, Jones 1461a; Sugar Loaf Gap, on understorey shrubs and small trees, Harrington 651. Loma Mountains, forest above Sokurela, 910-1220 m (3000-4000 ft), a slender and probably depauperate plant, named with doubt, Jones 1471a. Kambui Hills, Bambawo, on low vegetation in forest, Harrington 481. GHANA. Krokosua Hills F.R., 520-550 m (1700-1800 ft), on stems of small undershrubs, Jones 1379b; on liane, Jones 1389 p.p. Tarkwa, Saxby (CGE). Subri F.R., on base of liane, Jones 1357. Atewa Hills F.R., bases of shrubs by stream, Jones 1324; bole of Vernonia, Jones 1325b; at base of stems in swamp forest, Jones 1329; stems of understorey shrubs, Jones 1293. Kade A.R.S., in swamp forest, Jones s.n. Kpedze, on cocoa, Jones s.n. Plagiochila strictifolia is widely distributed and often abundant in the lowland rain forest of West Africa and the Congo Basin, and there are also records for Uganda, Tanzania (Usambara Mountains), and Angola.* It is very variable and some forms are difficult to distinguish from P. praemorsa; confusion with forms of P. fusifera is also possible. P. strictifolia usually grows near the bases of stems quite close to the ground, with other species of Plagiochila at higher levels. Thus on an old mango on Leicester Peak, Freetown, P. strictifolia grew at the base, with P. moenkemeyeri c. one m (3-4 ft) above (Jones 1461b). It often grows in deep shade, and often also on the bases of trees close to water (e.g. Jones 1324), though not where it is liable to submergence. XLIV. PORELLA L. 1. Porella subdentata (Mitten) E. Jones in Trans. Br. bryol. Soc. 4: 456 (1963), var. subdentata SIERRA LEONE. Musaia Agricultural Station, pumping station, Haswell B1 p.p. Near road-bridge over tributary of the Rokel River, c. 32 km (20 miles) south-west of Kabala, on boulder, Harrington 406. Lake Sonfon, on exposed tree roots in forest, Harrington 38. Loma Mountains: forest between Kurubonla and Seradu, 610 m (2000 ft), Jones 1520a; Seradu, in forest by the village, Jones 1516; tree-bough in forest above Sokurela, 1070 m (3500 ft), Jones 1467; near ‘Camp 1’, 1220 m (4000 ft), on boulder, Harrington 243, and on tree, Harrington 250, 266 p.p.; on sloping rocks and tree boles, very abundant, 1310 m (4300 ft), Jones 1508; forest below Dawule, Morton (Harrington 295a p.p., 295b p.p.); Jaeger 1358 (PC). GHANA. Agogo, Foote s.n. Amedzofe, 700 m (2300 ft), above the waterfall on dry rocks and tree boles in deep shade, Jones 1275a, b. var. camerunensis E. Jones in Trans. Br. bryol. Soc. 4: 456 (1963). GHANA. Begoro, draping horizontal bough in open place in forest below the waterfall, Jones 1313. Porella subdentata is widely distributed but local in West Africa; it is usually abundant where it occurs. It is probably the most widely distributed Porella in Africa, extending through the Congo Basin to Uganda, Kenya, and Angola. It shows montane tendencies; all the sites listed above are probably near or above 300 m (1000 ft), but it reaches its greatest abundance at moderate altitudes. Thus in the Loma Mountains it is uncommon below 900 m, and abundant in the forest at 1200-1300 m, but has not been recorded from the forest of the plateau at c. 1500 m, where it must be rare, if indeed it is present. Probably it does not migrate readily into new habitats, so that freedom from fires and disturbance may play a large part in determining its distribution. Var. subdentata has leaves, lobules, and underleaves that are entire or very weakly dentate, and var. camerunensis has leaves which are sharply dentate at the apex and lobules also sharply dentate, some of the teeth being spiniform; it also has broader lobules than the type variety. Jones 1313 from Begoro shows that var. camerunensis is not a high-altitude form, as might have * All records of Plagiochila spp. that are based solely on ancient and scanty herbarium specimens should be accepted with reserve. HEPATICS OF SIERRA LEONE & GHANA Zio been deduced from earlier records, but may be a phenotype produced by high atmospheric humidity, combined, perhaps, with good illumination. In their lobules, the Loma Mountain plants are intermediate between the two varieties; the leaf apices are mostly slightly dentate but without the spiniform teeth of var. camerunensis, so that they are best assigned to the type variety. XLV. PRIONOLEJEUNEA (Spruce) Schiffner 1. Prionolejeunea grata (Gottsche) Schiffner in Engl. & Prantl, Nat. Pflanzenfam. 1 (3): 127 (1893); Grolle in Wiss. Z. Friedrich-Schiller-Univ. Jena 27: 14 (1978). Prionolejeunea serrula (Mitten) Stephani, Sp. hepat. 5: 202 (1913); Vanden Berghen in Revue bryol. lichén. 39: 383 (1973). GHANA. Ankasa River F.R., mixed with Bazzania decrescens subsp. molleri on tree bole by the river, Jones 1372 p.p. Subri F.R., on stipe of Raphia in swamp forest, Jones 1352 p.p. Begoro, on bole and roots of Musanga near the foot of the waterfall, Jones 1308. Atewa Hills F.R., on boles of trees in swamp forest, mixed with Plagiochila praemorsa, Jones 1332 p.p. Prionolejeunea grata is widely distributed in tropical Africa, from Madagascar to Tanzania and eastern Nigeria, though apparently very local; it has not hitherto been recorded from further west. XLVI. PTYCHANTHUS Nees 1. Ptychanthus striatus (Lehm. & Lindenb.) Nees, Naturgesch. europ. Leberm. 3: 212 (1838); Vanden Berghen in Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8 (1): 92 (1972). SIERRA LEONE. Loma Mountains: forest between Seradu and Kurubonla, 610 m (2000 ft), Jones ’ 1520d; forest above Sokurela, near ‘Camp 1’, c. 1220 m (4000 ft), on rock, Jones 1506, on trees, Harrington 252, 265, etc.; forest below Dawule, Morton (Harrington 295a p.p., 295b p.p.). Ptychanthus striatus is one of the most characteristic bryophytes of mountain forest throughout tropical Africa, where it often drapes the trunks and branches of trees with its long pinnate fronds. It extends into South Africa, and is widely distributed in the mountains of the Indo-Malayan region. XLVII. PYCNOLEJEUNEA (Spruce) Schiffner 1. Pycnolejeunea contigua (Nees) Grolle in J. Hattori bot. Lab. 45: 179 (1979); E. Jones in J. Bryol. 10: 397 (1979). SIERRA LEONE. Gola North F.R., boughs in crown of large tree, rare and in small patches where no other bryophytes are present, Jones 1563. GHANA. Ankasa River F.R., boughs in crown of Glutea, one mile south of the Ankasa bridge, Jones 1380a. Atewa Hills F.R., boughs in crown of Lophira, Jones 1319d. Aiyaola F.R. near Kade, boughs in crown of large Piptadeniastrum, Jones 1234, 1235 p.p. Pycnolejeunea contigua, originally described from Brazil, has only recently been recognized in Africa, though it is evidently widely distributed in the rain forest. It has doubtless been overlooked and rarely gathered because it grows only on boughs in the crowns of large forest trees where it is inconspicuous, although not a small plant, because it forms small, very closely applied patches of a greyish-green or pale chocolate-brown colour. Moreover it tends to grow where the bryophyte cover is sparse; it was associated in Jones 1235 with the moss Calymperopsis disciformis (C. Mueller) P. Tixier, and in Jones 1319d with Cheilolejeunea trifaria. XLVIII. RADULA Dumort. 1. Radula appressa Mitten in Phil. Trans. R. Soc. 168: 397 (1879); E. Jones in J. Bryol. 9: 479 (1977). 276 E. W. JONES & A. J. HARRINGTON Radula guineensis Stephani in Hedwigia 23: 133 (1884). SIERRA LEONE. Loma Mountains, c. 1680 m (5500 ft), on Vernonia in scrub on the west flank of Bintimani, Jones 1499. Giema, on tree in secondary forest, Harrington 547. Gola North F.R., on small pole in exploited forest, Jones 1561. GHANA. Pampramasi, on smooth bole of small tree in forest, rare and depauperate, determination uncertain, Jones 1408. Ankasa River F.R., base of Parinari, Richards & Hossain (GC 36677); base of smooth bole on edge of small gap, c. per., Jones 1373a, b. Subri F.R., bole of small tree in Raphia swamp, Jones 1354. Begoro waterfall, bole of tree in deep shade, Jones 1310. Atewa Hills F.R., boughs in crown of Cassipourea, Jones 1299b; on bole of Cassipourea, Richards & Hossain (GC 36662). Kibi [‘Kibbi’], in evergreen forest, /rvine 29. Esukawkaw F.R., Jenik (OP). Kade A.R.S., amongst Plagiochila praemorsa on bole of tree in swamp forest, rare, Jones s.n. Osenasi (16 km south-east of Kade), on cocoa, Jones s.n. Radula appressa is the commonest species of corticolous Radula in the lowland rain forests of tropical Africa, where it is very widely distributed. If, as is probable, it proves synonymous with R. javanica Gottsche, it is also widely distributed in the Indo-Malayan region. It usually occurs in small amounts, however, often mixed with other larger bryophytes. The tough, dark brown stem will usually distinguish it in the field from corticolous forms of R. flaccida and from R. holstiana; in critical cases the cross section of the stem provides a clear distinction. 2. Radula boryana (Fried. Weber) Nees, Syn. hepat.: 254 (1845); E. Jones in J. Bryol. 9: 495 (1977). GHANA. Atewa Hills F.R., very abundant on tree boles in swamp forest (the ‘Kaolin Swamp’), Jones 1322b; on base of Gilbertiodendron limba, Richards & Hossain (perhaps the same locality) (GC 36672 p.p.). Amedzofe, in damp shady position near the waterfall, /rvine 416. Radula boryana is widely distributed throughout the mountains of tropical and South Africa, usually at altitudes of 1200 m and more. It is probably pantropical. The Ghana localities are exceptionally low, though it grows as low as 300 m in the Mascarenes. It doubtless requires high atmospheric humidity. 3. Radula flaccida Lindenb. & Gottsche, Syn. hepat.: 726 (1847); E. Jones in J. Bryol. 9: 499 (1977). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., epiphyllous, Harrington 86 p.p., 87 p.p., 103 p.p., Jones 1423 p.p.; York Pass, forest understorey by stream, Harrington 613 p.p., 615 p.p. Kasewe F.R., Richards R7170 p.p. Kambui Hills, Bambawo, on low vegetation in forest, Harrington 442 p.p.; on shrubs in stream gully, Jones 1552. Giema, epiphyllous in secondary forest, Harrington 512 p.p., 528 p.p., 529 p.p., 532 p.p. Gola Hills: in forest near Lalehun, Marshall s.n.; by the river, Lalehun, Jones s.n.; Gola North F.R., on herbs in unexploited forest, Jones 1568 p.p. GHANA. Krokosua Hills F.R., 490-550 m (1600-1800 ft), epiphyllous on low shrubs in forest, Jones 1396 p.p. Ankasa River F.R., by the river, Jones 1375 p.p.; in moist gully, Richards R7054; Jenik s.n. Aiyinasi, by the river Fiakpole, Jones 1364b. Begoro waterfall, Jones ! Southern Scarp F.R., Jones ! Bunso, on boles of cocoa, Jones 1317. Radula flaccida is the commonest Radula in the African lowland rain forest. It is usually epiphyllous, and is one of the most abundant epiphyllous hepatics. It is usually readily recognized in the field by the distinctive large reniform gemmae, which are, however, not invariably present. In their absence, the closely appressed shoots with their regular bipinnate branching and slender green stems help to distinguish it from all except R. stenocalyx. It is not uncommon on tree boles and branches, but in this habitat it has usually been either overlooked or misidentified. 4. Radula ?holstiana Stephani in Bot. Jb. 22: 320 (1895); E. Jones in J. Bryol. 9: 483 (1977). SIERRA LEONE. Loma Mountains, western flank of Bintimani, c. 1680 m (5500 ft), on Vernonia in scrub, Jones 1503b. GHANA. Atewa Hills F.R., on the large branches of a Cassipourea, Richards & Hossain (GC 36665); in HEPATICS OF SIERRA LEONE & GHANA A i | the crown of a Cassipourea, Jones 1299b; in the crown of a Lophira, Jones 1319. Aburi Botanical Gardens, on an old Cycas, Jones s.n. All the above specimens are very scanty and sterile, and cannot be named with certainty. The shoots are mingled with Lejeuneaceae and mosses, closely applied to twigs and small branches. They agree with Radula holstiana in the structure of the stem. The lobules have a long insertion and are slightly ampliate, but have a very strongly developed rhizoidiferous mamilla, and therefore often a very convex keel. A similar plant has been collected in Guinea in the Macenta Massif by S. Lisowski (‘apud flumen Loffa’, 25 January 1962, OP). R. holstiana is abundant in the East African mountains, and also occurs on Mont Cameroun (Cameroon Mountain), so that on geographical grounds its occurrence in montane situations in Ghana and Sierra Leone is likely. Moreover, R. holstiana is the only species of Radula with this form of stem and lobule known from West Africa, though others occur in East Africa and the Mascarenes. 5. Radula stenocalyx Montagne in Annis Sci. nat. (Bot.) IV, 3: 315 (1855); E. Jones in J. Bryol. 9: 502 (1977). SIERRA LEONE. Freetown Peninsula, York Pass, forest understorey by stream, Harrington 613 p.p., 615 p.p. Masimo, Marmo 160 p.p. Kambui Hills, Bambawo, in forest by stream, Harrington 440 p.p., 441b, 442 p.p. Gola North F.R., in unexploited forest, Jones 1568 p.p. GHANA. Ankasa River F.R., on leaves of undergrowth in moist gulley near mile 18 from Mpataba, abundant, and often mixed with Radula flaccida, Richards R7054. Atewa Hills F.R., by stream in forest, abundant, Jones 1294; on leaves of herbs in the ‘Kaolin Swamp’, abundant, Jones s.n. Radula stenocalyx resembles R. flaccida in habit, but differs in the form of the gemmae, which are circular and attached by one edge to the antical margin of the leaf, and in the shape of the lobule; the gemmae are not as freely produced as in R. flaccida, and are often inconspicuous. It is a montane species, and usually occurs at higher altitudes than R. flaccida; it is unusual to find both species together, though in Harrington 613 p.p., 615 p.p. from York Pass, Harrington 442 p.p. from the Kambui Hills, Jones 1568 from the Gola Hills, and Richards R7054 from the Ankasa River F.R. in Ghana both species are mixed on the same leaves. The Ankasa River F.R. is also abnormally low for the occurrence of R. stenocalyx. XLIX. RECTOLEJEUNEA A. W. Evans 1. Rectolejeunea ?arnellii E. Jones in J. Bryol. 8: 71 (1974). SIERRA LEONE. Freetown Peninsula: S. Arnell, in a letter to E. W. Jones dated 31 May 1953, wrote that he had found ‘a small specimen of Rectolejeunea rhodesiae’ amongst his collections from Freetown; he did not publish the record. The relevant packet (in S) is labelled ‘Freetown, S. Leo. S. Arnell 20 XII 1951’. It contains only a few fragments of bark-debris amongst which are two minute fragments of a Rectolejeunea each bearing two to three leaves. Kabala, near the District Officer’s house, on old Citrus, Jones 1524d. Kambui Hills, Bambawo, on smooth stem of a small Drypetes in young-growth forest near the Rest House, Jones 1532c. In Jones 1532c a few stems were scattered in a very mixed sheet of Archilejeunea africana, Lopholejeunea subfusca, Lejeunea caespitosa, Cheilolejeunea intertexta, Microlejeunea africa- na, and Lejeunea camerunensis, but they show the deciduous leaves with their uniformly thickened hexagonal cells 20-22 wm diam. and indications of the outgrowths of marginal cells characteristic of the genus; a single perianth was seen. Jones 1524d is less mixed but is also a very small gathering; the cells are thinner walled, but it is clearly the same species. The perianths are convex dorsally, not plane as described for East African Rectolejeunea arnellii (Jones, op. cit.) (and are thus nearer to R. brittoniae A. W. Evans), but they have only a very short and low dorsal keel. Neither collection is adequate for certain identification of the species. Schuster (1980) regards Rectolejeunea arnellii as falling within the range of variation exhibited by R. brittoniae. 278 E. W. JONES & A. J. HARRINGTON L. RICCARDIA Gray 1. Riccardia angusticosta [‘angusticostata’| (Stephani) Grolle in J. Hattori bot. lab. 36: 550 (1972). Riccardia stephanii (Bescher.) E. Jonesin Trans. Br. bryol. Soc. 3: 81 (1956), non S. Hatt. in Bull. Tokyo Sci. Mus. 11: 164 (1944). SIERRA LEONE. Gola North F.R., on fallen log, Jones s.n. GHANA. Ankasa River F.R., on decaying bark in swampy forest, mixed with Arachniopsis, Richards R7061 p.p. Aiyinasi, in forest north of the Agricultural Research Station, on large rotten log, Jones 1361c; on rotten log in cut-over forest near the river Fiakpole, Jones 1364c. Pra-Suhien F.R., abundant on rotting logs, Jones 1339a. Begoro, in forest below the waterfall, on rotting log, Jones 1309. Riccardia angusticosta differs from R. limbata in being smaller with branches consisting mostly of unistratose wing, only three cells thick in the middle, while R. erosa (Stephani) E. Jones differs in being even smaller. Jones 1309 and 1361c both include fronds that are small enough for R. erosa, mixed with fronds that are typical R. angusticosta; there seems to be no reason for considering these collections to be mixtures of two species. Berrie (1966) found chromosome numbers of n = 10, 20, and 40, supplemented by a variable number of chromosome fragments, in a population of R. stephanii (= R. angusticosta). These observations suggest that R. limbata, R. angusticosta, and R. erosa are at most varieties of a single species, with much ill-defined intraspecific variation. 2. Riccardia limbata (Stephani) E. Jones in Trans. Br. bryol. Soc. 3: 79 (1956). SIERRA LEONE. Freetown Peninsula: in forest above Bathurst, 460 m (1500 ft), covering low stone in stream, constantly wet, Jones 1433; by stream three km north-east of Toke, covering moist sheltered well-lit rock, Jones 1447a & b; near York, Richards R7094. Loma Mountains, c. 1520 m (5000 ft), moist earth bank of streamlet in shade, Jones 1496b. Kambui Hills, Bambawo, steep earth bank by path, Harrington 443, 445; damp silt in floor of old mine workings, Jones 1542b. GHANA. Begoro, below the waterfall, in wet well-lit site on tree boles, Jones 1308b. Atewa Hills F.R., earth side of old pit, Jones 1300. Amedzofe, in shallow pool in very shady rocky spot in waterhole below Gemini, Irvine 402 & 421a (sterile; an aquatic form of either Riccardia limbata or R. longispica (Stephani) E. Jones); on dripping wet cliff in deep shade, in ravine above the waterfall, Jones 1280 (sterile). Much more study of populations in the field is needed before the taxa of the complex that includes Riccardia limbata can be satisfactorily delimited. There seems to be little other than habit to differentiate Riccardia limbata from R. longispica, and little other than size and the lesser development of wings to differentiate it from R. angusticosta; both also are very close to R. multifida (L.) Gray. Of the plants recorded above, Jones 1447a & 1496b, Richards R7094 and Irvine 402 & 421a might equally well be referred to R. longispica, while Jones 1300 might equally well be large R. angusticosta. Pearson (1931) recorded Aneura travisiana Pearson, collected by Saxby from Tarkwa, though he stated that it differed from the type in being less winged and more digitately branched. Jones (1956) considered that the type of A. travisiana was Riccardia stephanii (= R. angusticosta) but that Saxby’s plant (Tarkwa, on dead wood, CGE - it has not been possible to trace a specimen at MANCH) was different. The Tarkwa plant may be a xeromorphic form of R. limbata, of which it has the dimensions (thallus 0-6-0-8 mm wide, five cells thick in the middle, biconvex in cross section, or, according to Pearson, plano-convex), differing only in the wings being only one cell wide and in the rather irregular subpinnate or more or less palmate branching. 3. Riccardia sp. GHANA. Between Vane and Amedzofe, rocks in roadside bank, Jones 1262. Amedzofe, earth bank near the Training College, Hall (GC47071). Fronds bright green, crowded, imbricate, ascending shoots 2-4 mm long and 0-4-0-5 mm wide, irregularly shortly pinnate, in cross section plane dorsally and convex ventrally or often slightly concave dorsally, thus channelled above, three cells thick and, where not gemmiferous, with a HEPATICS OF SIERRA LEONE & GHANA 279 unistratose wing two to three cells broad; usually producing abundant gemmae from the marginal and many cortical cells around the apices of the shoots; gemmae two-celled, 20-28 x 35-40 wm. Median cortical cells (25)-35-45 x 50-60-(95) um, inner cells of the wings 35 x 55-70 um, marginal cells 22-30 x 25-35 wm. Oil bodies in all cells, mostly one per cell, occasionally two or three, dark grey, opaque, granular, oval with rounded or occasionally pointed ends, those in the cortical cells 8 x 10-14 wm, in the medullary cells 9 x 24—30 wm. Only male plants seen. Jones kept GC 47071 (kindly sent by air mail by Mr J. B. Hall) in cultivation at Kirtlington for two years. It produced copious male inflorescences but no female, so that there is little doubt that it is a dioecious species. It produced few gemmae in cultivation, and continued to make prostrate pinnate shoots with narrow parallel-sided fronds. What is apparently the same species has been collected by De Sloover (13338, NAM) in the Forét de Gishwati, Rwanda, by Demaret (6471, BR) at Yaselia in the Parc National Albert of the former Belgian Congo, and in Mauritius by Vaughan (MAU 11730); De Sloover 13338 is female. The plant from Mauritius has also been cultivated at Kirtlington, and like that from Amedzofe it is persistently male. It is tempting to name this species Riccardia incurvata Lindb., though the geographical distribution of R. incurvata, so far as it is known at present, makes this identity improbable. European R. incurvata differs chiefly in having a thallus five to six cells thick, with the contrast in size between cortical and medullary cells less pronounced; it is also more consistently lunate in cross section. LI. RICCIA L. 1. Riccia fluitans L., Sp. p/.: 1139 (1753); Vanden Berghen in Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8 (1): 186 (1972). GHANA. Nsuta, in the Pra River, Jones ! Riccia fluitans is frequent throughout Africa in fresh water, and will doubtless prove to be widely distributed in both Sierra Leone and Ghana. 2. Riccia lanceolata Stephani in Hedwigia 27: 110 (1888); E. Jones in Trans. Br. bryol. Soc. 3: 218 (1957). GHANA. Near Kumasi, on an inselberg at mile 13 on the Mampong road, S. R. Woodell, June 1971 (Herb. Jones). Riccia lanceolata is frequent on thin earth over rocks in the southern savannas of Nigeria, and is likely to be frequent in similar situations in Ghana. The spores of the Kumasi plant differed from those described by Jones (op. cit.) in being larger (90-100 um diam.), and in having the outer face areolate over the whole of its surface. 3. Riccia membranacea Gottsche & Lindenb., Syn. hepat.: 608 (1846); E. Jones in Trans. Br. bryol. Soc. 3: 210 (1957). GHANA. Nsuta, banks of the Pra River, on heavy rich clay in the flood zone, Jones 1351. Riccia membranacea is widely distributed in tropical Africa on ground that is liable to inundation by rivers, and will doubtless be frequent in both Sierra Leone and Ghana. 4. Riccia moenkemeyeri Stephani in Bot. Jb. 8: 95 (1886 [‘1887’]); E. Jones in Trans. Br. bryol. Soc. 3: 211 (1957). Riccia undulata §. Arnell in Bot. Notiser 105: 307 (1952). SIERRA LEONE. Freetown, streamside, Arnell 2252 (as Riccia undulata). Musaia, in very damp beds of forest nursery, D. Small 272, September 1951. Koinadugu village, on bare ground, Sellar B46. GHANA. Cape Coast (5°06’N, 1°15’W), mud near stream, Hall 1536. Mpraeso Scarp, probably this species (sterile), in ditch mixed with Notothylas decurva, in damp forest, Hall 2630. 280 E. W. JONES & A. J. HARRINGTON According to the description provided by Arnell (op. cit.), Riccia undulata differed from R. moenkemeyeri only in its greater size (thallus up to seven mm wide). The type specimen in S is much smaller, and well within the normal range of size of R. moenkemeyeri; the spores are identical with those of R. moenkemeyeri. Riccia moenkemeyeri is widely distributed in tropical Africa from Nigeria eastwards into the Congo Basin. It has also been recorded from Gambia (rice fields near Keneba, D. H. Murphy, 30 December 1958, BM). Belonging to the subgenus Ricciella, it tends to occur in wetter country than most species, which belong to the subgenus Riccia, and is frequent in rain forest districts. 5. Riccia nigrosquamata E. Jones in Trans. Br. bryol. Soc. 3: 222 (1957). SIERRA LEONE. Freetown Peninsula, on laterite soil by the Fourah Bay College Guest House, G. K. Berrie s.n., August 1971 (Herb. Jones). Musaia town, on house foundation, Donald s.n. Koinadugu village, on bare ground, Sellar B48. Riccia nigrosquamata is known from Tanzania and from the neighbourhood of Ibadan in Nigeria; it is to be expected in Ghana. 6. Riccia rhodesiae S. Arnell in Bot. Notiser 105: 313 (1952); E. Jones in Trans. Br. bryol. Soc. SP 22061957 )5 GHANA. Kibi [‘Kibbi’], evergreen forest, /rvine 29. Achimota College, in lawn, Boughey (Herb. Jones). Legon University, in the Botanical Gardens, and in turf around the Botany Department, Jones 1386. LII. RICCIOCARPOS Corda 1. Ricciocarpos natans (L.) Corda in Opiz, Beitr. Naturgesch.: 651 (1829); Vanden Berghen in Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8 (1): 184 (1972). GHANA. Dabala, near Sogakofe, Lower Volta (5°59'N, 0°41’E), in newly flooded swamp, with Azolla, Salvinia, etc., Hall B10. Widely distributed in fresh water throughout tropical and temperate regions. LIT. SCHIFFNERIOLEJEUNEA Verd. 1. Schiffneriolejeunea fragilis Gradst. & E. Jones in J. Bryol. 12: 45 (1982). SIERRA LEONE. Loma Mountains, Seradu, on boles of trees in forest by the village, Jones 1515a, 1516a, 1517. Kambui Hills, Bambawo, bole of tree in forest near the Rest House, Jones 1550b. GHANA. Kade A.R.S., on main trunk and branches of Aningeria, Hossain (GC 39306, 39318 et al.); on upper branches of Parinari, Hossain (GC 36700, 39339). Schiffneriolejeunea fragilis is the plant alluded to by Jones (1954a: 400) as allied to Ptychocoleus pappeanus (Nees) Stephani (= Schiffneriolejeunea pappeana (Nees) Gradst. in J. Hattori bot. Lab. 38: 335 (1974)). It is a species of the lowland rain forest of West Africa and the Congo Basin, in contrast to S$. pappeana, which occurs in South Africa and the mountains of tropical Africa, including Mont Cameroun (Cameroon Mountain). 2. Schiffneriolejeunea occulta (Stephani) Gradst. in J. Hattori bot. Lab. 38: 333 (1974). Ptychocoleus occultus (Stephani) Stephani, Sp. hepat. 5: 25 (1912); Vanden Berghen in Bull. Jard. bot. Etat Brux. 19: 47 (1948). SIERRA LEONE. Freetown Peninsula: Mount Aureol, Arnell 2309, 2328; Fourah Bay College B.R., near Heddle’s Farm, on bole of tree with Marchesinia excavata, Jones 1420 p.p.; Toke, in scrub behind the beach, on boles of Terminalia scutifera and Chrysobalanops, Jones 1444b, 1446b; Black Johnson Beach near York, in thicket behind the beach, Harrington 22. Rokupr, West African Rice Research Station (9°01'N, 12°57’W), c. 15 m (50 ft), on tree buttress, Harrington 77. Njala, upper part of trunk of Nauclea diderrichii, Deighton 5313 p.p. Jawo, near Kenema, smooth bole of tree on bank of the river Moa, Jones 1538a. HEPATICS OF SIERRA LEONE & GHANA 281 GHANA. Aiyinasi A.R.S., on Citrus, Jones 1383. Schiffneriolejeunea occulta is known elsewhere from the lowland rain forest of western and eastern Nigeria, and the Congo Basin. 3. Schiffneriolejeunea polycarpa (Nees) Gradst. in J. Hattori bot. Lab. 38: 335 (1974). Ptychocoleus molleri (Stephani) Stephani, Sp. hepat. 5: 29 (1912); Vanden Berghen in Bull. Jard. bot. Etat Brux. 19: 44 (1948). SIERRA LEONE. Freetown Peninsula: Havelock Plateau, on branches of bushes in scrub, Jones s.n.; Leicester Peak, 460 m (1500 ft), on old mango below the Rest House, Jones 1464b p.p., in evergreen scrub, 520 m (1700 ft), Jones 1457f, in both collections mixed with Acrolejeunea emergens var. confertissima. Five km south of Kabala, in secondary forest, Harrington 120. Kabala, near the District Officer’s bungalow, on mango, Harrington 133, 140 p.p. Lake Sonfon, on Erythrophleum near stream, Harrington 28 p.p., 29 p.p.,52 p.p. Loma Mountains, Seradu, in forest by the village, on tree boles mixed with Schiffneriolejeunea fragilis, Jones 1515c. Kambui Hills, Bambawo, on large tree by road, Harrington 569 p.p. GHANA. Kade A.R.S., in crown of Entandophragma, Richards R6307 — probably this species, but determination uncertain as the female inflorescences are all eroded and the male not found. Aburi Botanical Gardens, on roots of Hura crepitans, and on bole of cabbage palm, Jones s.n. Amedzofe, ‘on tree trunks in cloud forest’, /rvine 412 p.p.; on shade tree in old coffee farm, 730 m (2400 ft), Jones 1245. Schiffneriolejeunea polycarpa extends from Guinea to Natal and the Mascarenes, but its main area of distribution seems to be the Congo Basin and the Tanzanian mountains, where it is frequent at altitudes of 800-1700 m. It is also known from tropical America. LIV. SPRUCELLA Stephani 1. Sprucella succida (Mitten) Stephani in Hedwigia 30: 214 (1891); Vanden Berghen in Bull. Jard. bot. Etat Brux. 18: 91 (1946). SIERRA LEONE. Freetown Peninsula: Fourah Bay College B.R., clay bank by path, Harrington 595, Jones !; Charlotte Falls, bank by road above the falls, Harrington 640 p.p. Lake Sonfon, dry watercourse, Harrington 51 p.p. Loma Mountains: near ‘Camp 1’, 1220 m (4000 ft), stream bank in forest, Harrington 268; walls of small cave in plateau grassland, south-west of ‘Camp 2’, 1520 m (5000 ft) Harrington 172; western flank of Bintimani, moist earth bank of streamlet, in shade, Jones 1496 p.p. Kambui Hills, Bambawo, on termite mound in forest, Harrington 413; on earth bank by path near the upper bungalows, Harrington 561, Jones 1529 p.p. GHANA. Ankasa River F.R., covering bole of Vitex to height of a metre, Jones 1370; log by the river, Richards R7049. Tarkwa, Saxby (CGE). Subri F.R., Jones ! Pra-Suhien F.R., abundant, Jones ! Begoro, abundant on shady wet rocks below the waterfall, Jones 1308 p.p. Near Kibi [‘Kibbi’], /rvine 31. Atewa Hills F.R., abundant on ironstone rocks on the plateau, Jones ! Amedzofe, shady cliff just below the summit of the hill, Jones s.n.; bank near the Training College, Hall (GC 47071 p.p.). Sprucella succida is widely distributed in the lowland rain forest of West Africa and the Congo Basin, where it often covers shady banks of moist earth or rock in thick sheets; it also occurs on tree boles and on decaying wood. It extends eastwards into Uganda (East Mengo, K. A. Lye B1, Herb. Jones). It is not a montane species, and its occurrence on the Loma plateau at c. 1500 m is surprising. LV. STICTOLEJEUNEA (Spruce) Schiffner 1. Stictolejeunea balfourii (Mitten) E. Jones in J. Bryol. 9:50 (1976). Stictolejeunea africana Vanden Berghen in Bull. Jard. bot. Etat Brux. 20: 15 (1950). GHANA. Krokosua Hills F.R., 490 m (1600 ft), on liane, Jones 1392b, and on tree roots in deep shade, Jones 1395. Ankasa River F.R., Jones s.n. (GC); on base of Parinari glabra, Richards (GC 36677 p.p.). Pra-Suhien F.R., on side of tree root in deep shade, Jones 1343a. Kade A.R.S., in forest in deep shade on roots of trees, bases of smooth boles and old lianes, abundant, Jones 1217, 1219. Stictolejeunea balfourii has been rarely collected, but is widely distributed and probably frequent 282 E. W. JONES & A. J. HARRINGTON in the lowland rain forest of West Africa and, as it was first described from the Island of Rodrigues, it probably extends throughout the Congo Basin. It is certainly frequent in parts of Bendel State, Nigeria, and in Ghana. It makes thin, very closely applied patches on smooth bark, of a blackish green with a characteristic oily appearance which, once recognized, is unmistakable, but it would be passed over as one of the Archilejeunea spp. amongst which it often grows. It usually grows very close to the ground. LVI. STREPSILEJEUNEA (Spruce) Schiffner 1. Strepsilejeunea brevifissa (Gottsche) Stephani, Sp. hepat. 5: 275 (1913); Vanden Berghen in Bull. Soc. r. Bot. Belg. 92: 136 (1960). SIERRA LEONE. Loma Mountains: above Sokurela, 940 m (3100 ft), on tree near lower edge of the forest, Jones 1512; gallery forest in centre of plateau, on large branch, Morton (Harrington 374 p.p.); near ‘Camp 2’, c. 1650 m (5400 ft), on tree on edge of gallery forest, Harrington 146; on liane in open place near ‘Camp 2’, Jones 1495; ‘sur rameaux d’Eugenia, forét relique d’altitude vers 1850 [sic] m’, Jaeger 1188 p.p. (PC), Strepsilejeunea brevifissa is widely distributed in the African mountains from Madagascar, the Mascarenes, and the Cape to Mont Cameroun (Cameroon Mountain); it has not hitherto been reported from further west. In tropical Africa it occurs from c. 1000 to 2500 m, usually straggling amongst other bryophytes on trees. It has often been collected accidentally, so that the specimens which are available in herbaria are mostly scanty. It is extremely variable, particularly in respect of the leaf apex, which may vary from rounded to acuminate, and the carination of the perianth. Gottsche (1882) described the perianths of the original collection from Madagascar as having ‘ventre carina lato (fructu maturo fere evanescente) notato’; such plants occur not only in Madagascar but also in East Africa; at maturity the perianth may be pyriform, terete, and completely eplicate. Plants with quinqueplicate perianths that are bicarinate ventrally are frequent, and every intermediate condition between this and the eplicate perianth occurs. Plants on Mont Cameroun have a triplicate perianth with a deep narrow ventral keel. The Loma Mountain plants have quinqueplicate perianths with two long well-defined ventral keels. It is not clear how much, if any, of this variation is genotypic; that it is not entirely due to age is shown by the fact that the perianths of e.g. Harrington 146 remain distinctly carinate even when the sporangia have discharged. LVII. SYMPHYOGYNA Nees & Montagne 1. Symphyogyna lehmanniana Montagne & Nees, Syn. hepat.: 483 (1846); Vanden Berghen in Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8 (1): 152 (1972). SIERRA LEONE. Loma Mountains, c. 1520 m (5000 ft), on moist earth bank of streamlet in deep shade, mixed with Riccardia limbata and Calypogeia fissa, Jones 1496 p.p. GHANA. Amedzofe, steep dripping cliffs near the waterfall, in deep shade, /rvine 417 (named with doubt). Vanden Berghen (1965) found that Symphyogyna lehmanniana and S. volkensii Stephani differed chiefly in dimensions, S. volkensii being a larger plant, and suggested that more copious material might show that the two should be treated as synonymous. He described the central strand of small cells in S. Jehmanniana in transverse section as 75-120 x 60-90 um, and consisting of 45—60 cells, that of S. volkensii as 50-110 x 80-100 um, and consisting of c. 30 cells. However, an original specimen of S. Jehmanniana (leg. Lehmann) in BM has a central strand only 40 wm broad, consisting of only 30 cells in section. Vanden Berghen recorded S. lehmanniana only from South Africa, and S. volkensii from Kilimanjaro and Zaire at altitudes of 2500 m upwards. The Loma plant, Jones 1496 p.p., agrees in dimensions with Symphyogyna lehmanniana, and though it has a small central strand only 50-70 wm broad x 45-50um deep, composed of 25-30 cells in section, this is clearly within the range of variation of S. lehmanniana. Irvine 417 from Ghana is sterile and badly preserved and cannot therefore be named with certainty. It may be HEPATICS OF SIERRA LEONE & GHANA 283 Pallavicinia lyellii (Hook.) Carruth., which is, however, not yet known with certainty from West Africa, whereas S. lehmanniana is known from Cameroon (Richards R5297 p.p., Bamenda— Banso road, 1675 m). A plant from Mont Cameroun (Cameroon Mountain), with its much greater rainfall (Jones 411, 1890 m alt., also leg. Savory, 1830-2130 m alt.) differs in having an even smaller central strand, in section of only 15 cells which are thin walled. It should probably be considered as being a hygromorphic form rather than a different taxon. Grolle (1980: 330) regards Symphyogyna lehmanniana as being identical to the neotropical S. brasiliensis Nees & Montagne. LVIUI. TARGIONIA L. 1. Targionia hypophylla L., Sp. pl.: 1136 (1753); Vanden Berghen in Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8 (1): 164 (1972). GHANA. Mole Game Reserve, Hall (GC 47203). Targionia hypophylla is widely distributed throughout those parts of tropical Africa which have a long severe dry season, growing especially on the banks of watercourses where it is liable to temporary inundation or kept moist by seeping water in the wet season. It is likely to be frequent in northern Ghana, and is to be expected in northern Sierra Leone. LIX. TAXILEJEUNEA (Spruce) Schiffner 1. Taxilejeunea conformis (Nees & Montagne) Stephani, Sp. hepat. 5: 456 (1913); E. Jones in Trans. Br. bryol. Soc. 5: 294 (1967). Lejeunea conformis Nees & Montagne in Annis Sci. nat. (Bot.) I, 19: 260 (1843). SIERRA LEONE. Loma Mountains: forest near ‘Camp 1’, 1220 m (4000 ft), Harrington 256 p.p.; valley-head west of Bintimani, 1680 m (5500 ft), epiphyllous in scrub, Jones 1498a. GHANA. Amedzofe, in ravine above the waterfall below dripping cliff in deep shade, on rocks, Jones 1285a, and epiphyllous, Jones 1285b. Taxilejeunea conformis is widely distributed in the wetter parts of tropical Africa. It is somewhat montane, and is usually found at altitudes of 1000-1600 m. 2. Taxilejeunea pulchriflora Pearson in Ark. Bot. 19 (5): 15 (1925); E. Jones in J. Bryol. 9: 50 (1976). SIERRA LEONE. Freetown Peninsula: valley above Kongo Dam, Bathurst, 490 m (1600 ft), on barkless rotten log in forest, Jones 1434b p.p. Loma Mountains, valley to west of Bintimani, c. 1520 m (5000 ft), on Cyathea, Jones 1492 (form with obtuse leaves and no subgynoecial innovations). Kambui Hills, Bambawo, on shaded stonework beneath bridge, Harrington 562 p.p. Gola North F.R., on bole of Vitex, with Lejeunea confusa in cushions of Calymperes lonchophyllum Schwaegr., Jones 1558 (form with obtuse leaves). GHANA. Pra-Suhien F.R., on tree root in bank, Jones 1340. Southern Scarp F.R., near Begoro, on boulder, Hall (Jones 1316). Taxilejeunea pulchriflora is widely distributed in West and East Africa but local, and rarely, if ever, abundant. It is to be expected in the Amedzofe district of Ghana, as it has been collected only a short distance to the east in Togo (Campement de Klouto, 720 m, on sandstone, Hiepke & Schultze-Motel 464, B). LX. TELARANEA Spruce ex Schiffner 1. Telaranea nematodes (Gottsche ex Austin) M. Howe in Bull. Torrey bot. Club. 29: 284 (1902). SIERRA LEONE. Loma Mountains, head of the Kongbundu Valley, north-west of Bintimani, c. 1520 m (5000 ft), on vertical banks of moist humic earth in grassland, mixed with Calypogeia longifolia, Jones 1502 p.p. Kambui Hills, Bambawo, on steep shaded earth bank, Harrington 445 p.p. 284 E. W. JONES & A. J. HARRINGTON GHANA. Tarkwa, on rotting wood, Saxby (CGE). Atewa Hills F.R., on base of Gilbertiodendron limba, Richards & Hossain (GC 36673); on very large fallen rotting tree bole, Jones 1321. Telaranea nematodes is widely distributed but local in tropical Africa. It is to be expected in the Amedzofe district, as, like Taxilejeunea pulchriflora, it has been collected a short distance to the east in Togo (between Palimé and Klouto, 650-750 m, Hiepke & Schultze-Motel 409, and on the walls of Chateau Vialle, 750 m, Hiepke & Schultze-Motel 427, in both mixed with Sprucella succida, B). LXI. THYSANANTHUS Lindenb. 1. Thysananthus spathulistipus (Reinw., Blume & Nees) Lindenb., Syn. hepat.: 287 (1845); Vanden Berghen in Revue bryol. lichén. 19: 35 (1950). Lejeunea triquetra Mitten in J. Proc. Linn. Soc. 7: 167 (1863 [‘1864’]). Mastigolejeunea tarkwana Pearson in Annls Cryptog. exot. 4: 63 (1931). SIERRA LEONE. Bagru [‘Bagroo’] River, Mann s.n. (BM, isotype of Lejeunea triquetra). GHANA. Ankasa River F.R., on bole of tree by the rapids, Jones 1376a; in crown of Glutea on bole and larger branches, Jones 1378b. Tarkwa, Saxby (CGE, isotype of Mastigolejeunea tarkwana). Thysananthus spathulistipus, a species widely distributed in the Indo-Malayan region, has also been recorded from Cameroon (Augier, 1978), Nigeria, Zaire, Madagascar, the Mascarenes (Vanden Berghen, op. cit.), and Seychelles (Grolle, 1978). The scarcity of records from West Africa may be due partly to the fact that it usually occurs in the crowns of large trees. It is probably confined to ‘old-growth’ forest. LXII. TRACHYLEJEUNEA (Spruce) Schiffner 1. Trachylejeunea serrulata Stephani, Sp. hepat. 5: 300 (1913). GHANA. Ankasa River F.R., epiphyllous, Jenik s.n.; on low shrubs near the river, Jones 1375, 1381; on undergrowth in moist gully near mile 18 from Mpataba, Richards R7054; on palm leaves in swamp a mile south of the Ankasa bridge, Jones 1382d. This species, hitherto known only from Cameroon, is evidently abundant in the Ankasa River district. The numerous, conspicuous, often reddish ocelli, scattered throughout the leaves, underleaves, and perianths, make it easily recognisable. It is probably wrongly placed in the genus Trachylejeunea; Dr R. Grolle (pers. comm.) considers it to be a Lepidolejeunea. LXIII. TYLIMANTHUS Mitten 1. Tylimanthus ?ruwenzorensis S. Arnell in Ark. Bot. I, 3: 560 (1957); E. Jones inJ. Bryol. 11: 317 (1980). GHANA. Subri F.R., in swamp forest on stipe of Raphia, Jones 1352. This plant was sterile and only a very small amount was found, so that identification of the species is impossible. The genus includes few species. Its occurrence in West Africa at a low altitude is of considerable interest, as hitherto it has been found in tropical Africa only at much higher altitudes in the mountains of the east. Acknowledgements We are indebted to the Royal Society for the financial support that made our collecting possible; E.W.J. for a Grant-in-Aid and A.J.H. for the award of a Leverhulme Scholarship. Our thanks are due to Professor J. K. Morton and Professor G. K. Berrie, both formerly of the Botany Department, Fourah Bay College, University of Sierra Leone, and Professor G. W. Lawson, formerly of the Botany Department, University of Ghana, Legon, for providing us with accommodation and working facilities. We also wish to record our gratitude to the staff of the aforementioned departments, in particular Professor Morton and Mr J. B. Hall, for their hospitality and companionship in the field, and for placing their knowledge of the local flora at our disposal. HEPATICS OF SIERRA LEONE & GHANA 285 We are most grateful to the curators of the following herbaria for the loan of specimens: B, BR, CGE, FH, G, MANCH, NAM, NY, OP, PC, S, and UPS. Finally, we wish to thank Miss L. G. M. Hosking for her care and patience in typing the manuscript. References Arnell, S. 1958. New Hepaticae from Cameroon Mountain. Svensk bot. Tidskr. 52: 63-67. —— 1963. Hepaticae of South Africa. 411 pp. Stockholm. Augier, J. 1972. Groupements de bryophytes terricoles sur le campus universitaire de Yaoundé. Annls Fac. Sci. Univ. féd. Cameroun 9: 73-85. —— 1974. Bryophytes corticoles al’étage submontagnard dans l’ouest camerounais. Annis Fac. Sci. Univ. féd. Cameroun 17: 67-93. — 1978. Sur les bryophytes au Cameroun dans le secteur forestier littoral. Annis Fac. Sci. Yaoundé 25: 95-129. Berrie, G. K. 1966. Polyploidy in some West African species of Riccardia (Gray). Revue bryol. lichén. 34: 302-308. Bizot, M. & Pécs, T. 1974. East African bryophytes I. Acta Acad, paedag. Agriensis II, 12: 383-449. —— 1979. East African bryophytes, III. Acta bot. hung. 25: 223-261. Cole, N. H. A. 1968. The vegetation of Sierra Leone. v+198 pp. Nijala. Duda, J. & Vanden Berghen, C. 1967 [‘1966’]. Hépatiques épiphylles récoltées en Guinée — 1. Revue bryol. lichén. 34: 928. Fulford, M. H. 1968. Manual of the leafy Hepaticae of Latin America — Part III. Mem. N.Y. bot. Gdn 11: 277-392. Gottsche, C. M. 1882. Lebermoose. Jn Reliquiae Rutenbergianae. V. (Botanik, 3. Fortsetzung). Abh. naturw. Ver. Bremen 7: 338-365. Gregory, S. 1965. Rainfall over Sierra Leone. Dep. Geogr. Univ. Lpool Res. Pap. 2. 58 pp. Liverpool. Grolle, R. 1976. Drepanolejeunea subgen. Kolpolejeunea—eine neue Untergattung aus der Palaotropis. J. Hattori bot. Lab. 40: 191-216. — 1978. Die Lebermoose der Seychellen. Wiss. Z. Friedrich-Schiller-Univ. Jena 27: 7-17. —— 1979. Miscellanea hepaticologica 191-200. J. Hattori bot. Lab. 46: 337-355. —— 1980. Miscellanea hepaticologica 201-210. J. Bryol. 11: 325-334. Hall, J. B. 1980. Gazetteer of plant collecting localities in Ghana cited in Flora of West tropical Africa. 122 pp. Accra. Hall, J. B. & Swaine, M. D. 1976. Classification and ecology of closed-canopy forest in Ghana. J. Ecol. 64: 913-951. Hepper, F. N. & Neate, F. 1971. Plant collectors in West Africa. xvi+96 pp. Utrecht. (Regnum veg. 74). Herzog, T. 1952. Miscellanea bryologica. III. Memo. Soc. Fauna Flora fenn. 27: 92-110. Holmgren, P. K., Keuken, W. & Schofield, E. K. 1981. Index herbariorum. Part I. The herbaria of the world. 7th ed. 452 pp. Utrecht. (Regnum veg. 106). Hutchinson, J. & Dalziel, J. M. 1954-72. Flora of West tropical Africa. 2nd ed. (Revised by R. W. J. Keay & F. N. Hepper). 3 vol. London. Jones, E. W. 1954a. African hepatics. IX. Some species of Ptychocoleus. Trans. Br. bryol. Soc. 2: 396—407. —— 1954b. African hepatics. X. Leptocolea and Cololejeunea. Trans. Br. bryol. Soc. 2: 408-438. —— 1956. African hepatics. XI. The genus Riccardia in tropical Africa. Trans. Br. bryol. Soc. 3: 74-84. —— 1962. African hepatics. XV. Plagiochila in tropical Africa. Trans. Br. bryol. Soc. 4: 254-325. —— 1968a. African hepatics. XIX. The Lejeunea flava complex. Trans. Br. bryol. Soc. 5: 548-562. 1968b. African hepatics. XX. Some little-known species and some extensions of range. Trans. Br. bryol. Soc. 5: 563-572. 1973. African hepatics. XXIV. Lejeuneaceae: some new or little-known species and extensions of range. J. Bryol. 7: 545-561. —— 1976. African hepatics. XXIX. Some new or little-known species and extensions of range. J. Bryol. 9: 43-54. 1979. African hepatics. XX XI. Some new or little-known Lejeuneaceae and extensions of range. J. Bryol. 10: 387-400. Kuwahara, Y. 1976. Studies of genus Metzgeria of Colombia collected by Mme. Hélén Bischler, 1956-59. J. Hattori bot. Lab. 40: 259-290. —— 1978. Synopsis of the family Metzgeriaceae. Revue bryol. lichén. 44: 351-410. Paton, J. A. 1973. Taxonomic studies in the genus Fossombronia Raddi. J. Bryol. 7: 243-252. 286 E. W. JONES & A. J. HARRINGTON Pearson, W. H. 1931. Notes on a collection of Hepaticae made by Mr. Saxby on the west coast of Africa. Annls Cryptog. exot. 4: 61-71. Pécs, T. 1975 [‘1974’]. Epiphyllous liverworts from Zambia. Bull. Acad. pol. Sci. Sér. biol. 22: 851-853. — 1980. Central African Hepaticae collected by M. Assel. Folia hist.-nat. Mus. Matraensis 6: 73-81. Potier de la Varde, R. 1948. Contribution 4 la flore bryologique de Sierra Leone. Revue bryol. lichén. 17: 16-23. Schuster, R. M. 1980. The Hepaticae and Anthocerotae of North America east of the hundredth meridian 4. xviii+ 1334 pp. New York. Taylor, C. J. 1960. Synecology and silviculture in Ghana. xit+418 pp. Edinburgh. Tixier, P. 1975. Contribution a l’étude de ’hépaticologie africaine. 1. Récoltes en bordure du Golfe de Guinée (Cameroun et Gabon). Annis Fac. Sci. Yaoundé 20: 3-10. — 1979 [‘1977’]. La famille des Cololejeunoideae (Grolle) dans l’Océan Indien occidental. Essai monographique. Bull. trim. Acad. Malgache 55: 173-247. Vanden Berghen, C. 1948. Contribution a l’étude des espéces africaines du genre Metzgeria. Bull. Jard. bot. Etat Brux. 19: 187-204. ; — 1951. Contribution a l’étude des espéces africaines du genre Ceratolejeunea. Bull. Jard. bot. Etat Brux. 21: 61-81. —— 1960a. Hépatiques récoltées par le Dr J.-J. Symoens dans la région péri-tanganyikaise. Bull. Soc. r. Bot. Belg. 92: 111-138. — 1960b. Hépatiques récoltées en Afrique par M. Monod. Revue bryol. lichén. 29: 50-67. —— 1961. Hépatiques récoltées par le Dr J.-J. Symoens dans la région péri-tanganyikaise. Bull. Soc. r. Bot. Belg. 93: 55-74. — 1965. Hépatiques récoltées par le Dr J.-J. Symoens dans la région péri-tanganyikaise. Bull. Soc. r. Bot. Belg. 98: 129-174. — 1972. Hépatiques et Anthocérotées. Résult. scient. Explor. hydrobiol. Bassin Lac Bangweolo & Luapula 8(1): 1-202. — 1973. Quelques hépatiques récoltées au Gabon par G. Le Testu. Revue bryol. lichén. 39: 365-385. — 1977. Hépatiques épiphylles récoltées par J. L. De Sloover au Kivu (Zaire), au Rwanda et au Burundi. Bull. Jard. bot. natn. Belg. 47: 199-246. Walter, H. & Lieth, H. 1967. Klimadiagramm-Weltatlas. Jena. White, F. (Ed., in press). Vegetation map of Africa (1:5.000.000). Paris, U.N.E.S.C.O. Index Accepted names are given in roman and synonyms in italic; new names and principal page references are in bold. An asterisk (*) denotes a figure. Acrolejeunea emergens (Mitten) Stephani var. Asterella abyssinica (Gottsche) Grolle ex confertissima (Stephani) Gradst. 224, 281 Vanden Berghen 226 emergens var. emergens 218, 223, 224 pycnoclada (Taylor) Schiffner 224 Alobiella chevalieri Stephani 243 Aneura pinguis (L.) Dumort. 224 travisiana Pearson 278 Anthoceros buettneri Stephani 219, 224 crispulus auct. 224 Bazzania decrescens (Lehm. & Lindenb.) Trev. St. Léon subsp. molleri (Stephani) E. Jones 217,226, 230, 250, 275 Brachiolejeunea camerunensis E. Jones & Vanden Berghen 227 nigra (Stephani) Stephani 265 tristis Stephani 217, 226 mandonii Stephani 224 Arachniopsis coactilis Spruce 225 Calypogeia fissa (L.) Raddi 227, 282 diacantha (Montagne) M. Howe 218, 224, longifolia Stephani 227, 283 262 Caudalejeunea africana (Stephani) Schiffner 227, Archilejeunea abbreviata (Mitten) Vanden 235 Berghen 225, 226 dusenii Stephani 227 africana Stephani 225, 226, 277 hanningtonii (Mitten) Schiffner 227 apiculata Pearson 264 lehmanniana (Gottsche) A. W. Evans 228 autoica Vanden Berghen 225 tricarinata E. Jones 228 linguaefolia Stephani 226 Cephalozia fissa Stephani 218, 225, 228, 262 HEPATICS OF SIERRA LEONE & GHANA Cephaloziella abyssinica Gola 243 atroviridis (T. Sim) S. Arnell 243 retusa E. Jones 243 ugandica E. Jones 243 vaginans Stephani 228 vaginans var. camerunensis S. Arnel] 228 Ceratolejeunea belangeriana (Gottsche) Stephani 230 beninensis E. Jones & Vanden Berghen 228, 229. calabariensis Stephani 215, 228, 229 cornuta (Lindenb.) Schiffner 230 cornutissima Stephani 229 diversicornua Stephani 228, 229 jungneri Stephani 229 maritima (Spruce) Stephani 230 saxbyi Pearson 215, 229 variabilis (Lindenb.) Schiffner 230 zenkeri Stephani 229 Chandonanthus hirtellus (Fried. Weber) Mitten 217, 230 Cheilolejeunea africana (Stephani) E. Jones 232 aneogyna (Spruce) A. W. Evans 231 decursiva (Sande Lacoste) R. M. Schuster 230 exinnovata E. Jones 231 inflata Stephani 231 intertexta (Lindenb.) Stephani 218, 231, 232, 255,211 newtonii Stephani ex Schiffner 231, 232, 255 principensis Stephani ex Paris 232 rigidula (Montagne) R. M. Schuster 232 serpentina (Mitten) Mizutani 232, 233, 255, 267 silvestris (Gottsche) E. Jones 232 surrepens (Mitten) E. Jones 232, 235, 247 tisserantii Vanden Berghen & Jovet-Ast 230 trifaria (Reinw., Blume & Nees) Mizutani 219, 229; 23a;215 Chiloscyphus dubius Gottsche 233, 262 Cololejeunea adhaesiva (Mitten) R. M. Schuster 234 africana (Stephani) R. M. Schuster 234 androphylla E. Jones 234 appressa (A. W. Evans) Benedix 234 auriculata (E. Jones) R. M. Schuster 234 bolombensis (Stephani) Vanden Berghen 234 calcarata E. Jones 215, 235, 236*, 237* cardiocarpa (Montagne) R. M. Schuster 237 cornuta E. Jones 218, 228, 237 crenatiflora (Stephani) Vanden Berghen 235, 238 cuneifolia Stephani ex A. W. Evans 236, 238, 241 dentata (E. Jones) R. M. Schuster 219, 235, 239 dissita E. Jones 240 duvignaudii E. Jones 238 elegans (Stephani) E. Jones 239 floccosa (Lehm. & Lindenb.) Schiffner 239 hyalino-marginata (Nees ex Montagne) Grolle 239 lanceolata E. Jones 239 leloutrei (E. Jones) R. M. Schuster 239 malanjae Stephani 236 minutissima (Smith) Schiffner 240 minutissima subsp. myriocarpa (Nees & Montagne) R. M. Schuster 240 minutissima subsp. utriculifera Vanden Berghen 240 myriantha (Herzog) S. Arnell 240 myriocarpa (Nees & Montagne) A. W. Evans 240 nigerica (E. Jones) R. M. Schuster 235, 240 peraffinis (Schiffner) Schiffner var. elegans Benedix 241 plagiochiliana P. Tixier 236 pusilla Stephani var. obtusifolia E. Jones 241 pusilla var. pusilla 241 zenkeri (Stephani) E. Jones 238, 241 Colura calyptrifolia (Hook.) Dumort. subsp. tenuicornis (A. W. Evans) Vanden Berghen 242 digitalis (Mitten) Stephani 235, 242 dusenii (Stephani) Stephani 242 tenuicornis (A. W. Evans) Stephani 242 Conoscyphus trapezioides (Sande Lacoste) Schiffner 217, 230, 242 Cyathodium africanum Mitten 242 cavernarum Kunze 243 Cylindrocolea atroviridis (T. Sim) Vana 243 chevalieri (Stephani) R. M. Schuster 243 gittinsii (E. Jones) R. M. Schuster 243 madagascariensis (Stephani) R. M. Schuster 243 ugandica (E. Jones) R. M. Schuster 243 Dicranolejeunea madagascariensis (Gottsche) Stephani var. madagascariensis 217, 243 Diplasiolejeunea aulae E. Jones 244 brachyclada A. W. Evans 244 cavifolia Stephani 244 cornuta Stephani 235, 244* harpaphylla Stephani 244, 247 incurvata Jovet-Ast & P. Tixier 244 Drepanolejeunea ankasica E. Jones 215, 245*, 246* clavicornis Stephani 248 claviformis Stephani 247, 248 cultrella (Mitten) Stephani 247 friesii Vanden Berghen 247* Dumortiera hirsuta (Sw.) Nees 248 Eulejeunea microclada Pearson 254, 255 camerunensis Stephani 254 Euosmolejeunea brachytoma (Gottsche) Stephani 233 Fimbriaria abyssinica Gottsche 226 287 288 E. W. JONES & A. J. HARRINGTON Fossombronia crozalsii Corbiére 249 husnotii Corbiére 217, 248 occidento-africana S. Arnell 218, 248 Frullania africana Stephani 249 apicalis Mitten var. apicalis 250, 251 apiculata (Reinw., Blume & Nees) Dumort. 250 arecae (Sprengel) Gottsche 217, 250 borgenii Pearson 252 depressa Mitten 217, 251 diptera (Lehm. & Lindenb.) Gottsche, Lindenb. & Nees 251 ecklonii (Sprengel) Gottsche & Lindenb. 250 eplicata Stephani 251 ericoides (Nees) Montagne 218, 251, 252 nodulosa (Reinw., Blume & Nees) Nees 252 obscurifolia Mitten 252 purpurea Stephani 252 spongiosa Stephani 252 squarrosa (Reinw., Blume & Nees) Dumort. Pails trinervis (Lehm & Lindenb.) Gottsche, Lindenb. & Nees 252 Gongylanthus ericetorum (Raddi) Nees 217, 253 richardsii E. Jones 217, 253 Jungermannia borgenii Gottsche ex Pearson 253 dusenii (Stephani) Stephani 253 Lejeunea abbreviata Mitten 225 acuta Mitten 217 autoica R. M. Schuster 253, 255 brenanii E. Jones 253, 259 caespitosa Lindenb. 231, 254, 255, 259, 277 camerunensis (Stephani) E. Jones 254, 268, 277 conformis Nees & Montagne 283 confusa E. Jones 253, 255, 283 corbieri Stephani 257 eckloniana Lindenb. 231, 254, 255, 268 eplicata Stephani 215, 259, 260 flava (Sw.) Nees 235, 253 flava subsp. flava 256 flava subsp. tabularis (Sprengel) S. Arnell 256 grossecristata (Stephani) E. Jones 215, 256 ibadana A. Harrington & E. Jones 257 jungneri Stephani 255 kamerunensis (Stephani) Vanden Berghen 254, 268 lomana E. Jones 215, 257, 258* longifissa Stephani 254 longirostris (Stephani) E. Jones 256 papilionacea (Stephani) E. Jones 215, 259 pililoba Spruce 260 ramosissima Stephani 215, 217, 254, 259 saccatiloba Stephani 254 setacea Stephani 254, 257, 260 triquetra Mitten 215, 284 tuberculiflora E. Jones ex Pécs 254 ulicina (Taylor) Gottsche, Lindenb. & Nees subsp. africana (Stephani) Vanden Berghen 267 Leptocolea africana (Stephani) E. Jones 234 bolombensis (Stephani) E. Jones 234 cardiocarpa (Montagne) A. W. Evans 237 crenatiflora Stephani 238 cuneifolia (Stephani ex A. W. Evans) A. W. Evans 238 dentata E. Jones 239 leloutrei E. Jones 239 nigerica E. Jones 240 Leptolejeunea astroidea (Mitten) Stephani 260, 261 maculata (Mitten) Schiffner 260, 261 quintasii Stephani 261 thomeensis (Stephani) Stephani 260 truncatiloba Stephani 260 Lophocolea concreta Montagne 261, 262 congoana Stephani 262 devexa Mitten 242 difformis Nees 261 martiana Nees subsp. newtonii (Stephani) R. M. Schuster 225, 233, 262 molleri Stephani 261 newtonii Stephani 262 subrotunda Mitten 261 Lopholejeunea abortiva (Mitten) Stephani 262 fragilis Stephani 262, 263 jonesii Vanden Berghen 227, 263 subfusca (Nees) Schiffner 263, 277 Marchantia chevalieri Stephani ex Bonner 264 parviloba Stephani 263, 264 planiloba Stephani 263, 264 wilmsii Stephani 264 Marchesinia excavata (Mitten) Stephani 229, 264, 280 moelleriana Pearson 217, 227, 264 Mastigolejeunea auriculata (Wilson) Schiffner 217, 218, 251, 265 carinata (Mitten) Stephani 265 florea (Mitten) Paris 265 nigra Stephani 217, 265 tarkwana Pearson 215, 284 turgida Stephani 266 Metzgeria camerunensis Stephani 266 conjugata Lindb. 266 saxbyi Pearson 266 thomeensis Stephani 266, 267 Microlejeunea africana Stephani 267, 277 ankasica E. Jones 268 jungneri Stephani 255 kamerunensis Stephani 268 ulicina (Taylor) A. W. Evans 267 Notothylas angolensis Stephani 268 decurva (Mitten) Stephani 268, 279 indica Kashyap 268 HEPATICS OF SIERRA LEONE & GHANA 289 Odontolejeunea tortuosa (Lehm. & Lindenb.) Stephani 217, 269 Pallavicinia lyellii (Hook.) Carruth. 283 Phaeoceros laevis (L.) Proskauer subsp. carolinianus (Michaux) Proskauer 269 Physocolea elegans Stephani 239 Plagiochasma eximium (Schiffner) Stephani 217, 269 Plagiochila africana Stephani 218, 269 barteri Mitten 269, 272 divergens Stephani var. divergens 270 divergens var. myriocarpa (Pearson) E . Jones 270 fusifera Taylor 270, 271*, 272, 273, 274 integerrima Stephani 270 moenkemeyeri Stephani 270, 271* , 274 neckeroidea Mitten 272, 273 pectinata (Willd.) Lindenb. 219, 272 pinniflora Stephani 272 praemorsa Stephani 270, 273, 274, 275, 276 salvadorica Stephani 272, 273 strictifolia Stephani 270, 271, 273, 274 Porella subdentata (Mitten) E. Jones 217 subdentata var. camerunensis E. Jones 274 subdentata var. subdentata 254, 274 Prionolejeunea aberrans Stephani 215, 259 grata (Gottsche) Schiffner 219, 275 serrula (Mitten) Stephani 275 Ptychanthus striatus (Lehm. & Lindenb.) Nees 217,275 Ptychocoleus confertissimus (Stephani) Stephani 224 emergens (Mitten) Stephani 223 molleri (Stephani) Stephani 281 occultus (Stephani) Stephani 280 pappeanus (Nees) Stephani 280 Pycnolejeunea contigua (Nees) Grolle 275 Radula appressa Mitten 275 boryana (Fried. Weber) Nees 219, 250, 276 flaccida Lindenb. & Gottsche 276, 277 guineensis Stephani 276 holstiana Stephani 219, 276 javanica Gottsche 276 stenocalyx Montagne 219, 276, 277 Rectolejeunea arnellii E. Jones 277 brittoniae A. W. Evans 277 rhodesiae (T. Sim) S. Arnell sensu S. Arnell 277 Riccardia angusticosta (Stephani) Grolle 278 erosa (Stephani) E. Jones 278 incurvata Lindb. 279 limbata (Stephani) E. Jones 278, 282 longispica (Stephani) E. Jones 278 multifida (L.) Gray 278 pinguis (L.) Gray 224 stephanii (Bescher.) E. Jones 278 Riccia fluitans L. 279 lanceolata Stephani 279 membranacea Gottsche & Lindenb. 279 moenkemeyeri Stephani 215, 279 nigrosquamata E. Jones 280 rhodesiae S. Arnell 219, 280 undulata S. Arnell 215, 279, 280 Ricciocarpos natans (L.) Corda 219, 280 Schiffneriolejeunea fragilis Gradst. & E. Jones 280, 281 occulta (Stephani) Gradst. 280 pappeana (Nees) Gradst. 280 polycarpa (Nees) Gradst. 263, 281 Sprucella succida (Mitten) Stephani 281, 284 Stictolejeunea africana Vanden Berghen 281 balfourii (Mitten) E. Jones 281 Strepsilejeunea brevifissa (Gottsche) Stephani 217, 282 Symphyogyna brasiliensis Nees & Montagne 283 lehmanniana Montagne & Nees 282 volkensii Stephani 282 Targionia hypophylla L. 218, 283 Taxilejeunea conformis (Nees & Montagne) Stephani 256, 283 nicholsonii Pearson 215, 229, 256, 257 pulchriflora Pearson 219, 283, 284 Telaranea nematodes (Gottsche ex Austin) M. Howe 283 Thysananthus spathulistipus (Reinw., Blume & Nees) Lindenb. 215, 218, 229, 284 Trachylejeunea serrulata Stephani 218, 228, 284 Tylimanthus ruwenzorensis S. Arnell 284 ~— British Museum (Natural History) Seaweeds of the British Isles The synthesis of many years’ research carried out by the British Museum (Natural History) and the British Phycological Society, this is the second in a series of books that will be published under this title covering all the British and the majority of northern Atlantic seaweeds. Volume 1 Rhodophyta Part 2A Cryptonemiales (sensu stricto), Palmariales, Rhodyme- niales. Linda M. Irvine Following the style and format laid down in Part 1, this book deals with the species attributed to the orders Cryptonemiales, Palmariales and Rhodymeniales. Each species description incorporates notes on ecology and distribution and is supported by one or more line illustrations. Keys to aid identification are also included. When complete, this title will be the standard work of reference in its field; it will provide students and researchers with a digest of the most up-to-date and compre- hensive information available on the marine algae of Great Britain and Ireland which will be indispensable throughout the north Atlantic region and beyond. ISBN 0 565 00871 4 128 pagesillustrated £13.00. Titles to be published in Volume 11 The algae of Lightfoot’s Flora scotica. By Peter S. Dixon. A taxonomic study of the lichen genus Micarea in Europe. By B. J. Coppins. The hepatics of Sierra Leone and Ghana. By E. W. Jones and A. J. Harrington. Studies in the Corallinaceae with special reference to Fosliella and Pneophyllum in the British Isles. By Y. M. Chamberlain. Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, Dorchester Bulletin of the Nc British Museum (Natural Histors) Studies in the Corallinaceae with special reference to Fosliella and Pneophyllum in the British Isles Y.M. Chamberlain Botany series Vol 11 No 4 29 December 1983 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), 1983 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 11 No 4 pp 291-463 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 29 December 1983 + GENER, = 2 JAN LIBRAR} PAL W\S > I> 4/ Studies in the Corallinaceae with special reference t Fosliella and Pneophyllum in the British Isles Y. M. Chamberlain* The Marine Laboratory, Portsmouth Polytechnic, Ferry Road, Hayling Island, PO11 0DG Contents MPN R MMS 5p c es cotbevtN As esaedes cue va veiaae vats a Uauiny vie veamy elporindulennbyN erOne ye Nee se a dea Bess 292 RTPA P ORICON conn ce csocr anc eiceca saree nc uaanecudn nd ipais Mo Meei ees annedee ream enh COs Tad ndeyn a Moonie 293 PRUIEG VIAUNOTAR is Sc ste ca cs cao cpa osyaks kash Sie ip odoen ey sexu sate ye cpveteanunreg doi sal Uoaconbimaaeneee 294 Weer Am is UNC TINE UENCNCS sc 200 sunk ao Foy dade dels peaMoms dk yoo eur ndeetandueh tas ank pocerue sou 295 BRE OMTNC HOE 555s os ie ie wah cate ak we Ped leharh ote Hews Cua phe ete cue ties ena oaae tee rook 295 MMU IAM OSCODY 0020 cas ee as sce oe snes aoe cae ss cesta earns saa Saka enas a lagad den meee 295 DOANE CLECIION MUCEOSCODY. << foo soc cnc dedin ve rcuiecn es cacsouece ss eae ancaciecs Supe 295 SRGUEH TAI NCIC OOSELVANONS 6200 ose e554 ccvvatin es ves vicers uc nosbash caviuncvanlaatueronnieere 296 BBDOPAUHY CUNT Oise case aie ond are bee ances nodet etued ab useee Usb venus savslnbuesetactuneees 296 Le PSLOLY OF eG OF AUIMACEAG voce a cocassiscsasconesscadeatacedy psi csosstradiattevaheesya eaceaie? 296 ae CHIME ST DOCEAR sedan dc ens topped secaGe «woe ore dewesecmein ea ang adse ante aun eTe 296 He SAM DLO CHUISLOSG CG OLIACCAC: 525. c esas fo ack sie iss venissashn isu oeeensaeeiasteeataentne es 297 EIRP nee cere kee dees on eaacate Lovonmnsea aids smenawior aus eaeemror racer aanest 297 BSCR const Uacras sa Seer atuce wemevea age ta sa cseesents decd sn sans ey eae Re NORMS aU 297 BN Co PME OUL his vcr el gaat capa u dsc nen eh on tava iumenerdadtc onus eee nes ree 298 Be coke WRONG cies dad cate ss eens eee te cate dirs atten vIn Ge eign unison Body ey dee < oa ens 298 WEA AMEE WSS cot ch cka gun eeaer asta sore pay ae ndduioe Lanne eeny core SoU aI NC xt ne cost T ESTE 301 Bs ANG Fate ROWAIN Fos aeiiiics cove cad on tes ope de socteek ve oee eee waa eee aETUT ERIS Oe 301 By hI Aeis ORME OEN ys sate htt ec uahen tte cok nda Stok ohonus ko uecetenean ada ey nasety reamed 302 OE GS Tas rs REINER ose RAST HOPE Meas 2h PR LEAM NRA EE NOR AMIR EEN i RES een ema r 308 Re INC ies OP aso cea re eeu enamine vai mncten Lava tes mie aerneensed sees 308 DVDR ORE 5 oe cay Face Poti n oats esa See nee rae ete eseaee ney eae ee ase 310 RLS Ae PARGUC TOE ctor yya' cite raisin usin Paes tte nN uae ees Cosa een Seewtonse oe eek ees 310 OIE NO oo Pe nO et eR NA re SE Rie eM ry Meter re ec aa ren tee oie WA. pete ana i. Rs Masonite ics tacs coz ses seaskcsastevcsteie yesnssuanisenaaaesuaeieesoes 312 US PCAC MAG Ee PROV oy oeg cp oesyaces eck vay eee fei ngs te bamea tape att niece eoueed dieeee 318 Ly RIOR RER EE Biecorueh Sepeaey Soren ere in carte Ye rh err ee ELE Hey VE YY Portes ge eee ye cee 319 EEE CSR ORR EE ae OPe PSB SAUD Sa ce Dpe eter Pa RCRA, SATAY geese apm Ain eRe are 319 pe re Tie AaB cP OP Ut eA PAE De Pry NEE SUS OE Ue PATE TOTNES RAR oe 319 Ne TORE SETAE Le I RBI ae OR Ba Oar OR ca ne ea Or ge ec er fy ren ate eo PA 319 hoe ns at bral) 1 LET CMI So a RP See OEE ects pte vert AER RE ARORA OANA Sr SR EPL E 320 PEMA PRA ACMMAAOMC TREE S100, os aetgnns oie roc t ha. ry eea et Ainae coay Ve eweueeerien aed s ce reNenreisde 320 PAP OOMORCISOIIE: hoot 5 ener dani ay ue eassou tela teases sor adareuneadaet ae vata veab eee meal ppm o's Sega Dae amen Aree ED, ttt Our Oia Astana pA eae enny RRRE bays SRA SiR ry ear 323 eG CON ee oiccascracyins oiuue car voce sce stem nan eke cen ion tude sewase acne decdeu nee vamaseulicess 323 PER OUVE VOTIMICMIOEE 5555 ei Csccno bys bved canes wee sus eanuer esas oe eeseewe outa oe uaiekontee a 323 SeeAMBS ENE MOINS coca ae chase hd eet octal deansmeneu ene conceed paicgaiy sae vance esemeenu easier ener taeanTes 323 Gametangia and the carposporangial phase .................:.cseceeceeeeeeceeeeeeeeeeeeees 327 Me PME IS BIRD UIST AIGA aoc nico ccs sce saeaat ocdu cies sh ccsesacaey etre ee ateeed 328 ESE ESSERE Os ete ET Ae eT ON REE Pare Pen DONA Ue Ee CORN ARES ee 328 NT ESE DEN AAS EES OOD LID EOE EO Ta EPEAT SN ATT IE SOD R Se De 329 Ee eeeCetO ON ie COTAINACERE 22. <6 Lyles ces centecicervonsc¢csenecadedvervedeswuvgonce¥s 330 sassy Gn Postielia and Prheophy lus «3... ices cccsscosesecsenssecuscnsascascdiséectansssecesees gaz Ca SEE LSE REY Ae ONIN HORE at PREIER Fete dese en eR NIRA oat SO eae RI ORT COP REY, * Mrs Y. M. Butler Bull. Br. Mus. nat. Hist. (Bot.) 11 (4): 291-463 Issued 29 December 1983 292 Y. M. CHAMBERLAIN The Organisation OL Species Gescr ny tONs os ice occ c ea rcestcnn cig: scsvsys vavekoneeshony eos sees 332 ATRTOGUCTIOR cae csocgse snc veecceeonsaccie ys re4ha4iraearnvastigenieraupaveasy bacescusedeCeverses 332 FFaxONOMC CALE SOMES ANG NOMENCIANUINS 5.225 dein sone e's sknchx00ssebpasdecovunwaeeaelte rach 332 Ey iasnosis BN COSCHINION och svar acrncetaes cerasedeni anes kereveeasictsaueebiosavicny stad, 333 IWMCASUTO IONS science So ctigs ew vos see ec nna eens Oeics eee Culpa Uses bay ecs Comey Ree ES 333 VeESELAUIVE CHATACLENS 00 /2522ia bo anus fed sal odepee as tein tun eueds erat eabeun rence tnercs hun ee 333 REDIOCUCHIVE CRATACIEES cfivsiviucc cs ey care ciduinss cebh eo sacess wren naceduseeedecesan decckaes 335 Culture methods for separating epilithic species of Pneophyllum ..............0..00065 336 Distpibutionand ther Teatrves esos cys iexccs tec. svwcavees sy canes sv eas uci any cuenstaseses 336 SSUTIEATY co7 toed te cen ee eels Wet scenes vonavanacnc teal rerees sound ceeebias a vuvecr rans ae re 336 SUISELMEA So cence Cu nencntecies saaee uel oie cpa beea sa DaES DR anWse rapbavag Raven adeeb sounaa oh hed veaerareeces 336 COONOTIC ROY’ 355.205; s eecsetecs asians ecwaoackerpsdasede%sanceanuaczisecsiaaa des adeasaateqoncyeedtan és 338 TS OSIENG INE TIQWO.< 90. dar eassssdacucaacer te soseeee leis oe aes ear ace eens 340 la. Fosliella farinosa (Lamouroux) M. Howe f. farinosa.............0.ccseceeeeeneeeeees 343 1b. F. farinosaf. callithamnioides (Foslie) Y. Chamberlain ....................0..00c00e 351 EE APHECOpRYV AN NG ot ooes Gack ia tice d sete esi visarseg asc sasanceta os Ceveuse tea baton caseerkeen tty 352 Key to species of Preophyllum in the British Isles 3. «sis wvcccascssedsesesasescetvesse sees 355 Ls SE REOPAY HUTT TARTE UIT e sce teave ence dentcise sata sonavaddiwed tae ti eee cases heaven 356 2. P léejolish tRosanoll) Y. CRAMberanni Gees co525 horns vcsa te onece ine vesvasecaniun 358 De, th FOSANO/ FEY CAIN OCT IAIN coo caendct ais sss sks anv oes a blanshedovde eet i cnecoeeoases 367 A] EP. lenttation (Postie yy « Chamber laie o.o7,s0ss0tsees sass scecseves Livan eueatane ares 374 5a. P. confervicolum (Kiitz.) Y. Chamberlain f. confervicolum ................060005 383 5b. P. confervicolum f. minutulum (Foslie) Y. Chamberlain ..................::000008 392 GLP miicrosporiut (ROSENV.) X, CNAMDETI AI: 0 .cc2. nd dacecegrssccnycnyeoeo eee ine 395 Tih COMCOULIN 1. KS RADIDOEIAIN oc en co sea tine bast one ce ren aerate ae eae aaa 402 8. P.myriocarpum (P. Crouan & H. Crouan) Y. Chamberlain ..................... 410 9. Po lobescens ¥: Chamberlain: occ) voce ae iotiue: serene aecenes sede teveondedscwencthens 420 10.4, plarivalidian YC ami Dera ss ss2:5 caacisnese nics segs cdavee teateane tone ssc bexense eae 428 11... P. zonale(P. Crouan & H. Crouan) Y. Chamberlain. ......33.5..04.-<:255005 NE uy - WAAAY WARN MALL AATTATS bg N325a% IA is | Ad nat. del.ethth. S Rosanoff. [up Becquet a Paris. lOmm 311 312 Y. M. CHAMBERLAIN Florideophyceae. I examined material of this species at both Concarneau (CO) and Paris (PC) and found it to be the creeping form Pneophyllum confervicolum f. minutulum (q.v.). M. A. Howe Howe considered that Melobesia had to be typified by M. membranacea (Esper) Lamouroux and was, therefore, a lithothamnioid genus with many-pored roofs on the sporangial conceptacles. A new generic name had to be found for the genus of simple crustose corallines, such as M. farinosa, with secondary cell fusions and sporangial conceptacle roofs with a single pore. Howe (1920) proposed the name Fosliella to designate Foslie’s Eumelobesia (Foslie, 1905b) which (in that publication) contained only M. farinosa sensu Lamouroux. However, although Foslie restricted Eumelobesia to monostromatic plants bearing trichocytes, Howe also included M. lejolisii and M. chamaedoris, both of which were considered to be without trichocytes, while M. chamaedoris was also ‘polystromatic’. It is proposed, therefore, that Howe’s genus Fosliella be emended to include only species with secondary lateral cell fusions, single-pored sporangial conceptacles, and terminal hypothallial trichocytes. As has been discussed, species with intercalary or no trichocytes are classified in Pneophyllum Kitz. W. A. Setchell and L. R. Mason Setchell & Mason (1943) investigated an American alga which had been identified as Dermato- lithon pustulatum, and concluded that it was not a Dermatolithon at all; they reclassified it as Heteroderma nicholsii. This led them to redefine the genus Heteroderma Foslie to include all oligostromatic species (i.e. with a thin thallus), without either trichocytes or a palisade hypothallus and with single-pored conceptacles. They designated H. subtilissima (Foslie) Foslie as the type species, without actually seeing a specimen. Recent examination of the type specimen from Herb. Foslie (TRH) shows it to be a minute Pneophyllum growing on Corallina from Papua. Mason (1953) did not propose any further name changes although she suggested that the generic name Fosliella be restricted to species with trichocytes whether terminal or intercalary. She gave a clear account of the nomenclatural problems connected with the simple crustose Fig.5 From Rosanoff (1866). PLANCHE IV. Fig. 1.— Une partie du bord de la fronde représentée dans la fig. 10 de la pl. III. (Grossissement de 320 diamétres). Fig. 2. — Jeune fronde du Melobesia pustulata. (Gross. de 40 diam.). Fig. 3. — Etat plus développé de la méme espéce. (Gross. de 40 diam.). Fig. 4. - Frondes de Melob. macrocarpa (sur Phyllophora rubens). On voit que la plupart des frondules réniformes ont leur incisure dirigée du cété inférieur de la fronde du substratum. (Grandeur naturelle). Fig. 5. - Coupe perpendiculaire au substratum d’une fronde de Melob. macrocarpa. (Gross. de 350 diam.). Fig. 6. — Une pareille coupe, menée tangentiellement. (Gross. de 350 diam.). Fig. 7. - Une partie de la membrane cellulaire du Melob. macrocarpa, montrant une ponctuation réguliére. (Gross. de 1050 diam.). Fig. 8.- Trois cellules frondales du M. macrocarpa, vues d’en haut. Leurs parois supérieures sont trés nettement ponctuées. L’incrustation trés forte et la couche de chaux carbonatée qui tapisse la face intérieure des cellules frondales, rend leurs cavités plus ou moins arrondies. (Gross. de 1050 diam). Fig. 9. - Coupe verticale et centrale d’un cystocarpe du M. coronata, (sur Pollexfenia pedicellata). (Gross. de 200 diam.). Fig. 10. - Coupe verticale et centrale d’un cystocarpe de M. farinosa. Fig. 11. — Etat jeune d’un sporange de M. macrocarpa, entouré des paraphyses. La cellule-mére est remplie d’un contenu incolore plein de vacuoles, et on voit le commencement de la division transversale. (Gross. de 600 diam.). Fig. 12. — Etat plus avancé du sporange; le contenu est coloré en rose brunatre clair; les petites vacuoles ont disparu, et on ne voit que la disposition du contenu granuleux en réseau; Dans le voisinage de la cloison, qui se forme, se trouve (dans chaque loge) une vacuole elliptique, avec un nucleus excentrique. (Gross. de 600 diam.). Fig. 13. - Un sporange, qui s’est déja divisé en deux loges dont le contenu est granuleux et rose intense. (Gross. de 600 diam.). Fig. 14. — Plusieurs paraphyses d’un cystocarpe de Melob. macrocarpa; leur sommet est €paissi, mamilliforme et leur contenu offre une disposition en spirale. Deux des paraphyses offrent des trous a leurs sommets. (Gross. de 600 diam.). Fig. 15. - Coupe verticale et centrale d’un cystocarpe de Melob. macrocarpa. (Gross. de 200diam.). Fig. 16.—a, b. Sporanges, quicommencment as’élever du fond d’un cstocarpe. Fig. 17.— Sporanges du M. macrocarpa, pris dans un échantillon desséché; a—vus de face, b— vus de cété. Fig. 18.—a, b, deux sporanges de la méme espéce extraits d’un échantillon desséché. Fig. 19. - Une partie de la fronde désincrustée du M. macrocarpa; les cellules corticales sont trés apparentes. Fig. 20.— Formation des paraphyses dans le Melob. macrocarpa; l\es cloisons inclinées transversales commencent a se dédoubler, les longues cellules montrent la ponctuation réguliére: a avant, et b aprés la désincrustation. Les cellules de la couche, qui sert de base aux futures paraphyses, ont les, parois verticales trés €paisses dans leur moitié inférieure et minces dans la moitié supérieure. La partie mince correspond, selon moi, al’étendue dans laquelle s’est fait l’accroissement dans le sens vertical. (Gross. de 600 diam.). FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 313 Mém. de la Soc. Imp. des sc.nat.de Cherb. T. XII. PL_IV. x \ i ai erect Tee, ex ake) See eee suki gy Mi iV y ee ANG: Adnat. del et Lth S.Rosanoff. ee imp Becquet a Paris. 314 Y. M. CHAMBERLAIN Mem. de la Soc.imp.des sc.natde Cherb. T. XII. 2 1 “PORS SE SSES ern Shion pee. Oe 3 SN eeeeee a Ad nat. del. et Lith. S. Risa 1Omm Fig.6 From Rosanoff (1866). PLANCHE VII. Fig. 9. — La ligne de rencontre de deux ramules de la fronde du Melobesia Lejolisii. du Melob. Lejolisii. Fig. 12. -— Cystocarpe atrophié du Melobesia farinosa. Inp Becguet a Farts. Fig. 10 et 11. — Partie des frondes FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES fa Table 2 To show names under which specimens pertaining to Pneophyllum rosanoffii, P. lejolisii, and P. concollum have been published. (Initials indicate herbaria examined.) P. rosanoffii (with ostiolar P. concollum (with ostiolar P. lejolisii (with flat-topped Publication filaments) conceptacles) collar) Areschoug, Melobesia membranacea 1847, 1852 CHE PC Areschoug, 1875 M. lejolisii CHE PC Kiitzing, 1869 M. farinosa ie Harvey, 1849, 1851 M. membranacea? Le Jolis, 1863 M. farinosa CHE Rosanoff, 1866 Weber van Bosse, 1886 Lloyd no. 339 (1886) M. lejolisii pro parte CHE Johnson and Hensman M. lejolisii (1899) Foslie (1905a) Rosenvinge (1917) Suneson (1937, 1943) M. lejolisii f. typica M. lejolisii pro parte c M. lejolisii pro parte M. lejolisii pro parte? CHE M. rosea CHE M. lejolisii L M. membranacea CO M. lejolisii f. typica M. lejolisii pro parte Cc M. lejolisii Table 3 Foslie’s principal systems of classifying simple crustose corallines.* Genus Subgenus Species FOSLIE 1898a Systematical survey of the Lithothamnia Melobesia Lamour. emend Single orifice, with paraphyses Choreonema Schm. emend. Single orifice, no paraphyses Lithothamnion Phil. emend. Numerous orifices Heteroderma Fosl. mscr. Thallus crust-like Chaetolithon Fosl. mscr. Parasitic, with rhizoids Eulithothamnion Fosl. M. pustulata Lamour. C. Notarisii (Duf.) Fosl. mscr. C. Thuretii (Born.) Schm. L. corticiformis (Kitz.) Fosl. mscr. FOSLIE 18985 List of species of the Lithothamnia Melobesia Lamour. emend. (Gen. Choreonema in 1898a) Single orifice, no paraphyses * Eumelobesia Fosl. mscr. Monostromatic all authorities as written by Foslie. M. farinosa Lamour. M. callithamnioides Cr. M. coronata Rosan. M.confinis Cr. M. confervoides (Kitz.) M. hildenbrandtioides (Cr.) M. zonale (Cr.) M. coccinea (Cr.) 316 Table 3 — cont. Y. M. CHAMBERLAIN Genus Subgenus Species Choreonema Schm. Parasitic Dermatolithon Fosl. mscr. (Gen. Melobesia in 1898a) Single orifice, with paraphyses Lithothamnion Phil. emend. Heteroderma Fosl. Polystromatic Epilithon (Heydr.) Fosl. mscr. Monostromatic M. inaequilaterata Solms. M. Novae Zelandiae Heydr. M. rosea Kitz. M. Notarisii Duf. M. prototypus Fosl. M. Laminariae Cr. M. Corallinae Cr. M.? myriocarpa Fosl. M. Cystosirae Hauck. M.? rhizomae (Heydr.) C. Thuretii (Born.) Schm. D. pustulatum (Lamour.) Fosl. mscr. f. bispora Fosl. mscr. (M. pustulata Lamour. s.s.) f. macrocarpa (Rosan.) Fosl. mscr. D. Lejolisii (Rosan.) Fosl.mscr. D. hapalidioides (Cr.) Fosl. mscr. L. membranaceum (Esper) Fosl. mscr. L. corticiforme (Kitz.) Fosl. FOSLIE 19006 Revised systematical survey of the Melobesieae Melobesia Lamour. emend. Conceptacles superficial or slightly immersed or conical, disc plain and connected to roof only at first Lithophyllum Phil. emend. Conceptacles scarcely raised, sporangial disc overarched, initially attached to roof then freed, roof usually finally decorticated Dermatolithon Fosl. Conceptacles sub-immersed or conical, disc plain, club-shaped free paraphyses Eumelobesia Fosl. Monostromatic Heteroderma Fosl. Polystromatic Carpolithon Fosl. Conceptacles scarcely raised, roof finally decorticated, disc less overarched Lepidomorphum Fosl. Conceptacles immersed or sub-prominent-convex, roof finally decorticated, disc + overarched and connected to roof M. farinosa Lamour. M. callithamnioides Falkbg. nec. Crn. (M. farinosa f.?) M. lejolisii Rosan. M. confervicola (Kitz.) Fosl. M. caspica Fosl. M.? Novae Zelandiae Heydr. M. Corallinae Solms (nec. Crn.?) M. coronata Rosan. M. canescens Fosl. M. zonalis (Crn.) Fosl. M.? Cystoseirae Hauck L. decipiens Fosl. L. discoideum Fosl. L. zostericolum Fosl. f. tenuis Fosl. f. mediocris Fosl. L. papillosum (Zan.) Fosl. D. pustulatum (Lamoutr.) Fos. f. typica f. crinita Mob. D. macrocarpum (Rosan.) Fosl. f. faeroénsis Fosl. FOSLIELLA AND PHEOPHYLLUM IN THE BRITISH ISLES 317 f. Laminariae (Crn.) Fosl. D. hapalidioides (Crn.) Fost. f. typica (Inc. Melob. simulans Crn.) f. confinis (Crn.) Fosl. D.? adplicitum Fosl. (D. hapalidioides f.?) D.? prototypus Fosl. Lithothamnion Phil. emend. Epilithon (Heydr.) Fosl. L. membranaceum (Esper) Fosl. L. corticiforme (Kiitz.) Fosl. FOSLIE 1905a Remarks on northern Lithothamnia FOSLIE 1905b New Lithothamnia and systematical remarks (19055 additions in brackets) Melobesia Lamour. Eumelobesia Fosl. M. farinosa Lamour. monostromatic, f. typica epithallium present, f. borealis Fosl. mscr. heterocysts present f. callithamnioides (Falkbg.) Fosl. (f. mauritiana Fos. mscr.) Heteroderma Fosl. M. lejolisii Rosan. monostromatic, f. typica epithallium present or f. limitata Fos. mscr. absent, M. minutula Fosl. heterocysts absent f. typica f. lacunosa Fosl. mscr. (M. Cymodoceae Fosl.) (M. triplex Heydr. = M. Cymodoceae Heydr.?) (M. coronata Rosan.) (M. rugulosa Setch. et Fosl.) (M. subtilissima Fosl.) (M. Novae Zelandiae Heydr.?) Lithophyllum Phil. Dermatolithon Fosl. ee Ls ba L. pustulatum (Lamour.) Fos. f. australis Fosl. mscr. f. intermedia Fosl. mscr. f. macrocarpa (Rosan.) Fosl. f. Corallinae (Crn.) Fosl. mscr. f. Laminariae (Crn.) Fosl. mscr. p. 128 changed this classification to:— L. pustulatum (Lamour.) Fosl. L. macrocarpum (Rosan.) Fosl. mscr. f. intermedia Fosl. mscr. f. faerdensis Fosl. mscr. f. Laminariae (Crn.) Fosl. mscr. L. hapalidioides (Crn.) Fosl. f. typica f. confinis (Crn.) Fosl. (L. canescens Fosl.) (L. prototypum Fosl.) (L. tumidulum Setch et. Fosl.) (L. polycephalum Fosl.) corallines, and in particular of the history of all the changes that had occurred to Heteroderma in the course of Foslie’s publications. W. H. Adey and P. J. Adey On the basis of his examination of Foslie’s herbarium in Trondheim, Adey (1970) considered that simple crustose corallines with lateral cell fusions and single-pored conceptacles should not 318 Y. M. CHAMBERLAIN Table 4 Genera to which species listed by Foslie now belong. D. Adplicitum D. macrocarpa D. bermudense M. marginata P. mauritiana P. Callithamnioides M. mediocris D. Canescens L. melobesioides *D.? Caspica (or Litholepis?) M. membranacea P. Caulerpae P. minutula P. Chamaedoris P. myriocarpa M. coccinea P. confervicola M. Notarisii D. confinis P.? Novae Zelandiae P. confervoides D. conspectum D. papillosum D. corallinae D. polycephalum P. coronata D. prototypus D. corticiformis D. pustulatum P. Cymodoceae D. Cystoseirae D. rasile P.? rhizomae Ps. decipiens M. Rosanoffii Ps. discoideum P. rosea D. dispar P. rugulosa P. explanata P. Sargassi P. subtilissima F. farinosa C. Thuretii M. galapagense P. triplex P. Gibbsii D. tumidulum D. hapalidioides P. zonale M._hildenbrandtioides P. zostericola F.? inaequilaterata D. Laminariae P. lejolisii P. lepturum PP. limitata C. = Choreonema Schmitz D. = Dermatolithon Foslie F. = Fosliella Howe M. = Melobesia Lamouroux N. = Neogoniolithon Setchell & Mason P. = Pneophyllum Kitzing Ps. = Pseudolithophyllum sensu Lemoine L. = Lithoporella Foslie be divided into separate genera on the basis of the presence or absence of trichocytes. He referred them all to Heteroderma Foslie. In 1973, however, Adey & Adey described two new species of Fosliella, both of which possessed intercalary trichocytes. * usually given as Litholepis but Zinova (1967) refers it to D. caspicum (Foslie) Zaberzh. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 319 Summary The solutions arrived at in the present investigation are: (1) Melobesia membranacea (Esper) Lamouroux should remain the type species of Melobesia Lamouroux (1812) since I consider that the drawing by Esper (1806) which Lamouroux cited in his description illustrates the present species. (2) The name Fosliella M. Howe (1920) should be restricted to species with terminal trichocytes and be typified on F. farinosa (Lamouroux) M. Howe. Species with intercalary trichocytes should be classified in Pneophyllum Kiitz. (1843) and be typified on P. fragile Kiitz. Coralline structure Introduction Spore germination characteristics, thallus structure, and development as well as reproductive features of coralline algae have recently been considered in a number of publications. In particular Chamberlain (1978) discusses simple crustose corallines, while Johansen (1981) summarises current knowledge for the entire Corallinaceae. Only a brief account will, there- fore, be given of those principal features which particularly appertain to Fosliella and Pneophyllum. Spore germination Newly released coralline spores of all types (ie. carpospores, tetraspores, and bispores) are approximately globular, deep red bodies encased in a clear coating which is presumably mucilaginous (Jones & Moorjani, 1973). As described in Chamberlain (1982) the spores, on settling, become attached to the substratum by means of an adhesive mucilage and segment to produce up to 32 cells within the confines of the original spore wall before growth of the sporeling beyond this wall commences (Figs 7, 8). This type of segmentation was described by Chemin (1937) as the Dumontia-type and is seen almost throughout the Corallinaceae when spores germinate under favourable conditions. A filamentous type of germination, the Naccar- la-type, is, however, seen in parasitic and semiendophytic species (Cabioch, 1972) and in other species such as Fosliella farinosa when germination occurs under unfavourable conditions (Chamberlain, 1982). Fig. 7 Spore segmentation in Pneophyllum and Fosliella. A, Second division of Pneophyllum lobescens carpospore showing adhesive mucilage (arrow). Scale = 10 um. B, Fosliella farinosa sporeling (from bispore) showing four-celled central element (1) and young peripheral growth (2). Scale — 20 wm. 320 Y. M. CHAMBERLAIN The sequence of spore segmentation was further studied by various authors such as Chihara (1973, 1974a, b), Notoya (1974, 1976a, b), and Bressan (1980) who all recognised that particular sequences could generally be related to taxonomic groupings, particularly to the subfamily. These authors all found, however, that the simple mastophoroid species (variously attributed to the genera Fosliella and Heteroderma) showed a wide diversity of pattern. Chamberlain (1982) considered that these patterns could all be related to two fundamental types, those with a four-celled central element (Fig. 7B) to the germination disc, as found in Fosliella, and those with an eight-celled central element (Fig. 8), as found in Pneophyllum (previously including Fosliella spp. and Heteroderma spp.). This is considered to be the most consistent and fundamental character at present recognised as separating these two genera (Chamberlain, 1982). After segmentation, peripheral growth commences (Figs 7B; 8), either all round the segmented spore to form immediately a circular disc, or from one or two points on the periphery to form fan-shaped discs, which usually, however, eventually unite into a more or less circular disc. The vegetative thallus. Thallus components As with the majority of red algae, the coralline thallus is fundamentally composed of filaments. The basic structure of the thallus may be seen most easily in the simple crust, but this form is not necessarily primitive and may, in some cases at least, be neotenic (cf. Cabioch, 1971b, 1972). The mature simple crust is generally a disc-like structure which starts as a single basal layer of radiating, calcified filaments, called the hypothallium, which is initiated from the periphery of the segmented spore (Figs 8; 9A) and continues to grow by the activity of peripheral apical initials (Figs 9; 10A). Each differentiated cell of the hypothallium cuts off a small, lightly calcified or uncalcified cell on the upper surface of its distal end called the epithallial cell (Figs 8; 9A, B; 10B, C). In the simplest forms, including all currently known species of Fosliella and ~ ‘ Fig. 8 Scanning electron micrograph of germinating bispore of Pneophyllum lobescens to show eight- celled centre (arrow) giving rise to radiating filaments of hypothallial cells produced by the apical initial which forms the first meristem; each hypothallial cell cuts off an epithallial cell. Scale = 25 wm. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 321 some Pneophyllum spp., no further cell types are formed in the vegetative thallus. Many simple crustose species and all complex crusts, however, develop a third component, the perithallium< which lies between the hypothallium and the epithallium (Figs 9B; 10B; 11C). At its simplest, the perithallium comprises single calcified meristematic cells (subapical initials) cut off by a periclinal division from the upper side of each cell of the hypothallial filaments. In many species, however, the initials continue dividing periclinally, cutting off successive perithallial cells from their lower side (Fig. 10B) which form perithallial filaments; these may be up to twenty cells deep in Pneophyllum depending on the species. Perithallial and hypothallial cells often contain abundant floridean starch grains (Fig. 11B). In addition to cutting off perithallial cells towards the base of the crust the subapical initials continue (possibly throughout the life of the crust), to cut off successive epithallial cells on the crust surface, the older cell usually being sloughed off as the new one develops (Bressan et al., 1981). In some complex crustose species, and very occasionally in Pneophyllum, the epithallial cells are not sloughed off as the new ones develop and short filaments of epithallial cells consequently occur. It can be seen, therefore, that crust growth is accomplished by the activity of two types of meristem (Fig. 9). The hypothallium expands peripherally by the division of apical initials which collectively comprise a meristem that will be called the first meristem since it is the first to be initiated after spore segmentation. The epithallium and perithallium subsequently develop from the activity of intercalary, or subapical, initials which are situated between these two thallus components and which will collectively be called the second meristem, since it is always initiated after the hypothallial meristem. Woelkerling (1980), Johansen (1981), and Turner & Woelkerl- ing (1982) refer to the first meristem as the primary meristem and the second meristem as the secondary (Woelkerling, 1980; Turner and Woelkerling, 1982) or intercalary (Johansen, 1981) meristem, by analogy with phanerogams. I think it preferable, however, to limit the analogy solely to the sequence of development without making precise terminological comparisons with higher plants, since the basically filamentous structure of the algal thallus is quite different from the phanerogam thallus organisation. Cell connections The successive cells of hypothallial, epithallial, and perithallial filaments are joined by normal red algal pit connections (Figs 9B; 10B, C; 11C), the pit plugs of which (Pueschel & Cole, 1982) have recently been shown to be of a particularly elaborate nature (Fig. 11A). A characteristic of the Corallinaceae is the development of secondary lateral synapses between contiguous cells of Fig. 9 Thallus features in Fosliella and Pneophyllum. A, Diagrammatic surface view of Pneophyllum lobescens sporeling (arrow = original spore). Scale = 50 wm. B, Diagrammatic radial VS of Pneophyl- lum crust periphery (redrawn after Cabioch, 1972). Heh Figures 8 and 9 represent the following features: h = hypothallial cells forming the prostrate radiating filaments; p = perithallial cells of erect filaments; e = epithallial cells; ai = apical initials; M1 = first meristem composed of apical initials; sai = subapical initial; M2 = second meristem composed of subapical initials; pc = primary pit connection; cf = cell fusion. Y. M. CHAMBERLAIN Fig. 10 Thallus components of Fosliella and Pneophyllum. A, Fosliella farinosa sporeling to show peripheral initiation of terminal trichocyte (tr), apical initial (ai) and onset of calcification (ca). Scale = 20 um. B, Vertical section of Pneophyllum plurivalidum crust to show perithallial (p) and hypothallial (h) filaments, epithallial cells (e), pit connections (pc) between successive perithallial cells, fusions (cf) between contiguous perithallial cells, and hypothallial cells and subapical initials (sai). Scale = 10 um. C, Surface view of Pneophyllum rosanoffii crust to show hypothallial cells (h), epithallial cells (e), branch cell intercalary trichocyte (btr), intrafilament intercalary trichocyte (itr), pit connections (pc), and cell fusion (cf). Scale = 25 wm. hypothallial and perithallial filaments. These synapses are described in detail by Cabioch (19706, 1971b) and may be in the form of secondary pit connections similar to the primary pit connections mentioned above, or fusions in which the walls of adjoining cells decompose to form coenocytes (Figs 9B, 10B, C). The type of secondary synapse present is considered to be of phylogenetic significance and in Fosliella and Pneophyllum only cell fusions are seen. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 320 Trichocytes A feature of many members of the Corallinaceae is the presence of trichocytes (i.e. hair-bearing cells) in the thallus; Cabioch (1968, 1970a, b, 1971b) comprehensively describes the diversity of trichocytes in the Corallinaceae so that only the forms found in simple crusts (as investigated by myself) are considered here. The trichocytes which occur in hypothallial cells in the crust periphery are of two types: (1) in Fosliella the terminal trichocyte develops at the first meristem when an apical initial (Fig. 10A) becomes somewhat swollen and, when mature, divides by a tangential, more or less anticlinal wall; the centrifugal part then develops a hair. The hair is often shed quite quickly, but the trichocyte persists in the thallus and terminates the growth of the filament which bears it; contiguous thallus filaments subsequently grow round the trichocyte to continue the growth of the thallus; (2) in Pneophyllum trichocytes usually develop from a differentiated thallus cell (or occasionally in an apical initial) and in this case the cell divides by a radial, anticlinal wall into a pigmented part and a colourless, hair-bearing part (Fig. 10C). Since this type of trichocyte develops within a filament which grows on beyond it, it is known as an intercalary trichocyte: it may develop either within an unbranched length of filament (Fig. 10C) here called an intrafilament type, or in a filament branch cell (Fig. 10C) here designated as a branch cell type. In addition, large, sac-like, terminal cells bearing hairs occasionally develop in Pneophyllum, probably as a response to specific environmental conditions; these have been observed in P. limitatum by Suneson (1943, as Melobesia) and in P. myriocarpum (Chamberlain, 1982) and can be regarded as abnormal structures clearly distinguishable from the ‘normal’ terminal trichocytes of Fosliella by their large size (at least twice the length of crust cells) and lack of cross walls. The taxonomic significance of trichocytes in these genera has, as Johansen (1981) remarks, frequently been questioned, especially as their occurrence appears to depend on environmental conditions; I have confirmed (Chamberlain, 1982) that their occurrence depends on a sufficiently high temperature and light intensity and they are, for example, usually absent in winter and in deeper subtidal habitats. However, while I have never found intercalary trichocytes in the hypothallium of Fosliella, in Pneophyllum the trichocytes, when present, are normally intercalary. At the very least, therefore, I consider this a useful feature for practical generic identification. Reproduction Vegetative reproduction As Johansen (1981) remarks, many coralline algae are readily capable of vegetative reproduc- tion in that small fragments of thallus are able to reattach themselves to a substratum and regenerate. Specific vegetative reproductive organs are rare however, although propagules have been described in Fosliella farinosa (Solms-Laubach, 1881, as Melobesia callithamnioides; Coppejans, 1978) and F. paschalis (Hollenberg, 1970), while Suneson (1943) observed con- tracted protoplasts in marginal cells of Pneophyllum limitatum (as Melobesia), which he suggested might be organs of vegetative reproduction. Conceptacles The non-vegetative reproductive cells of all corallines are borne in protruding, more or less hemispherical, or immersed, flask-like, chambers known as conceptacles (Fig. 11F) which are formed by the growth of the vegetative thallus. Four different conceptacle types occur, containing tetrasporangia, bisporangia, carpogonia, and spermatia respectively. The carposporophyte develops within the carpogonial conceptacle after the carpogonium has been (presumably) fertilised by a spermatium although direct evidence that fertilisation occurs has yet to be obtained. Details of the mode of conceptacle development in various coralline groups are summarised by Johansen (1981). Almost all conceptacle types release spores through a single common pore, the ostiole, in the conceptacle roof; in the subfamily Melobesioideae (Fig. 16) however, tetra- and bisporangia are released through individual pores, and the conceptacle roof in this case is therefore pierced by numerous pores (see Chamberlain, 1978). 324 Y. M. CHAMBERLAIN The conceptacles of Fosliella and Pneophyllum are prominent structures in species with a thin vegetative thallus but may be entirely immersed in the thicker species such as P. plurivalidum. Gametangial, tetrasporangial, and bisporangial conceptacles are known for most species, but bisporangial conceptacles have not yet been recorded in some of the present species such as P. rosanoffii and P. sargassi; two distinct forms of bisporangial conceptacle occur in P. limitatum, both bearing uninucleate bisporangia. Garbary (1978) examined the appearance of the conceptacle in a range of articulated and crustose corallines under the S.E.M. He found that, while the conceptacle surface resembled the thallus surface in most species, in some species such as Dermatolithon litorale and Fosliella farinosa, uneven calcification of the outer cell walls occurs causing the cells to bulge outwards, thus orientating the epithallial cell in an almost horizontal position. Garbary also noted distinctive rosette-like cell patterns round the ostiole in some species. In the present investigation, the structure of the conceptacle, particularly of the ostiolar area, has been found to be of systematic significance in separating species of Pneophyllum. Studies on decalcified, sectioned material, in conjunction with S.E.M. studies of air and critical point dried material of the same taxon, have made it possible to follow conceptacle development and this has led to a better understanding of how the mature features should be interpreted. Many aspects of the developmental process are common to all conceptacle types and the basic principles are demonstrated for the carpogonial/carposporangial conceptacle of Pneophyllum rosanoffii (Figs 12A-G). A carpogonial conceptacle of P. rosanoffii is initiated (Fig. 12A) when the subapical initials cut off additional perithallial cells over a circular area of crust and these perithallial cells (called cavity cells in Johansen, 1981) become vertically stretched. Fertile initials (Fig. 12A) develop centrally on the floor of the central disc and the perithallial cells above them disintegrate. In P. rosanoffii a ring of persisting perithallial cells surrounding the ostiole become meristematic apically (Fig. 12B) and give rise to upward and downward ostiole filaments (Fig. 12F). Eventually the perithallial cells at the periphery of the conceptacle rupture, their outer walls form a lining to the chamber which is evident in section as a thin hyaline layer (Fig. 12E) and can be seen as a smooth surface in S.E.M. preparations (e.g. in P. myriocarpum, Fig. 11D). The filaments surrounding the ostiole (papillary cells in Masaki, 1968) develop in characteris- tic ways in different species. In the thin crusted P. /ejolisii and deep crusted P. microsporum, for example, development is minimal. Upward ostiole filaments are particularly well developed in species such as P. limitatum, P. zonale, and P. rosanoffii (q.v.); these filaments are probably uncalcified but they nevertheless persist during S.E.M. preparation, particularly in critical point dried material. In most species a ring of downward pointing filaments surrounds the ostiole (Fig. 12F); in some species such as P. concollum the filament tips may become swollen. Under the S.E.M. the inside surface of the conceptacle roof of P. zonale (Fig. 11E) shows that while the basal parts of these filaments are heavily calcified, the tips presumably remain thin-walled and uncalcified since they have disappeared in air dried material. Some species, in addition to producing ostiole filaments, produce filaments which form a new roof below the original crust surface as can be seen in P. plurivalidum (e.g. Fig. 77A). When this occurs the original crust surface sloughs off to reveal the new roof cells (e.g. in P. zonale, Fig. 11F), which are more or less isodiametric in most species. The upper parts of the perithallial cells which produce the ostiole filaments can often be seen as irregularly shaped structures (in P. lobescens for example). In air dried, S.E.M. preparations these irregular cell remnants disintegrate, leaving a pattern of pit connections (e.g. in P. zonale, Fig. 11E) showing where the Fig. 11 Scanning electron micrographs to show morphological features of Pneophyllum. A, Pit plugs from primary pit connection of Pneophyllum zonale. Scale = 2 um. B, Vertical section of Pneophyllum zonale crust to show the globular grains of floridean starch. Scale = 10 wm. C, Vertical section of Pneophyllum sp. showing heavily calcified cell walls with calcite crystals orientated at right angles to the lumen (1) and a pit connection between perithallial cells (2). Scale = 10 wm. D, Vertical section of tetrasporangial conceptacle of Pneophyllum myriocarpum showing the smooth lining formed from remnants of perithallial cell walls (arrow). Scale = 50 wm. E, View from below of the roof of a FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES bisporangial conceptacle of Pneophyllum zonale showing the calcified parts of downward ostiole filaments (1) and rows of pit connections marking the place at which ostiole filaments are given off by the perithallial cell (2). Scale = 25 wm. F, Bisporangial conceptacles of Pneophyllum zonale (critical point dried). Scale = 250 um. 326 Y. M. CHAMBERLAIN new filaments were given off. Whether or not a new conceptacle roof is formed, and the way in which the old thallus surface is shed in species which produce a new roof, has proved to be a useful taxonomic character (see page 335). While the features can only be seen clearly under the S.E.M. it is often possible to deduce information about them from vertical sections. ° tgeia. 2 9 B 0 g 0° s> eatin Com — i agin AR aliastial-eeee® maralela' FTEs \| aueNavecScecetenes WAG nce een = 20um Fig. 12 Vertical sections of Pneophyllum rosanoffii to show structure and development. A-C, Car- pogonial conceptacles. A, Initiation and stretching of perithallial cells and appearance of fertile initials (shaded). B, enlargement of perithallial cells and initiation of roof and ostiole filaments (arrow); young procarps developing (shaded). C, Mature procarps (shaded) and further growth of roof. D, Appearance of fusion cell (fc) and initiation of peripheral gonimoblast filaments (shaded); early stage in growth of upper ostiole filaments (arrow). E-G. Carposporangial conceptacles. E, Growth of peripheral goni- moblast filaments (shaded); maturation of upward ostiole filaments (1). Perithallial cells now form a conceptacle lining (2). F, Maturation of carposporangia (c) and downward ostiole filaments (arrow). G, Dispersal of final carpospore through ostiole and senescence of fusion cell and remaining procarp remanants (arrow). H, Spermatangial conceptacle showing escaping spermatia and ring of cells (1) round conceptacle neck which produces the spout (2). FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES a2) As mentioned previously, the development and structure of the conceptacle is essentially similar for carpogonial/carposporangial, tetra- and bisporangial conceptacles. Spermatangial conceptacles have basically the same process of development but the mature conceptacles are simple, flask-like structures (Fig. 12H) usually immersed in the thallus at the edge of carpogonial conceptacles. The ostiole filaments are reduced to a ring of cells round the neck of the conceptacle and produce a hyaline spout through which the spermatia are released. In one species, P. concollum, a similar spout develops on top of some of the collar-like ostioles of the other conceptacle types. Gametangia and the carposporangial phase Gametangial crusts are almost always monoecious, with spermatangial conceptacles occurring adjacent to the carpogonial/carposporangial conceptacles. Under certain environmental con- ditions, however, carpogonial conceptacles may fail to develop and crusts bearing only spermatangial conceptacles occur; this has been seen, for example, in Fosliella farinosa collected from the shore in winter (Chamberlain, 1977b). Spermatangial structure shows little variation among Fosliella and Pneophyllum species. Fertile initials (spermatangial mother cells) develop from the floor of the conceptacle only; they produce elongated processes (Fig. 12H), usually from two sites on each initial, and these are eventually cut off and released into the chamber of the conceptacle. Here they round off to form more or less oval spermatia. The odgonium is borne on a procarp (Figs 12C; 13) which comprises a supporting cell bearing one or two carpogonial branches and often a sterile filament. Each carpogonial branch consists of a hypogynous cell and a carpogonium, which has a swollen basal portion containing the odgonium and an extended, tube-like, upper portion, the trichogyne, through which the spermatium enters to fertilise the odgonium. In all species of Pneophyllum fertile procarps, bearing one or two carpogonial branches, occur in the centre of the fertile disc, while simpler structures without carpogonia occur at the periphery (Fig. 12C); a similar gradation has been noted in most coralline algae (Lebednik, 1977). In Fosliella only a single carpogonium has been observed on each procarp in the present investigation, and this is corroborated by the observations of Balakrishnan (1947, as Melobesia farinosa) and also appears to be the case in the illustrations of F. farinosa in Gordon etal. (1976); on the other hand, Masaki & Tokida (1960, pl. VII, Fig. 2) show two carpogonia on some procarps of F. farinosa (as Melobesia). While further investigation is needed, it seems possible that the presence of two carpogonia in Pneophyllum, and one in most samples of Fos/liella, is a character of generic significance. Suneson (1937) put sterile filament trichogyne | : | eee carpogonium lbranch PROCARP ~hypogynous cell | supporting (auxiliary) cell LL. = = perithallium hypothallium Fig. 13 An hypothetical coralline procarp. 328 Y. M. CHAMBERLAIN forward the hypothesis that in the Corallinaceae procarps of ‘primitive’ taxa bear more carpogonia than those of ‘advanced’ types, which suggests, therefore, that Fosliella may represent a more ‘advanced’ evolutionary line than Pneophyllum. As in all corallines, once a carpogonium has been fertilised the usual sequence of events leading to the formation of a fusion cell occurs. No evidence to prove the path taken by the zygotic nucleus in this process was uncovered in the present investigation, but Suneson (1937) figured a tube leading from the carpogonium to the auxiliary cell in P. lejolisii (as Melobesia). In Pneophyllum there seems to be no cellular connection between the fusion cell and subtending thallus cells which usually wither away; in British material of Fosliella farinosa (see p. 347) however, cell connections may appear between the fusion cell and the persistent subtending cells, and these were also observed by Balakrishnan (1947, as Melobesia). This is a further feature of possible significance. In the Pneophyllum species studied, the fusion cell varies from a thin, wide, pancake-like structure, closely appressed to the conceptacle floor (as in P. concol- lum) to a short, dumpy structure which is often situated high in the conceptacle cavity (as in P. sargassi). In all cases, gonimoblast filaments bearing terminal carposporangia develop from the periphery of the fusion cell (Figs 12E, F), and the remaining procarps in the centre of the fusion-cell surface (Fig. 12G) gradually disintegrate. Previously (Chamberlain, 1977a) it seemed that in British material of P. limitatum, gonimoblast filaments develop from the entire upper surface as well as the periphery of the fusion cell, as observed by Suneson (1937) in Swedish plants. Further examination has not confirmed this and although mature conceptacles are so packed with gonimoblast filaments and carposporangia that they occupy the whole area in surface view, almost all the gonimoblast filaments are initiated peripherally. However, on one occasion a filament appeared to be borne on the upper surface of the periphery of a fusion cell. Tetrasporangia and bisporangia Tetrasporangia develop peripherally in all species (e.g. Fig. 14A). In most taxa sterile paraphyses have not been seen to develop among the sporangia, although remnants of old perithallial cells are often observed in the centre of the conceptacle. In P. concollum (q.v.), however, a conspicuous central columella develops. All tetrasporangia develop in a manner characteristic of the Mastophoroideae (Chamberlain, 1982). The tetrasporangial initial (Fig. 15A) enlarges from the conceptacle floor, divides horizontally into a lower, stalk cell, which persists and develops no further, and an upper cell which enlarges to form the tetrasporangium. In species such as P. /ejolisii, the young tetrasporangia are pigmented and the sporangium remains farctate throughout development, with the eventual septa appearing as thin lines in side view. In other species such as P. rosanoffii and Fosliella farinosa (q.v.), the young sporangium is pale, thread-like, deeply invaginated, and enclosed in a loose, sac-like casing; the sporangia expand in this pale, invaginated form, and only in the final stages of development do they become pigmented and farctate. Variations in the structure (Fig. 14B) and development of bisporangia resemble those of the tetrasporangium in the same taxon; bisporangial development in F. farinosa, for example, is described (Chamberlain 19775). Bisporangial plants with binucleate bispores are known in both genera (e.g. in F. farinosa— Balakrishnan, 1947, as Melobesia) but plants with uninucleate (and presumably diploid) bisporangia are probably more common, especially in colder climates (see Bauch, 1937, and Suneson, 1950). Life history A self-perpetuating, bisporangial life history (Fig. 15C) has been shown in plants of F. farinosa (Chamberlain, 1977b) with uninucleate bispores. In the present study (Chamberlain, 1982) the morphological sequences of a Polysiphonia-type (Fig. 15B) life history (Dixon, 1973) have been fully established for Pneophyllum lobescens and P. myriocarpum, and partly for P. plurivalidum and Fosliella farinosa, while a self-perpetuating bisporangial life history is demonstrated in Pneophyllum zonale, P. lobescens, and P. plurivalidum. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 329 Summary The most fundamental and consistent character separating Fosliella and Pneophyllum is the presence of a four-celled central element to the germination disc in Fos/iella, and an eight-celled central element in Pneophyllum. In practice, however, this feature is not usually observable, and the two genera are, therefore, most easily distinguished by the presence of terminal hypothallial trichocytes divided by a tangential wall in Fosliella, as opposed to intercalary hypothallial trichocytes divided by a radial wall in Pneophyllum. However, abnormal, very large terminal trichocytes (at least twice the length of crust cells), lacking a dividing wall, have been observed in Pneophyllum, although intercalary trichocytes are so far unknown in Fosliella. Two further features which may possibly prove of significance in distinguishing the genera are: (a) the presence of two carpogonia on at least some of the procarps of all species of Fig. 14 The structure of tetrasporangial and bisporangial conceptacles in Pneophyllum zonale. A, Vertical section of tetrasporangial conceptacle. B, Vertical section of bisporangial conceptacle. 330 Y. M. CHAMBERLAIN A M ASTOPHOROIDEAE LB te) ty ee - - . E BEEP gh RTE pees fi =Sc p o h = B 2n oe ae n tetrasporangial fUeesis 5 ——_ gametangial plant plant n tetraspores | | Carpogonium spermatium Oo. (2") - \/ bs < 2n carpospores zygote carposporangial phase Cc Qn 2n bispores bisporangial >} plant ii Bei Fig.15 Tetrasporangial development and life-history phases in Fosliella and Pneophyllum. A, Tetraspor- angial development: p = perithallial cell, h = hypothallial cell, ti = tetrasporangial initial, sc = stalk cell. B, Polysiphonia-type life history. C, Bisporangial life history. Pneophyllum, as opposed to the single carpogonium almost always observed in Fosliella; and (b) the presence of a cellular connection between the fusion cell and subtending conceptacle floor cells in Fosliella, but not in Pneophyllum. A detailed survey of these features throughout the genera would be valuable. The classification of the Corallinaceae The Corallinaceae belong to the order Cryptonemiales, an order which Kylin (1956) character- ised as having auxiliary cells which are present before fertilisation, and which occur as accessory filaments either adjacent to, or remote from, the carpogonium. Traditionally the members of the Corallinaceae have been divided into two groups, containing the crustose and the geniculate forms respectively, in a scheme of classification initiated by Areschoug (1852). More recently, 331 FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES ‘(Z7Q6T) UlejJoquieyD UI pajiejop pue 7/6] VOUIs poysi[qnd uONeWIOJUI Jo SIseq dy} UO JOYINe JUdsoI1d ay} Aq poppe ud0q dARY YSL9Ise uv Aq poysew Solu “(Z76[) ussueyor 2 Aopy Joye xnoINoOWeK’T sesdeUTT[e1O_ A[IWLJ IY} JO UOTJBOYIsse|D OUT, OT “SY sajoeidaouoo jeisuesods poiod-ajsuls suonoauuod-jid A1z payepnoie sajoeidaouoo jeisuesods posod-ay3uls suonsauuos-jid A1Z aso}snId po191) ajsuls eynoiuad sajoejdaouos jeisueiods poiod-aj3uls suolsauuod9-}id AZ Ou paiejnonie DNRW9}SIIW ByNIIUIS S aipridsoude yesuerod? sajovjdaouoo [eisuesods poiod-aj3uls ; posod-ajsuis suonsauuods-}id AZ Ou ; : : suonsauuos-jid A1z ou asojsnio : : payepnonie sajoejdaauos jeisuesods poiod-Auew suonsouuos-jid Az Ou asoysnio aeopionyduy aeaproy|Aydoyy] avaplouly|e1og aeaplooydojyseyy dvaployIOIUOse}IJ dRIPIOIsaqo]ay «ny homme van Reine) and have observed the same modification in herbaria (e.g. Titius, no. 855 from Spalato, identified by Rosanoff as ‘Melob. farinosa f. macrocarpa’, in CN). This form corresponds in size with that recorded from Venezuela by Ganesan (1971). The thin, vegetative thallus and abruptly-domed conceptacles are common to all descriptions, with the exception of the drawing by Masaki & Tokida (1960, Pl. VI, Fig. 7) which shows a tetrasporangial conceptacle immersed in a deep thallus. Terminal trichocytes are nearly always present except in winter and on material collected from deeper than about 5 m (Lemoine, 1923a; Chamberlain, 1977b), but sometimes crusts with and without trichocytes are found growing side by side (Lemoine, 1917; Ganesan, 1971). Lemoine (1917) also depicted differences in cell size and shape according to the host which is another factor to be taken into consideration regarding species variability. The creeping forms, particularly callithamnioides, occur quite frequently, especially in warmer climes and I have observed f. callithamnioides in Ireland where the crusts may also show a transition from a uniform crust to a creeping filamentous form. The variety chalicodictya has been recorded from the Caribbean (Taylor, 1939) and the Mediterranean (Coppejans, 1976) while a further Mediterranean form (unnamed) was described by Boudouresque & Cinelli (1971) which bears a strong resemblance to the early germination filaments of Amphiroa verruculosa as depicted by Cabioch (1972, p. 147). One further form, Melobesia farinosa f. borealis, was described by Foslie (1905a) from Ireland. This has proved to be the bisporangial phase of F. farinosa (Chamberlain, 1977b), which is the commonest phase of this species in the British Isles and has also been recorded from northern France (Lemoine, 1923a). Fosliella paschalis was described from Easter Island by Lemoine (1923b, as Melobesia) as a small, greyish plant in which the thalli never superimposed; it had loosely coalescent hypothal- lial filaments, the conceptacles were smaller than in F. farinosa and were each borne on a mound of perithallial cells. Subsequent records of F. paschalis (e.g. Dawson, 1960; Masaki, 1968; Hollenberg, 1970) have confirmed the small conceptacle size and absence of superimposition of thalli, but none has mentioned the presence of perithallial mounds. Vegetative propagules have been described in Fosliella — such structures occur very rarely in the Rhodophyta. Solms-Laubach (1881) described multicellular, triangular propagules growing on F. farinosa f. callithamnioides (as Melobesia callithamnioides) in the Mediterranean and the same type was described from f. farinosa by Coppejans (1978). Much simpler, unicellular, pear-shaped propagules were depicted by Hollenberg (1970) growing on F. paschalis. F. cruciata occurs commonly in the Trieste area (Bressan et al., 1977) and its distinctive characters were described as the germination disc pattern and the elongated terminal tricho- cytes, Bressan & Tomini (1983) suggest that it may be a polyploid species. It seems certain that there are more Fosliella species than have at present been described. Difficulties arise, however, because the plant shows such a wide range of structure and growth pattern even when growing on the same host. Thus normal and giant crusts, with and without trichocytes, with solid and/or creeping crusts, may all grow on one small area of host which would seem to preclude the possibility that they are environmental forms but, at the same time, they are not separable into clearly defined species. In addition, many forms of small Pneophyl- lum spp., such as P. confervicolum, may grow in such close association with Fosliella, particularly in the tropics, that it becomes impossible to distinguish the separate thalli and identify the entities. It seems that a world wide survey of Fosliella based on extensive shore collections coupled with laboratory culture will be essential before the complexity of this genus can be understood. Polyploidy is a possible mechanism which may contribute to the observed variation in giving rise to the giant form. It is also possible that some forms of Fosliella may be neotenic forms of more complex mastophoroid genera such as Neogoniolithon and Porolithon. A more puzzling situation is, as mentioned, the strong resemblance of the germination stages of Amphiroa to creeping forms of Fosliella even though mature Amphiroa has secondary pit connections and belongs to the Amphiroideae. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 343 Eco.ocy: Fosliella occurs principally as an epiphyte on other algae and on sea-grasses and may form a dense, floury covering over the host plant. It also grows on pebbles and shells, it is probably less common on these substrata than on plants but few data are available. DIsTRIBUTION: Fosliella occurs in tropical and temperate parts of the world (Chamberlain, 1977b). It is, however, particularly abundant in warmer climates and absent from colder areas; the British Isles, for example, seems to represent the northern limit of its European distribution. la. Fosliella farinosa (Lamouroux) M. Howe forma farinosa BasionyM: Melobesia farinosa Lamouroux, Histoire des polypiers coralligénes flexibles, vulgairement nommés zoophytes: 315 (1816). NOMENCLATURAL SYNONYM: Fosliella farinosa (Lamouroux) M. Howe, Jn N. L. Britton and C. F. Millspaugh (Eds) The Bahama flora: 587 (1920). REFERENCES: Melobesia farinosa Lamouroux; (?) Harvey, A manual of British algae 2nd ed.: 109 (1849); Areschoug, Jn J. G. Agardh (Ed.) Species genera et ordines algarum: 512 (1852), in Nova Acta Reg. Soc. scient. Upsala III, 10: 366 (1875); Rosanoffin Mem. Soc. imp. Sci. nat. Cherbourg 12: 69 (1866); Foslie in K. nor. Vidensk. Selsk. Skr. 1898 (3): 10 (1898), K. nor. Vidensk. Selsk. Skr. 1900 (5): 20 (1900), K. nor. Vidensk. Selsk. Skr. 1905 (3): 69 (1905) — as f. typica and f. borealis; Heydrich in Ber. dt. bot. Ges. 15: 44 (1897); Lemoine in Archs Mus. natn. Hist. nat. Paris V, 5: 140 (1913), In C. Skottsberg (Ed.), The Natural History of Juan Fernandez and Easter Island 2: 19 (1923b); Rosenvinge in K. dansk. Vidensk. Selsk. Skr. VII, 7: 240 (1917); Mazza, Saggio di Algologia Oceanica 3: 1113 (1922); Newton, A handbook of the British seaweeds: 300 (1931); Balakrishnan in J. Indian bot. Soc. 1946: 305 (1947 [‘1946’]); Hamel & Lemoine in Archs Mus. natn. Hist. nat. Paris VII, 1: 102 (1953 [‘1952’]); Funk in Pubbl. Staz. zool. Napoli 25 (Suppl.): 97 (1955); Masaki & Tokida in Bull. Fac. Fish. Hokkaido Univ. 11: 39 (1960); Edelstein & Komarovsky in Bull. res. Coun. Israel D, 1-4: 56 (1961); Lewalle in Pubbl. Staz. zool. Napoli 32: 265 (1961); Zinova, Check-list of green, brown and red algae of the southern seas of the U.S.S.R.: 229 (1967); van der Ben in Proc. int. Seaweed Symp. 6: 79 (1969); Pankow, Algenflora der Ostsee, I. Benthos: 249 (1971); South in J. mar. biol. Ass. U.K. 56: 820 (1976). Melobesia farinosa var. borealis Lemoine [sic]; Newton, A handbook of British Seaweeds: 301 (1931). Melobesia granulata Meneghini, Jn Zanardini, Saggio di classificazione naturale delle Ficee: 44 (1843), Kiitzing, Species Algarum: 696 (1849), Tabulae Phycologicae: 34, Tab. 95a, b (1869); Areschoug, Jn J. Agardh, Nova Acta reg. Soc. scient. Upsala III, 10: 518 (1852). Melobesia zonalis sensu Lemoine in Archs Mus. natn. Hist. nat. Paris V, 5: 140 (1913). Melobesia minuta Schiffner in Oesterr. Bot. Zeit. 82: 291 (1933). Fosliella farinosa (Lamouroux) Howe; Feldmann in Rev. algol. 11: 316 (1942 [‘1939’]); Taylor, Marine algae of the northern eastern coast of North America — revised edition: 252 (1957), Marine algae of the eastern tropical and subtropical coasts of North America: 388 (1960); Masaki in Mem. Fac. Fish. Hokkaido Univ. 16: 21 (1968); Ganesan, In: Symposium on investigations and resources of the Caribbean Sea and adjacent regions, UNESCO: 411 (1971); Bressan in Boll. soc. Adriat. Sci. nat. 59: 74 (1974); Parke and Dixon in J. mar. biol. Ass. U.K. 56: 534 (1976); Gordon, Masaki and Akioka in Micronesica 12: 255 (1976); Chamberlain in Br. phycol. J. 12: 343 (1977) — but excluding Figs 5, 6 and 7. SPECIMENS: Melobesia farinosa sensu Le Jolis, no. 282, Cherbourg, coll. 30 Nov. 1853 on Rhodymenia palmata (CHE!). Melobesia membranacea sensu P. Crouan & H. Crouan, exsiccata no. 244 on Zostera (CO!). Lithothamnion membranaceum sensu E. M. Holmes, Sea Mill, Ayrshire, August 1890 on Zostera (BM!). EXCLUDED RECORDS: Melobesia farinosa auct. div.; Le Jolis, Liste des algues marines de Cherbourg: 150 (1863); Kiitzing, Tabulae Phycologicae: 34, Tab. 95c, d (1869) (both = Pneophyllum rosanoffii). Fosliella farinosa sensu Y. Chamberlain in Br. phycol. J. 12: 343 (1977) — Figs 5, 6 and 7 only (= Pneophyllum concollum). Ho ortyre: CN!, Herb. Lamouroux, from the Mediterranean, growing on Sargassum linifolium (Fig. 19). Diacnosis: Species mainly epiphytic, of thin, bright mauvy-pink crusts, often bleached, terminal hypothallial trichocytes frequent, conceptacles abruptly hemispherical: differing from Pneophyllum species by having terminal hypothallial trichocytes and a ring of trichocytes round the base of conceptacles. 344 Y. M. CHAMBERLAIN DEscriPTION: Mainly epiphytic bright mauvy-pink crusts (often bleached in summer), probably not individually exceeding 5 mm diameter but confluent and covering large areas of host, often much superimposed, individal crusts up to 20 wm deep, perithallium absent except in immediate vicinity of conceptacles; crust cells in surface view elongate, 14-28-5 um long x 5—16-5 wm wide, epithallial cells rounded, 7-13-5 wm long x 3-10 wm wide, hypothallial trichocytes terminal, bulbous, 30-43 um long* x 12-20 um wide; conceptacles abruptly hemispherical, all (except spermatangial) with simple or slightly elaborated ostiole, carposporangial conceptacles 62-100 pm internal diameter X 52-83 um high, ring of trichocytes round the base, ostiole sometimes with small collar, spermatangial conceptacles 33-91 wm internal diameter X 26-44 um high, ostiole prolonged into a spout, tetrasporangial conceptacles as carposporangial conceptacles, 65-94 ym internal diameter x 52-78 um high, up to 8 tetrasporangia per conceptacle, 36-65 wm long X 23-39 um wide, some central sterile cells present, young tetrasporangia deeply invaginate, bisporangial conceptacles hemispherical, considerably larger than other types, 166-208 um internal diameter X 45-125 yum high, rarely with ring of trichocytes, 20 or more bisporangia per conceptacle, 54-78 uw long x 26-42 yu wide, central sterile paraphyses present. HIstTorIcAL: The taxon now called Fosliella farinosa was first described by Lamouroux (1816) as Melobesia farinosa, a species of the genus which he had described (Lamouroux, 1812) as a polypore and which was based on Esper’s (1806) Corallina membranacea. The reasons for the erection of a new genus, Fosliella, to accommodate this species are discussed on page 312. Lamouroux’s holotype (CN) of Melobesia farinosa (Fig. 19) is an epiphyte on Sargassum linifolium from the Mediterranean which is so thickly covered with crusts as to appear floury, hence, presumably, the epithet farinosa. I have examined the material and found that many of the crusts agree well with the present concept of Fosliella farinosa, these are growing together with crusts of Pneophyllum and Dermatolithon. Areschoug (1852) further expanded the description of Fosliella farinosa (as Melobesia) and pointed out that it was the same taxon as Meneghini’s (1843) Melobesia granulata. The first detailed anatomical investigation of Fosliella farinosa, however, was made by Rosanoff (1866) who observed and illustrated the terminal trichocytes (which he termed heterocysts) for the first time. Throughout the works of Foslie (1898b-1905a) F. farinosa retained its position in the genus Melobesia since he accepted Heydrich’s (1897b) generic name Epilithon for membranaceum, and it was generally maintained in Melobesia thereafter until Howe’s (1920) publication, after which the generic name Fosliella was accepted by many authors, although Balakrishnan (1947), Hamel & Lemoine (1953), Kylin (1956), and Masaki & Tokida (1960) are notable exceptions. Examination of BM material of Melobesia minuta Schiffner, growing on Corallina virgata, showed it to be Fosliella farinosa. As this collection (no. 835) was cited by Schiffner (1933) in the description, it would seem that Melobesia minuta must be subsumed in Fosliella farinosa. CRUST APPEARANCE: The bright, mauvy-pink crusts are thin and often become very irregular as parts of them fall off the host. The plants are usually epiphytic on algae (Fig. 38) and Zostera but also occur quite frequently on pebbles, glass and shells. The crusts usually show concentric markings and are often confluent covering large areas of the substratum; they may also superimpose (Fig. 22F) to form multi-layered colonies. Individual crusts probably do not exceed 5 mm diameter but are often impossible to measure. In fresh material the hemispherical conceptacles have a delicate and bubble-like appearance, the bisporangial being noticeably larger than the tetra- and carposporangial conceptacles. Under the S.E.M. mature carposporangial (Fig. 20A) conceptacles have a smooth or regularly stepped pattern of roof cells (sometimes with a small spout round the ostiole) and a ring of trichocytes round the base. Spermatangial conceptacles (Fig. 20E) show a spout surrounded by elongated cells. Observations on developing carpogonial conceptacles under the light micro- scope (Figs 23A, B) show that the original thallus surface peels off in strips. The bisporangial * including the length of the basal cell. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 345 Fig. 20 Scanning electron micrographs to show external features of Fosliella farinosa. A, Gametangial crust showing spermatangial conceptacles (s) and carposporangial conceptacles (c) encircled by tricho- cytes (tr). Scale = 50 wm. B, Creeping filaments of Fosliella farinosa f. callithamnioides (1) and Pneophyllum confervicolum f. minutulum (2). Scale = 25 wm. C, Bisporangial crust showing domed conceptacles and ridged thallus (arrow). Scale = 250 um. D, Ostiole of bisporangial conceptacle. Scale = 10 um. E, Roof and spout of spermatangial conceptacle. Scale = 10 wm. conceptacle (Fig. 20C) shows a honeycomb-like surface under the S.E.M., with a rosette of cells (Fig. 20D) surrounding the ostiole (c.f. Garbary, 1978) and each conceptacle is borne on an individual segment of crust which forms a ridge in contact with confluent segments. The crust cells within these segments are relatively shorter than the characteristically elongated peripheral crust cells. VEGETATIVE ANATOMY: The crusts are composed of hypothallial and epithallial cells, the hypothallial cells are usually radially elongated in surface view while the large, rounded epithallial cells (Fig. 21A) frequently show a conspicuous central nucleus surrounded by radiating crooked canals. In VS the crust cells vary from square to considerably wider than high depending on the angle of the section while the epithallial cells vary from flat to dome-shaped or triangular. The terminal hypothallial trichocytes are bulbous (Figs 10A; 21B), and are usually 346 Y. M. CHAMBERLAIN divided by a wall which appears tangential and anticlinal in surface view but which is somewhat oblique in VS. The centrifugal part of the trichocyte usually bears a long hair (which may branch once, dichotomously) with a thin cell lumen. GAMETANGIAL PLANTS: Gametangial plants are usually monoecious although dioecious plants are found particularly during the winter. When a spermatangial or carpogonial conceptacle is starting to grow a small patch of thickened cells, some of which bear hairs, appears on the thallus surface. As this disc expands, both in width and depth, the original thallus surface peels off in strips (Fig 23A) until the surface of the mature conceptacle is composed entirely of newly developed cells (Fig. 23B) with a ring of trichocytes round the base. In monoecious plants the spermatangial conceptacles are adjacent to the carpogonial conceptacles (Figs 23A, B), their roof is composed of a layer of cells each of which bears an epithallial cell (Figs 22C; 23C), and the cells immediately surrounding the ostiolar spout (Fig. 20E) are radially elongated. Spermatan- gial initials are borne on the single layer of hypothallial cells which forms the conceptacle base (Fig. 23C) and each gives rise to spermatia from two points. There appear to be two types of spermatangial conceptacle: the larger type (Fig. 22C) measures c. 60-90 um diameter and has elongated spermatia measuring 7—13-5 wm long, while the smaller type (Fig. 23C) measures c. 30-50 wm diameter with rounded spermatia c. 2-3 wm diameter. Insufficient evidence is available to know whether these types are distinct or extremes of a variation range, but the larger form often occurs on dioecious plants. Carpogonial conceptacles (Figs 22A; 23A, B, D) are raised and when seen from above (Fig. 23B) the roof cells have a stepped appearance; trichogynes can be seen protruding from the ostiole. The procarps are borne in a bunch in the centre of the conceptacle floor (Figs 22A; 23D) and do not each bear more than a single carpogonium. CARPOSPORANGIAL CONCEPTACLES: Carposporangial conceptacles are regularly hemispherical (Figs 20A; 22B; 23E) and may have a smooth or stepped surface once the original thallus has been shed; in VS (Figs 22B; 23E) the roof is composed of a single layer of cells, each with an Fig. 21 Surface views of Fosliella farinosa crusts. A, Calcified crust showing hypothallial (h) and epithallial (e) cells. Scale = 20 um. B, Decalcified crust showing terminal trichocyte divided radially into a hair-bearing (arrow) and basal part. Scale = 20 wm. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 347 epithallial cell. The ostiole is sometimes entirely unelaborated but there is usually some development of downwardly-orientated ostiolar filaments (Fig. 23E) and sometimes a small hyaline collar develops round the ostiolar pore. In VS the fusion cell is usually narrow and deep (Figs 22B; 23E), it is often in open contact with the subtending cell (cf. Balakrishnan, 1947) and bears substantial remains of carpogonia on its upper surface. Gonimoblast filaments are borne peripherally and are about five cells long. Fig. 22 Vertical sections of Fosliella farinosa conceptacles. A, Carpogonial conceptacle. Scale = 20 um. B, Carposporangial conceptacle showing deep fusion cell (arrow). Scale = 50 wm. C, Large type spermatangial conceptacle. Scale = 20 um. D, Mature tetrasporangial conceptacle. Scale = 25 um. E, Immature bisporangial conceptacle. Scale = 25 wm. F, Mature bisporangial conceptacle (arrow) borne on one of five superimposed crusts. Scale = 100 wm. 348 Y. M. CHAMBERLAIN Fig. 23 Gametangial and tetrasporangial conceptacles of Fosliella farinosa. A and B, Surface views of developing gametangial conceptacles grown in culture — as seen under the light microscope. A, Young carpogonial (1) and spermatangial (2) conceptacle showing strips of original thallus surface covering parts of the conceptacles. B, Somewhat older conceptacles, after all the thallus has been stripped off, showing the stepped appearance of conceptacle roof cells, the triochocytes round the conceptacle base, and trichogynes protruding from carpogonial ostiole. C-—G, Vertical sections. C, Small type of spermatangial conceptacle. D, Carpogonial conceptacle. E, Carposporangial conceptacle. F, Young conceptacle showing tetrasporangial initials (shaded). G, Immature conceptacle showing incompletely divided tetrasporangia within hyaline sacs. All scales = 50 um. t = trichogyne; cg = carpogonium; s = sterile cell; tr = trichocyte. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 349 TETRASPORANGIAL PLANTS: Conceptacle shape and ostiole structure (Fig. 22D) are similar to that seen in the carposporangial conceptacle. The tetrasporangia are borne peripherally on stalk cells; after initiation (Fig. 23F) immature tetrasporangia (Fig. 23G) become deeply invaginated and do not stain deeply, but when mature they are farctate and stain deeply (Fig. 22D). There are up to about 8 tetrasporangia per conceptacle and there is usually some development of sterile cells in the centre of the conceptacle. BISPORANGIAL PLANTS: Bisporangial conceptacles are considerably larger than tetra- and carpo- sporangial conceptacles. They have a hemispherical roof (Figs 22E, F), composed of a layer of cells each with a large epithallial cell, the concavity of which gives the roof surface a honeycomb-like appearance under the S.E.M. (Fig. 20C), (see also Garbary, 1978, Figs 16 and 18). A ring of trichocytes has only occasionally been observed round the base of the bisporangial conceptacles and no ostiolar filament development takes place, although a characteristic rosette of cells can be seen round the ostiole in surface view (Fig. 20D). Twenty or more bisporangia are borne peripherally on stalk cells in each conceptacle and the young sporangia have a curious structure like two toadstools joined by their stalks (Fig. 22E, see also Chamberlain, 1977b). Mature tetrasporangia are farctate and stain deeply (Fig. 22F). PHENOLOGY: Fosliella farinosa occurs throughout the year on the shores of the British Isles but, while bisporangial plants occur at all seasons gametangial plants are less common and occur mainly in August and September in warmer areas such as Dorset and southern Ireland. When they occur in winter, gametangial plants are usually abnormal and apparently infertile (Cham- berlain, 1977b, Figs 8, 9) — a phenomenon also observed by Rosanoff (1866, see Fig. 6 [12]). These observations reflect the geographical variations in this species in that in warmer areas such as the Mediterranean only gametangial/tetrasporangial plants occur, while bisporangial plants (with diploid bispores) are known only from northern France and the British Isles. ForM RANGE: In addition to the reproductive variations with the season, the crust cells of F. farinosa show seasonal differences. Typical crust cells are considerably elongated and the epithallial cells are large and rounded; such cells occur during the summer when the crusts are growing rapidly. In winter, however, the crust cells are smaller in all dimensions (Tab. 6), relatively shorter, and more heavily calcified, while the epithallial cells are smaller and more ovate. Trichocytes are abundant during warm, light conditions but absent in winter and in deeper subtidal habitats. DISTRIBUTION: British Isles: Fosliella farinosa occurs throughout the British Isles, most commonly in southern England and Ireland. The most northerly record of tetrasporangia/gametangia is from Sea Mill, Ayrshire, Scotland where it was collected in August 1890, growing on Zostera, by E. M. Holmes (BM). In general, northern populations are probably mainly bisporangial. World: Recorded from most temperate and tropical parts of the world although, as already mentioned, the taxonomy is somewhat confused. The British Isles records represent the most northerly locality in Europe since the species appears to be absent from Denmark, Sweden, and Norway. HasiratT: In warmer areas of the world F. farinosa is an abundant epiphyte on a wide range of algae and marine angiosperms on which it forms a dense powdery covering of closely crowded crusts smothered with conceptacles; in the intertidal zone the crusts are usually bleached. This type of covering is not usually seen in the colder parts of Britain, but has been noted in warmer localities such as Kimmeridge and Lough Ine. F. farinosa is also common on many algal hosts as well as on Zostera in the warm waters of the Fleet in Dorset. F. farinosa is, however, present throughout the year in most parts of the British Isles as mauvy-pink, often extensive, confluent crusts on many intertidal algae, but particularly on the tophules of Cystoseira nodicaulis and the fronds of Chondrus crispus. It also frequently forms a 350 Y. M. CHAMBERLAIN Table6 Dimensions of vegetative structures (in zm unless stated). British Hamel & Masaki & Ganesan Fosliella farinosa Isles Lemoine (1953) Tokida (1960) (1971) crust diameter up toc. 5mm 24mm 1-5mm crust depth ca. 20 crust cell length (14) 20-5 (28-5) (9) 10-20 (22) 12-20 (8-5) 14 (20)* surface width (5) 10 (16-5) (5) 7-12 (18) 5-9 (5) 7:5 (11) crust cell height 9-12 8-14 5-13 epithallial cell length (7) 9 (13-5) rounded (2-5) 4 (6)* surface view —_ width (3) 7 (10) (2) 4 (6)* epithallial cell height c. 5-8 hypothallial cell width 7-13 15-30 VS height 5-13 (6) 9-12 (15) perithallial width cell height VS number trichocyte type terminal terminal terminal terminal trichocyte length (30) 35 (43) 17-30 (43) _ 20-25 14-25 (35) surface view width (12) 16 (20) 12-18 (30) 7-10 (6) 8-12 (15) RIDGWAY COLOUR CODE: dry crust — light vinaceous lilac, Pl. XLV Table 7 Dimensions of reproductive structures (um unless stated). Fosliella farinosa British Hamel & Masaki & Ganesan f. farinosa Isles Lemoine (1953) Tokida (1960) (1971) CARPOSPORANGIAL CONCEPTACLE external diameter 140-200 (250) (29) 105-126 218-300 surface diameter (91) 99 (104) VS internal diameter (62) 85 (110) (29) 105-126 110-145 VS height (52) 63 (83) (25) 42-63 (91) 58-87 VS roof height (13) 16 (18) roof type irregular no. basal cell rows 1 ostiole simple or small tiny spout corona no. gonimoblast filaments 5-10 diameter carpospore c. 30 SPERMATANGIAL CONCEPTACLE VS/surface width (33) 76 (91) 60-80 (42) 55-84 VS height (26) 37 (44) (29) 33-46 (55) length spermatium (7) 10 (13-5) 3-7 width spermatium 1-5-3 2 (or rounded 2-3 wm diam.) * dimensions in winter-collected material FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES Jol TETRASPORANGIAL CONCEPTACLE external diameter 140-200 (250) (84) 126-168 (189) 230-300 surface (78) 114 (130) VS internal diameter (65) 86 (94) VS height (52) 66 (78) 63-126 VS roof height (10) 15 (20) roof type irregular no. basal cell rows 1 ostiole simple or small tiny spout corona no. tetrasporangia c. 8 3-7 4-7 length tetrasporangium (36) 42 (65) 50-90 40-80 32-58 width tetrasporangium (23) 30 (39) 30-50 (20) 25-50 (63) 35-44 BISPORANGIAL CONCEPTACLE external diameter 140-200 (250) surface diameter (156) 193 (216) VS internal diameter (166) 187 (208) VS height (45) 89 (125) VS roof height (18) 30 (39) roof type zonal no. basal cell rows 1 ostiole simple rosette no. bisporangia 20+ length bisporangium (54) 65 (78) 45-55 width bisporangium (26) 34 (42) bloom, together with Pneophyllum limitatum, on a range of intertidal algae such as Palmaria palmata, Laminaria spp. and Fucus serratus from September until about November, depending on the weather conditions (Chamberlain, 1982). Herbarium specimens such as Le Jolis’s (exsiccata no. 282 on Rhodymenia palmata 30 November 1853 [CHE]) indicate that a similar bloom occurs in northern France. Lire History: A self-perpetuating bisporangial life history was established for Fosliella farinosa in laboratory culture (Chamberlain, 1977b). In addition, tetraspores were germinated and grown in von Stosch culture medium to produce monoecious crusts bearing spermatangial and carpogonial conceptacles (Figs 23A, B), but the carpogonia apparently remained unfertilised and the carposporangial phase did not develop. 1b. Fosliella farinosa forma callithamnioides (Foslie) Y. Chamberlain, comb. nov. BasionyM: Melobesia farinosa f. callithamnioides Foslie in K. nor. Vidensk. Selsk. Skr. 1905 (3): 96 (1905), non M. callithamnioides (P. Crouan & H. Crouan) Falkenb. NOMENCLATURAL SYNONYMS: Melobesia solmsiana Falkenb. in Fauna Flora Golf Neapel 26: 109 (1901). Melobesia farinosa f. solmsiana (Falkenb.) Foslie in K. nor. Vidensk. Selsk. Skr. 1908 (7): 16 (1908). REFERENCES: Melobesia callithamnioides Falkenb. in Mitt. zool. Stn Neapel 11: 265 (1879); Hauck, Die Meeresalgen: 262 (1885); Solms-Laubach in Fauna Flora Golf Neapel 4: 11 (1881); Heydrich in Ber. dt. Boi Ges. 15: 44 (1897). Melobesia farinosa f. callithamnioides (Falkenb.) Foslie; Zinova, Check-list of green, brown and red algae of the southern seas of U.S.S.R.: 229 (1967). Melobesia farinosa var. callithamnioides Foslie; Newton, A handbook of the British seaweeds: 301 (1931). Melobesia farinosa f. solmsiana (Falkenb.) Foslie; Lemoine in Dansk. bot. Arkiv. 3: 171 (1917); Hamel & Lemoine in Archs Mus. natn. Hist. nat. Paris VII, 1: 102 (1953[‘1952’]); Cabioch, Jn Feldmann in Trav. Stn biol. Roscoff 11, 15: 17 (1964). Fosliella farinosa var. solmsiana (Falkenb.) Feldmann in Rev. algol. 11: 316 (1942 [‘1939’]); Taylor, Marine ra 04 Y. M. CHAMBERLAIN algae of the eastern tropical and subtropical coasts of North America: 389 (1960); Cabioch in Bull. Soc. phycol. Fr. 19: 78 (1974); Bressan in Boll. Soc. adriat. Sci. nat. 59: 79 (1974). EXCLUDED RECORDS: Hapalidium callithamnioides P. Crouan & H. Crouan in Annals Sci. nat. Bot. TV, 12: 287 (1859). Guerinia callithamnioides (P. Crouan & H. Crouan) Picquenard in Trav. scient. Lab. Zool. Physiol. marit. Concarneau 4 (3): 1 (1912) (= Pneophyllum confervicolum f. minutulum q.v.). TYPE SPECIMEN: See text. Diacnosis: Differing from f. farinosa in having filamentous thalli. DEscriPTION: Forming creeping, filamentous thalli, with gametangial and tetrasporangial conceptacles only. Historica: Fosliella farinosa f. callithamnioides was first recorded from the Gulf of Naples by Falkenberg (1879) as Melobesia callithamnioides (Crouan frat.) since he thought it was the taxon described by the Crouans (1859) as Hapalidium callithamnioides. Solms-Laubach (1881) subsequently recorded and figured the taxon under the name Melobesia callithamnioides. Later, Falkenberg (1901) concluded that his own and Solms-Laubach’s material was a different taxon from that described by the Crouans and he therefore erected a new species, Melobesia solmsiana, to accommodate the entity. Foslie (1905a), however, considered Falkenberg’s species to be a form of Melobesia farinosa and recorded it as ‘M. farinosa f. callithamnioides (Falkenb.) Foslie’. The attribution to Falkenberg is, however, incorrect, because he was simply using the Crouans’ taxon, as explained above. Since Foslie’s (1905a) publication was the first use of the name for the rank of form (and which excluded Hapalidium callithamnioides as a synonym) the basionym should be cited as Melobesia farinosa f. callithamnioides Foslie (1905a). Foslie refers to Falkenberg’s and Solms- Laubach’s publications but apparently did not see their material, and I have not been able to locate either of their specimens. The lectotype could therefore either be chosen from among the specimens which Foslie had seen (Wille-Mediterranean, Sauvageau—Tenerife; Howe- Caribbean) or Solms-Laubach’s illustration could be used for this purpose. Since Falkenberg (1901) clearly agreed with Solms-Laubach’s identification and publication, the last solution seems preferable. GENERAL COMMENTS: As previously remarked (p. 333) it is possible that the creeping filaments (Fig. 20B) of f. callithamnioides may not be genetically distinct from f. farinosa, nevertheless this growth form has been widely reported, particularly from warm water areas such as the Mediterranean. Since its appearance is so distinctive it is considered valuable to retain its identity as anamed forma. DISTRIBUTION: British Isles: Co. Cork. World: Mediterranean; Caribbean; southern U.S.S.R. Hasirar: Plants of f. callithamnioides (Fig. 20B) have been noted on only one occasion — at Lough Ine growing on turf-like communities of Gelidium pusillum and other algae. The creeping forms of Fosliella farinosa occur principally in warmer waters and Lough Ine shore conditions bear some resemblance to those found in the Mediterranean, as indicated by the fact that. southern Ireland is the only British Isles habitat for the sea urchin Paracentrotus lividus (Lamarck). This sea urchin is common in the Mediterranean (Goss-Custard et al., 1979). II. Pneophyllum Pneophyllum Kiitz., Phycologia generalis: 385 (1843). Type species: Pneophyllum fragile Kiitz., Phycologia generalis: 385 (1843). Hotortype: L! no. 941.241.152, Herb. Kiitzing, from the Mediterranean, growing on Sphaerococcus coronopifolius (Fig. 24). FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 353 Synonymy: There are no nomenclatural synonyms of Pneophyllum, but Hapalidium sensu P. Crouan & H. Crouan (1859) non Kitz. , and Heteroderma Foslie (1909) are taxonomic synonyms. Description: Thallus calcified, crustose, either epiphytic on algae and marine angiosperms, or epilithic on stones, glass and rocks, or epizoic on shells; vegetative thallus either composed of hypothallium and epithallium only or with perithallial filaments up to 12 or more cells deep, cell fusions frequent, hypothallial trichocytes (when present) normally intercalary, occasional abnormal very large terminal trichocytes present. Gametangial plants monoecious, carpogonial ZING. ' Fig. 24. Type specimen of Pneophyllum fragile (L no. 941.241.152); the frond bearing the Pneophyllum crusts is on the left. Scale = 10 mm. 354 Y. M. CHAMBERLAIN conceptacles raised to immersed, with central ostiole through which trichogynes protrude, fertile procarps bearing one or two carpogonia; spermatangial conceptacles immersed, ostiole prolonged into a spout. Carposporangial conceptacles prominent or immersed, ostiole simple, or with corona of free filaments, or with filaments fused into a collar, funnel or dome, roof uniform, irregular or zonal; fusion cell not in open connection with subtending thallus cells. Tetrasporangial plants with single-pored conceptacles similar to carposporangial conceptacles, tetrasporangia borne peripherally on stalk cells, usually some development of central sterile cells. Bisporangial plants with uninucleate (presumably diploid) bispores, conceptacles similar to tetrasporangial ones. Spore germination disc with central element of eight cells, the four central cells initiating one hypothallial filament each, the four peripheral cells initiating two hypothallial filaments each. HistToricAL: The genus Pneophyllum, in the family Spongitae (i.e. crustose corallines), was proposed by Kiitzing (1843) to accommodate the single species P. fragile, which had a thallus of two cell layers; the upper, cortical cells (i.e. epithallium) were much smaller than the lower layer of somewhat rounded cells (i.e. hypothallium). This distinguished it from: a) Hapalidium which Kiitzing thought had only a single cell layer; b) Melobesia (i.e. Dermatolithon) which had very elongated lower cells; and c) Spongites (i.e. complex crusts including Lithothamnium and Lithophyllum) with many cell layers. For some reason the name Pneophyllum has never been taken up, possibly because the status of the single species has remained obscure. However, Kitzing’s specimen (Fig. 24) is in good condition, and can be assigned without doubt to the taxon under discussion. On grounds of priority, therefore, the name Pneophyllum is applied to the present genus. Further discussion on the interpretation of Kiitzing’s work on simple crustose coralline algae is given on p. 298, while his specimen of P. fragile is described here in detail for the first time. Species belonging to Pneophyllum, as now defined, have been mainly assigned to Melobesia since 1843; other genera to which they have been attributed include Hapalidium, Derma- tolithon, Lithocystis, Heteroderma, Guerinea, and Fosliella (Tab. 1). The nomenclature of species now assigned to Pneophyllum suffered numerous changes in the course of Foslie’s publications and these are demonstrated in Table 3. Foslie’s (1909) final circumscription of Heteroderma, however, as simple crustose species lacking trichocytes is essentially the same as that of Pneophyllum. Foslie recognised the terminally situated hyaline cells of Fosliella farinosa (which he assigned to the monotypic genus Melobesia) as trichocytes; he did not consider the hyaline intercalary cells (which he observed in some species of Heteroderma) to be essentially the same type of structure. SYSTEMATICS: About 20 species of Pneophyllum have been described mostly under the generic names Fosliella and Heteroderma. The species vary from those with only a hypothallium and epithallium in the vegetative parts of the thallus (e.g. P. lejolisii and P. confervicolum) to thalli up to 12 or more cells deep (e.g. Fosliella chamaedoris (Foslie & M. Howe) E. Dawson (1960, p. 54) and P. plurivalidum (q.v.). The relationship of Pneophyllum to the more complex crustose mastophoroideaen genera, such as Neogoniolithon and Porolithon, which have many hypothallial layers and a deep perithallium, is not yet clear. There is, however, a group of species which may represent an intermediate stage between Pneophyllum and these genera. These species are simpler than Porolithon and Neogoniolithon, have a single or a few hypothallial layers and a more developed perithallium than is seen in Pneophyllum. The species are at present grouped in Pseudo- lithophyllum sensu Adey (see Fig. 16). Adey (1970) lists 22 species in the taxon, but information about many of these is sparse, and there is a lack of knowledge as to whether the tetrasporangial conceptacles are single- or many-pored. Until more information is available and the nomenc- latural tangle between Pseudolithophyllum sensu Lemoine, a lithophylloid genus, and the mastophoroid Pseudolithophyllum sensu Adey is resolved, it is not beneficial to theorise further on the possible relationships amongst these genera. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES Sake, Eco.ocy: Pneophyllum probably occurs mainly as an epiphyte on other algae and sea-grasses; it also, however, occurs epilithically and epizoically, and may be common in these forms but very few data are at present available. Probably the most familiar association which includes Pneophyllum species is the sea-grass epiphyte association, which has been studied by workers such as van der Ben (1969) in the Mediterranean and Brauner (1975) in North Carolina. The species most commonly found on sea-grasses is P. lejolisii which probably occurs in most parts of the world. DIsTRIBUTION: Pneophyllum probably occurs in seas throughout the world outside the polar regions. Key to species of Pneophyllum in the British Isles 1 (1) 302) 43) 5(4) 6(2) 7(6) 8(7) 9(6) 10(9) 11(1) 12(11) 13(11) 14(13) MN tae sh cece a sae nec eh aang Ss enone ep ainaN Maas nien es sec aaaas LAA fate ei pana nate fe Bpilitisie OF GOLZOIC ccs scooter ack ls ta nh cess daeem ceva Spe eT Pena oes an atnE dence na teT ate Vegetative crust of hypothallium and epithallium only ............0...0cceceeeeeeeeeeeeeeeeeeeeen ees Vegetative crust, at least in older parts, with perithallium .................. ce ceeeeeeceeeeeeeeeee ees Conceptacles flattened, ostiole simple, occurring on Zostera only ........... 2. P. lejolisii(p. CONCED PACES PI OTC INE ee cr cos casites sees eno ns oh stants Cae de cup ate alg adc Rau eas nese oa peee Conceptacles with long, free, ostiolar filaments ..................... cee 3. P. rosanoffii(p. Conceptacles not with long: tree, ostiolar filamentsi.i0 0032.20.00. a ashss eee sac suse deneee oes cew ease Conceptacles less than 100 wm internalin diameter, ostiolesimple 5. P. confervicolum (p. Conceptacles more than 100 um internal in diameter, ostiole funnel- REBTIO ck Fase tees ace tie opines cae peated tus Cece eee at ta tyne Rake 4. P. limitatum (p. Conceptacles with simple GstOles 5 ccna coves ees ces aoe molen tps ven vie va cccr ed anee ne borauntenNet Conceotacies with ostiolar Rlaments Or collar 304.2) ync 20% —o.- S00 e a hy ie. es e. w, allUEG \ LIIEL DU = 38 = See 5S OO i Ce —_———— Fig. 32 Vertical sections of Pneophyllum lejolisii conceptacles. A, Carposporangial conceptacle. B, Tetrasporangial conceptacle. C, Young and mature bisporangial conceptacles with bisporangia lying horizontally. D, Large bisporangial conceptacle with bisporangia + vertical. tetrasporangia are borne peripherally on stalk cells and their development is described and illustrated by Suneson (1937); he shows that central, sterile cells are present in the young conceptacle, but may be absent in mature conceptacles. Mature tetrasporangia are large in relation to the small size of the conceptacle. In epilithic crusts the conceptacles measure 70-80 um internal diameter, with 5—6 tetrasporangia per conceptacle which measure about 45-50 wm long. BISPORANGIAL PLANTS: Bisporangial plants are similar to carposporangial and tetrasporangial plants. Bisporangial conceptacles (Figs 30B; 32C, D) are the largest type found in P. /ejolisii (see Chamberlain, 1982) and show a general gradation in size, with no suggestion of forming two 366 Y. M. CHAMBERLAIN groups as is seen with tetrasporangial and carposporangial conceptacles. The bisporangia are somewhat globular and, like the tetrasporangia, are very large in relation to conceptacle size. In the smaller conceptacles the bisporangia lie horizontally (Fig. 32C), while they are more or less vertical in the larger conceptacles (Fig. 32D). Bisporangia and tetrasporangia are sometimes found in the same conceptacle (Fig. 30E), particularly in the autumn (Chamberlain, 1982). PHENOLOGY: P. lejolisii is the only species of Pneophyllum investigated which has a pronounced seasonal cycle of presence and absence. Crusts start to appear on intertidal Zostera at Bembridge in July and disappear again between January and April depending on the weather. The only exceptions are the epilithic crusts collected in May which had presumably been cast from the subtidal zone; Rosenvinge’s (1917) and Suneson’s (1943) results also indicate that crusts appear earlier in the subtidal in Denmark and Sweden since they record their presence in May and June respectively. Reports from other parts of the world show that P. lejolisii occurs seasonally elsewhere; van der Ben (1969, as Melobesia) records its presence on Posidonia in the Mediterranean from January till October, and Brauner (1975, Tab. I, as Heteroderma) shows that, while it is present throughout the year except in June on Zostera marina in North Carolina, it is relatively scarce from March till May. Very little difference was seen within populations in the relative abundance of different conceptacle types throughout the season, although bisporangial conceptacles increased in number slightly at two sites at Bembridge as the season progressed (Chamberlain, 1982). There was, however, a sharp difference in the proportion of conceptacle types at different sites at Bembridge, with two populations having mainly tetrasporangial and carposporangial concep- tacles and a third site predominantly bisporangial conceptacles. This difference was maintained throughout three successive seasons. ForM RANGE: As remarked by Suneson (1943) and by Bressan et al. (1977), P. lejolisii shows much variation even at one locality and like these authors, I have found considerable differences in crust cell size both within and between populations. The dimensions given for crust cell size in Table 10 were made on material from Ireland and England collected in August and September, which is probably the optimum season for growth and reproduction. Cells from material collected in November at Bembridge, however, are relatively longer and thinner and measure (7) 11 (16) wm long x (5) 8 (13) wm wide; in addition trichocytes are abundant under warm, well-illuminated conditions, but disappear as winter progresses. As has been discussed, there is a very considerable range of form and size within each conceptacle type and differences are seen in other areas. Masaki (1968, as Fosliella), for example, describes the conceptacles of Japanese samples as being convex or hemispherical. DISTRIBUTION: British Isles: Northumberland, Essex, Hampshire, Dorset, Devon, Cornwall, Anglesey, Ayr- shire, Orkney, Jersey, Co. Clare, Co. Cork. World: Norway, Sweden, Denmark, Netherlands, France (north and Mediterranean), Italy, Corsica, Canary Islands, Canada, U.S.A. (east and west coasts), Caribbean, Pacific Mexico, Japan, southern U.S.S.R., Australia. Hasirat: As has been discussed, P. lejolisii occurs predominantly as an epiphyte on Zostera and other sea-grasses; during the present study it has been found twice growing epilithically on glass and china. Records of its occurrence as an epiphyte on algae require confirmation. During the present investigation, detailed study (Chamberlain, 1982) has been restricted to crusts growing on Zostera marina from the intertidal zone. Here the Zostera clumps have either been situated in pools or runs which never dry out completely at low tide since, in common with all other Pneophyllum species examined, P. /ejolisii cannot withstand desiccation. For this reason, it is also more common on Zostera blades growing at the edge of clumps than on those on the surface which are more likely to dry out. P. lejolisii also grows on subtidal angiosperms in the British Isles and elsewhere (e.g. Weber van Bosse, 1886, Netherlands; van der Ben, 1969, Mediterranean). FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 367 At Bembridge, P. limitatum occurs with P. lejolisii on Zostera, but is very much less abundant and mostly recorded early in the season, occasional crusts of Fosliella farinosa have also been found. In localities such as Falmouth in the west of England, Pneophyllum rosanoffii may grow with P. lejolisiion Zostera, while in Ireland the two species seem to be equally common, growing either together or in pure stands. The occurrence of Pneophyllum lejolisii in northern Europe suffered a check in the 1930s when a wasting disease destroyed many of the Zostera populations (Tutin, 1938). However, in Sweden the epiphyte apparently became plentiful again as soon as the Zostera started to recover (Suneson, 1943), and Davey’s (1943) record from Wales indicates that this had probably also occurred in the British Isles. Table 10 Dimensions of vegetative structures (in wm unless stated). British Hamel & Suneson Masaki Pneophyllum lejolisii Isles Lemoine (1953) (1943) (1968) crust diameter up to2mm 0-5 mm 2-4 mm crust depth €.30) crust cell length (3-5) 7-5 (11-5) 6-9 (15) (6) 8-15 8-13 surface width (6:5) 9 (12) 6-10 (5) 6-10 5-10 crust cell height co15 7-11 7-11 epithallial cell length (1-5) 2-5 (3-5) 2-3 surface view = width (3-5) 5-5 (8) 3-7 epithallial cell height 4-5 hypothallial cell width VS height perithallial width 7-10 6-7 cell height 17-20 7-20 VS number trichocyte type mainly intrafilament intrafilament intrafilament and branch cell and branch cell trichocyte length (11) 13 (16) 13 11-16 10-12 surface view width (8-5) 10 (13-5) 6-10 6 3. Pneophyllum rosanoffii Y. Chamberlain, sp. nov. REFERENCES: Melobesia farinosa auct. div., Le Jolis, Liste des algues marines de Cherbourg: 150 (1863); Kiitzing, Tabulae phycologicae: 34 Tab. 95c, d (1869). Melobesia lejolisii Rosanoff pro parte in Mem. Soc. imp. Sci. nat. Cherbourg 12: 62 Pl. I, Figs 1-3, 5-7, 11 (Fig. 2 of present study) (1866). Melobesia lejolisii auct. div.; Johnson & Hensman in Scient. Proc. r. Dubl. Soc. Il, 9: 27 (1899); Rosenvinge in K. danske Vidensk. Selsk. Skr. I1, 7: Figs 158D, E only (1917); Mazza, Saggio di algologia oceanica 3: 1116 pro parte (1922); Wuitner in Ann. Assoc. naturalistes de Levallois-Peret 20: 58 (1931). SPECIMENS: Pneophyllum fragile Kiitz.; Herb. Kiitzing (L no. 941.241.153!), on Zostera from Adriatic, mixed with other species. Melobesia lejolisii; Herb. Rosenvinge (C!), no. 6246, Sanden, Denmark, 5 July 1895, on Zostera. Hotoryre: CHE! Herb. Le Jolis no. 1200, 31 December 1857, Cherbourg, growing on Zostera marina! (It is to be found in Le Jolis’s Melobesia lejolisii folder and is illustrated in Fig. 33). Duplicates of no. 1200 have also been labelled Le Jolis 283 (CHE!), but were apparently never distributed. Diacnosis: Species epiphytica perithallio prope conceptacula profunditate usque ad 5 cellulas et filamentis ostiolaribus longis liberisque; a P. sargassi, P. lejolisii, P. limitato et P. concollum filamentis ostiolaribus liberis, et a P. zonali crusta tenuiore et conceptaculis minoribus differt. Descriptio: Crustae laeves, roseae ad pallido roseae, diametro usque ad 4 mm, profunditate 50 um, perithallium prope conceptacula profunditate usque ad 4 cellulas, cellulae perithallii (in sectione verticali) 9-18 wm latae, 12-5—29 wm altae, crusta vegetativa ex hypothallio et epithallio modo constata, 368 Table 11 Dimensions of reproductive structures (4m unless stated). W. M. CHAMBERLAIN British Hamel & Suneson Masaki Pneophyllum lejolisii Isles Lemoine (1953) = (1937, 1943) (1968) CARPOSPORANGIAL CONCEPTACLE external diameter surface diameter (35) 77 (99) 100-175 VS internal diameter (59) 70 (91) 60-100 60-80 VS height (22) 34 (40) 35-60 30-60 (25) 38-50 VS roof height (6) 8 (9) 10-13 roof type uniform no. basal cell rows 1 1 1 1 ostiole simple simple simple simple no. gonimoblast filaments 5-16 c.6 diameter carpospore (20) 26 (32) SPERMATANGIAL CONCEPTACLE VS/surface width (13) 34 (41) 40-55 40-55 21-33 VS height (12) 16 (21) 21-33 length spermatium 344 2-4 width spermatium 1-5-2 1 TETRASPORANGIAL CONCEPTACLE external diameter surface diameter (52) 102 (135) VS internal diameter (75) 92 (125) (90) 125-200 (250) 65-130 75-100 (120) VS height (30 42 (52) 45-80 40-90 (24) 43-63 VS roof height (6) 9 (13) 10-13 roof type uniform no. basal cell rows 1 | 1 i ostiole simple simple simple simple no. tetrasporangia 4-11 1-10 1-10 (20) length tetrasporangium (31) 50 (70) 45-80 (25) 30-50 width tetrasporangium (18) 33 (49) 30-50 (13) 23-30 (48) BISPORANGIAL CONCEPTACLE external diameter surface diameter (70) 114 (140) VS internal diameter (91) 101 (130) 65-130 VS height (33) 49 (65) 40-90 VS roof height (7) 10 (13) roof type uniform no. basal cell rows 1 1 ostiole simple simple no. bisporangia 3-14 5-20 length bisporangium (31) 51 (65) width bisporangium (17) 35 (52) cellulae crustae aspectu superficiali rectangulares, 5-5—15-5 wm longae X 4-5-10-5 wm latae, cellulae epithallii 2-5 wm longae X 2-9-5 wm latae, trichocyti et intra filamenta et ramos facientes frequentes, amplitudine eadem ac cellularum crustae; conceptacula, praeter ea spermatangialia, prominentia, ostiola in statu vivo nitentia, filamentis ostiolaribus longis liberis circumcincta, conceptacula carpospor- angialia diametro interno 84-156 wm, altitudine 45-117 wm, conceptacula spermatangialia immersa, 25-50 um lata x 19-46 wm alta, ostiolo in canalem producto, conceptacula tetrasporangialia diametro interno 104-164 um, altitudine 70-164 wm, usque ad 20 tetrasporangiis in unoquoque conceptaculo, tetrasporangia 41-5—78 wm longa X 26-45-5 um lata. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 369 - Z z 4 SP. de tn ! * ‘e esd i «. oi @ ial 6 ’ a Dy ae Fo Fig. 33 Type specimen of Pneophyllum rosanoffii (= Melobesia lejolisii Rosanoff pro parte). A, Melobesia lejolisii, Le Jolis No. 1200, Herb. Le Jolis, Cherbourg; Rosanoff’s handwriting (1), Le Jolis’s handwriting (2). B, Scanning electron micrograph of crusts from A; ostiole filaments indicated by arrows. Scale = 100 um. 370 Y. M. CHAMBERLAIN DiaGnosis: Epiphytic species with perithallium near conceptacles up to 5 cells deep and long, free ostiolar filaments: differing from P. sargassi, P. lejolisii, P. limitatum, and P. concollum in having free ostiolar filaments, from P. zonale in having a thinner crust and smaller conceptacles. DESCRIPTION: Crusts deep to pale pink, smooth, up to 4 mm diameter, 50 wm deep, perithallium up to 5 cells deep near conceptacles, perithallial cells (in VS) 9-18 wm wide X 12-5-29 um high; vegetative crust of hypothallium and epithallium only, crust cells in surface view rectangular, 5-5-15:5 wm long X 4-5-10-5 wm wide, epithallial cells 2-5 wm long X 2-9-5 um wide, intrafilament and branch cell trichocytes common, of same size as crust cells; conceptacles (except spermatangial) raised, ostiole shiny when living, surrounded by long, free ostiolar filaments, carposporangial conceptacles 84-156 jm internal diameter X 45-117 um high, spermatangial conceptacles immersed, 25-50 xm wide X 19-46 ym high, ostiole prolonged into a spout, tetrasporangial conceptacles 104-164 um internal diameter x 70-164 um high, up to 20 tetrasporangia per conceptacle, 41-5—78 wm long x 26—45-5 wm wide. HistoricaL: This taxon was first described by Le Jolis (1863, p. 150) as Melobesia farinosa growing on Zostera, having flattened conceptacles with ostioles surrounded by hairs. Le Jolis published it as Melobesia farinosa because he had sent some of the material to Kiitzing who had identified it as this species. The collection on which Le Jolis’s description was based was ‘A Le Jolis, no. 1200, Cherbourg, 31 Décémbre 1857’ (Fig. 33). Several samples from this collection are in Herb. Le Jolis (CHE) and the sample sent to Kiitzing is in Leiden, no. 941.156.75; all have been examined during the present investigation. The species was illustrated by Kiitzing (as Melobesia farinosa) in Tabulae Phycologicae (Kiitzing, 1869, p. 34, Tab. 95c, d). Rosanoff (1866: 62) considered that the entity differed from M. farinosa and described a new species M. lejolisii, based partly on this material. As discussed on p. 305 however, three distinct components were included in Melobesia lejolisii Rosanoff, and as one of these (the entity with flat-topped conceptacles lacking ostiolar filaments) has come to be widely accepted as lejolisii, it seems preferable to continue to apply the epithet to this entity and give a new name to the present species. The name rosanoffii has been chosen to commemorate the author of the first detailed description of the species. Although this appears to be a common European alga, the only records (since Rosanoff’s) of its occurrence which have been traced are those of Johnson & Hensman (1899: 27) from Ireland, Rosenvinge (1917, Fig. 158D, E only) from Denmark, Mazza (1922) from Italy, and Wuitner (1931) from France (all as M. lejolisii). Further discussion of Rosanoff’s original description is given on p. 304. Le Jolis’s specimen no. 1200 is chosen as the holotype and the species description is based on this material, together with British material. Information is also given (Tab. 12) about material gathered from Roscoff (Brittany) in order to gain further insight into the variation of Pneophyllum rosanoffii in France. CRUST APPEARANCE: P. rosanoffii forms rather thin crusts (up to 50 wm deep) which grow epiphytically on Halopitys incurvus, Palmaria palmata, and Zostera marina. When fresh, crusts are a somewhat translucent brownish pink, while dried crusts appear solid and bright rosy pink to brownish or greyish pink depending on the conditions under which they had been growing. The crusts measure up to about 4 mm diameter. They adhere firmly to Palmaria and Zostera, but on dried Halopitys they maintain their rigidity as the host plant shrinks, so that they are liable to flake off or form a sleeve encircling the host but becoming detached. The somewhat raised conceptacles are easily distinguished in fresh material by having a conspicuously shiny ostiole. Under the S.E.M. (Fig. 34B) the crust cells are rectangular with conspicuous epithallial concavities; branch cell and intrafilament (Fig. 34B) trichocytes are present and bear epithallial concavities. The tetra- and carposporangial conceptacles are somewhat raised, they have uniform roofs and long ostiolar filaments which are clearly visible in mature crusts (Fig. 34A); it is these that impart the characteristically shiny appearance to the ostiole in fresh material. Figures 34C-E show the development of the ostiolar filaments in air dried material, while Figure FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES ee hl : : te - a ill ” + 4 Fig. 34 Scanning electron micrographs of Pneophyllum rosanoffii. A, View of critical point dried crust surface showing spermatangial (s) ostioles and carposporangial conceptacles with ostiole filaments (arrow). Scale = 250 um. B, Critical point dried crust cells with intercalary trichocyte (arrow). Scale = 25 um. C-E, Stages in the development of ostiole filaments. C, Ostiole filaments starting to emerge (arrow). Scale = 100 um. D, Ostiole filaments emergent but still fused. Scale = 10 um. E, Ostiole filaments free. Scale = 10 wm. F, Critical point dried ostiole filaments from above, showing their terete structure. Scale = 5 wm. G, View looking up into the conceptacle roof showing the basal parts of ostiole filaments (arrow). Scale = 25 wm. H, Spermatangial conceptacle with spout. Scale = 10 wm. 34F shows the terete nature of the filaments in critical point dried material. Spermatangial conceptacles (Figs 34A, H) can be seen beside the carposporangial conceptacles. VEGETATIVE ANATOMY: The vegetative crust is composed of epithallium and hypothallium only. In surface view the crust cells (Fig. 35A) are robust, rectangular, and particularly well pigmented, and the epithallial cells are conspicuous and rectangular; the germination disc is sometimes visible in mature crusts. Intrafilament trichocytes occur quite commonly; they appear always to bear epithallial cells. In Irish and Roscoff material both intrafilament and branch cell trichocytes are present and may bear long hairs. Trichocytes on British material are of a similar size to the surrounding crust cells, but those in the Irish and Roscoff material are noticeably 372 Y. M. CHAMBERLAIN longer. A perithallium up to five cells deep develops in the region of the conceptacles, the cells vary from square to vertically elongated in vertical section (Figs 12A—H). GAMETANGIAL PLANTS: The crusts are monoecious with spermatangial conceptacles adjacent to carpogonial conceptacles (Figs 34A; 36C). The spermatangial conceptacles are small, im- mersed, flask-like chambers and the ostiole is prolonged into a spout (Figs 12H; 34H). The carpogonial conceptacles (Fig. 36A) are shallow and wide; their development (Figs 12A—C) follows the normal pattern and the fertile procarps (Fig. 12C) bear one or, more rarely, two carpogonia, with long trichogynes. CARPOSPORANGIAL CONCEPTACLES: The stages in the development of carposporangial concep- tacles are shown in Figures 12E-G and 36B. The conceptacles of P. rosanoffii are neat and regularly symmetrical. The roof is composed of a single, rather regular layer of cells each with an epithallial cell; it is gently rounded (Figs 34A; 36B) and has.a uniform upper surface. The downward ostiole filaments are quite well developed, whilst the upper filaments are prolonged into a corona of tentacle-like filaments (e.g. Figs 12F; 36B) which are about 50 wm long when mature. This is very characteristic of the species, it is important, however, to appreciate that young and senescent plants may not necessarily show this feature. The filaments can also be seen in whole crust mounts by focusing up in the ostiole area (Fig. 35B) while their basal parts are visible in the conceptable roof under the S.E.M. (Fig. 34G). The fusion cell (Figs 12E, F) is quite broad, and the peripheral carposporangial filaments are composed of small cells, while the carposporangium is relatively large. TETRASPORANGIAL PLANTS: Tetrasporangial conceptacles are similar in structure and develop- ment to carposporangial ones. The tetrasporangia are borne peripherally on stalk cells and the centre of the conceptacle is occupied by a columella of sterile cells. The young tetrasporangia are pale, thread-like and deeply invaginated zonately into four (Fig. 37A); they are contained in a loose, balloon-like, hyaline sac. The tetrasporangium swells and becomes pigmented, remaining deeply invaginated (Figs 37B, E) and four nuclei can often be seen at this stage. Before final maturation, however, the sporangium swells and apparently loses its divisions (Fig. 37C); it subsequently redivides zonately into four cells and rounds off to become the characteristic farctate tetrasporangium (Figs 37E, F). Mature conceptacles in surface view (Fig. 37F) show the tetrasporangia to be numerous and markedly peripheral. On one occasion, a bisporangium (Fig. 37D) was seen in an otherwise tetrasporangial crust. BISPORANGIAL PLANTS: Unknown. PHENOLOGY: P. rosanoffii occurs throughout the year on Halopitys at Bembridge and Beer, but is most abundant during autumn and early winter. Only sporadic records are available as regards its occurrence on Zostera; it will be interesting to know whether it occurs throughout the year on this host, or is absent for a season as with P. /ejolisii, for example. FORM RANGE: Regular, rather thin, crusts are typical of P. rosanoffii growing under optimum conditions. Under unfavourable conditions (e.g. on Palmaria at Beer in a particularly cold winter) the crusts become much thicker and reproduce more slowly. Trichocytes occur during warmer, long day-length seasons of the year. In England only intrafilament trichocytes have been observed; they are quite frequent and are about the same size as surrounding crust cells. In Brittany and Ireland intrafilament and branch-cell trichocytes occur abundantly and are appreciably larger than the crust cells. Material from Brittany is generally larger than English material, while the Irish samples have somewhat shorter crust cells than English specimens (see Tabs 12; 13). DISTRIBUTION: British Isles: Hampshire, Dorset, Devon, Cornwall, Jersey, Co. Galway, Co. Clare. World: Denmark, France (Channel, Atlantic and Mediterranean coasts), North Africa, Italy. Although this is a newly recognised species, by searching through BM herbarium material of FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 373 Fig.35 Decalcified crusts of Pneophyllum rosanoffii. A, Crust cells with intrafilament (1) and branch-cell (2) trichocytes. Scale = 20 um. B, View of ostiolar filaments from above. Scale = 10 wm. Me 6 Fig. 36 Gametangial/carposporangial crusts of Pneophyllum rosanoffii. A, Vertical section of carpogo- nial conceptacle. B, Vertical section of carposporangial conceptacle. C, Surface view of young carposporangial (c) and spermatangial (s) conceptacle. Scale = 50 um (all figures). one of its characteristic hosts, Halopitys incurvus, it has been possible to elucidate the distribution of the epiphyte. By this means it has been found to be common throughout the Mediterranean. It also occurs on Zostera marina in the Mediterranean according to Mazza’s (1922) description (as Melobesia lejolisii), and a specimen of Zostera in Kiitzing’s herbarium labelled Pneophyllum fragile (L no. 941.241.153) bears P. rosanoffii among other species. 374 Y. M. CHAMBERLAIN oe oe ae a Fig. 37 Sporangial conceptacles of Pneophyllum rosanoffii. A, Vertical section of young tetrasporangia in hyaline sacs. Scale = 50 wm. B, Vertical section of immature tetrasporangium. Scale = 50 wm. C, Vertical section of nearly mature tetrasporangial conceptacle in which the zonate divisions have become inconspicuous. Scale = 25 wm. D, Vertical section of bisporangium from an otherwise tetrasporangial crust. Scale = 50 um. E, Surface view of immature (arrow) and mature tetrasporangia. Scale = 20 um. F, Surface view of tetrasporangial conceptacles. Scale = 100 um. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 375 Table 12 Dimensions of vegetative structures (in wm unless stated). Pneophyllum rosanoffii England Ireland Brittany crust diameter up to4mm crust depth up to 50 crust cell length (5:5) 11-5 (15-5) = (8) 9-5 (11-5) (7:5) 12 (15) surface width (4-5) 7-5 (10-5) (6) 8 (11-5) (6) 8-5 (11) crust cell height (7) 11 (16) epithallial cell length (2) 3-5 (5) (1-5) 3 (4-5) (3) 4-5 (6-5) surface view width (2) 5-5 (9-5) (4) 5-5 (7:5) (3-5) 7-5 (8) epithallial cell height 3-6 4-9 hypothallial cell width VS height perithallial width (9) 12 (18) (10-5) 13 (18) cell height (12:5) 20 (29) (10-5) 17-5 (27) VS number 1-5 14 trichocyte type intrafilament intrafilament intrafilament and branchcell and branch cell trichocyte length as crust cells (12:5) 14 (7-5) (8-5) 9-5 (10-5) RIDGWAY COLOUR CODE: dry crust — light grayish vinaceous, pl. XX XIX surface view width as crust cells Table 13 Dimensions of reproductive structures (um unless stated). Le Jolis no. 1200 Pneophyllum rosanoffii England CARPOSPORANGIAL CONCEPTACLE external diameter surface diameter VS internal diameter VS height VS roof height roof type no. basal cell rows ostiole no. gonimoblast filaments diameter carpospore SPERMATANGIAL CONCEPTACLE VS/surface width VS height length spermatium width spermatium TETRASPORANGIAL CONCEPTACLE external diameter surface diameter VS internal diameter VS height VS roof height roof type no. basal cell rows ostiole no. tetrasporangia length tetrasporangium width tetrasporangium (110) 137 (170) (84) 110 (156) (45) 78 (117) (13) 28 (52) uniform 1-3 long terete filaments up to 12 (25) 39 (50) (19) 36 (46) 3:5-5:5 353 (104) 147 (182) (104) 138 (164) (70) 109 (164) (13) 32 (57) uniform 1-2 long terete filaments up to 20 (41-5) 62 (78) (26) 40 (45-5) 90-130 c. 80 ree 810) 155-170 122 50 15-6 uniform 35-45 26-35 376 Y. M. CHAMBERLAIN This is almost certainly one of the most common European species of Pneophyllum, and now that it has been separated from P. Jejolisii, it should be easy to identify because of the very characteristic ostiolar filaments which can readily be seen in both fresh and dried material. It seems probable that southern Britain and Denmark are its northern limits of distribution. Hasirat: In the British Isles, P. rosanoffii occurs most frequently on intertidal plants of Halopitys incurvus along the south coast of England and in the Channel Islands. It usually grows in association with Melobesia membranacea and Pneophyllum sargassi. It also grows occasion- ally on Palmaria and Zostera in the inter- and subtidal zones. At Beer (Fig. 18) it forms a regular association with Pneophyllum concollum on intertidal Palmaria palmata between about Octo- ber and April. In Ireland P. rosanoffii occurs commonly on Zostera in the inter- and subtidal zones, often growing together with P. lejolisii. In France P. rosanoffii appears to be common on Zostera and Halopitys in Normandy, and is probably very abundant on Zostera in Brittany. It is curious that, although P. rosanoffii grows so commonly on Zostera in France and Ireland and is also found on this host in Devon and Cornwall, it has never been found during extensive investigations on Zostera growing at Bembridge, despite its abundance on Halopitys at this locality. 4. Pneophyllum limitatum (Foslie) Y. Chamberlain, comb. nov. Basionym: Melobesia lejolisii forma limitata Foslie in K. nor. Vidensk. Selsk. Skr. 1905 (3): 102 (1905). NOMENCLATURAL SYNONYMS: Melobesia limitata (Foslie) Rosenv. in K. dansk. Vidensk. Selsk. Skr. I, 7: 245, Figs 163-167 (1917). Fosliella limitata (Foslie) Ganesan in Phykos 2: 41 (1963). Heteroderma limitata (Foslie) Adey in K. nor. Vidensk. Selsk. Skr. 1970 (1): 16 (1970). REFERENCES: Melobesia limitata (Foslie) Rosenv.; Suneson in Acta Univ. lund. II, 33: 14 (1937), in Acta Univ. lund. I, 39: 24 (1943); Kylin in Acta Univ. lund. II, 40: 44 (1944); Hamel and Lemoine in Archs Mus. natn. Hist. nat. Paris VII, 1: 105 (1953 [‘1952’]); Sundene in Skr. nor. Vidensk. — Akad. mat.-nat. Kl. 1953 (2): 192 (1954); Cabioch Jn Feldmann in Trav. Stn biol. Roscoff I, 15: 17 (1964). Fosliella limitata (Foslie) Ganesan; Parke & Dixon in J. mar. biol. Ass. U.K. 56: 534 (1976); Chamberlain in Br. phycol. J. 12: 69 (1977); Rueness, Norsk algeflora: 60 (1977); Garbary In D. E. G. Irvine and J. H. Price (Eds), Modern approaches to the taxonomy of brown and red algae: 205-222 Figs 13, 14 (1978); de Valéra, Pybus, Casley and Webster in Proc. r. Ir. Acad. 79 B: 265 (1979). LecrotyPE: TRH! Herb. Foslie, Coll. L. K. Rosenvinge, no. 3807, 22 August 1893, Lungfjorden, Lendrup Rgn, on an air bladder of Fucus vesiculosus. D1aGnosis: Mainly epiphytic with a well calcified thin thallus, trichocytes frequent, conceptacles conical and ostiole surrounded by long, segmented filaments united into a funnel: differing from P. lejolisii and P. concollum in having long ostiolar filaments, from P. rosanoffii in not having free, terete ostiolar filaments; and from P. zonale in not having a deep vegetative crust or free ostiolar filaments. DEscrIPTION: Well-calcified, regular, dull mauvy-pink, mainly epiphytic crusts up to 10 mm diameter, c. 75 um deep, perithallium absent except near conceptacles (up to 4 cells deep in crusts with flat bisporangial conceptacles), crust cells in surface view sharply rectangular, 6-5-20-5 um long xX 5-5-13-5 wm wide, epithallial cells 3-5-5 wm long X 5-5—10-5 wm wide, large branch cell or intrafilament trichocytes 14-26 wm long X 10-5—17 wm wide; conceptacles (except spermatangial and one bisporangial form) prominent, ostiole surrounded by funnel of fused segmented filaments, carposporangial conceptacles 117-169 um internal diameter X 52-109 um high, carposporangia densely crowded when mature, spermatangial conceptacles immersed beside carpogonial conceptacles, 33-59 wm wide x 21-30 um deep, ostiole prolonged into a spout, tetrasporangial conceptacles 151-229 ym internal diameter X 68-117 um high, tetraspor- angia crowded, up to c. 30 per conceptacle, 31-60 zm long x 29-39 wm wide, OR conceptacles FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 377 flat with simple ostiole, in thallus up to 6 cells deep, 65-96 um internal diameter X 29-61 um high, bisporangia peripheral, 23-39 wm long x 18-30 um wide. Historica: Foslie (1905a) described P. limitatum as a form of P. lejolisii (as Melobesia) with smoother, thicker crusts, and conceptacles which were more conical and widely spaced. The lectotype (Adey, 1970) is a Rosenvinge specimen (TRH) in which the crusts of P. limitatum are epiphytic on Fucus vesiculosus. According to Rosenvinge (1917, p. 245) Foslie came to regard Melobesia lejolisii f. limitata as mainly growing on algae as opposed to f. typica which mainly grew on marine phanerogams. Rosenvinge (1917) considered the features of forma limitata to be sufficiently distinct to warrant specific status as Melobesia limitata. Pneophyllum limitatum has proved to be one of the commonest simple crustose coralline epiphytes in northern Europe and is common on Scandinavian, British, and northern French shores. CRUST APPEARANCE: P. limitatum forms well calcified, rather regular, dull mauvy-pink crusts which are often crazed on the surface. The plants are predominantly epiphytic on algae, but are also found on marine phanerogams and occasionally on glass or china. When growing most vigorously, on large, intertidal algae in autumn, the crusts (Fig. 38) are regularly orbicular or confluent with other crusts, and may exceed 10 mm diameter; at other times the crusts are less regular and smaller. The conceptacles are usually well spaced out in the crust centre, with the crust border being vegetative only; sometimes two adjacent conceptacles fuse together. Even at low magnification the conceptacles are conspicuously conical, and the ostiolar funnel forms a characteristically pale ring (Fig. 39C) in the conceptacle centre. ee | 3 ‘RO-4 Fig. 38 Pneophyllum limitatum (1) and Fosliella farinosa (2) growing on Chondrus crispus. Scale = 10 mm. 378 Y. M. CHAMBERLAIN Under the S.E.M. the crust cells (Fig. 39D) are broadly rectangular with short, wide, epithallial concavities and large branch cell and intrafilament trichocytes. In all types of conceptacle the roof is uniform (Figs 39A, B, C; 40A, B), and the inconspicuous spermatangial ostiole can be seen at the side of the carpogonial conceptacle (Figs 40A, F). The small type bisporangial conceptacles (Figs 39A, B) have a gently raised roof and a simple ostiole. The young conical conceptacle type has a plateau-like roof and simple ostiole (Fig. 40A); this later develops into a conical structure and the top of this may acquire a coating (presumably mucilaginous) as it senesces (Fig. 40E). The fused filaments of the funnel-like ostiole are very conspicuous, and the stages in their development and senescence are seen in Figures 40C-E. In the centre of the funnel is a further ring of filaments which forms a diaphragm-like structure (Fig. 40C). VEGETATIVE ANATOMY: The crusts are composed of hypothallium and epithallium only, except near conceptacles, in all but the small type bisporangial conceptacles; these have 1-4 perithallial cells. The crust cells of all phases are sharply rectangular (Fig. 41A), with conspicuous epithallial cells. The trichocytes are usually noticeably larger than the surrounding crust cells, and are divided by a radial wall into a pigmented, hair-bearing part and a colourless part (see Chamberlain, 1977a); trichocytes usually lack epithallial cells. Suneson (1943) showed abnor- mal, very large, terminal trichocytes, and large, richly protoplasmic cells in some crusts of P. limitatum. I have not found these in this species, but have seen similar trichocytes in epilithic species such as P. myriocarpum (q.v.). Suneson (1943) considered the possibility that the rounded contents of the richly protoplasmic cells represented some form of vegetative reproduc- tive body. GAMETANGIAL PLANTS: Gametangial plants are monoecious; the spermatangial conceptacle (Figs 40A; 42C, F) is immersed in the thallus beside the carpogonial conceptacle; it is more or less oval in VS and about twice as wide as high, with the ostiole prolonged into a hyaline spout. Spermatangial initials usually produce elongated spermatangia from two sites; these round off into small, oval spermatangia when released. Spermatangium development is described and depicted by Suneson (1937). The carpogonial conceptacle (Fig. 42C), is gently raised; its structure and development have been investigated and illustrated in detail by Suneson (1937). The conceptacle cavity develops by the elongation and final rupturing of the perithallial cells, while the upper surface of the perithallial cells gives rise to the roof filaments and the extensively developed ostiolar filaments. Suneson (1937, Figs 6E and 7) shows that the procarps in the centre of the fertile disc bear one or two carpogonia, while the peripheral procarps bear one or no carpogonia. CARPOSPORANGIAL CONCEPTACLES: The typical carposporangial conceptacle (Fig. 42E) has one or, occasionally, two basal cell layers, a fairly thin roof with rather irregular cells, and a substantial fusion cell immediately above the basal cells. Suneson (1937) found that, while in most carposporangial conceptacles gonimoblast filaments were initiated only from the periphery of the fusion cell (as in all other known species of Pneophyllum), in some they were initiated over the entire fusion cell surface, which seemed to be unique for this genus. I previously concurred with this observation (Chamberlain, 1977a), but on further examination have found that while carposporangial filaments are very occasionally produced on the surface of the extreme periphery of the fusion cell in British material, they have not been found across the entire surface, and are mainly borne in the usual way from the actual periphery. Nevertheless, the fact that they are borne on the surface of the fusion cell at all is a distinguishing character. The size of the conceptacles varies considerably (Tab. 15), and in practice the conceptacles are usually larger when the plants are growing under optimum conditions. The discrepancy in conceptacle height dimension between the present values and those given by Suneson (1937, 1943) and Chamberlain (1977a) is due to the fact that the length of the ostiolar funnel is excluded from the present values. In general, there are one or two perithallial cells at the periphery of the conceptacle (Fig. 42E), and beyond these the crust quickly reverts to being composed of hypothallium and epithallium only. Another form has been found, however, which has a deep "WIM CT = afBdg “satyAvoUOD [eTT]eYIIdo pure s[[9o Jsn41o Sulmoys Aroydiad ysndg ‘q ‘wa —E¢z = a]R9S ‘(MOLIB) s[OUUNJ BJONSO JINJeW YA aus ‘safoejdaouOd [eoIUOD BUIMOYS JsNIO JeIsURIOdseIIOT, ‘D “wi! YG = a[kOg “(MOIIR) sa]AD0Y9I) Axeyeo19}Ul pue soso ajduis oy} Sulmoys YW Wo UsWOsIe[Uq ‘g “WIT! YZ = B]Bdg “soTONsO [[eWIS YIIM sopoe}daoUOD pastes APYSIs dy} SUIMOYS }sNJo [eIsueIOdsiq adA} [JeUIg ‘WW “wnwynUy wnpjXydoaug JO SdIN{eIJ JSNID MOYS 0} sydeIZOIOIUI UOIDE[a BuTUURIS HE “BIA i Nn es me 2 x 2 = m ea 12) en B rz = » 3 S a & z Q Z < << — _ = S & Y. M. CHAMBERLAIN Fig. 40 Scanning electron micrographs to show conceptacle features of Pneophyllum limitatum. A, Carpogonial (1) and spermatangial (2) conceptacles at an early stage before the roof has become conical. Scale = 100 wm. B, Tetrasporangial conceptacle with developing ostiole funnel. Scale = 50 um. C, Young carposporangial ostiole with outer ostiole filaments (1) developing and inner filament ring (2) visible. Scale = 25 wm. D, Mature funnel-like tetrasporangial ostiole. Scale = 25 wm. E, Senescent tetrasporangial conceptacle roof in which the ostiole has disintegrated. Scale = 50 um. F, Spermatangial ostiole. Scale = 10 um. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 381 Fig. 41 Crust and ostiole features of Pneophyllum limitatum. A, Crust in surface view with trichocytes (tr). Scale = 50 um. B, Vertical section of inner and outer rings of tetrasporangial ostiole filaments. Scale = 20 um. thallus and larger conceptacles; a similar form was illustrated by Rosenvinge (1917, Fig. 164). This form occurs in the two epilithic samples found at Bembridge and occasionally in epiphytic material. The structure of the ostiolar filaments in VS is shown in Figure 41B, where it can be seen that they are segmented for much of their length. This differs from the unsegmented filaments of P. zonale and P. rosanoffii, with which this species may otherwise be confused. However, the fusion of the filaments to form a funnel is not obvious in decalcified sectioned material of P. limitatum although it can be seen clearly under the S.E.M. (Fig. 40D). The presence of an inner ring of filaments (Figs 40C; 41B) is also, as far as is known, unique to P. limitatum. Large quantities of carposporangial filament cells and carposporangia are produced by P. limitatum, and the mature conceptacle is packed tightly with them as was also shown by Suneson (1937). TETRASPORANGIAL PLANTS: The structure and development of tetrasporangial conceptacles (Figs 39C; 40B; 42D) follows the same pattern as in the carposporangial conceptacles. The tetraspor- angia are borne peripherally on stalk cells (Fig. 42D); Suneson (1937) showed that young sporangia develop among sterile cells within a balloon-like sac, and are deeply divided into four from an early stage. As with the carposporangia, large numbers of tetrasporangia are produced in each conceptacle and these are piled high in the mature conceptacle. Sometimes the ostiolar filaments have been observed to develop deep within the conceptacle, and some sections of this type of structure resemble procarps (Chamberlain, 1982). BISPORANGIAL PLANTS: As has already been mentioned, two types of bisporangial conceptacle occur in P. limitatum. One form (Fig. 42B) is similar to the tetrasporangial conceptacles, except that it produces uninucleate bispores (which are presumed to be diploid) instead of tetraspores. This form has been found only occasionally in the British Isles and has not been recorded previously. The other form (Figs 39A, B; 42A) has a quite different aspect, although the vegetative crust border has crust cells similar to P. limitatum cells. The thallus of this form has perithallial filaments up to four cells deep and the perithallial cells are more or less isodiametric in VS. The conceptacles are immersed in the thallus and have a simple ostiole. The bisporangia are borne peripherally and are more rounded than in the other bisporangial conceptacle type, while relatively few bisporangia (probably not exceeding 10) occur in each conceptacle. This type of conceptacle was first recorded and figured by Suneson (1937); it is quite easy to identify under optimum conditions when it occurs among crusts with conical conceptacles, but crusts found in 382 Y. M. CHAMBERLAIN aes oe ee a ESS Fig. 42 Vertical sections of conceptacles of Pneophyllum limitatum. A, Flat type of bisporangial conceptacle. Scale = 20 wm. B, Domed type of bisporangial conceptacle. Scale = 50 um. C, Gametangial crust showing carpogonial (c) and spermatangial (s) conceptacles. Scale = 100 um. D, Tetrasporangial conceptacle. Scale = 50 wm. E, Carposporangial conceptacle. Scale = 50 um. F, Spermatangial conceptacle. Scale = 20 um. isolation are virtually impossible to distinguish from, for example, bisporangial crusts of P. microsporum. P. limitatum is the only species in which two bisporangial conceptacle types have been identified. PHENOLOGY: P. limitatum occurs throughout the year on the shores of the British Isles (see Chamberlain, 1977a), but it is most abundant and vigorous during autumn and early winter (September to November). At this time it has been observed at Bembridge on a wide range of intertidal algae, but particularly on the fronds of Laminaria digitata, L. saccharina, Fucus serratus, Chondrus crispus, and Palmaria palmata; it usually grows together with Fosliella farinosa (Fig. 38) at this season, but F. farinosa tends to become more abundant and Pneophyllum limitatum less so as winter deepens. However, the degree of abundance depends FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 383 on the weather and in 1975, 1976, and 1980 this bloom was much more in evidence than in the years between or since. A collection of Pneophyllum limitatum on Palmaria is present in Le Jolis’s herbarium (CHE). It was collected in the Cherbourg area (Le Jolis no. 240, 30 November 1853). Le Jolis originally identified the crusts as Melobesia pustulata, while Rosanoff (1866) reidentified them as M. farinosa and M. pustulata. It seems probable, therefore, that a bloom of Pneophyllum limitatum, similar to that seen in England, occurs in autumn in northern France, and this was corroborated by a collection made in October 1981 at Hercquemoulin on the northwestern side of the Cotentin peninsula, in which P. limitatum was abundant on several shore algae, particularly Sargassum muticum. ForM RANGE: As has been mentioned, the crusts of P. limitatum tend to be larger and more regular in autumn than at other seasons. Trichocytes are more abundant at this time, and absent from about November until June. Crust cells tend to be smaller and relatively longer under less favourable conditions; the dimensions given in Table 14 were made on material collected in September, while in material collected during November trichocytes were absent, crust cells measured (7) 10 (14) wm long x (4) 7 (9) wm wide, and epithallial cells were c. 3 um long X 2-5 pm wide. As noted previously (Chamberlain, 1977a), P. limitatum crusts growing on Zostera tend to have more abundant trichocytes than those growing on algal hosts. As has been discussed, the conical type of bisporangial conceptacle occurs very rarely in Britain, while the flat roofed type is common here and in Sweden (Suneson, 1937); it has also been observed in material from Normandy. In a single collection made in Brittany conical bisporangial conceptacles were frequent, but no flat-roofed ones were seen, nor have flat-roofed conceptacles been observed in Irish material. DISTRIBUTION: British Isles: Recorded throughout the British Isles with the exception of the east coasts of England and Scotland. P. limitatum is the commonest epiphytic Pneophyllum on algae in south and south-west England, as it is in Scandinavia, but it seems possible that its place is taken to some extent by P. concollum in Ireland, where P. limitatum is generally less common, although it was abundant in one collection of algae from Co. Clare made in November 1981. World: Norway, Sweden, Denmark, northern France. Table 14 Dimensions of vegetative structures (in wm unless stated). British Rosenvinge Suneson Pneophyllum limitatum Isles (1917) (1937, 1943) crust diameter up to 10 mm up to5 mm up to 5(10) mm crust depth a pe crust cell length (6:5) 10-5 (20-5) (7) 8-10 (11) 13-16 surface width (5-5) 10 (13-5) (5)-12 (5) 6-10 crust cell height c. 15-20 epithallial cell length (3) 4 (5:5) surface view width (5:5) 7:5 (10-5) epithallial cell height c. 5-8 hypothallial cell width VS height perithallial width cell height VS number trichocyte type branch cell and intercalary trichocyte length (14) 21 (26) up to 22 surface view width (10-5) 12 (17) 384 Y. M. CHAMBERLAIN Table 15 Dimensions of reproductive structures («m unless stated). British Rosenvinge Suneson Pneophyllum limitatum Isles (1917) (1937, 1943a) CARPOSPORANGIAL CONCEPTACLE external diameter 210-325 90-210 surface diameter (143) 165 (177) VS internal diameter (117) 141 (169) 90-115 VS height (52) 78 (109) 100-180 VS roof height (8) 17 (26) roof type uniform no. basal cell rows 1 (rarely up to 4) ostiole funnel shaped no. gonimoblast filaments diameter carpospore SPERMATANGIAL CONCEPTACLE VS/surface width (33) 44 (59) 56 70 VS height (21) 26 (30) 56 length spermatium width spermatium TETRASPORANGIAL CONCEPTACLE external diameter (170) 230-325 140-270 surface diameter (138) 162 (182) VS internal diameter (151) 177 (229) 150-240 VS height (68) 92 (117) VS roof height (10) 17 (26) roof type uniform no. basal cell rows 1 (rarely up to 4) ostiole funnel shaped no. tetrasporangia up toc. 30+ length tetrasporangium (30) 53 (60) 46-77 width tetrasporangium (29) 32-5 (39) 21-46 (61) BISPORANGIAL CONCEPTACLE external diameter 75-125 surface diameter VS internal diameter (65) 82 (96) [156 210 234] VS height (29) 45 (61) 60-70 60-110 [65 80 91] VS roof height (10) 13 (18) [13 17 21] roof type uniform [uniform] no. basal cell rows 1-3 | (1 ostiole simple simple simple [funnel-shaped] no. bisporangia up toc. 10 [c. 30+] length bisporangium (23) 30 (39) 33 [39 48 52] width bisporangium (18) 22-5 (30) 23 [23 30 36] [] large type bisporangial conceptacle FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 385 HapsitatT: Epiphytic on a wide range of intertidal algae, but particularly on Palmaria palmata, Fucus serratus, Laminaria saccharina, and L. digitata, where it usually grows together with Fosliella farinosa. It also occurs quite commonly on Zostera, usually together with Pneophyllum lejolisii, and has very occasionally been found occurring epilithically. In Denmark (Rosenvinge, 1917) P. limitatum was found most commonly on Fucus vesiculosus (an alga which rarely supports crustose coralline epiphytes in the British Isles), but also grew on other algae and frequently occurred together with Dermatolithon pustulatum (as Lithophyllum macrocarpum). LiFE HISTORY: No experimental evidence is yet available regarding the life history of P. limitatum. It would be particularly interesting to obtain information on the life history of this species in view of the two types of bisporangial crust which occur. 5a. Pneophyllum confervicolum (Kitz.) Y. Chamberlain comb. nov. Forma confervicolum Basionym: Phyllactidium confervicolum [‘confervicola’| Kitz., Phycologia generalis: 295 (1843). NOMENCLATURAL SYNONYMS: Hapalidium phyllactidium (Kitz.) Kitz., Species algarum: 695 (1849). Hapalidium confervicolum (Kitz.) Areschoug Jn J. G. Agardh (Ed.), Species, genera et ordines algarum: 509 (1852). Hapalidium confervoides (Kiitz.) P. Crouan & H. Crouan pro parte (?), Florule du Finistére: 149 (1867). Melobesia confervicola (Kitz.) Foslie in K. nor. Vidensk. Selsk. Skr. 1900 (5): 21 (1900). REFERENCES: (?) Lithocystis allmanni Allman In Harvey, Phycologia Britannica: pl. CLXVI (1948). Hapalidium phyllactidium (Kitz.) Kitz.; Kiitzing, Tabulae phycologicae: 33, tab. 92 c—d (1869); Crouan P.-L. and H.-M. in Annals Sci. nat. Bot. 1V, 12: 286, pl. 21, figs 14-20 pro parte? (1859) [see Fig. 44]. (?) Melobesia inaequilaterata Solms-Laub. in Fauna Flora Golf Neapel 4: 12 (1881). (?) Heteroderma inaequilaterata (Solms-Laub.) Foslie in K. nor. Vidensk. Selsk. Skr. 1909 (2): 56 (1909). Melobesia minutula f. typica Foslie in K. nor. Vidensk. Selsk. Skr. 1905 (3): 107 (1905); Rosenvinge in K. danske Vidensk. Selsk. Skr. 11, 7: 252 (1917); Knight & Parke, Manx algae: 102, 108 (1931); Suneson in Acta Univ. lund. II, 39: 27 (1943); Hamel & Lemoine in Archs Mus. natn. Hist. nat. Paris VII, 1: 107 (1953 [‘1952’]); Feldmann in Trav. Stn biol. Roscoff Supplement 6: 78 (1954); Zinova, Check-list of green, brown and red algae of the southern seas of the U.S.S.R.: 232 (1967). Melobesia fosliei Rosenvinge in K. danske Vidensk. Selsk. Skr. 11, 7: 249 (1917). Melobesia fosliei sensu Kylin in Acta Univ. lund. II, 40: 45 (1944). Heteroderma minutula (Foslie) Dawson in Pacif. Sci. 10: 47 (1956); Adey in K. nor. Vidensk. Selsk. Skr. 1970 (1): 16 (1970). Fosliella minutula (Foslie) Ganesan in Phykos 2: 38 (1963); Parke & Dixon in J. mar. biol. Ass. U.K. 56: 534 (1976); Rueness, Norsk algeflora: 60 (1977). Ho totyee: L! 941.156.120. Herb. Kiitzing, near Trieste, on ‘Conferva vasta’ (Figs 43A, B; 45A). Diacnosis: Epiphytic species with small, square crust cells, very short, wide epithallial cells and abruptly hemispherical conceptacles: differing from Fos/iella farinosa in having intercalary (not terminal) trichocytes and lacking trichocytes round conceptacle base, from Pneophyllum lejolisii in having domed conceptacles and isodiametric crust cells. DEscRIPTION: Crusts pale to mauvy-pink, not exceeding 2 mm diameter, perithallium absent except near conceptacle in some forms, crust cells in surface view typically isodiametric, 5-5—13-5 um long X 4-14-5 wm wide, epithallial cells short and wide, 1-5-4 wm long Xx 2-5-8 wm wide, intrafilament or branch cell trichocytes similar to crust cells in size or sometimes longer; conceptacles abruptly hemispherical, all except spermatangial conceptacles with simple ostiole, or with a small beak, or occasionally with ostiole filaments, carposporangial conceptacles 47-104 fm internal diameter X 39-68 wm high, spermatangial conceptacles c. 30 wm wide X 23 wm high, ostiole prolonged into a spout, tetrasporangial conceptacles 52-91 wm internal diameter x 30-74 wm high, up to 10 tetrasporangia per conceptacle, 26-46 um long Xx 15-5-27:5 wm wide; bisporangial conceptacles 68-107 um internal diameter X 52-65 um high, bisporangia 26-39 um long X 14-5—21 wm wide. 386 Y. M. CHAMBERLAIN Fig. 43 Scanning electron micrographs of Pneophyllum confervicolum f. confervicolum. A, Type specimen of Phyllactidium confervicolum Kiitz. (L.). Scale = 50 wm. B, Crust cells from A; trichocytes (arrow) present. Scale = 10 wm. C, Crusts from Ardkeen growing on Cladophora rupestris. Scale = 50 um. D, Crust cells from C; trichocytes (arrow) present. Scale = 25 wm. E, Crusts from Bembridge growing on Furcellaria lumbricalis. Scale = 50 um. F, Beaked ostiole. Scale = 5 wm. Historica: P. confervicolum was originally described by Kiitzing (1843, p. 295) as Phyllacti- dium confervicola, a green calcareous alga epiphytic on Chaetomorpha vasta from Trieste. Later, Kiitzing (1849) realised that the five species he had included in Phyllactidium (Kitzing, 1843) belonged to several different algal groups and he moved confervicolum to Hapalidium, a genus of coralline algae he had described earlier (Kiitzing, 1843) as having a monostromatic thallus and no cortical cells. Rosanoff (1866), however, later showed cortical cells to be present. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 387 The type specimen (Figs 43A, B; 45A) of Pneophyllum confervicolum f. confervicolum (L-—no. 941.156.120) agrees with the present concept. The Crouan brothers (1859) included Hapalidium phyllactidium Kitz. in their notes on the genus Hapalidium. They recorded it as being epiphytic on various algae or Zostera and epilithic on glass and china. Material seen in Herb. Crouan (CO) was all on glass or china; it comprised a mixture of epilithic species of Pneophyllum but did not include P. confervicolum as now circumscribed. The Crouans’ drawings, however, (Fig. 44C) may illustrate epiphytic material, since the crust shown in their Figure 16 appears to arise from a cellular substrate; the crust has square cells and prominent tetrasporangial conceptacles and may represent the present species. The Crouans (1867) included P. confervicolum in the Florule du Finistére; they had changed the specific epithet from Hapalidium phyllactidium to H. confervoides (Kiitz.) J. Ag., although Areschoug (1852) in J. Agardh had spelled it confervicolum. Foslie (1900a) discussed the publications of the Crouans (1859, 1867) concerning Hapalidium confervoides and described the Crouans’ epilithic material of it which he had obtained from PC: he concluded that the material was identical with Melobesia zonalis (P. Crouan & H. Crouan) Foslie and different from Hapalidium phyllactidium Kitz. It seems possible that Lithocystis allmanni Allman in Harvey (1848) is conspecific with Pneophyllum confervicolum judging by the small, square cells in Harvey’s illustration. Unfortu- nately the type specimen cannot be found at present so it is not possible to be sure. Foslie (1905a) discussed Epilithon van heurckii which Heydrich (in Chalon, 1905) described on the basis of Chalon’s material from Jersey. Foslie had received material from Chalon and he considered it to be conspecific with Melobesia minutula. However, the illustration (Heydrich in Chalon, 1905, p. 207) shows a bisporangial crust with a multiporate conceptacle roof. It cannot, therefore, belong to the genus Pneophyllum, and is now regarded as a species of the lithotham- nioid genus Melobesia. Danish material of Melobesia minutula was described by Rosenvinge (1917) who showed, contrary to the opinions of Kiitzing and Foslie, that the species usually bore cortical cells (epithallium). Rosenvinge (1917) described another species, M. fosliei, which differed from M. minutula only in possessing (intercalary) trichocytes. Suneson (1943) subsumed M. fosliei in M. minutula. Kylin (1944) used the epithet M. fosliei as asynonym of M. minutulaf. typica and restricted M. minutula Foslie to forma lacunosa. Ganesan (1963) described material from India which agrees with the present concept and he transferred the species to the genus Fosliella. CRUST APPEARANCE: The crusts of P. confervicolum vary considerably in appearance depending on their host. On Zostera and algae with flat thalli the small crusts (up to about 2 mm diameter) are pale and delicate; they may be regularly discoid, lobed, or show stages between this and the creeping filaments of f. minutulum. The conceptacles are hemispherical and scattered. On Cladophora and other algae such as Plocamium with narrow thalli, the conceptacles are so densely crowded (Fig. 43C) as to give the host a knobbly appearance. It is difficult to see the extent of the crusts in these circumstances, but many individual crusts probably bear only one or a few conceptacles; these crusts are often bright pink. Under the S.E.M., the shape of the broad crust cells and very short, wide epithallial cells are shown in Figure 43D, while intercalary trichocytes are shown in Kiitzing’s type material (Fig. 43B) and English material (Fig. 43D). The domoid conceptacles may have a stepped appearance (Figs 43A, E), or a somewhat irregular but smoother surface (Fig. 43C). In this, as in all other features, this species shows considerable variation. VEGETATIVE ANATOMY: The vegetative crust is composed of epithallium and hypothallium only, and the crust cells (Figs 45D; 46A, B) are characteristically small and almost isodiametric with extremely short, broad epithallial cells. Sometimes, however, the cells are longer and narrower as shown by Suneson (1943). Intrafilament and branch cell trichocytes occur. Usually these are of similar length to the crust cells but bulge somewhat. Sometimes they are longer than the crust 388 Y. M. CHAMBERLAIN inn.des Setene. nat. 4° Serie. } “a Bot. Toners. Plt oan ile Marpalum 3 a” Doutiot Fes ———— NM Hemond wy 7 belle -Batrapude th, Lares 10mm Fig.44 Plate from Notice sur le genre Hapalidium by Crouan & Crouan (1859). (‘Harpalium’ at the foot of the plate is presumably a printer’s error). A, Hapalidium zonale. B, Hapalidium coccineum. C, Hapalidium phyllactidium. D, Hapalidium callithamnioides. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 389 cells with a large pigmented part and a narrow unpigmented strip down one side (Figs 45D; 46B). GAMETANGIAL PLANTS: The crusts are monoecious. The spermatangial conceptacles are borne in the thallus at the edge of the carpogonial conceptacles and have the ostiole prolonged into a spout (Suneson 1943: 30 as Melobesia minutula). The raised carpogonial conceptacle (Fig. 45E) contains procarps bearing up to two carpogonia. CARPOSPORANGIAL CONCEPTACLES: Carposporangial conceptacles are more or less hemispherical (e.g. Figs 45F-H). They may rise very abruptly from the thallus as in material from Lough Ine giowing on Gelidium, and plants growing on Corallina, while on Plocamium, for example, they rise more gradually (Figs 45H; 46F). On Cladophora (Fig. 45G) the conceptacles are so crowded that sections often appear to show a thallus with perithallial development which is really the walls of adjacent conceptacles. The ostiole structure is very variable as was found also by Rosenvinge (1917, as M. fosliei) and Suneson (1943). Sometimes the ostiole is very simple with no development of either upper or lower filaments. Sometimes the lower ostiole filaments become quite well developed (Figs 45H; 46F), while the upper filaments may be prolonged into a small collar (Fig. 45F) or a spout (Fig. 45G), (see also Rosenvinge, 1917, fig. 169; Suneson, 1943: 30). On one occasion a conceptacle with well developed, upper ostiole filaments (Fig. 46G) was found among others with simple or slightly beaked ostioles. The fusion cell (Fig. 46F), is rather flat and bears carposporangial filaments peripherally. TETRASPORANGIAL CONCEPTACLES: Tetrasporangial conceptacles show a similar range of structure and size to carposporangial conceptacles. Abruptly hemispherical conceptacles are seen in Lough Ine material (Fig. 46D) and on Furcellaria from Bembridge (Fig. 45B), and more gently raised ones from Fanore (Fig. 45C). Beaked ostioles occur as illustrated by Suneson (1943: 29). Young tetrasporangia are thin, deeply divided into four, and enclosed in a hyaline, balloon-like sac (Fig. 46D). The farctate mature tetrasporangia are large in relation to the size of the conceptacle and usually entirely fill the cavity. BISPORANGIAL CONCEPTACLES: Bisporangial conceptacles show the same variety of structure as carposporangial and tetrasporangial conceptacles; an abruptly hemispherical form from Lough Ine is shown in Figure 46C, while a more gently rounded one with more developed ostiolar filaments is seen in Figure 46E of material from Fanore, Co. Clare, growing on Plocamium. PHENOLOGY: P. confervicolum has been found throughout the year in the British Isles but is most common from May to November. FORM RANGE: It is apparent from the species description that P. confervicolum shows a wide range of variation in the structure of the conceptacles and that sometimes the crust cells are not the very characteristic, isodiametric shape with very short, wide epithallial cells. In addition one Irish population growing on Zostera has the characteristic isodiametric crust cells (Fig. 46B), but they measure on average 10-5 um long x 11 wm wide which is considerably larger than usual (Tab. 16). P. confervicolum is usually recognisable by its external appearance: the thin, delicate crust and minute bubble-like conceptacles are tiny compared with most other crustose, epiphytic corallines. Figure 45F shows P. confervicolum growing on a conceptacle of Melobesia membra- nacea in comparison with which it looks very small, although M. membranacea itself is not large. Pneophyllum confervicolum is most easily confused with the gametangial and tetrasporangial plants of Fosliella farinosa which may be very similar, although when trichocytes are present the two species are readily separated. In addition both species produce filamentous forms, but the difference in cell shape is usually apparent and can be seen in Figure 20B, where both are growing on Gelidium at Lough Ine. It remains possible, however, that distinct varieties, or even species, are present among the forms described here and further investigation of this taxon is needed. 390 Y. M. CHAMBERLAIN FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 391 Fig. 45 Thallus features and conceptacle variation in Pneophyllum confervicolum f. confervicolum. A, Vertical section of the type specimen Phyllactidium confervicolum Kitz. (L). Scale = 20 um. B, Vertical section of tetrasporangial conceptacles growing on Furcellaria lumbricalis at Bembridge; ostiole plug arrowed. Scale = 50 um. C, Vertical section of tetrasporangial conceptacle growing on Plocamium coccineum at Fanore, Co. Clare. Scale = 50 wm. D, Thallus cells growing on Zostera marina at Black Head, Co. Clare, with trichocyte (arrow). Scale = 20 wm. E, Vertical section of young carposporangial conceptacle growing on Plocamium coccineum at Fanore, Co. Clare. Scale = 20 um. F, Vertical section of carposporangial conceptacle growing on conceptacle of Melobesia membranacea (arrow) at Bem- bridge. Scale = 50 wm. G, Vertical section of carposporangial conceptacles with spout growing on Cladophora rupestris at Ardkeen. Scale = 50 wm. H, Vertical section of carposporangial conceptacle growing on Plocamium coccineum at Fanore, Co. Clare. Scale = 50 wm. Fig. 46 Pneophyllum confervicolum. A, Decalcified crust in surface view to show typically small, isodiametric crust cells with very short epithallial cells. B, Decalcified crust from Irish material with larger crust cells, showing intrafilament trichocytes (arrows) with very narrow colourless parts. C, Vertical section of bisporangial conceptacle of abruptly hemispherical type (Lough Ine, Ireland). D, Vertical section of tetrasporangial conceptacle of abruptly hemispherical type, with young tetrasporan- gia in balloon-like sacs (1) and nearly mature tetrasporangium (2) (Lough Ine, Ireland). E, Vertical section of bisporangial conceptacle of rounded type with downward ostiole filaments (3) and small beak (4) (Fanore, Ireland). F, Vertical section of carposporangial conceptacle of similar type to E (Fanore, Ireland). G, Vertical section of carposporangial conceptacle with elaborate ostiole filaments found among normal hemispherical conceptacles (Bembridge). 392 Y. M. CHAMBERLAIN DISTRIBUTION British Isles: Hampshire, Dorset, Devon, Cornwall, Isle of Man, Anglesey, Mull, South Uist, Shetland, Channel Islands, Co. Antrim, Co. Down, Co. Wexford, Co. Cork, Co. Clare, Co. Galway. World: Sweden, Norway, Denmark, France (northern and Mediterranean), Madeira, Italy, southern U.S.S.R., India, Pacific Mexico, central Pacific. As Dawson (1960) has remarked, it is probable that this species occurs widely in tropical and temperate regions of the world, but its small size has caused it to be overlooked in many collections. Hapsirat: P. confervicolum has been found on a wide range of algal hosts and on Zostera marina; it also grows on bryozoans and hydroids. It is most abundant on warm, sheltered shores, like those of Lough Ine and Kimmeridge (Chamberlain, 1982). 5b. Pneophyllum confervicolum forma minutulum (Foslie) Y. Chamberlain, comb. nov. BasionyM: Melobesia minutula Foslie in K. nor. Vidensk Selsk. Skr. 1904 (2): 8 (1904). NOMENCLATURAL SYNONYM: Melobesia minutula f. lacunosa Foslie in K. nor. Vidensk. Selsk. Skr. 1905 (3): 107 (1905), nom. illeg. (Art. 63.1). REFERENCES: Hapalidium callithamnioides P. Crouan & H. Crouan in Annis Sci. nat. Bot. IV, 12: 289 (1859); Foslie in K. nor. Vidensk. Selsk. Skr. 1899 (7): 7 (1900 [‘1899’]). Guerinea callithamnioides (P. Crouan & H. Crouan) Picquenard in Trav. scient. Lab. Zool. Physiol. marit. Concarneau 4(3): 1 (1912). Melobesia callithamnioides sensu Batters in Grevillea 21: 53 (1892). Table 16 Dimensions of vegetative structures (in wm unless stated). Suneson Suneson (1943) Hapalidium British British (1943) f callitham- Isles Isles f. typica lacunosum nioides Pneophyllum f. fi: (= f. con- (=f. Crouan confervicolum confervicolum minutulum fervicolum) minutulum) material crust diameter up to2 mm crust depth crust cell length (5:5)9(13-5) (6) 8 (10) 8-12 6-12 (7) 8-5 (10-5) surface width (4) 7-5 (14-5) (6-5) 11 (15-5) 5-9 6-14 (8) 11 (16) crust cell height 6. 10 epithallial cell length (1:5) 2:5(4) (1:5)2(3-5) very short, veryshort, (2) 2-5 (4) lentiform lentiform surface view width (2-5) 5 (8) (4:5)6:5(9) (6) 7-5 (10) epithallial cell height hypothallial cell width VS height perithallial width cell height VS number trichocyte type intrafilament and branch cell trichocyte length as crust cells surface view width as crust cells FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 393 Table 17 Dimensions of reproductive structures («um unless stated). Suneson (1943) f. confervicolum — as f. typica British (f. minutulum — as Ganesan Pneophyllum confervicolum Isles f. lacunosa) (1963) CARPOSPORANGIAL CONCEPTACLE external diameter 75-120 surface diameter (57) 62 (65) 75-100 (35-60) VS internal diameter (47) 72 (104) 45-70 VS height (39) 55 (68) 50-100 40-65 VS roof height (8) 13 (16) roof type uniform — irregular no. basal cell rows 1-2 ostiole simple, beaked simple or simple or or filaments beaked beaked no. gonimoblast filaments c.8: diameter carpospore SPERMATANGIAL CONCEPTACLE VS/surface width 30 (1 only) (28) 25-40 VS height 23 (18) 25-35 length spermatium width spermatium TETRASPORANGIAL CONCEPTACLE external diameter (90) 100-120 surface diameter (67) 75 (91) 85-100 (45-75) VS internal diameter (52) 69 (91) 60-80 VS height (30) 56 (74) 85-100 50-60 VS roof height (8) 14 (18) roof type uniform — irregular no. basal cell rows 1 ostiole simple or simple beaked no. tetrasporangia up to 10 6-12 (3-10) length tetrasporangium (26) 33 (46) 30-35 width tetrasporangium (15-5) 22 (27-5) 15-20 BISPORANGIAL CONCEPTACLE external diameter surface diameter i (60-90) VS internal diameter (68) 83 (107) VS height (52) 57 (65) 60 VS roof height 9-10 roof type uniform — irregular no. basal cell rows 1 ostiole simple or beaked no. bisporangia (5-15) length bisporangium (26) 34 (39) width bisporangium (14-5) 18 (21) 394 Y. M. CHAMBERLAIN Melobesia minutula f. lacunosa Foslie; Suneson in Acta Univ. lund. I1, 39: 31 (1943); Feldmann in Trav. Stn. biol. Roscoff supplement 6: 78 (1954); Zinova, Check-list of green, brown and red algae of the southern seas of the U.S.S.R.: 232 (1967). Melobesia minutula sensu Kylin in Acta Univ. lund. II, 40: 56 (1944). Fosliella minutula f. lacunosa (Foslie) Cabioch in Bull. Soc. phycol. Fr. 19: 78 (1974). Ho.otyPe: TRH! Herb. Foslie, 25 August 1903, south Norway, Hvalgerne ‘Galtene’ ved Kirkgen c. 5-8 fathoms, on Corallina officinalis (Figs 47B; 48A). Diacnosis: Differing from f. confervicolum in having a creeping, filamentous thallus and smaller conceptacles. DEscriPTION: With creeping filaments of broad crust cells, crust cells 6-10 wm long X 6-5-15-5 pm wide, epithallial cells 1-5—3-5 wm long X 4-5-9 wm wide, conceptacles hemispherical. Fig. 47 Scanning electron micrographs of Pneophyllum confervicolum f. minutulum. A, From Lough Ine. Scale = 50 wm. B, From type specimen of Melobesia minutula Foslie (TRH). Scale = 25 wm. Fig. 48 Features of Pneophyllum confervicolum f. minutulum. A, Vertical section of conceptacle from type specimen of Melobesia minutula Foslie (TRH). Scale = 25 wm. B, Crust growing on Palmaria palmata at Bembridge. Scale = 10 um. C, Crust from type specimen of Hapalidium callithamnioides P. Crouan & H. Crouan (CO). Scale = 10 wm. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 395 Historicac: Foslie (1904b) described Melobesia minutula from Hvaléerne, south Norway, where it grew on Corallina officinalis; this was a creeping form (Fig. 47B) which he subsequently (Foslie, 1905a) named f. lacunosa at the same time applying the epithet f. typica to a discoid form found on hydroids from Norway and Denmark. Since the f. /acunosa included the type of the species, it should have been called f. minutula according to the International Code (Art. 26.1). Suneson (1943) described and figured Swedish material of Melobesia minutula f. lacunosa. Kylin (1944) correctly restricted Melobesia minutula Foslie f. lacunosa, but Ganesan (1963) wrongly indicated that this usage was incorrect. Foslie (1905a: 108) included M. inaequilaterata Solms-Laub. in the synonymy of M. minutula f. lacunosa; later (Foslie, 1909, p. 57), he recombined M. inaequilaterata in Heteroderma, and noted that he considered this synonymous with Melobesia minutula. I have not seen Solms- Laubach’s material of M. inaequilaterata and the description and drawing of young plants (Solms-Laubach, 1881) are not sufficient to enable identification. Other authors (e.g. Mazza, 1922) give it as a synonym of Fosliella farinosa (as Melobesia). A further species, described by the Crouan brothers (1859) as Hapalidium callithamnioides (see Fig. 44D) appears to be sterile material of Pneophyllum confervicolum f. minutulum; type material from Concarneau (Fig. 48C) agrees with this form in appearance and cell size (Tab. 16). The material was dredged off Brest and a similarly subtidal collection of f. minutulum was reported by Cabioch (1974, as Fosliella minutula f. lacunosa) growing on maérl at a depth of 45 m off Madeira. Both Foslie (1900a) and Picquenard (1912a) concluded that the Crouans’ material (PC and CO respectively) was not a calcareous alga, but was possibly allied to Rhodochorton. However, I found the specimens at Corncarneau to be markedly calcified and composed of broad crust cells bearing epithallial cells (Fig. 48C) which Foslie (1900a) and Picquenard (1912a) had thought to be absent. Picquenard (1912a) proposed a new genus, Guerinea, to accommo- date this species. CRUST APPEARANCE: The thallus of f. minutulum forms filaments (Fig. 20B) which creep, often for considerable distances, over other algae and sea-grasses; Foslie (1905a) and Suneson (1943) describe plants growing on Corallina and Jania. The filaments usually produce rather fewer conceptacles than the crusts of f. confervicolum. VEGETATIVE ANATOMY: The cells of f. minutulum (Fig. 48B) are usually considerably broader than wide, with very broad, short epithallial cells; these features are also apparent in the type material of Hapalidium callithamnioides (Fig. 48C). CONCEPTACLES: Few conceptacles have been observed in the present investigation. In general, however, they are similar to but smaller than those of f. confervicolum. Suneson (1943, p. 32) shows adjacent carpogonial and spermatangial conceptacles. PHENOLOGY: Forma minutulum has been recorded from May to November in the present investigation. DISTRIBUTION: British Isles: Hampshire, Dorset, Anglesey, Co. Cork, Co. Galway, Co. Clare. World: Sweden, France, Madeira, southern U.S.S.R. 6. Pneophyllum microsporum (Rosenv.) Y. Chamberlain, comb. nov. BasionyM: Melobesia microspora Rosenv. in K. danske Vidensk. Selsk. Skr. VU, 7: 256 (1917). Hotortyre: C! Herb. Rosenvinge no. 4670, 23 April 1894, Ryes Flak, Denmark, 2-5 fathoms, on Furcellaria lumbricalis (Fig. 49A). Diacnosis: Epiphyte with deep, crumbly crust up to 14 cells deep and immersed conceptacles with a simple ostiole: differing from P. sargassi in having uniform conceptacle surface, from P. plurivalidum in not having domed ostiole, from P. concollum in not having an ostiolar collar and in having fewer sporangia and smaller conceptacles, from P. /ejolisii in having a deep perithal- lium and from P. confervicolum in having flat conceptacles and a deep perithallium. 396 Y. M. CHAMBERLAIN DescriPTIONn: Deep, heavily calcified, crumbly, brownish-pink crusts up to 5 mm diameter, 140 jm deep, perithallium up to 14 cells deep in crust centre, one to a few cells deep at periphery, perithallial cells (in VS) 5-14 wm wide X 5-21 um high; crust cells in surface view square to somewhat elongate, 5—13-5 wm long X 4-5—12 wm wide; intrafilament and branch cell trichocytes of same size range as crust cells; conceptacles immersed, all except spermatangial conceptacles with simple ostiole, carposporangial conceptacles 45-104 um internal diameter X 33-78 wm high, spermatangial conceptacles 15-49 4m wide X 23-34 um high, ostiole prolonged into a spout, tetrasporangial conceptacles 54-130 mm internal diameter xX 49-70 wm high, 6-12 peripheral tetrasporangia per conceptacle, 22-48 wm long xX 18-29 um wide, central sterile paraphyses sometimes evident, bisporangial conceptacles 62-98 ym internal diameter x 42-63 um wide, 3-8 peripheral bisporangia per conceptacle, 25-38 um long x 19-5-29 um wide, central sterile paraphyses present. HistoricaL: P. microsporum was described by Rosenvinge (1917) as an epiphyte growing on Furcellaria lumbricalis in Denmark. It was characterised by having perithallial filaments up to 5 cells deep, very small tetrasporangia, and no epithallial cells. Examination of the type material under the S.E.M. (Fig. 49A) and decalcified whole mounts, however, show that short, wide epithallial cells are present, while the tetrasporangia on slides made by Rosenvinge appear to be young; mature ones would probably be larger than the dimensions of 17—24 wm long x (9) 11-12 (16) wm wide recorded by Rosenvinge (1917). P. microsporum has not been recorded since 1917, but an entity found commonly in southern England and south-west Ireland is now referred to this species; although crusts occur quite frequently they are rather inconspicuous and difficult to characterise, which no doubt accounts for the previous lack of records. The principal difference between Rosenvinge’s material and that from the British Isles lies in the appearance of the crust cells under the S.E.M. Rosenvinge’s material (Fig. 49A) has broad but very short epithallial concavities, while these are relatively longer in material from the British Isles (Fig. 49E). Decalcified crusts from the British Isles, however, have short, wide epithallial cells (Fig. 50) similar to those observed on Rosenvinge’s material. CRUST APPEARANCE: This species forms thick, heavily calcified crusts (up to 140 ~m deep) which crumble and detach easily. It grows epiphytically on a number of algal species, most frequently on Furcellaria lumbricalis (as did the type) and Chondrus crispus. The crusts are deep pink or brownish-pink and often have a somewhat granular, glistening appearance; they are irregularly shaped and measure up to about 3 mm diameter on Furcellaria and 5 mm on Chondrus. The small crowded conceptacles (Figs 49B-D) are scarcely raised; when old they often become very pale and form conspicuous circles on the darker crust. Under the S.E.M. (Fig. 49E) the crust cells vary from square to elongate with conspicuous epithallial concavities which are shorter than wide. The crust edge is thick since perithallial filaments usually start to develop immediately behind the first meristem. Intercalary trichocytes occur only very occasionally in English material, possibly because it grows there mainly in winter; they are abundant, however, on material collected in Ireland in May (Fig. 49E). The conceptacles have a uniform surface often with a smooth covering in the ostiole area (Figs 49B, C). The ostiole is small and simple when young (Figs 49B, C), but the edges cave in to produce a large hole in old conceptacles (Figs 49F, G). Fig. 49 Scanning electron micrographs of Pneophyllum microsporum from various localities. A, Type specimen of Melobesia microspora Rosenv. (Ryes Flak, Denmark, April 1894) showing carposporangial (1) and spermatangial (2) ostiole. Scale = 25 ym. B, Side view of conceptacle (Bembridge, 13 January 1979). Scale = 50 um. C, Ostiole from same crust as B (Bembridge, 3 January 1979). Scale = 10 wm. D, Vertical section of conceptacle (Bembridge, 13 January 1979). Scale = 25 wm. E, Crust periphery with trichocytes (arrow) (Finavarra, 3 May 1981). Scale = 10 wm. F, Mature ostiole (Finavarra, 3 May 1981). Scale = 10 wm. G, Senescent ostiole (Finavarra, 3 May 1981). Scale = 10 wm. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 398 Y. M. CHAMBERLAIN VEGETATIVE ANATOMY: In surface view (Fig. 50) the crust cells vary from almost square to somewhat elongate, and the epithallial cells are relatively short and wide. The intercalary trichocytes are mainly intrafilament but occasionally occur in branch cells; they are the same size as normal crust cells. In vertical section (e.g. Figs 51; 52A; 54B) the crusts are composed of regular, mainly almost isodiametric perithallial cells; the perithallial filaments may be up to 13 cells deep and are usually initiated at, or near, the crust periphery. GAMETANGIAL PLANTS: Gametangial plants are monoecious, the spermatangial conceptacles (Figs 49A; 51A; 52C) are adjacent to the carpogonial conceptacle, they are completely immersed and have a spout (Figs 51C; 52D). Carpogonial conceptacles are immersed and bear procarps in a bunch in the centre of the conceptacle floor (Figs 52A, B; 54A), but the conceptacles are so small that their structure has not been fully elucidated. In material from Ireland the crusts are very deep, there are usually two or three cells below the conceptacle base, and the conceptacles appear entirely flat in surface view (Figs 51A). This is sometimes the case in English material but conceptacles may also be initiated on thinner, peripheral parts of the crust in which case they have only a single basal cell layer and may appear raised. CARPOSPORANGIAL CONCEPTACLES: The mature conceptacle is either entirely immersed, asin Irish and some English material (Figs 51A; 54B), or somewhat raised as in other English material (Fig. 51B). The roof is composed of small, irregular filaments, and the ostiole is simple externally but with some development of the downward filaments (Fig. 51A). The small fusion cell bears carposporangial filaments up to seven cells long from the periphery Fig. 50 Surface view of decalcified crust of Pneophyllum microsporum. Scale = 10 wm. TETRASPORANGIAL PLANTS: The shape, size, and ostiole structure of tetrasporangial conceptacles (Figs 53A; 54E, F) are similar to the carposporangial conceptacles. The farctate sporangia are borne peripherally, and there is some development of central, sterile paraphyses. The sporangia are not particularly small in relation to the conceptacle; the small dimensions given by Rosenvinge (1917), after which the species is named, were probably obtained from immature sporangia. BISPORANGIAL PLANTS: These are similar to the tetrasporangial plants and likewise have central sterile paraphyses (Figs 53B; 54C, D). The bisporangial plants may be difficult to distinguish from the small type bisporangial form of P. /imitatum found in the British Isles. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 399 Fig.51 Gametangial/carposporangial conceptacles of Pneophyllum microsporum. A, Vertical section of carposporangial and spermatangial (arrow) conceptacles from Finavarra, Co. Clare. Scale = 25 um. B, Vertical section of carposporangial conceptacle from Bembridge. Scale = 25 um. C, Vertical section of spermatangial conceptacle from Finavarra. Scale = 20 um. PHENOLOGY: Apart from sporadic records in other parts of southern England, Ireland, and France, the principal population of P. microsporum investigated was that at Bembridge where the species is present mainly during the winter and is most abundant at this time on Chondrus crispus (Chamberlain, 1982). Otherwise the principal host is Furcellaria lumbricalis and samples were collected at all times of year. All the Irish material was collected in May and no winter collections are available for comparison. FORM RANGE: There are considerable differences in the structure and appearance of P. microsporum from Ireland and that mostly seen in England. The Irish plants have a deep thallus with the conceptacles situated near the surface; the conceptacles are noticeably larger than in the English material, tetrasporangial conceptacles, for example, measure 54-99 um internal diameter in English plants and 117-130 um in Irish plants. In addition the English plants often have somewhat raised conceptacles (Fig. 49B) among which much detritus collects, while the Irish plants have completely flat conceptacles externally. English material may also have flat conceptacles and very deep thalli, but the predominance of the larger form in Ireland suggests a geographical variation. DISTRIBUTION: British Isles: Hampshire, Dorset, Devon, Mull, Channel Islands, Co. Galway, Co. Clare, Co. Down. World: Denmark, France. Insufficient data are available to assess the distribution range of P. microsporum. On present evidence it would appear to be restricted to northern Europe. Hasirat: Epiphytic on the following algae: Furcellaria lumbricalis, Polyides rotundus (once only), Phyllophora crispa, P. pseudoceranoides, Chondrus crispus, Palmaria palmata, Gastroc- 400 Y. M. CHAMBERLAIN O) DO ws BUS CAINS Pel @ Diba 50 um é ees ey, 20 um Fig. 52 Carpogonial/carposporangial and spermatangial conceptacles of Pneophyllum microsporum from Bembridge and Ireland. A, Vertical section of young carpogonial conceptacle (Ireland). B, Vertical section of carpogonial conceptacle (Bembridge). C, Surface view carposporangial (1) and spermatangial (2) conceptacles (Bembridge). D, Vertical section of spermatangial conceptacle (Bem- bridge). FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 401 _ ii ‘ — _ MS | ’ | Fig. 53 Tetra/bisporangial conceptacles of Pneophyllum microsporum. A, Vertical section of tetraspor- angial conceptacle from Bembridge. Scale = 20 um. B, Surface view of bisporangial conceptacles from Bembridge. Scale = 50 um. lonium ovatum, Halopitys incurvus, Laurencia pinnatifida, Polysiphonia elongata, Laminaria digitata, Fucus serratus, Cystoseira nodicaulis, Sargassum muticum. The hosts on which Pneophyllum microsporum are most commonly found are Chondrus crispus on which it may form extensive, brownish-pink colonies and Furcellaria lumbricalis on which it is frequent but usually in scattered individual crusts. Table 18 Dimensions of vegetative structures (in wm unless stated). Pneophyllum microsporum British Isles Rosenvinge (1917) crust diameter up to 5mm 1-2 mm crust depth up to 140 crust cell length (5) 8-5 (13-5) surface width (4-5) 8-5 (12) crust cell height C$ epithallial cell length (1:5) 3 (4-5) surface view — width (4) 5-5 (8-5) epithallial cell height 3-6 hypothallial cell width (5) 9-5 (15) 5-8 VS height (5) 8-5 (12-5) c. 5-8 perithallial width (5) 11 (14) 6-8 cell height (5S) 12 (21) 6-16 VS number up to 14 cells up to 6 cells trichocyte type intrafilament (few branch cell) trichocyte length as crust cells surface view width as crust cells RIDGWAY COLOUR CODE: dry crust — brownish vinaceous fresh crust — pale brownish vinaceous 402 Y. M. CHAMBERLAIN Table 19 Dimensions of reproductive structures (4m unless stated). Pneophyllum microsporum British Isles Rosenvinge (1917) CARPOSPORANGIAL CONCEPTACLE external diameter 120-140 surface diameter (58) 87 (102) VS internal diameter (45) 86 (104) VS height (33) 54 (78) VS roof height (7) 18 (31) roof type uniform uniform no. basal cell rows 1-4 ostiole simple simple no. gonimoblast filaments 8-14+ diameter carpospore SPERMATANGIAL CONCEPTACLE VS/surface width (15) 38 (49) c. 40 VS height (23) 29 (34) length spermatium 3-5-5-5 6 width spermatium 2-3 2 TETRASPORANGIAL CONCEPTACLE external diameter 120-140 surface diameter VS internal diameter (54) 103 (130) 70-110 VS height (49) 60 (70) VS roof height (13) 23 (31) roof type uniform uniform no. basal cell rows 1-5 1+4 ostiole simple no. tetrasporangia 6-12 length tetrasporangium (22) 37 (48) 17-24 width tetrasporangium (18) 23 (29) (9) 11-12 (16) BISPORANGIAL CONCEPTACLE external diameter surface diameter (39) 76 (117) VS internal diameter (63) 80 (98) VS height (42) 54 (63) VS roof height (12) 17 (20) roof type uniform no. basal cell rows 1-3 ostiole simple no. bisporangia 3-8 length bisporangium width bisporangium (25) 32-5 (38) (19-5) 24 (29) 7. Pneophyllum concollum Y. Chamberlain, sp. nov. REFERENCES: Melobesia rosea sensu Rosanoff in Mem. Soc. imp. Sci. nat. Cherbourg 12:77 P|. II, Figs 1, 2 (1866), non Hapalidium roseolum Kitz. (See Fig. 3 of present study). Melobesia lejolisii Rosanoff pro parte in Mem. Soc. imp. Sci. nat. Cherbourg 12: P1. 1, Figs 8, 9 only (1866). (See Fig. 2 and p. 305 of present study.) Ho.otyPe: BM! Y. M. Chamberlain no. 76/100, 4 April 1976, Ballyandreen, Co. Cork, southern Ireland, growing on Laurencia pinnatifida coll. M. D. Guiry (Fig. 55A). D1aGnosis: Species epiphytica, filamentis perithallis profunditate usque ad 8 cellulas et ostiolis ad instar colli; a P. rosanoffii differt filamentis ostiolaribus libris carentibus, a P. sargassi et P. microsporo ostiolis ad instar colli, et a P. myriocarpo (specie epilithica) perithallio profundo. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 403 Fig. 54 Reproductive crusts of Pneophyllum microsporum from Bembridge. A, Vertical section of very young carpogonial conceptacle. B, Vertical section of mature carposporangial conceptacle. C, Vertical section of mature bisporangial conceptacle. D, Surface view of mature bisporangial conceptacle. E, Surface view of mature tetrasporangial conceptacle. F, Vertical section of mature tetrasporangial conceptacle. DEscrIPTION: Crustae leaves, malvinae, diametro usque at 5 mm, profunditate 75 wm, lobis parum superpositis; perithallium media in crusta profunditate usque ad 8 cellula, ad marginem carens vel profunditate 1-4 cellularum, cellulae perithallii (in sectione verticali) 6-5—14-5 wm latae x 4-5—-28-5 wm altae, cellulae crustae aspectu superficiali rectangulares, 6-5—22-5 xm longae x 4-15-5 ym latae, cellulae epithallii 3-6-5 wm longae X 3-8 um latae, trichocyti non visi; conceptacula immersa vel parum elevata, omnia, praeter ea spermatangialia, ostiolo ad instar colli, nonnumquam canale superimposito, concepta- cula carposporangialia diametro interno 91-156 um, altitudine 52-83 wm, cellula conjungens lata et tenuis, conceptacula spermatangialia immersa, 20-65 um lata X 19-37 um alta, ostiolo in canalem producto, conceptacula tetrasporangialia diametro interno 104-208 um, altitudine 39-104 wm, 16-20 tetrasporangiis in unoquoque conceptaculo, tetrasporangia 30-65 wm longa X 18-41 um lata, para- physes centrales steriles adsunt, conceptacula bisporangialia diametro interno 104-182 um, altitudine 52-104 um, 16-20 bisporangiis in unoquoque conceptacula, bisporangia 33-5-52 wm longa Xx 15-542 um lata, paraphyses centrales steriles conspicuae. D1aGnosis: Epiphytic species with perithallial filaments up to 8 cells deep and collar-like ostiole: differing from P. rosanoffii in not having free ostiolar filaments; from P. sargassi and P. Y. M. CHAMBERLAIN 404 ‘pury S,siof oT Ul PST SuIzINYy AW ‘puey s,suizny Ul “JeUpe J ‘Inowe’y] vunisiqiyg sisdodug ‘(GHO) sUIZINY Wnjoasos UnipyodvyY — uautizads 4amoTT ‘pury S,St[of a] Ul (JJouRsOY) ‘JUsY vasou visaqojayp ‘puey sjyoursoy ur (“WI Ul) j"zINy wunjoasod unipyvdvy] :(1ap{O} Vavuvaquiau visaqojap Ul) (AHO) SOL a] ‘q1aH Xe Jyoursoy vaso. visaqojapw jo 3dAyo399] — uaunoads 4addyr ‘g (Wd) Mnjjoouos uinzjkydoaug Jo adAjoOH ‘YW “wnjjoou0s un Aydoaug pur JJoursoy vasos visaqojayw jo suouitosds sdk], gg ‘Bi <¢) /9L. OA 0 Cup ab) who 91 ' Ponet)' mg9‘seermo hog K ot aes Aa a ee as * i hla c id le | k | wey? MP mon Pda] } vowed ah \ r. Sy gh - << # i “sy . Sri ss ian sere) L< 4 eyfos Lt oF i a BY, ER rye pee wry -y0:h & , bd yor, y 7? FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 405 microsporum in having a collar-like ostiole, and from P. myriocarpum (epilithic) in having a deep perithallium. DEscrIPTION: Smooth, mauvy-pink crusts up to 5 mm diameter, 75 wm deep, with slightly overlapping lobes, perithallium up to 8 cells deep in crust centre, absent or 1-4 cells deep at periphery, perithallial cells (in VS) 6-5-14-5 wm X 4-5—28-5 wm high; crust cells in surface view rectangular, 6-5—22-5 wm long X 4-15-5 wm wide, epithallial cells 3-6-5 wm long X 3-8 wm wide, trichocytes not seen; conceptacles immersed or somewhat raised, all except spermatangial conceptacles with collar-like ostiole sometimes surmounted by a spout, carposporangial con- ceptacles 91-156 wm internal diameter < 52-83 wm high, fusion cell wide and thin, spermatangial conceptacles immersed, 20-65 xm wide xX 19-37 um high, ostiole prolonged into a spout, tetrasporangial conceptacles 104-208 zm internal diameter x 39-104 um high, 16-20 tetraspor- angia per conceptacle, 30-65 um long x 18-41 wm wide, central sterile paraphyses present, bisporangial conceptacles 104-182 ym internal diameter x 52-104 wm high, 16-20 bisporangia per conceptacle, 33-5—52 wm long x 15-5—42 um wide, central sterile paraphyses pronounced. Historica: The identity of P. concollum has only become clear during a prolonged study of Pneophyllum species in the British Isles. It is characterised by the collar-like ostiole, somewhat developed perithallium, and frequent presence of balloon-like paraphyses in the tetra- and bisporangial conceptacles. None of these characters is very conspicuous, however, and the ostiole structure, which is the most clearly diagnostic character, may easily be missed in sectioning. For these reasons the taxon has not previously been distinguished, although it is now clear that it is the taxon which Rosanoff (1866), with some hesitation, described as Melobesia rosea growing on Bornetia secundiflora (J. Agardh) Thuret at Cherbourg (Fig. 55B). The material in Le Jolis’s herbarium (CHE) on which Rosanoff’s description is based agrees in surface features and ostiole structure with the present taxon. In addition I believe that material of Pneophyllum concollum was included originally by Rosanoff (1866) in his concept of Melobesia lejolisii. It was on this material that Rosanoff did his conceptacle drawing shown in Figures 2 [8, 9] of the present study, and on which his record of M. /ejolisii growing on Bornetia, was based. Since no Bornetia specimens are now present in the folders of Melobesia lejolisii (CHE) it seems that Rosanoff later changed his mind about this material and transferred it to M. rosea. Further discussion of Rosanoff’s concept of Pneophyllum concollum is given on pages 306-308. CRUST APPEARANCE: The mauvy-pink crusts appear opaque but rather thin (up to 75 um deep). They may have a spotted appearance and the surface is usually cracked in dried material. The plants are epiphytic on algae and are characteristically composed of a number of slightly overlapping crusts (Fig. 56A). It is not usually possible to measure individual crusts, but they probably do not attain more than 5 mm diameter. Under the S.E.M. (Fig. 56D) the crust cells are regularly rectangular and the epithallial concavities vary from rounded rectangular to short and broad; trichocytes have not been seen. In fresh material the conceptacles appear almost flat while in dried material they are somewhat rounded. Under the S.E.M. they mostly have a uniform surface (Figs 56A—C) which may be covered with a smooth coating (Fig. 56C) which is probably mucilaginous; in critical point dried material (Fig. 56B) the smooth surface may crack irregularly to reveal the newly developed roof cells. Under the S.E.M. the ostiole is mostly simple (Figs 56B, C) or surrounded by a small collar. VEGETATIVE ANATOMY: The crusts may have an extended margin composed of hypothallium and epithallium only or may initiate perithallial filaments immediately behind the periphery. In surface view (Fig. 57B) the crust cells are regularly rectangular; the size varies considerably both within and between crusts. Plastids are plentiful and conspicuous but starch grains are not evident. The epithallial cells vary from rounded rectangular to short and broad, and are dome-shaped or flattened in VS. In VS (e.g. Figs 59B; 60B) the crusts appear rather regularly zoned horizontally; the "wi! QT = aR9¢ ‘Apun’] WJ s[[99 ISNID ‘q ‘Wi YOT = BJBdg ‘UsdIpURATTeEg WO doRJINS BJR}da0NUOD ‘_D “WIN CZ = BIBIg “YW WIJ 9]01)SO ‘g ‘WI YSZ = BTROS ‘(ad4}) usorpuedT[eg WO $}SN.IO PoLIp JUIOd [BOYLID JO MAIA [BIOUIH “W ‘wnjjoouos uinyjAydoaug Jo sydeiso1NIW UOIDE]e SuUURIS 9g “BIA Z < = Es a ES = < <>) S 5 > FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 407 perithallial filaments vary from 1-8 cells deep and their cells vary from short to very elongated, the uppermost cell (subapical initial) often markedly tapers upward. GAMETANGIAL PLANTS: Gametangial plants are monoecious. The spermatangial conceptacles, which are usually adjacent to the carpogonial conceptacles, are relatively broad and shallow (Figs 57C; 58B). They are completely immersed and have a spout. Carpogonial conceptacles are usually gently raised (Fig. 58A), with the fertile disc being surrounded by a wide ring of stretched perithallial cells. There are from 1+ rows of cells below the conceptacle base and the procarps bear one or two carpogonia (Fig. 58A). In some populations such as that at Beer, south Devon, the carpogonial conceptacles are narrower and more abruptly dome-shaped. CARPOSPORANGIAL CONCEPTACLES: The mature conceptacle is somewhat raised (Figs 57A; 58C, D), the roof is quite deep, and there are 1-4 cell rows below the conceptacle. The ostiole is surrounded by filaments which are united laterally to form a collar (Fig. 57D), and sometimes this is surmounted by a spout (Figs 57A; 58D) similar to that seen in the spermatangial conceptacle; the downward ostiolar filaments are well developed and may become balloon-like. The fusion cell is characteristically wide and very thin (Figs 58C, D), and the gonimoblast filaments, which are produced peripherally, are usually seven cells long. conceptacle from Ballyandreen. Scale = 20 um. D, Characteristic collar-like ostiole. Scale = 20 um. 408 Y. M. CHAMBERLAIN aoe S TRO OY fe) carpe wv ° ay 9" ae: SQO 0 Soo oD Ne pes ; : av? Ly Nie ce sehaee ce Aas a ; : CEO" yy is) | je © O00 Bate i> \ S SS ae la = = ie Z FORE See, GARE? ey SS] BESS Sees, eae? Sees OSCR eee PUR ITS Fig. 58 Vertical sections of gametangial crusts of Pneophyllum concollum. A, Carpogonial conceptacle. B, Spermatangial conceptacle. C, Carposporangial conceptacle with ostiole collar. D, Carposporangial conceptacle with ostiole prolonged into a spout. Scale = 20 um (all figures). TETRASPORANGIAL PLANTS: Conceptacle shape and ostiole structure (Figs 59A; 60A, B) are similar to that seen in the carposporangial conceptacle. The tetrasporangia are borne peripher- ally, they remain pigmented and farctate throughout development, and are not invaginated. The mature sporangia are rather small in relation to the size of the mature conceptacle and appear as a peripheral ring in surface view; sterile cells develop centrally on the conceptacle base (Figs 59A; 60B) to form a columella and these cells may become balloon-like. BISPORANGIAL PLANTS: The development and structure of bisporangial conceptacles (Figs 59B; 60C-E) is the same as for tetrasporangial conceptacles. The bisporangia are even smaller in relation to conceptacle size than the tetrasporangia. The central sterile filaments and downward ostiolar filaments may be balloon-like (Fig. 60E) in bisporangial as in tetrasporangial plants. PHENOLOGY: Insufficient evidence is available to generalise on the seasonal periodicity of P. concollum. At present it is known that at Beer, south Devon the species occurs in the intertidal zone only in winter (October—April) and the intertidal records from Wembury (south Devon) are also mainly of winter material. In the subtidal, however, P. concollum is abundant in July at Lundy Island and the Scilly Isles. In southern and western Ireland the species is common in both intertidal and subtidal habitats from April to August but no records are yet available for the winter months. All reproductive phases occur at the different localities. Le Jolis’s samples on Bornetia are in excellent condition; both were collected in February (1857 and 1859) which suggests that in that locality, on that host, this was the optimum season. The indications are, therefore, that Pneophyllum concollum is able to grow throughout the year except in very cold conditions, but that its actual occurrence depends on local circum- stances. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 409 ” Fig. 59 Vertical sections of sporangial conceptacles of 2% Pneophyllum concollum. A, Tetrasporangial conceptacle from Ballyandreen. B, Bisporangial conceptacle from Wembury. Scale = 25 wm. Cte Bagh Rid Roe « eee io a Sati il ome es SePo na ea \ | q ‘ (f\ () O Napa he ni (Vesna abRa n° Asan 9908 eee poe nee! (0000S a) . 2 fa ” O eae cette pee é eg is Oog00 UDA Bas hs Fig. 60 Vertical sections of tetra/bisporangial conceptacles of Pneophyllum concollum. A, bisporangial initials (shaded). B, Mature bisporangial conceptacle. C, Tetrasporangial initials (shaded). D, Imma- ture tetrasporangial conceptacle. E, Mature bisporangial conceptacle with distended columella and ostiole cells (arrow). FoRM RANGE: Plants collected in Ireland between April and August and subtidal material collected from the Scilly Isles and Lundy in July have noticeably thinner crusts with flatter conceptacles than material collected at Beer during winter. The thickness of the Beer crusts, and often heavy infestation of the conceptacles with bacteria and general detritus (Fig. 57A), suggest that the plants were growing under unfavourable conditions, while the thinner crusts from elsewhere were probably growing healthily and rapidly under favourable conditions. DISTRIBUTION: British Isles: Hampshire, Dorset, Devon, Cornwall, Jersey, Co. Wexford, Co. Clare, Co. Galway. World: Northern France. 410 Y. M. CHAMBERLAIN The frequency with which this species occurs in collections from Ireland suggests that it is one of the commonest simple crustose epiphytes there; it becomes much less common towards the eastern British Isles and has been collected, for example, only occasionally at Kimmeridge and Bembridge. Other than Le Jolis’s samples on Bornetia collected from the Cherbourg area only one specimen has been collected from France (Y. M. Chamberlain, Gatteville, Normandy, 25 July 1977). Hasitat: Pneophyllum concollum occurs as an epiphyte on the following algae: Furcellaria lumbricalis, Calliblepharis ciliata, Gracilaria verrucosa (Huds.) Papenf., Chondrus crispus, Palmaria palmata, Bornetia secundiflora (in France), Gastroclonium ovatum (Huds.) Papenf., Laurencia pinnatifida, Polysiphonia elongata, Desmarestia aculeata (L.) Lamouroux, Laminaria hyperborea (Gunnerus) Foslie, L. ochroleuca Bach. Pyl., Cladostephus spongiosus (Huds.) Agardh, Cystoseira foeniculacea (L.) Grev.; it was also found once growing on the leathery egg case of a nursling fish. The species has been found intertidally in pools and areas of shore which do not dry out and subtidally to a depth of about 8 m. A population of Pneophyllum concollum growing together with P. rosanoffii on Palmaria palmata at Beer is discussed in Chamberlain (1982). TAXONOMIC NOTE: It is possible P. concollum may belong to the same taxon as P. myriocarpum (q.v.). Table 20 Dimensions of vegetative structures (in wm unless stated). Pneophyllum concollum British Isles crust diameter up toc. 5mm crust depth up toc. 75 crust cell length (6:5) 12-5 (22-5) surface width (4) 8-5 (15-5) crust cell height 65 epithallial cell length (3) 4-5 (6:5) surface view width (3) 5-5 (8) epithallial cell height c. 5-7 hypothallial cell width (4-5) 11 (18) VS height (5-5) 8-5 (16) perithallial width (6-5) 10 (14-5) cell height (4-5) 11-5 (28-5) VS number up to 8 trichocyte type _ trichocyte length surface view width RIDGWAY COLOUR CODE: dry crust — pale laelia pink 8. Pneophyllum myriocarpum (P. Crouan & H. Crouan) Y. Chamberlain, comb. nov. BasionyM: Melobesia myriocarpa P. Crouan & H. Crouan, Florule du Finistére: 150 (1867). NoMENCLATURAL SYNONYM: Melobesia myriocarpa P. Crouan & H. Crouan in Bull. Soc. bot. Fr. 1860: 4 (1860), nomen nudum; Foslie in K. nor. Vidensk. Selsk. Skr. 1899 (7): 10 (1900 [‘1899”}). REFERENCES: Melobesia (subgenus Pliostroma) zonalis forma myriocarpa, Foslie; Foslie in K. nor. Vidensk. Selsk. Skr. 1908 (11): 4 (1908); Hamel and Lemoine in Archs Mus. natn. Hist. nat. Paris VII, 1: 108 (1952 [‘1953’]). Melobesia (subgenus Pliostroma) zonalis sensu Lemoine in Bull. Lab. marit. Mus. Hist. nat. St. Servan 7: 20 (1931). FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES Table 21 Dimensions of reproductive structures (um unless stated). Pneophyllum concollum British Isles CARPOSPORANGIAL CONCEPTACLE external diameter surface diameter VS internal diameter VS height VS roof height roof type no. basal cell rows ostiole no. gonimoblast filaments diameter carpospore SPERMATANGIAL CONCEPTACLE VS/surface width VS height length spermatium width spermatium TETRASPORANGIAL CONCEPTACLE external diameter surface diameter VS internal diameter VS height VS roof height roof type no. basal cell rows ostiole no. tetrasporangia length tetrasporangium width tetrasporangium BISPORANGIAL CONCEPTACLE external diameter surface diameter VS INTERNAL DIAMETER VS height VS roof height roof type no. basal cell rows ostiole no. bisporangia length bisporangium width bisporangium (96) 159 (182) (91) 136 (156) (52) 65 (83) (13) 20 (29) uniform or irregular 1-4 collar-like (+ spout) up toc. 20 (20) 47 (65) 19-37 (130) 148 (182) (104) 151 (208) (39) 75 (104) (10) 21 (29) uniform or irregular 1-3 collar-like (+ spout) c. 16-20 (30) 47-5 (65) (19) 28-5 (41) (104) 136 (182) (104) 135 (182) (52) 91 (104) (13) 29 (104) uniform or irregular 1-4 collar-like (+ spout) c. 16-20 (33-5) 45-5 (52) (15-5) 27-5 (42) Melobesia zonalis sensu Batters pro parte inJ. Bot. Lond. 40 (suppl.): 96 (1902). Fosliella (subgenus Heteroderma) zonalis forma myriocarpa sensu Bressan in Boll. Soc. adriat. Sci. nat. 59: 78 (1974). 411 SPECIMEN: Melobesia zonalis sensu Foslie (1900a). Coll. Brebner, Cawsand Bay, 11.11.1895, on rissoids (BM Algal Box Collection no. 506!) (Fig. 62E). 412 Y. M. CHAMBERLAIN EXCLUDED RECORD: Melobesia myriocarpa sensu Holmes, Alg. Br. rar. 218. (= mixture of species but mainly Pneophyllum confervicolum) (BM!). Lectotype: CO! Herb. P.-L. and H.-M. Crouan, 12 December 1856, Rade de Brest (Figs 61A, B) Slide: Chamberlain no. 1391. DiaGnosis: Epilithic species with thallus composed of hypothallium and epithallium only, ostiole forming a collar: differing from P. plurivalidum, P. lobescens, P. zonale, and P. concollum in having a vegetative thallus of hypothallium and epithallium only, and from P. zonale in not having long ostiolar filaments. DESCRIPTION: Smooth, brownish-pink crusts composed of hypothallium and epithallium only, up to c. 50 mm diameter, crust cells in surface view robust, oblong, 11-5—15-5 wm long x 6-12 wm wide, epithallial cells 2:5—5-5 wm long X 4-9 wm wide, intrafilament and branch cell trichocytes of similar size to crust cells; conceptacles prominent, domed, crowded, all except spermatangial conceptacles with beaked collar-like ostioles, carposporangial conceptacles 83-130 um internal diameter, 78-104 um high, spermatangial conceptacles small, prominent or immersed beside carpogonial conceptacles, with ostiole prolonged into a spout, 27-55 wm wide x 39-47 wm high, tetrasporangial conceptacles 109-156 ym internal diameter X 96-143 um high, up to 16 tetrasporangia per conceptacle, 44-64 wm long x 2442 um wide; bisporangial conceptacle (one only) 117 wm internal diameter x 91 wm high, bisporangium 48 um long xX 26 um wide. HistoricAL: This taxon was first described by the Crouan brothers (1867, as Melobesia) on material dredged from the ‘Rade de Brest, 12 Décémbre 1856’ (Figs 61A, B). Two sheets bearing this species are present in Herb. Crouan (CO); the samples comprise similar pieces of porcelain which bear crusts of Pneophyllum and are presumably part of the same collection. The sheet shown in Figures 61A, B has been selected for lectotype material, however, since the other Pee apes Pts = Ce ee eee as | | Pap io Bhi: PPI LI ¢ aasarere , (ae peal Prclobedio ep Sy aa. buy Drage cabs ik PAlo a Lr2 % ~- ase | Seer porestainns oT Verte : 4 Fig.61 Type specimen of Pneophyllum myriocarpum and appearance of cultured tetrasporangial crusts. A, Lectotype specimen of Melobesia myriocarpa P. Crouan & H. Crouan, Herb. Crouan, Concarneau. Packet closed. B, As A, packet open. Scale = 10 mm. C, Mature tetrasporangial crust with crust centre starting to become detached (seen in transmitted light). Scale = 5 mm. D, Young tetrasporangial crust (seen in reflected light). Scale = 1 mm. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 413 sheet is one of the Crouans’ decoratively mounted specimens lacking collection details. Crusts of more than one species are present on the lectotype sheet, but crusts of the present species predominate. The P. myriocarpum crusts have thin thalli with large crust cells and prominent conceptacles. A permanent slide (Chamberlain 1391) was made from a tetrasporangial crust. Annotations on the Crouans’ specimen (Fig. 61A) show that they initially classified the present alga as a species of Hapalidium (i.e. with a single layer of thallus cells), but they eventually published it as Melobesia (i.e. with more than one layer of thallus cells) despite its thin thallus. The species was described (P.-L. & H.-M. Crouan, 1867) as having thin, pink discs which were often confluent, with lobed edges; the conceptacles were very small, hemispherical, very numerous and covering the entire crust; the spores were elliptic and transversely divided into four. Two further Crouan specimens are present in Paris (PC); one, in the Herbier de France, is another piece of the same crust-bearing porcelain as that at Concarneau. The second specimen is in the Melobesia myriocarpa (zonalis) folder in Herb. Thuret and has been annotated by Thuret as follows: ‘Cet échantillon, que j’ai regu du frére des algologues brestois et qu’il a ainsi* nommé, “parce que la plante est vert jaune’, ne parait pas répondre a l’éspéce que MM Crouan ont désignée sous ce nom. Semble plutét devoir étre raprochée du Melobesia myriocarpa’. I did not examine either of these specimens microscopically because they are very sparse and firmly attached to the substrate, but presumably one or both of them were sent to Foslie when he was preparing his analysis of Crouan species (Foslie, 1900a). In this publication Foslie concluded that M. myriocarpa was conspecific with M. zonalis. Later Foslie (1908a) gave it the status of M. zonalis f. myriocarpa and then commented (Foslie, 1908) ‘it corresponds to M. farinosa f. solmsiana and M. minutula f. lacunosa with a very irregular, often subdichotomous thallus’. Foslie (1909) transferred M. zonalis to Heteroderma subgenus Pliostroma but did not list forma myriocarpa in this publication. Holmes (Alg. Br. rar. 218) identified crutose coralline epiphytes on Laminaria blades (BM: Ferry Bridge (Weymouth), August 1893 and Weymouth, August 1895) as Melobesia myriocarpa (listed as M. zonalis in Batters (1902)). Foslie (1905a) reidentified these crusts as a mixture of Fosliella farinosa (as Melobesia) and Pneophyllum limitatum (as Melobesia lejolisii+). 1 ex- amined some of these specimens (BM) and found mainly Fosliella farinosa and Pneophyllum confervicolum with some P. limitatum. A further collection included in Batters’ (1902) records of Melobesia zonalis is from Cawsand Bay, Plymouth (BM Algal Box Collection no. 506, coll. Brebner, 11.11.1895) and comprises numerous crust-covered rissoid shells (Fig. 62E). The crusts were initially named Melobesia lejolisii on the specimens, but were reidentified by Foslie (1900a) as M. zonalis. I have examined these crusts and find them to be mainly Pneophyllum myriocarpum. In common with epilithic species of Pneophyllum generally, no description of P. myriocarpum has been published since Foslie’s work. It has been recorded only rarely and seems to have been regarded as a form of P. zonale characterised by having crusts smothered with small, hemispher- ical conceptacles (e.g. Hamel & Lemoine, 1953, as Melobesia zonalis f. myriocarpa; Bressan, 1974, as Fosliella zonalis f. myriocarpa). I have not examined the samples on which any of these records were based. The written description of Fosliella tenuis (Adey & Adey, 1973, p. 398) as having extremely thin, nearly transparent crusts composed of hypothallium and epithallium only, possibly refers to the present species, although the illustrative diagrams (Adey & Adey, 1973, Tab. XIV) refer to their slide of the type which is material of Pneophyllum zonale (Fig. 82E). The rock specimen (BM, Algal Box Collection no. 1171) which bears crusts identified as Fosliella tenuis by W. H. Adey bears some crusts of the present species. CRUST APPEARANCE: In laboratory culture (Chamberlain, 1982) P. myriocarpum forms thin (up to c. 25 um deep), translucent, brownish-pink, epilithic crusts (Figs 61C, D) which are strongly * presumably it was sent as M. zonalis by the Crouans’ brother. + accepting Rosenvinge’s (1917) statement that Foslie came to regard all M. lejolisii growing on algae as forma limitata. 414 Y. M. CHAMBERLAIN adherent. The crust edge may produce fan-like outgrowths under unfavourable conditions and the crusts may grow up to about 30 mm diameter before the central part loosens and becomes detached (Fig. 61C). Under the S.E.M. (Fig. 62F) the crust cells are regularly rectangular, with epithallial concavities about twice as wide as long. Intrafilament (and very occasionally branch cell) trichocytes occur; they are the same size as normal crust cells, and an epithallial concavity is ae o Fig. 62 Scanning electron micrographs of Pneophyllum myriocarpum. A, Tetrasporangial crust (critical point dried). Scale = 250 um. B, Ostiole of tetrasporangial conceptacle. Scale = 25 um. C, Gametangial crust showing carposporangial (c) and spermatangial (s) ostioles. Scale = 50 wm. D, Spermatangial ostiole with remains of spout (arrow). Scale = 5 wm. E, Conceptacle of crust growing on Rissoa sp. (coll. Brebner). Scale = 50 ym. F, Crust cells with trichocytes (arrow). Scale = 10 wm. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 415 usually present. The lower surface shows tube-like hypothallial filaments at the periphery, anda flattened surface towards the centre. The conceptacles are abruptly prominent (Figs 62A, C, E), conical, or domed, and they often become extremely crowded. The conceptacle roof (Figs 62B, C, E) shows a jumble of filaments running at various levels and in different directions, which may also slough off. The ostiole is composed of filaments united into a small collar (Figs 62B, C, E). When growing on rissoid shells on the shore (Fig. 62E) the thallus curves round the shell and the conceptacles are particularly crowded. VEGETATIVE ANATOMY: P. myriocarpum is characterised by the thin, translucent crusts which are composed of robust, brick-shaped cells (Figs 63A, B) which in living material are brownish-pink in transmitted light; in cultured crusts the cells are full of floridean starch grains. Branch cell, and rarely intrafilament, trichocytes (Fig. 63B) are common under warm, long daylength conditions, and are of similar size to the crust cells. Large, abnormal, terminal trichocytes occur occasionally (Chamberlain, 1982); they appear to be similar to those recorded for P. limitatum by Suneson (1943, as Melobesia). : ws “ , ‘ 4 ’ Fig. 63 Crust cells of Pneophyllum myriocarpum. A, Focused to show epithallial cells (arrow). B, Focused to show hypothallial cells and trichocyte (arrow). Scale = 10 wm. GAMETANGIAL PLANTS: The crusts are monoecious with spermatangial conceptacles closely adjacent to carpogonial conceptacles (Figs 62C, D; 65E); they can clearly be seen in surface view on crusts growing on glass (Fig. 64F). The spermatangial conceptacles may be immersed in the edge of the carpogonial conceptacles (Fig. 64A) or prominent and dome-shaped (Fig. 65E). Elongated spermatangia are cut off at two sites on each initial (Fig. 655E) and when released the spermatangia are oval. The ostiole is prolonged into a spout which is occasionally forked. The carpogonial conceptacles (Fig. 64A) are raised and develop by the growth, stretching and rupture of perithallial cells; the centrally-borne procarps bear one or two carpogonia (Fig. 65A). As the conceptacle grows, filaments develop round the ostiole (Fig. 65A), and substantial downward-growing filaments are evident in many carpogonial conceptacles. Although the conceptacles rise abruptly from the thin thallus, a considerable amount of perithallial develop- ment contributes to the structure of the conceptacle wall. 416 Y. M. CHAMBERLAIN CARPOSPORANGIAL CONCEPTACLES: In carposporangial conceptacles the downward ostiolar fila- ments (Figs 64D, E; 65D) are well-developed, while the upper ones fuse and extend into an hyaline collar (Figs 64D, E; 65C, D). The fusion cell is usually rather flat. A fusion cell (Fig. 65B) was extruded from a conceptacle and the cell is seen to be somewhat segmented; there are the remnants of a number of procarps on the upper surface and the carposporangial filaments are initiated peripherally. TETRASPORANGIAL PLANTS: Tetrasporangial plants bear crowded conceptacles (Fig. 62A) which are similar to carposporangial conceptacles in structure and development, but usually somewhat larger. The tetraspores are borne peripherally and there is often considerable development of columella-like, central sterile paraphyses (Figs 64C; 66B). Young tetrasporangia are shown in Figure 66A. Tetrasporangia are very crowded in mature conceptacles and up to about 16 occur per conceptacle. BISPORANGIAL PLANTS: Only one bisporangial plant was found in British Isles collections of P. myriocarpum and this was seen only as a single section made on shore-collected material. Bisporangial conceptacles are common, however, among the crusts collected from the aquarium glass at Roscoff, and it can be seen (Fig. 64B) that the conceptacles are similar to tetrasporangial conceptacles. Bisporangia are more numerous than tetrasporangia, however, with up to about 40 being found in a single conceptacle. PHENOLOGY: Very little is known about the periodicity of P. myriocarpum on the shore although all the verified collections (Crouan - December; Brebner —- November; Yealm — October; Kimmeridge — throughout the summer; Roscoff aquarium — June; Co. Clare — June) are fertile. Tetrasporangial and gametangial crusts of P. myriocarpum have, however, been grown through several generations in laboratory culture (Chamberlain, 1982) under normal, seasonal day- length/temperature conditions and it has been found that, while it grows and reproduces at all seasons, growth and maturation are quicker during summer (April-October) than during the rest of the year. At Kimmeridge, P. myriocarpum grows on rissoid shells; studies by Wigham (1975) show that only a few individuals in such rissoid populations survive the winter so that P. myriocarpum crusts must, therefore, mainly grow and reproduce during the summer. ForM RANGE: The dimensions of British Isles and Roscoff samples of P. myriocarpum (Tabs 22; 23) suggest that there are no noticeable variations between the populations at these two localities. When grown in culture (Chamberlain, 1982) it was found that crust cells tended to be smaller in winter than summer and also that under very cold conditions areas of gametangial crust bearing only spermatangial conceptacles developed; adjacent carposporangial concep- tacles were, however, initiated as conditions became warmer. In laboratory culture trichocytes were produced in summer but not in the winter months (Chamberlain, 1982). DISTRIBUTION: British Isles: Kent, Dorset, Devon, Co. Clare. World: France, Italy. It is probable that P. myriocarpum occurs widely at least in Europe, but very little is known at present about epilithic simple, crustose Corallinaceae. Hasirat: P. myriocarpum is known to grow on fragments of glass and china and on shells, typically rissoids; it probably also grows on rocks. As an epilith, P. myriocarpum probably grows mainly in subtidal areas where it is not in danger of drying out since laboratory culture experiments have shown that it is unable to tolerate more than a very few minutes of desiccation. When growing on shells, however, P. myriocarpum is found in the intertidal zone. The only population which has been studied of this type is at Kimmeridge (Fig. 18). At this locality, Halopitys incurvus is acommon intertidal alga growing in shallow pools on the ledges; during the summer this alga houses large populations of rissoids many of which are covered with crusts of P. myriocarpum. Although many plants of Halopitys dry out considerably during hot weather since Fig. 64 Reproductive features of Pneophyllum myriocarpum. A, Vertical section of gametangial crust with carpogonial (ca) and spermatangial (s) conceptacles. Scale = 50 um. B, Vertical section of bisporangial conceptacle. Scale = 50 um. C, Vertical section of tetrasporangial conceptacle. Scale = 50 um. D, Vertical section of carposporangial conceptacle with simple ostiole. Scale = 50 wm. E, Vertical section of carposporangial conceptacle with ostiolar spout. Scale = 50 um. F, Surface view of growing crust with two carposporangial and one spermatangial (arrow) conceptacles. Scale = 100 um. 418 Y. M. CHAMBERLAIN Fig. 65 Gametangial and carposporangial structures of Pneophyllum myriocarpum. A, Vertical section of carpogonial conceptacle. B, Fusion cell and two carposporangial filaments extruded from concepta- cle. C, & D, Vertical section of two carposporangial conceptacles to illustrate differences in size and degree of immersion. E, Vertical section of spermatangial conceptacle. Scale = 20 ym (all figures). FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 419 OS QO osc RRO Y Fig. 66 Vertical sections of tetrasporangial conceptacles of Pneophyllum myriocarpum. A, Tetrasporan- gial initials (shaded). B, Mature conceptacle. they are in shallow pools, the rissoids move into the central parts of the plant which usually remain moist, and their attendant Pneophyllum crusts are saved from desiccation in this way. Lire HIsTory: A regular alternation of gametangial/carposporangial and tetrasporangial phases has been found in this species growing in laboratory culture (Chamberlain, 1982). TAXONOMIC NOTE: Pneophyllum myriocarpum and P. concollum appear very distinct when at their most characteristic. The epiphytic crusts of P. concollum have a well-developed perithal- lium and broad, slightly raised conceptacles, while epilithic crusts of P. myriocarpum lack a perithallium and have abruptly conical conceptacles. However, the collar-like ostiole is essen- tially similar in both species; in addition, crusts of P. concollum may have only a slightly developed perithallium, while Irish samples of P. myriocarpum growing on shells have deeper crusts than usual. It is possible, therefore, that further research may indicate that both species must be referred to the same taxon: if so, P. concollum will have to be subsumed in P. myriocarpum. 9. Pneophyllum lobescens Y. Chamberlain, sp. nov. REFERENCE: Melobesia zonalis sensu Batters pro parte in J. Bot., Lond. 40 (Suppl.): 96 (1902) (this refers to the specimens below). SPECIMENS: Melobesia myriocarpa sensu Batters, Duke’s Rock, Plymouth, 7 April, 1897 and Plymouth 12 March, 1900 (both in BM Algal Box Collection no. 505!). Ho.ortyrPe: BM! Y. M. Chamberlain no. 77/228, 29 October 1977, mouth of River Yealm, south Devon, at a depth of 8 m. Culture material nos. 104/4/Sa gametangial, 104/27 tetrasporangial and 104/4/3/b bisporangial. D1acnosis: Species epilithica cellulis crustae latioribus, filamentis perithallii profunditate usque ad 6 cellulas et ostiolis tholiformibus: a P. plurivalidio differt trichocytis non modo intra filamenta sed etiam ramos facientibus et absentia cellularum crustae longarum et sinu osarum, a P. zonali absentia filamen- torum ostiolarium longorum, et a P. myriocarpio perithallio profundo differt. Descriptio: Crustae phloginae ad ferruginescentes, diametro usque ad 10 mm, profunditate 90 wm, margine regulari, sed saepe lobescenti quum condiciones infavorabiles sunt, perithallium media in crusta profunditate usque ad 6 cellulas, cellulae perithallii (in sectione verticali) 8-16 wm latae x 8-5—23 wm altae, margo crustae ex hypothallio et epithallio modo ‘constatus, cellulae crustae aspectu superficiali relative latae, 6-5-16-5 um longae Xx 4-5—11 um latae, cellulae epithallii 34-5 wm longae x 4-5-11 um latae, trichocyti et intra filamenta et ramos facientes frequentes, amplitudine eadem ac cellularum crustae; conceptacula immersa, velut circuli indefiniti pallidi in crusta magis atrata, omnia, praeter ea spermatan- 420 Y. M. CHAMBERLAIN Table 22 Dimensions of vegetative structures (in ~m unless stated). Roscoff Crouan type Pneophyllum myriocarpum British Isles aquarium specimen crust diameter 30-50 mm crust depth c. 50 crust cell length (11-5) 14-5 (15-5) (10) 13 (14-5) (10-5) 13 (15) surface width (6) 10-5 (12) (6:5) 9-5 (12) (6-5) 8 (10) crust cell height c.13 epithallial cell length (2-5) 4 (5-5) (3) 3-5 (4-5) (4) 4-5 (5) surface view width (4) 6-5 (9) (5:5) 7-5 (9:5) (5S) 6 (7) epithallial cell height 344 hypothallial cell width VS height perithallial width cell height VS number trichocyte type intrafilament one branch cell (occasionally type seen branch cell) trichocyte length as crust cells surface view width as crust cells RIDGWAY COLOUR CODE: dry crust — dull indian purple, pl. XLIV fresh crust — dark vinaceous, pl. XX VII gialia, tholo ostioli in superficiei superiore, conceptacula carposporangialia diametro interno 88-112 um, altitudine 46-68 um; conceptacula spermatangialia immersa, 34-52 wm lata, 35-48 wm alta, ostiolo in canalem producto, conceptacula tetrasporangialia diametro interno 104-138 um, altitudine 65-91 um, usque ad 18 tetrasporangiis peripheralibus in unoquoque conceptaculo, paraphysibus centralibus sterilibus instructa, tetrasporangia 43-54-5 wm longa Xx 24-5—41-5 wm lata, conceptacula bisporangialia diametro interno 82-134 um, altitudine 59-93 um, usque ad 20 bisporangiis peripheralibus in unoquoque concepta- culo, paraphysibus centralibus sterilibus instructa, bisporangia 36-5—51 wm longa x 19-5-35 wm lata. Diacnosis: Epilithic species with rather broad crust cells, perithallial filaments up to 6 cells deep and domed ostiole: differing from P. plurivalidum in having branch cell as well as intrafilament trichocytes and in lacking long, sinuous crust cells, from P. zonale in not having long ostiolar filaments and from P. myriocarpum in having a deep perithallium. DescriPTIon: Crusts bright pink to brownish-pink, up to c. 10 mm diameter, 90 um deep, margin regular or often becoming lobed under unfavourable conditions, perithallium up to 6 cells deep in crust centre, perithallial cells (in VS) 8-16 wm wide X 8-5—23 um high; crust border of hypothallium and epithallium only, crust cells in surface view relatively broad, 6-5—16-5 wm long x 4-5-11 um wide, epithallial cells 3-4-5 wm long X 4-5-11 wm wide, intrafilament and branch cell trichocytes common, of similar size to crust cells; conceptacles immersed, appearing as pale indefinite circles in darker crust, all except spermatangial conceptacles with ostiole dome, carposporangial conceptacles 88-112 wm internal diameter X 46-68 wm high, spermatangial conceptacles 34-52 wm wide X 35-48 wm high, with ostiole prolonged into a spout, tetrasporan- gial conceptacles 104-138 mm internal diameter xX 65-91 mum high, up to 18 peripheral tetrasporangia per conceptacle, 43-54-5 wm long X 24-5—41-5 wm wide, central sterile paraph- yses present, bisporangial conceptacles 82-134 jm internal diameter X 59-93 wm high, up to 20 peripheral bisporangia per conceptacle, 36-5—51 wm long xX 19-5-35 wm wide, central sterile paraphyses present. Historica: P. lobescens is a newly described, epilithic species of Pneophyllum. It was originally collected from the mouth of the River Yealm, south Devon, where it was growing, together with P. zonale, P. plurivalidum, and P. myriocarpum, on glass bottles at a depth of about 8 m and it FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES Table 23 Dimensions of reproductive structures (jm unless stated). Pneophyllum myriocarpum British Isles Roscoff aquarium CARPOSPORANGIAL CONCEPTACLE external diameter surface diameter (70) 114 (156) (117) 128 (143) VS internal diameter (83) 101 (130) (99) 117 (130) VS height (78) 97 (104) (50) 87 (117) VS roof height (13) 15 (31) 13-15-6 roof type irregular no. basal cell rows 1 (rarely 2) 1 ostiole beaked beaked no. gonimoblast filaments €12 diameter carpospore SPERMATANGIAL CONCEPTACLE VS/surface width (27) 49 (55) (27) 36 (42) VS height (39) 42 (47) 22-29 length spermatium width spermatium TETRASPORANGIAL CONCEPTACLE external diameter surface diameter (106) 146 (171) (120) 143 (156) VS internal diameter (109) 131 (156) (143) 150 (156) VS height (96) 118 (143) (117) 139 (161) VS roof height (13) 18 (26) cris roof type irregular no. basal cell rows 1-2 1 ostiole beaked beaked no. tetrasporangia 8-16 8-14 length tetrasporangium (44) 53 (64) (57) 58 (63) width tetrasporangium (24) 29 (42) (31) 35 (42) BISPORANGIAL CONCEPTACLE one only external diameter surface diameter (101) 111 (156) VS internal diameter 117 (107) 122 (130) VS height 91 (65) 97 (117) VS roof height 21 (13) 15 (18) roof type no. basal cell rows 2 1 (rarely 2) ostiole beaked beaked no. bisporangia c. 20-c. 40 length bisporangium 48 (39) 44 (52) width bisporangium 26 (17) 23 (29) was subsequently grown in laboratory culture (Chamberlain, 1982). Two collections of Batters’s (BM) were found to be of the same taxon; these were growing on razor shells at Plymouth on 12 March 1900 and on glass at Duke’s Rock, Plymouth from a depth of 3-6 fathoms on 7 April 1897: Batters labelled them both ‘Melobesia myriocarpa Crouan’ but he later listed them (Batters, 1902) as ‘M. zonalis Fosl.’. CRUST APPEARANCE: In laboratory culture (Figs 67A, B; 69A) P. lobescens forms rather thin (up to 90 um deep), brownish or bright rosy-pink, epilithic crusts with pale edges which rapidly become irregularly lobed under unfavourable conditions. The crusts tend to detach easily and rarely grow to more than 10 mm diameter before the central part detaches. Under the S.E.M. (Fig. 67G) the crust cells can be seen to vary from elongated to broad, and the epithallial concavities vary from rounded to about twice as wide as long; branch cell and Y. M. CHAMBERLAIN Fig.67 Scanning electron micrographs of Pneophyllum lobescens. A, General view of crust. Scale = 500 um. B, Enlarged view of A showing crust lobes (1) and ostioles (2). Scale = 100 wm. C, Gametangial crust showing carposporangial (c) and spermatangial (s) ostioles. Scale = 50 um. D, Bisporangial crust. Scale = 100 um. E, Vertical fracture through tetrasporangial conceptacles. Scale = 50 wm. F, Lower crust surface. Scale = 10 um. G, Crust cells and trichocyte (arrow). Scale = 10 wm. H, Nodules from lower surface of crust. Scale = 5 wm. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 423 intrafilament trichocytes occur. On the younger parts of the lower surface (Fig. 67F) the hypothallial filaments are tube-like, and calcareous nodules are frequent (Fig. 67H). The conceptacles are immersed in the thallus (Figs 67C, D, E; 68A), and appear flat in fresh material; in dried material they are gently rounded, and appear as rather diffuse pale areas in a darker crust. The conceptacle roof is uniform and the ostiole is dome-like (Figs 68B, C, E). — i ae Fig. 68 Scanning electron micrographs to show conceptacle and ostiole features of Pneophyllum lobescens. A, Vertical section of carposporangial conceptacle. Scale = 25 wm. B, Young tetrasporangial ostiole.. Scale = 10 wm. C, Mature carposporangial ostiole. Scale = 10 wm. D, Spermatangial ostiole with ruptured spout (arrow). Scale = 10 wm. E, Bisporangial ostiole. Scale = 10 wm. F, Underside of tetrasporangial conceptacle roof. Scale = 25 wm. 424 Y. M. CHAMBERLAIN Fig. 69 Crust features of Pneophyllum lobescens. A, Bisporangial crust in culture showing thallus lobes (arrow). Scale = 50 um. B, Bisporangial crust cells with trichocytes (arrow). Scale = 10 wm. VEGETATIVE ANATOMY: The crust border is composed of hypothallium and epithallium only. In surface view (Fig. 69B) the crust cells vary from isodiametric to about one and a half times as long as wide, and usually contain many floridean starch grains. Epithallial cells vary from rounded rectangular to short and broad; they are dome-shaped to flattened in vertical section. Intrafilament and branch cell (Fig. 69B) trichocytes are common; they are about the same length as crust cells but often bulge sideways squashing adjacent cells. In vertical section (Figs 70E; 72B) the thallus appears regularly zoned horizontally. The perithallial filaments vary from 1-6 cells deep and their cells vary from short to markedly elongated; these cells are full of starch grains. GAMETANGIAL PLANTS: Gametangial plants are monoecious. The spermatangial conceptacles are adjacent to the carpogonial conceptacles (Figs 67C; 71A), and are flask-like chambers with a conspicuous ostiole and spout (Fig. 68D). The oval spermatia (Figs 70A; 71B) are larger (up to 8 um long) than in most of the described species of Pneophyllum. Carpogonial conceptacles (Fig. 71C) are immersed with one to two cell rows below the conceptacle. The procarps are often borne rather high in the conceptacle chamber, with conspicuous supporting cells and the central procarps bear one or two carpogonia with long trichogynes. CARPOSPORANGIAL CONCEPTACLES: The mature conceptacle (Figs 67C; 68A; 70B; 71A, D) is immersed in the thallus with only the dome-like structure, formed from the upper ostiolar filaments, protruding above the thallus surface (Fig. 68C). The downward ostiolar filaments are well-developed (Fig. 71D), and the development pattern formed by the remnants of the perithallial cells and the roof and ostiole filaments to which they give rise is often particularly clear in this species. The fusion cell is quite deep (Fig. 71D), and the peripheral gonimoblast filaments are about five cells long with the carposporangium considerably larger than the penultimate cell (Figs 70B; 71D). TETRASPORANGIAL PLANTS: Conceptacle shape and ostiole structure (Figs 67E; 68B, F; 70E; 72B) are similar to that seen in the carposporangial conceptacle. Young tetrasporangia (Fig. 72A) are interspersed with sterile cells, but mature tetrasporangia are borne peripherally (Figs 70E; 72B), and are plump and well-pigmented. A central columella of sterile cells is present, but the mature conceptacle is so full of tetrasporangia that the columella and the cells beneath the conceptacle are often squashed flat. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 425 . . ee Fig.70 Vertical sections of conceptacles of Pneophyllum lobescens. A, Spermatangial conceptacle. Scale = 20 um. B, Carposporangial conceptacle. Scale = 50 wm. C, Bisporangial conceptacle showing crowding of mature bisporangia. Scale = 20 um. D, Bisporangial conceptacle showing central sterile cells (arrow). Scale = 20 wm. E. Tetrasporangial conceptacle in a superimposed crust. Scale = 50 wm. 426 Y. M. CHAMBERLAIN 20um Fig.71 Vertical sections of gametangial crusts of Pneophyllum lobescens. A, Spermatangial and adjacent carposporangial conceptacle. B, Spermatangial conceptacle. C, Mature carpogonial conceptacle show- ing bicarpogonial procarps (arrow). D, Mature carposporangial conceptacle showing the fusion cell with gonimoblast filaments and growth of ostiole filaments from perithallial cells (arrow). BISPORANGIAL PLANTS: The development and structure of bisporangial conceptacles and ostioles (Figs 67D; 68E; 70C, D; 72C) is the same as for tetrasporangial conceptacles. The plump, oval bisporangia are also very numerous, and, when mature, burst out of the conceptacle when it is sectioned (Fig. 70C). PHENOLOGY: Collections from three areas have been seen of this species. In July 1973 subtidal bisporangial and gametangial material was collected in the Shetland Islands, in October 1977 material of all reproductive phases was collected on glass from the River Yealm at a depth of 8 m; while Batters collected gametangial plants from Plymouth in March and April. It is probable that the species grows throughout the year subtidally but in the laboratory growth and reproduction were slower in winter than in summer (Chamberlain, 1982). ForM RANGE: In laboratory culture, crusts grown in winter tend to be in poor condition, to have much more lobed edges, and more elongated cells than the smoother, more regular crusts, with large, more or less square cells, seen under warm but shaded summer conditions. Trichocytes (Figs 67G, 69B) occur abundantly in summer and may occasionally be present, particularly in an immature form, throughout the winter (Chamberlain, 1982). DISTRIBUTION: British Isles: Devon, Shetland Is. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 427 Hapsirat: This species is so far known only from glass fragments, stones and razor shells collected from the subtidal zone. Lire History: A regularly alternating Polysiphonia-type life history and a self-perpetuating bisporangial life history have both been demonstrated in laboratory culture. The time taken to complete life history phases varies with environmental conditions (Chamberlain, 1982). Table 24 Dimensions of vegetative structures (in wm unless stated). Pneophyllum lobescens British Isles crust diameter up toc. 10mm crust depth c. 90 crust cell length (6-5) 12 (16-5) surface width (4-5) 8 (11) crust cell height epithallial cell length (3) 4 (4-5) surface view width (4-5) 6-5 (11) epithallial cell height Cl hypothallial cell width (5) 13 (25) VS height 4-9 perithallial width (8) 10 (16) cell height (8-5) 13 (23) VS number up to 6 trichocyte type intrafilament and branch cell trichocyte length 11-5-13 surface view width 10-14-5 RIDGWAY COLOUR CODE: dry crust — vinaceous lilac, pl. XLIV tourmaline pink, pl. XXXVIII (lower sur- face) fresh crust — deep corinthian red, pl. XX VII 10. Pneophyllum plurivalidum Y. Chamberlain, sp. nov. Ho.otyPeE: BM! Y. M. Chamberlain no 77/228, 29 October 1977, mouth of River Yealm, south Devon, ata depth of 8 m. Culture slide no 104/21 (Fig. 73A) grown from spores from this original collection. DiaGnosis: Species epilithica perithallio profundo: a P. lobescenti differt cellulis longis sinuosis et a P. zonali conceptaculis immersis sine filamentis ostiolaribus. DESCRIPTION: Crustae leaves, roseae ad sanguineae, diametro usque ad 60 mm, profunditate 100 um, perithallium media in crusta profunditate usque ad 8 cellulas, cellulae perithallii (in sectione verticali) 6-18-5 wm latae xX 7-5-15 um altae, margo crustae ex hypothallio et epithallio modo constatus, cellulae crustae aspectu superficiali elongatae, sinuosae, 12-5—22-5 wm longae x 2-8 uum latae, cellulae epithallii 3-5-9 um longae x 1-5-6 wm latae, trichocytis intra filamenta bulbosis amplitudine cellulae crustae simili; conceptacula immersa, omnia, praeter ea spermatangialia, ostiolis tholiformibus, conceptacula carposporangialia diametro interno 65-81 wm, altitudine 39-73 wm, conceptacula spermatangialia immersa, diametro 39-52 um, altitudine 26-42 wm, ostiolo in canalem producto, conceptacula tetraspor- angialia incognita, conceptacula bisporangialia diametro interno 104-117 wm, altitudine 65-81 um, usque ad 16 bisporangii in unoquoque conceptaculo, bisporangia peripheralia, 36-547 wm longa x 23-5-31 um lata. Diacnosis: Epilithic species with deep perithallium; differing from P. /obescens in having long, sinuous crust cells and from P. zonale in having immersed conceptacles which lack ostiolar filaments. 428 Y. M. CHAMBERLAIN Table 25 Dimensions of reproductive structures (um unless stated). Pneophyllum lobescens British Isles CARPOSPORANGIAL CONCEPTACLE external diameter surface diameter (88) 105 (130) VS internal diameter (88) 99 (112) VS height (46) 60 (68) VS roof height (13) 21 (26) roof type uniform no. basal cell rows 1-2 ostiole domed no. gonimoblast filaments c. 14-18 diameter carpospore SPERMATANGIAL CONCEPTACLE VS/surface width 34-52 VS height 35-48 length spermatium 6-8:5 width spermatium 2-3-5 TETRASPORANGIAL CONCEPTACLE external diameter surface diameter (100) 122 (164) VS internal diameter (104) 120 (138) VS height (65) 76 (91) VS roof height (15) 20 (26) roof type uniform no. basal cell rows 1-2 ostiole domed no. tetrasporangia up toc. 18 length tetrasporangium width tetrasporangium BISPORANGIAL CONCEPTACLE external diameter (43) 51 (54-5) (24-5) 30 (41-5) surface diameter (91) 117 (143) VS internal diameter (82) 109 (138) VS height (59) 69 (93) VS roof height (8) 17 (26) roof type uniform no. basal cell rows 1-3 ostiole domed no. bisporangia up toc. 20 length bisporangium width bisporangium (36-5) 42-5 (51) (19-5) 26 (35) DEscriPTION: Smooth, reddish-pink to brown crusts up to 60 mm diameter, 100 um deep, perithallium up to 8 cells deep centrally, perithallial cells (in VS) 6-18-5 wm wide X 7:5—15 wm high, crust border composed of hypothallium and epithallium only, crust cells in surface view elongated, sinuous, 12-5—22-5 um long X 2-8 um wide, epithallial cells 3-5-9 wm long X 1-5-6 uum wide, bulbous intrafilament trichocytes of similar size to crust cells; conceptacles immersed, all except spermatangial conceptacles with dome-like ostioles, carposporangial conceptacles 65-81 wm internal diameter x 39-73 um high, spermatangial conceptacles immersed, 39-52 wm diameter X 26-42 um high with ostiole prolonged into a spout; tetrasporangial conceptacles unknown, bisporangial conceptacles 104-117 um internal diameter X 65-81 wm high, bisporan- gia peripheral, 36-5—47 um long X 23-5—31 um wide, up to 16 per conceptacle. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 429 Fig. 72 Vertical sections of tetra- and bisporangial conceptacles of Pneophyllum lobescens. A, Young tetrasporangial conceptacle showing tetrasporangia (shaded) developing between sterile cells. B, Mature tetrasporangial conceptacle with central sterile paraphyses. This also shows two superimposed crusts. C, Mature bisporangial conceptacle with central sterile paraphyses. Historica: P. plurivalidum is a long-lived epilithic species when grown in laboratory culture (Chamberlain, 1982). The same taxon was collected by Mme Lemoine from the glass of the aquarium tanks at the marine station at Roscoff, Brittany in August 1912 (PC) and identified as Melobesia zonalis. When I collected crusts from the same tanks in June 1979 only Fosliella farinosa and Pneophyllum myriocarpum were observed. P. plurivalidum appears to be similar to Fosliella valida (Adey & Adey, 1973). Both have a deep crust (Fig. 73C) and the immersed conceptacles are of a similar size (Tab. 27), but the perithallial cells of F. valida are much smaller than those of Pneophyllum plurivalidum. No details regarding the shape of the thallus cells in surface view or conceptacle structure and surface features are available for Fosliella valida. CRUST APPEARANCE: In laboratory culture (Chamberlain, 1982) P. plurivalidum forms substan- tial (up to 100 wm deep), epilithic crusts (Figs 73A; 74A) which are pinkish-red to dense brownish-purple, becoming darker in the older parts. The crusts are strongly adherent and usually have a smooth, regular periphery which does not become lobed. Crusts have grown to 60 mm diameter to date. The centre of older crusts becomes loosened and finally detaches. Under the S.E.M. (Fig. 75C) the crust cells are elongated with relatively large, rounded- rectangular, epithallial concavities. The cells are broadest at the epithallial concavity, which gives them a characteristically wavy outline. Intrafilament trichocytes occur (Fig. 75D); they appear swollen relative to adjacent cells and the epithallial concavity is absent. The lower surface of the crust (Fig. 75F) shows flattened filaments with circular holes at intervals. The conceptacles are immersed in the thallus (Figs 74C, D; 75E), but may become prominent in older dried material (Figs 74A, B). The conceptacle roof (Fig. 74 C, D) is smooth except in the immediate vicinity of the ostiole in younger conceptacles, while the thallus surface is irregularly shed in older conceptacles (Fig. 74B), when the ostiole dome (Figs 75A, B) becomes surrounded by rounded cells with large epithallial concavities. VEGETATIVE ANATOMY: In culture the crusts have a border composed of hypothallial and epithallial cells only, but this is probably not usually visible in shore-collected specimens. The 430 Y. M. CHAMBERLAIN Fig. 73 Vegetative features of Pneophyllum plurivalidum and Fosliella valida. A, Type specimen of Pneophyllum plurivalidum with bisporangial and gametangial (arrow) crusts. Scale = 5 mm. B, Surface view of Pneophyllum plurivalidum crust showing elongated crust cells and large, oval, epithallial cells. Scale = 10 um. C, Vertical section of Fosliella valida (Adey slide no. 70-2 10-30C 4) showing small, more or less isodiametric perithallial cells and edge of a carposporangial conceptacle. Scale = 25 wm. crust cells are long and narrow (Fig. 73B), and the epithallial cells are radially ovate in surface view and domed in vertical section. Intrafilament trichocytes are present but are not divided into a pigmented and non-pigmented parts so that they are difficult to locate except by the absence of an epithallium in decalcified crusts; they are of the same order of size as the crust cells. A perithallium up to eight cells deep develops, the cells of which vary from being more or less isodiametric to vertically elongated in VS (Figs 75E; 76A, B; 77). The perithallial cells usually contain floridean starch grains (e.g. Figs 76B-E) which increase in quantity and size from the crust surface downward. GAMETANGIAL PLANTS: The crusts are monoecious with spermatangial conceptacles usually adjacent to carpogonial conceptacles (Figs 74A—C; 76B). The young spermatangial conceptacle (Fig. 77D) comprises a fertile disc at the base of the conceptacle bearing elongate cells; the mature conceptacle (Figs 76E; 77E) is globose and immersed in the thallus. Although the ostiole is prolonged into a spout, it is not very conspicuous in surface view (Figs 74B, C). Young carpogonial conceptacles (Fig. 77A) develop within the perithallium and, when mature (Figs 76C; 77B), the fertile disc in the centre of the conceptacle floor bears procarps. The central ‘WM ODT = 2[89S “IdA09 Wo}eIp AAvoy Y}IM JsNIO eIsuvlodsig ‘q ‘win (OT = a[ed¢g ‘sajoe}daou09d (s) JeIsueJeuods pure (ed) [eTUOSOdIe9 SUIMOYs }sNJO [eIsULeJOWeS BUNOZ ‘Dd ‘wi! EYT = a]B9S ‘sajor}daou09 (s) yetsueyeutsods pure (9) [eIsuerodsodied SuImoys Y JO MOIA posielUY ‘g ‘wi! ESZ = ZBI “so[oe}da0U0d (s) [eIsUR}JeUIOdS pur (9) [eId -uvodsodies ZUIMOYS }sNJo [eIsUL}OUTLS INJLUI JO MIIA [PIDUDH ‘Yy ‘wnpyvatanjd wn dydoaug Jo sydesBOIIIU UOIDIT[A BuIUURIS «pL “BIA 4 orn i / x 4 f 4 al 2) im — 4 en a = 2 Fa = ee) = g = » 3 = Sy g z Q Zz < << 3 = — S & Y. M. CHAMBERLAIN Fig.75 Scanning electron micrographs of Pneophyllum plurivalidum. A, Bisporangial ostiole. Scale = 10 um. B, Carposporangial ostiole. Scale = 10 wm. C, Crust cells. Scale = 10 um. D, Trichocytes. Scale = 10 um. E, Vertical fracture of bisporangial crust. Scale = 100 um. F, Lower surface of thallus showing ‘pores’. Scale = 5 um. procarps bear one or two carpogonial branches, each with long trichogynes (Figs 76C; 77B). The fertile area is surrounded by vertically stretched perithallial cells which initiate roof and ostiolar filaments apically; as these filaments develop they start to push off the original thallus surface (Figs 76C; 77A). CARPOSPORANGIAL PLANTS: As the carposporangial conceptacles mature, the stretched perithal- lial cells disintegrate and the original thallus surface is sloughed off round the ostiole (Figs 76D; 77C). The downward ostiolar filaments are short and somewhat swollen at the tip (Figs 76D; 77C), while the upward ostiolar filaments are short, but immediately round the ostiole they form a dome-like structure of broad cells (Figs 75B; 76D; 77C), the top parts of which break off as the ostiole ages (Fig. 75B). The fusion cell (Figs 76D; 77C) is quite small and deep, it is closely FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 433 Fig. 76 Vertical sections of reproductive crusts of Pneophyllum plurivalidum. A, Bisporangial crust. Scale = 25 um. B, Gametangial crust showing carposporangial and spermatangial (arrow) conceptacles. Scale = 25 wm. C, Carpogonial conceptacle. Scale = 10 zm. D, Carposporangial conceptacle. Scale = 10 um. E, Spermatangial conceptacle. Scale = 10 um. Y. M. CHAMBERLAIN 434 ‘9]9ejda0u09 [eIsue}eULIads oINjeU JO UON}IES [BOT}IOA ‘q “9[oe}d90U09 JeISuLeULIOds SunoK Jo uordes [eonJoA ‘q “(MOLIe) s}UdWeTY sJONSO preMUMOP ‘UdTOMS 94} SULMOYsS pue (YoR]q) SoRyINs snjyey) Jo surewos YM gjoe}daouon [e1suesodsodies Jo uoNdes [eII9A ‘Dd *(Z) eIUOSodse9 OM} BuLIVAg dieso1d pue (T) 1199 femeyiied wo Suidojaaop syuoweyy ajoNso Surmoys a[oe}da0u09 [eIUOSOdIed oINJeU JO UONDeS [BONIOA ‘g “(MOIIR) dRJINS snyTey) 24} MOJaq SuIdo[aAep JooI Mou ZuIMOYs gjoejdaouoo [etuosodseo SuNOA Jo UONes [RONIDA “W ‘wnpyvarunjd wnykydoaug Jo sajse\daou09 jeisuerodsodieoyeisuejoury LL “BI i aus os O Sigg eG wee | O . a) OC) fk se5 Ooxuw FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 435 appressed to the basal cells, and carposporangial filaments up to seven cells long are borne peripherally. TETRASPORANGIAL PLANTS: Attempts to grow crusts initiated from carpospores are in progress; it is probable that these plants will be tetrasporangial. BISPORANGIAL PLANTS: These are similar to the carposporangial plants (Figs 74D; 75A,E; 76A) and the conceptacle and ostiole development and structure follow the same pattern. The bisporangia (Fig. 76A) are borne peripherally on stalk cells; there is no development of sterile filaments or columella. PHENOLOGY: Little information is available about the occurrence of this species in nature. In the laboratory, however, the crusts remained healthy throughout the winter, but grew and reproduced much more slowly at this season than during the summer (Chamberlain, 1982). ForM RANGE: In crusts grown in the laboratory crust cells are noticeably shorter and broader [(5) 8-5 (9-5) um long X (5) 6:5 (8) «wm wide] in winter conditions than the characteristically elongated cells seen under favourable conditions. Trichocytes are present under warm, well- illuminated conditions, but absent in colder darker conditions (Chamberlain, 1982); very large, irregular crust cells and trichocytes are seen occasionally as in P. myriocarpum. DISTRIBUTION: British Isles: Devon. World: France (Brittany). HasiraT: Known only on fragments of glass collected from the subtidal zone and the glass walls of aquarium tanks. Lire History: A self-perpetuating, bisporangial life history has been demonstrated. The time taken to complete life history phases varies with environmental conditions (Chamberlain, 1982). Table 26 Dimensions of vegetative structures (in jm unless stated). Adey & Adey (1973) Pneophyllum plurivalidum British Isles Fosliella valida crust diameter up to 60 mm crust depth up to 100 re (8 crust cell length (12-5) 17 (22:5) 7-14 surface width (2) 6-5 (8) 5-9-5 crust cell height epithallial cell length (3-5) 7 (9) 2°5-5:5 surface view width (1:5) 4-5 (6) 3-8 epithallial cell height (3) 4-5 (6) hypothallial cell width (5-5) 11 (18-5) VS height (5-5) 7 (8-5) perithallial width (6) 10 (18-5) 2-5-6°5 cell height (7-5) 10 (15) 3-5-8-5 VS number up to 8 up to 7 or more trichocyte type intrafilament rare trichocyte length as crust cells surface view width as crust cells RIDGWAY COLOUR CODE: dry crust — deep purplish vinaceous, pl. XLIV fresh crust — hydrangea red, pl. XX VII 11. Pneophyllum zonale (P. Crouan & H. Crouan) Y. Chamberlain, comb. nov. BasionyM: Hapalidium zonale P. Crouan & H. Crouan in Annals Sci. nat. (Bot.) IV, 12: 284 Pl. 21, Fig. A(1-7) (1859) (see Fig. 44A of present study). 436 Y. M. CHAMBERLAIN Table 27 Dimensions of reproductive structures («wm unless stated). Adey & Adey (1973) Pneophyllum plurivalidum British Isles Fosliella valida CARPOSPORANGIAL CONCEPTACLE external diameter surface diameter (70) 78 (86) VS internal diameter (65) 73 (81) (77) 89 (95) VS height (39) 54 (73) (37) 43 (47) VS roof height (13) 19 (26) (10) 13 (15) roof type irregular no. basal cell rows 2-3 ostiole domed no. gonimoblast filaments ey diameter carpospore SPERMATANGIAL CONCEPTACLE VS/surface width (39) 45 (52) (47) 53 (57) VS height (26) 33 (42) (25) 27 (30) length spermatium (4-5) 7 (9-5) width spermatium (2) 2-5 (3-5) TETRASPORANGIAL CONCEPTACLE external diameter surface diameter VS internal diameter (75) 87 (100) VS height (55) 60 (77) BS roof height (12) 14 (16) roof type no. basal cell rows ostiole no. tetrasporangia length tetrasporangium width tetrasporangium BISPORANGIAL CONCEPTACLE external diameter surface diameter (78) 110 (130) VS internal diameter (104) 113 (117) VS height (65) 76 (81) VS roof height (13) 16 (21) roof type smooth to irregular no. basal cell rows 2-4 ostiole domed no. bisporangia up toc. 16 (36-5) 43 (47) (23-5) 27 (31) length bisporangium width bisporangium NOMENCLATURAL SYNONYMS: Melobesia (subgenus Eumelobesia) zonalis (P. Crouan & H. Crouan) Foslie in K. nor. Vidensk. Selsk. Skr. 1898 (3): 11 (1898). Melobesia (subgenus Heteroderma) zonalis (P. Crouan & H. Crouan) Foslie in K. nor. Vidensk. Selsk. Skr. 1899 (7): 4 (1900 [‘1899"]). Lithophyllum (subgenus Carpolithon) zonale (P. Crouan & H. Crouan) Foslie in K. nor. Vidensk. Selsk. Skr. 1905 (5): 9 (1905). Lithophyllum (subgenus Pliostroma) zonale (P. Crouan & H. Crouan) Foslie in K. nor. Vidensk. Selsk. Skr. 1908 (7): 20 (1908). Melobesia (subgenus Pliostroma) zonalis (P. Crouan & H. Crouan) Foslie in K. nor. Vidensk. Selsk. Skr. 1908 (11): 4 (1908) — forma typica [Foslie (19085) said that Foslie (1908a) should have read Melobesia (Pliostroma) zonalis not Lithophyllum]. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 437 Heteroderma (subgenus Pliostroma) zonale (P. Crouan & H. Crouan) Foslie in K. nor. Vidensk. Selsk. Skr. 1909 (2): 56 (1909). Fosliella (subgenus Pliostroma) zonalis (P. Crouan & H. Crouan) Feldmann in Rev. algol. 11: 317 (1942 [‘1939]). Fosliella (subgenus Heteroderma) zonalis (P. Crouan & H. Crouan) Bressan in Boll. Soc. adriat. Sci. nat. 59: 398 (1974). Fosliella zonalis (P. Crouan & H. Crouan) Ganesan (sic); Parke & Dixon in J. mar. biol. Ass. U.K. 56: 534 (1976). REFERENCES: Hapalidium zonale P. Crouan & H. Crouan; P.-L. and H.-M. Crouan, Florule de Finistére: 149 (1867); Foslie in K. nor. Vidensk. Selsk. Skr. 1899 (7): 4 (1900 [‘1899’]). Melobesia (subgenus Heteroderma) zonalis (P. Crouan & H. Crouan) Foslie; Foslie in K. nor. Vidensk. Selsk. Skr. 1900 (5): 21 (1900). Melobesia (subgenus Pliostroma) zonalis (P. Crouan & H. Crouan) Foslie; Hamel and Lemoine in Archs Mus. natn. Hist. nat. Paris VII, 1: 108 (1953 [‘1952’]) — excluding Fig. 77; Feldmann in Trav. Stn. biol. Roscoff, suppl. 6: 79 (1954). Melobesia zonalis sensu Newton, A handbook of British seaweeds: 302 (1931) — pro parte. Melobesia trichostoma Rosenv. in K. danske Vidensk. Selsk. Skr. VII, 7: 253 (1917). Fosliella tenuis Adey & Adey in Br. phycol. J. 8: 398 (1973) (as to Pl. XIV and slide 78-1650-70E; description possibly includes Pneophyllum myriocarpum). EXCLUDED RECORDS: Melobesia (subgenus Pliostroma) zonalis sensu Lemoine in Archs Mus. natn. Hist. nat. Paris V, 5: 140 Fig. 7 (1913) (= Fosliella farinosa), in Bull. Lab. marit. Mus. Hist. nat. St. Servan 7: 20 (1931) (= Pneophyllum myriocarpum); Hamel and Lemoine in Archs Mus. natn. Hist. nat. Paris VI, 1: Fig. 72 (1953 [‘1952’]) (= Fosliella farinosa). Various other Pneophyllum species described or referred to by the Crouan brothers have been quoted (e.g. by Hamel & Lemoine, 1953, p. 108 and Bressan, 1974, p. 77) as synonyms of P. zonale. Examination of the Crouan collection (CO) showed that most of the samples on which these species were based were either epiphytes or such a mixture of epilithic crusts that it seems preferable to exclude them from the synonymy. They comprise: Hapalidium coccineum P. Crouan & H. Crouan in Annals Sci. nat. Bot. IV, 12: 285 (1859) (= Melobesia membranacea). Hapalidium phyllactidium sensu P. Crouan & H. Crouan in Annis Sci. nat. Bot. IV, 12: 286 (1859) — changed to H. confervoides in P. and H. Crouan, Florule du Finistere: 149 (1867) (= mixture of epilithic crusts). Hapalidium roseum sensu P. Crouan & H. Crouan, Florule du Finistére: 149 (1867) (= mixture of epilithic crusts including Pneophyllum zonale). Hapalidium callithamnioides P. Crouan & H. Crouan, Florule du Finistére: 149 (1867) (= Pneophyllum confervicolum f. minutulum q.v.). Melobesia myriocarpa P. Crouan & H. Crouan, Florule du Finistére: 150 (1867) (= Pneophyllum myriocarpum). LecrotyPe: CO! Herb. P.-L. & H.-M. Crouan, 24 January 1859, Rade de Camaret (dredged), France (Figs 78A, B, D). Chamberlain slide no. 1383. D1aGnosis: Plants mainly epilithic or epizoic, sometimes epiphytic, from intertidal pools or subtidal, with perithallium up to 8 cells deep and conceptacles with long, flattened ostiolar filaments: differing from P. lobescens, P. plurivalidum and P. myriocarpum in having larger conceptacles with ostiolar filaments, and from P. rosanoffii and P. limitatum in having larger conceptacles with a deeper roof of smaller cells and flattened ostiolar filaments. DEScRIPTION: Dull, reddish or mauvish-pink crusts up to a few millimetres in diameter, 170 wm deep, often superimposing, frequently mixed with other species, perithallium up to 8 cells deep in crust centre, crust border of hypothallium and epithallium only (at least in cultured material), perithallial cells (in VS) 7-14-5 wm wide X 10-28-5 um high; crust cells in surface view broadly rectangular, 10-17 wm long x 513-5 ym wide; epithallial cells 3-5-6-5 wm long X 4-9-5 wm wide, intrafilament and branch cell trichocytes 20-30 wm long X 11-14 wm wide; conceptacles gently raised, all except spermatangial conceptacles with long, wispy ostiolar filaments, carpos- porangial conceptacle (one only) 135 wm internal diameter x 73 wm high, tetrasporangial 438 Y. M. CHAMBERLAIN oa g pesohie “a ; Fig. 78 Type material and crust appearance of Pneophyllum zonale. A, Packet containing the type specimen. B, Packet opened: the lectotype crust of Hapalidium zonale P. Crouan & H. Crouan is on the smallest piece of glass (arrow). Scale = 10 mm. C, Bisporangial crust of Pneophyllum zonale grown in laboratory culture. Scale = 1 mm. D, Type crust showing thallus surface. Scale = 25 wm. conceptacle 156-240 um internal diameter x 78-99 um high, tetrasporangia 49-5—74 wm long x 2641-5 um wide, bisporangial conceptacles 96-260 jm internal diameter X 65-117 wm high, up to about 60 bisporangia per conceptacle, 41-5—73 um long xX 26-34 wm wide. Historica: Because of the technical difficulty in examining epilithic simple crustose corallines the identity of the Crouan brothers’ (1859) Hapalidium zonale has hitherto remained somewhat vague and the specific epithet has been generally applied to many simple epilithic crusts. However, when dimensions and descriptions have been given, they have usually indicated that a crust of up to 7-8 cells deep with large conceptacles (c. 300 ~m diameter) are features regarded as typical of this taxon (e.g. Feldmann, 1942; Hamel and Lemoine, 1953; Bressan, 1974). Examination of the Crouans’ herbarium in Concarneau revealed three collections labelled Hapalidium zonale by the Crouans. One collection is undated (Fig. 1) and displayed with the drawings as in the published description (P.-L. and H.-M. Crouan, 1859), whilst the other two are dated 1859 and 1862 respectively. The Crouans (1859) cited material collected up to and including January 1859 (p. 301) and it is from the collection dated 1859 that the lectotype (Figs 78A, B, D) has been selected. The collection comprises a packet containing a piece of mica on which nothing can be seen and three pieces of glass bearing a number of small, calcareous crusts. These crusts belong to various taxa. There is, however, one distinct and thicker tetrasporangial crust (Fig. 78B arrowed) which corresponds well with the Crouans’ illustration (see Fig. 44A) of the crust cells, conceptacle, and tetrasporangium of H. zonale. Scanning electron micrographs (Fig. 78D) show this crust to have a deep thallus and elongated crust cells; the conceptacles are somewhat raised and measurements of conceptacles and cells, made on a decalcified whole mount, are shown in Tables 28 and 29. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 439 Picquenard (19125) published a catalogue of the red algae of the Crouan herbarium in which he gave the following references for H. zonale: ‘H. phyllactidium Crouan excl. synon.; H. confervoides Crouan excl. synon. (non Batters), Melobesia myriocarpa Crouan’. Of these names M. myriocarpa is a separate species (see Pneophyllum myriocarpum), while the Hapalidium phyllactidium (syn. H. confervoides) collections in Herb. Crouan show a mixture of crust types probably belonging to several species (see also under Pneophyllum confervicolum). Foslie (1900a) discussed Hapalidium zonale on the basis of a Crouan collection sent to him from Paris. There are three such collections in Paris. One in Herb. Thuret is labelled Brest 1879, the other two are in the Herbier de France and are labelled as being ‘ex Herb. Crouan’ with a reference to the Crouans’ (1859, 1867) publications. There is no indication as to which material Foslie received and his slide (TRH) has now dried out. Examination of one of the Herbier de France collections shows it to be young material of the present species. Foslie (1900a) described the Paris material of Melobesia zonalis as having a thallus only one or two cells deep with conceptacles 150-200 4m diameter seen from above which agrees better with Pneophyllum myriocarpum than P. zonale as now defined. On the other hand, Foslie (1900a) mentioned some crusts on glass, collected at 2-5 fathoms at Plymouth, which Batters sent him and which Foslie describes as having a thallus up to eight cells deep with conceptacles 150-200 um from above. This is presumably the material identified by Batters as Melobesia myriocarpa (BM algal box collection no. 505) which is Pneophyllum lobescens. Foslie (1900a) also referred to some material collected by Batters at Plymouth growing on ‘small shells of snails’ which is presumably the collection on rissoid shells (BM algal box collection number 506) which is Pneophyllum myriocarpum (q.v.). Lemoine’s (1913) record and illustration of P. zonale (as Melobesia) from Clew Bay, Ireland, was based on Fosliella farinosa, material of which is in BM-K and DBN and shows superimposed bisporangial crusts. Rosenvinge’s (1917) description and slides (C) of Melobesia trichostoma agree closely with the present description. Pneophyllum zonale has most frequently been recorded from French localities and the records almost certainly represent more than one species. Lemoine (1931), for example, comments that material from the St Servan area (northern Brittany) was very thin and had conceptacles 175-200 wm diameter xX 120-150 wm high. The material on which the St Servan records were based is in PC; unfortunately the crusts are so sparse that identification is not feasible, although Lemoine’s description agrees well with P. myriocarpum. The dimensions given by Bressan (1974) for Italian material agree with P. zonale. Finally Fosliella tenuis Adey & Adey (1973) appears, from the written description, to be partly Pneophyllum myriocarpum (q.v.); however, the slide cited as being from the type specimen (67-16, 50-70E from Oslofjord) has been examined and it comprises sections of gametangial P. zonale with the characteristic zonale ostiole (Fig. 82E); the Adeys’ illustrative diagram and table of dimensions (Adey & Adey, 1973, Tab. XIV) is also based on this entity and was prepared from the same slide (Norris & Townsend, 1981). CRUST APPEARANCE: In culture (Chamberlain, 1982), crusts of P. zonale (Fig. 78C) grew to c. 10 mm diameter, and appeared brownish-pink with a border composed only of hypothallium and epithallium, which frequently became lobed because the crusts were growing on glass slides to which they did not adequately adhere. On the shore, individual crusts were most often seen on Littorina, Patella, and other shells in the intertidal zone, and such crusts were thick and brownish-mauve in colour. Layers of superimposed crusts as illustrated by Rosenvinge (1917) were not observed in the present investigation, but in cultured crusts it was noted that spores are often not liberated through the ostiole but released downwards when the bottom of the conceptacle falls out (Fig. 79B). The spores then germinate either on the substratum below the parent crust or on the under surface of the crust, and it is probably by this means that the crust layers are built up — with the older crusts eventually being pushed off by the younger ones below. Under the S.E.M. (Fig. 79A) the crust periphery is seen to be composed of broadly rectangular cells with conspicuous, ovate epithallial concavities, intrafilament and branch cell trichocytes. 440 Y. M. CHAMBERLAIN ee Fig.79 Scanning electron micrographs of laboratory cultured bisporangial crusts of Pneophyllum zonale. A, Crust cells with trichocyte (arrow). Scale = 10 um. B, Underside of crust showing detached conceptacle base (1) and sporelings (2) developing on lower crust surface. Scale = 500 wm. C, Senescent conceptacle. Scale = 50 wm. D, Young crust showing very early (1) and slightly later (2) conceptacle development. Scale = 500 wm. In fresh and dried material the conceptacles appear gently rounded, and under the S.E.M. the conceptacle surface is uniform in young material (Fig. 79D), but the thallus surface cracks off irregularly in older conceptacles (Fig. 79C) to reveal the more or less isodiametric roof cells with large, central, epithallial concavities. The ostiole (Figs 830A—E) becomes surrounded by long, flat, papery-looking filaments in the mature conceptacle; in the early stages these filaments appear as a rim round the ostiole; when they start to elongate they are fused to each other and only become free when mature. In senescent conceptacles the filaments wither away. Scanning electron micrographs of vertically fractured crusts show the deep conceptacle roof and perithallium (Fig. 80F). VEGETATIVE ANATOMY; The peripheral cells in cultured crusts (Fig. 81A) are broadly rectangular in surface view with conspicuous, more or less ovate epithallial cells, and the crust cells contain floridean starch grains. The perithallial filaments (Fig. 14) vary from being one to eight cells deep. Trichocytes (Fig. 81A) are frequent in summer months; they may occur within the filaments or in branch cells and are similar in size or somewhat larger than the surrounding crust FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES Fig.80 Scanning electron micrographs to show conceptacle structure and stages in ostiole development in bisporangial crusts of Pneophyllum zonale grown in laboratory culture. A, Early stage of conceptacle growth shown as a depression in the thallus surface. Scale = 50 wm. B, Ostiole filaments beginning to surround ostiole depression. Scale = 10 wm. C, Ostiole filaments opened out but still fused. Scale = 10 pm. D, Mature ostiole filaments, critical point dried, showing flattened shape. Scale = 10 um. E, Fused ostiole filaments starting to rise up. Scale = 25 wm. F, View from below of two adjacent conceptacles, showing deep conceptacle roof (1), ostiole in centre of underside of conceptacle roof (2), and lower surface of the thallus (3). Scale = 100 wm (both figures). 442 Y. M. CHAMBERLAIN i « .. " : : sshenieesdentarascmememmimanereteamen Fig. 81 Crust cells and characteristic ostiole filaments of Pneophyllum zonale. A, Crust cells with trichocyte (tr). B, Ostiole of bisporangial conceptacle showing upward (1) and downward-pointing (2) ostiole filaments. Scale = 10 um. cells. When mature they are divided by a radial wall into a pigmented part and a colourless, hair bearing part, and the epithallial cell is absent. GAMETANGIAL PLANTS: Only one gametangial plant, in poor condition, has been found in the present investigation; it was growing on Chondrus crispus at Bembridge on 6 February 1978. In VS it shows a young spermatangial conceptacle beside an empty, presumably carposporangial, conceptacle. From the descriptions and slides of Rosenvinge (1917, as Melobesia trichostoma) and Adey & Adey (1973, as Fosliella tenuis, see Fig. 82E) it is evident that the carposporangial and tetrasporangial conceptacles are similar to the bisporangial conceptacles in structure, and the bisporangial conceptacles will, therefore, be described in detail as they were the only type found in quantity in the present study. TETRASPORANGIAL PLANTS: Tetrasporangial plants (Fig. 82D) have been found on only two occasions, at Bembridge and Anglesey, and both samples were growing on Chondrus crispus. The Anglesey conceptacles measure 216-240 um internal diameter as compared with the Bembridge ones which measure 156-169 um, but the overall size range is similar to that found for the bisporangial conceptacles. The tetrasporangia develop peripherally on stalk cells, and sterile paraphyses are present in the centre of the conceptacle (Fig. 14A). BISPORANGIAL PLANTS: The young conceptacle is gently raised above the thallus surface (Figs 79D; 82A) with an internal cavity which is about twice as wide as high. The cultured conceptacles tend to be larger and more regularly formed than those of the shore material which are often humped and misshapen as were the tetrasporangial and carposporangial ones observed by Rosenvinge (1917). The conceptacle cavity develops by the vertical stretching (Fig. 82A) and eventual rupture of the perithallial cells. The roof filaments are initiated from the upper ends of the perithallial cells (Fig. 83) and form deep filaments of cells (Figs 14B; 80F; 82B), with small FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 443 - i alae * % = =" Fig. 82 Conceptacle features in decalcified crusts of Pneophyllum zonale. A, Vertical section of young bisporangial conceptacle grown in laboratory culture showing stretched perithallial cells (arrow). Scale = 50 um. B, Vertical section of mature bisporangial conceptacle grown in laboratory culture; note the deep roof filaments (arrow). Scale = 50 wm. C, Surface view of ruptured bisporangial conceptacle grown in laboratory culture, showing bisporangia at various stages of maturity and ostiole filaments (arrow). Scale = 50 wm. D, Vertical section of mature tetrasporangial conceptacle from Bembridge growing on Chondrus; note central sterile paraphyses (arrow). Scale = 50 wm. E, Vertical section of carposporan- gial ostiole from slide of type material of Fosliella tenuis (Adey & Adey, slide no. 67-16, 50-7E). Scale = 20 wm. lumina, the filaments terminate in domed epithallial cells. The remnants of the upper parts of perithallial cells which give off the roof filaments are shown in Figure 83, while the position which they held can be deduced in an S.E.M. preparation of the underside of the conceptacle roof (Fig. 11E), in which the rows of pit connections indicate the site at which each roof filament was given off; the elaborate nature of the pit plug can also be seen in this material (Fig. 11A). The ostiolar filaments (Figs 81B; 83) are very well developed in P. zonale; the downward filaments are robust and plentiful, while the upward filaments are long and characteristically wispy, appearing to taper off indefinitely at the apex as is indicated in their S.E.M. appearance previously described. In VS the ostiolar filaments appear to dip down before rising up through the ostiole, where they form a criss-cross pattern (Fig. 81B). These features are also seen in Rosenvinge’s (1917) drawings and specimens (C) of Melobesia trichostoma and are instantly recognisable features of Pneophyllum zonale. Young bisporangia (Fig. 82A, C) are more or less farctate and develop peripherally on stalk 444 Y. M. CHAMBERLAIN Fig. 83. Roof and ostiole structure of bisporangial conceptacle of Pneophyllum zonale. 1 = shrivelled remains of perithallial cell; 2 = deep roof filaments; 3 = upward ostiole filaments; 4 = downward ostiole filaments. cells on the conceptacle floor; the central sterile filaments may be swollen in young material but collapse and become tenuous in older specimens. Mature conceptacles are crowded with up to about 60 bisporangia which, on release, are noticeably pear-shaped, which is a further characteristic of P. zonale. PHENOLOGY: Little is known about the seasonal occurrence and reproduction of P. zonale on the shore. Specimens collected at various times of year were all bisporangial, except for the single epiphytic gametangial plant which was collected in February, and epiphytic tetrasporangial crusts collected in August and November. In laboratory culture (Chamberlain, 1982) P. zonale did not grow very well but a bispore which germinated in September formed a crust which took 160 days to produce mature spores over the winter months, while one which germinated in February took only 95 days. This suggests that warmer and/or longer daylength conditions stimulated growth and reproduction. ForM RANGE: Insufficient material has been collected to permit any observations on form range on the shore, although conceptacles grown in culture tended to be larger than those collected from the shore. In culture (Chamberlain, 1982), trichocytes were produced only between May and September and were absent during the winter. DISTRIBUTION: British Isles: P. zonale is known only from Hampshire, Dorset, Devon and Anglesey at present, but is probably widely distributed. World: Norway, Denmark, France, Italy, southern Australia (W. J. Woelkerling—pers. comm. ). Hasirat: P. zonale is predominantly an epilithic or epizoic species occurring on glass, pebbles, and mollusc shells. It occurs as an epiphyte fairly often on Chondrus crispus and has been recorded once on Halopitys incurvus. In Australia it is a sea-grass epiphyte. Lire History: The bisporangial plant is self-perpetuating (Chamberlain, 1982); the bispores are presumably diploid. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 445 Table 28 Dimensions of vegetative structures (in wm unless stated). Adey & Adey (1973) Hamel & (Fosliella Crouan British Lemoine tenuis pro type Pneophyllum zonale Isles (1953) parte) specimen crust diameter up to 10+ mm 2-10 mm crust depth up to 130 up to 70 crust cell length (10) 14 (17) 9-14 (20) 8-5-18-5 12-18 surface width (5) 9-5 (13-5) 7-10 (12) 8-16 3-6-10 crust cell height enl2 epithallial cell length (3-5) 5 (6-5) 4-5-6:5 5:5 surface view width (4) 6-5 (9-5) 4-5-6:5 5:5 epithallial cell height c. 6 hypothallial cell width VS height perithallial width (7) 10-5 (14-5) 10-14-5 cell height (10) 15 (28-5) 13-19 VS number up to 8 up to 9 trichocyte type branch cell intercalary branch cell and and intrafilament intrafilament trichocyte length 20-30 2 surface view width 11-14 12 RIDGWAY COLOUR CODE: dry crust — rocellin purple, pl. XXX VIII 12. Pneophyllum sargassi (Foslie) Y. Chamberlain, comb. nov. Basionym: Melobesia marginata forma sargassi [‘Sargassii’| Foslie in K. nor. Vidensk. Selsk. Aarsberetn. 1903: 22 (1904). NOMENCLATURAL SYNONYMS: Lithophyllum (subgenus Carpolithon) sargassi (Foslie) Foslie in K. nor. Vidensk. Selsk. Skr. 1906 (2); 26 (1906). Melobesia (subgenus Pliostroma) sargassi (Foslie) Foslie in K. nor. Vidensk. Selsk. Skr. 1908 (11): 6 (1908). Heteroderma (subgenus Pliostroma) sargassi (Foslie) Foslie in K. nor. Vidensk. Selsk. Skr. 1909 (2): 57 (1909). REFERENCES: Melobesia sargassi (Foslie) Foslie; Masaki & Tokida in Bull. Fac. Fish. Hokkaido Univ. 14: 4 (1963); Adey & Lebednik, Catalog of the Foslie herbarium: 37 (1967). Heteroderma sargassi f. sargassi Masaki in Mem. Fac. Fish. Hokkaido Univ. 16: 25 (1968); Adey in K. nor. Vidensk. Selsk. Skr. 1970 (1): 17 (1970); Park in Bull. natn. Fish. Univ. Busan 20: 19 (1980). Heteroderma sargassi f. parvula Masaki in Mem. Fac. Fish. Hokkaido Univ. 16: 26 (1968); Park in Bull. natn. Fish. Univ. Busan 17 (1, 2): 61 (1977), in Bull. natn. Fish. Univ. Busan 20: 20 (1980). SPECIMENS: Although no herbarium material of Pneophyllum sargassi as such was available, a search through BM specimens of the principal host Halopitys incurvus yielded a number of records: Pulteney, Plants of Dorset; Mrs Griffiths, November 1927, Sidmouth; Schiffner, Alg. Mar. 1402, July 1928, Trieste; S. Pignatti, 13 July 1966, Trieste; and many specimens from Brittany and Normandy. Hototyre: TRH Yendo, April 1903, Misaki, Japan, on Sargassum serratifolium (not seen, but see remarks below on the identity of this material). Diacnosis: Epiphytic species, almost exclusively on Halopitys, with thallus up to 13 cells deep, zonal conceptacles and simple ostiole: differing from P. rosanoffii in not having free ostiolar filaments; from P. microsporum in having zonal conceptacles; from Melobesia membranacea in having a deep thallus and single-pored sporangial conceptacles; and from Fostiella farinosa in having a deep thallus and lacking terminal trichocytes. 446 Y. M. CHAMBERLAIN Table 29 Dimensions of reproductive structures (zm unless stated). Rosenvinge Adey & Adey (1917) Hamel & (1973) Crouan British (Melobesia Lemoine (Fosliella type Pneophyllum zonale Isles trichostoma) (1953) tenuis) specimen CARPOSPORANGIAL CONCEPTACLE external diameter 210-300 surface diameter VS internal diameter 135 (1 only) 123 72-108 [130] VS height a3 83 100-200 53-78 VS roof height 16 roof type irregular no. basal cell rows 2 ostiole long long filaments filaments no. gonimoblast filaments diameter carpospore 30-40 x 20 SPERMATANGIAL CONCEPTACLE VS/surface width 45 32 [50] VS height length spermatium width spermatium TETRASPORANGIAL CONCEPTACLE external diameter 140-250 up to 300 surface diameter VS internal diameter (156) 202 (240) 154 106-135 180-200 VS height (78) 86 (99) 131 120-150 78-96 VS roof height (26) 29 (34) roof type irregular no. basal cell rows 1-2 ostiole long long filaments filaments no. tetrasporangia length tetrasporangium (49-5) 57-5 (74) 52:5 45-60 40 width tetrasporangium (26) 32:5 (41-5) 24-6 40 23 BISPORANGIAL CONCEPTACLE external diameter surface diameter (169) 189 (234) VS internal diameter (96) 193 (260) VS height (65) 92 (117) VS roof height (18) 29 (39) roof type irregular no. basal cell rows up to 3 ostiole long flattened filaments no. bisporangia up to 60+ length bisporangium (41-5) 57 (73) 85 width bisporangium (26) 29 (34) 30 FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 447 DESCRIPTION: Small, obscure, milky grey crusts up to 2 mm diameter, 200 um deep, perithallium up to 12 cells deep in crust centre and usually present at periphery, perithallial cells (in VS) irregular, 5-S—10-5 «4m wide x 7—16-5 wm high, crust cells in surface view oblong, 10-13-5 wm long X 3-5-7-5 wm wide, epithallial cells 3-5-5 wm long < 3-5-5 wm wide, trichocytes not seen; conceptacles somewhat raised, all except spermatangial conceptacles with simple ostiole and zonal roof, carposporangial conceptacles 72-130 wm internal diameter x 48-95 wm high, mature fusion cell narrow and deep, spermatangial conceptacles immersed (often deeply) beside carpogonial conceptacles, 28-59 wm wide x 19-44 um deep, ostiole prolonged into a spout, tetrasporangial conceptacles 78-122 ym internal diameter < 66-106 um high, tetrasporangia borne peripherally, up to c. 8 per conceptacle, 39-65 um long x 26-50 um wide, bisporangial conceptacles unknown. HistoricaL: Pneophyllum sargassi was described by Foslie (1904c, as Melobesia marginata f. sargassi) on material growing on Sargassum serratifolium in Japan. Masaki & Tokida (1963 as Melobesia) recorded it from Japan growing on Sargassum spp. and Laurencia intermedia. They noted that plants growing on Laurencia and the stems and leaves of Sargassum were 3-8 cells deep below the conceptacles, while those growing on Sargassum vesicles were only from one to a few cells deep. Masaki (1968 as Heteroderma) later described the plants growing on Laurencia as a distinct form (f. parvula) which differed from f. sargassi in having smaller tetrasporangial and carposporangial conceptacles. The taxon described by Masaki & Tokida (1963) is well defined, but the identity of the type specimen (TRH) is in some doubt; Adey (1970), for example, comments ‘Lectotype: in the original description, only Japan is given for a locality. However, this is the only specimen in the collection’, unfortunately he does not give data for the specimen. Adey & Lebednik (1967), on the other hand, catalogue only one specimen of the entity as ‘Yendo, Japan, Misaki, 4. 1903’. Masaki & Tokida (1963) received material from what was believed to be the type specimen in Trondheim which proved to be Fosliella farinosa; they gave no collection data but it would presumably have been the Yendo specimen. CRUST APPEARANCE: This species forms lightly calcified, deep (up to 200 wm) crusts which usually grow epiphytically on Halopitys incurvus (Fig. 84A) together with P. rosanoffii and M. membranacea. When fresh, the crusts are a milky grey colour which, despite their considerable depth, have an unsubstantial appearance; the conceptacles are small, rounded, and dark in the centre. In dried material the vegetative crust becomes invisible against the dark background of the host, and only dark-centred conceptacles can be seen. It is difficult to measure the crust diameter but it probably does not exceed 2 mm. Under the S.E.M. it was not possible to find the crust edge to describe cell shape, since the whole area of the crust was occupied by conspicuous, rounded conceptacles (Figs 84A, B); in air dried material (Figs 84B, D) the roof of the conceptacle is occupied by more or less isodiametric cells with calcified rims and large, central, epithallial concavities, the cells may be partially covered with a smooth coating and the ostiole is small and simple. In critical point dried material (Figs 84A, C), the central area of the roof with the isodiametric cells has a thick covering, while the original crust cells form a distinctive ring at the conceptacle base; the ostiole is closed by a thick, mucilage plug. The spermatangial conceptacle (Fig. 84A) is situated in the side of the carposporangial conceptacle. Material from Japan kindly sent by Dr Masaki has much larger crusts (up to 30 mm diameter), and under the S.E.M. the air dried conceptacles have more conspicuous central cells, although the roof appears to be zonal as in British material. VEGETATIVE ANATOMY: P. sargassi has a deep thallus which rarely has a thin border so that it is difficult to see crust cells in surface view, but when they are visible they are elongated and have rounded epithallial cells. The perithallium usually develops immediately behind the first meristem (Fig. 88A). It is composed of cells which vary from isodiametric to vertically elongated which tend, at least in the deeper crust, to be irregularly arranged rather than appearing to fall in horizontal rows. The perithallial filaments may be up to 12 cells deep; the uppermost (second Y. M. CHAMBERLAIN Fig.84 Scanning electron micrographs of Pneophyllum sargassi. A, Gametangial crust critical point dried showing spermatangial (1) and carposporangial (2) conceptacles and rim of original thallus surface round conceptacle base (3). Scale = 100 wm. B, Air-dried crust growing adjacent to Melobesia membranacea (arrow). Scale = 50 wm. C, Conceptacle roof from A with mucilage plug in ostiole. Scale = 25 um. D, Conceptacle roof from B showing isodiametric shape of roof cells. Scale = 25 wm. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 449 meristem) cell may be squat or tall and tapering depending upon how recently it has divided. The epithallial cell is flattened or triangular in vertical section and occasionally two to three-celled epithallial filaments develop — the only British species of Pneophyllum in which this occurs. Trichocytes have not been found in British material of P. sargassi, and Masaki (1968) did not find any in Japanese material. The cell dimensions and general shape and arrangement agree well with Masaki’s (1968) description. GAMETANGIAL PLANTS: English plants are monoecious. Spermatangial conceptacles (Figs 84A; 85A, F; 86C; 87) are immersed in the thallus beside the carpogonial conceptacles, the ostiole is prolonged into a spout, and often the conceptacles are so deeply buried within the perithallium that the spout traverses a considerable depth of crust before emerging on the surface. The spermatangial initials cut off elongate spermatangia (Fig. 87), which become more or less oval when released and are relatively large, measuring up to 6-5 wm long. The material described by Masaki & Tokida (1963) appears from their illustrations to be dioecious, as were the crusts on the slide loaned by Dr Masaki. The Japanese conceptacles are larger and relatively wider than the English ones taking into account the inclusion of the spout in the dimension given by Masaki & Tokida (1963) for conceptacle height. Stages in carpogonial conceptacle development observed in English material correspond closely with those described and illustrated by Masaki & Tokida (1963). When a conceptacle is initiated (Fig. 86A), the perithallial cells beside the fertile disc swell and elongate, while those above the disc disintegrate. The procarps (Figs 85B; 86B, C) are borne on a conceptacle base up to four cells deep; Figures 86E-H show three sections through the parts of the fertile disc as shown, which indicate that the central procarps (Fig. 86H) bear single carpogonia, while the peripheral structures (Figs 86F, G) are simpler and may lack fertile cells. Masaki & Tokida (1963) recorded one or two carpogonia per procarp, but their illustrations show only single ones. As the conceptacle develops, very pronounced growth occurs round the ostiole (Figs 85B; 86B), which becomes deeply invaginated, while the old thallus surface is pushed back from the conceptacle centre allowing the new roof structure to emerge. CARPOSPORANGIAL CONCEPTACLES: The carposporangial conceptacles (Figs 85E; 86D) are roun- ded and somewhat prominent. The centre of the roof emerges entirely through the thallus surface, the remains of which form a ring round the conceptacle base (Fig. 84A). The ostiolar area, which appeared very invaginated in the carpogonial conceptacle, finally rises to the surface, so that the mature ostiole is simple and often conspicuously plugged with mucilage (Figs 84C; 86D). The conceptacle roof in VS is composed of three rather regular cell layers (Fig. 86D):—the epithallium, a layer of tall cells, and an internal layer of flattened cells. The fusion cell is rather narrow and deep (Figs 85D; 86D), and often appears to be in open contact with the subtending cells (Fig. 85C); the carposporangial filaments are developed peripherally. In mature conceptacles (Fig. 85E) the fat fusion cell is often situated high in the conceptacle chamber. TETRASPORANGIAL PLANTS: Mature tetrasporangial conceptacles (Figs 88B; 89B) are similar to carposporangial conceptacles in appearance, but seem to develop much more simply without the deeply invaginate stages seen in carpogonial conceptacles. The tetrasporangia are borne peripherally on stalk cells; the young ones are deeply invaginate (Figs 88A; 89A) with the four nuclei often being clearly visible, and the tetrasporangium apparently goes through an unstruc- tured later phase (Masaki & Tokida, 1963, Pl. IX, Fig. 6) before finally maturing and becoming farctate. The ostiole (Fig. 89A) is simple. BISPORANGIAL PLANTS: Unknown. PHENOLOGY: Crusts are found throughout the year growing on intertidal plants of Halopitys incurvus, but P. sargassi is most abundant and in best condition between about September and December; at this time the crusts are thinner because they are growing more rapidly and the sporangia are clear and healthy looking. In very cold periods and in summer the crusts are ‘wn 07 = 9]B9¢ “9[9e}d90U09 [eISURJeULIOdS Jo VOTES [BIIOA ‘J “WIN CZ = BRIS ‘JOquIeYO o[oR}da0U009 UT YSIY [[90 UOIsNy MOYS 0} 919e}d99uU09 [eIsuLIOdsOdIed JO UOTIIES [ROI A “q ‘WI! EZ = [BIG *]]99 UOISNy MOYs 0} a[9e}da0uU09 [eIsUeIOdsOdIe9 AT Ie JO UOTIDS [eonI9A ‘q ‘Wi 07 = 9[k9g *(Z) [[99 SUIPUS}QNs puk [][99 UOIsNy U99Mj}9qQ UOT}IOUUOD pur (T) JUOWETY Jse[qouUOs SUNOA MoOYs 0} 9Joe}da90u09 je1suviodsodies Suidojaasp Jo UoTdas [BONIOA “DO ‘WIN OZ = B]R9S *(Z) sdreso1d pue ({) syuswWeTY Joo1 Suidojaasp Zuimoys 9oe}da90u09 [etuosodie9d Jo UOT}DES [BOTTID A “g “WI YC = 9]BIG “S[[90 JOOI 9[9e}d99U09 SIIJOWIPOs! 9Y} SUIMOYs Sofde}d90U09 (MOLI) [eIsUeJeULIOdS pue jersuviodsodies Jo MOIA ddRJING ‘Y ‘Issvsuvs inj] Xydoaug JO [el19}eU Poyloyedep Ul sofoejdoouos [eIsueIOdsodies pue jeisurjowReH sg °sIq si rth, ool oa ~ ee .. gq z Ss m a a = < em Ss) = > 451 FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES “soudSOyoL] pue eIuoSod1v9 a[SuIs YIIM Sdred01g ‘HY “S][99 a]119}s pue ‘souXSoyo11) psonpo ‘eruosodieo yim sdiesoig ‘5 ‘sdiesoid [eroydied poonpoy ‘J ‘pouonses o10m YY pur ‘dD ‘4 sdresoid oy} yorym Wo UOTIsod SuNeoIpUT |[99 UOIsNy Jo wesseiq ‘q ‘9}9e1d990U09 9y} UT YsIY BuIA] (MOLL) [[99 UOIsNy doap oy} SuLMOYs 2]9e}da0u09 [eIsURIOdsodie9 oINjyeY| ‘C “a[ONSO poyeUulseaut Ajdoap yyim sopoeidaouos (Mose) [ersuejeusods pur feruosodreD ‘D *(MorIe) s]]90 Joos SuIdojaaap YIM aJoe}da0U00 [eTUOsod sed one ‘g *(p) sq[o0 [eTTeyILIEd UsT[oms oY} pue (¢) ddeJINS snyTeY) 94) Jo UONeIdoJUISIP By} SUIMOYs (7) aJe}da0U0d [eTUOsodIe9 sunok pur (T) gjoejdaouos [eisueyeuntods 3uno x ‘y ‘issv8uvs unykydoaug JO Sojaedaou0d [eIsueI1Odsodied pure Jetuosodsed Jo sUOTIOIS [PINIOA 98 “BI witos & @ & 4 IH pe 1) ses a == 4 as & ee enue: C3 Se FD HY — ~ -s S 2 aS Ce Gras CEES Fe, & 452 Y. M. CHAMBERLAIN Fig. 87 Vertical section of spermatangial con- ceptacle of Pneophyllum sargassi. thicker, more heavily calcified, and the spores are often degenerate or absent. Differences are also noted from year to year and P. sargassi was, for example, more abundant in 1978 than in 4979: ForM RANGE: The British Isles material compares well with Japanese material as described by Masaki & Tokida (1963) and material kindly sent by Dr Masaki; of the two forms described by Masaki (1968) the conceptacle dimensions agree most closely with f. parvula (Tab. 31); however, until further European material is available for comparison it seems preferable not to distinguish forms in the present sample. DISTRIBUTION: British Isles: Hampshire, Dorset, Devon. World: France, Italy, Japan. It seems probable that P. sargassi is more widely distributed than is indicated above, but it is not an easy species to find and may have been missed elsewhere. The records from France and Italy were acquired by examining specimens of Halopitys incurvus in the British Museum; it was found on many plants from Normandy and Brittany, one plant from Biarritz, and two plants from Trieste. Hasirat: In England, P. sargassi is almost entirely restricted to the host Halopitys incurvus. On only one occasion it was found on Palmaria palmata, at a time (October 1978) when it was generally much more abundant than usual. When growing on Halopitys, Pneophyllum sargassi is always accompanied by Melobesia membranacea, which it frequently overgrows (Fig. 84B). Pneophyllum rosanoffii also commonly occurs on Halopitys together with the other two species. In Japan P. sargassi grows mainly on species of Sargassum and Laurencia. Although Sargassum muticum has become extremely abundant at Bembridge in recent years, Pneophyl- lum sargassi has never been found on this plant. TAXONOMIC NOTE: The taxon is here classified as a Pneophyllum with some doubt since it shows some features not entirely typical of that genus but which seem to bear some relationship to Fosliella. At present, almost all known Fosliella species lack a vegetative perithallium and bear terminal hypothallial trichocytes, while no trichocytes have been observed in Pneophyllum sargassi and a deep perithallium is present. However, the single recorded exception in the genus Fosliella is a drawing of Japanese material of F. farinosa by Masaki & Tokida (1960, Pl. VI, Fig. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 453 te = ‘7 f Fig. 88 Vertical sections of tetrasporangial conceptacles of Pneophyllum sargassi. A, Young conceptacle showing deeply invaginated tetrasporangium and deep crust periphery. B, Vertical section of mature tetrasporangial conceptacle. Scale = 50 wm. 50um Fig. 89 Vertical sections of tetrasporangial conceptacles of Pneophyllum sargassi. A, Immature tetras- porangia. B, Mature tetrasporangia. 7, as Melobesia) which shows a tetrasporangial conceptacle which strongly resembles those of Pneophyllum sargassi and which was probably also growing on Sargassum. In addition, the elongated crust cells and rounded epithallial cells seen in present and Japanese material (Masaki Table 30 Dimensions of vegetative structures (um). Masaki (1968) Pneophyllum sargassi British Isles (as Heteroderma) crust diameter up toc. 2mm 5-10 (30) crust depth up to 200 97-126 crust cell length (10) 11-5 (13-5) 12-20 surface width (3-5) 6 (7:5) 5-8 crust cell height 6-7 5S-7 epithallial cell length (3) 4 (5-5) 12-20 (?) surface view width (3) 4-5 (5:5) 5-8 epithallial cell height 24 5-7 hypothallial cell width 5-7 (10) VS height 9-12 (17) perithallial width (5-5) 8 (10-5) 7-11 cell height (7) 11 (16-5) 9-21 VS number up to 12 up to 12 trichocyte type trichocyte length surface view width 454 Y. M. CHAMBERLAIN Table 31 Dimensions of reproductive structures (um). Masaki (1968) Masaki (1968) Pneophyllum sargassi British Isles f. sargassi f. parvula CARPOSPORANGIAL CONCEPTACLE external diameter (42) 84-134 (172) 75-100 surface diameter (83) 101 (130) VS internal diameter (72) 98 (130) VS height (48) 72 (95) 50-80 50-75 VS roof height (15) 23 (33) 12-21 roof type zonal zonal zonal no. basal cell rows 1+4 ostiole simple simple simple no. gonimoblast filaments c. 8 diameter carpospore SPERMATANGIAL CONCEPTACLE VS/surface width (28) 43 (59) 60-92 VS height (19) 32 (44) 25-84 length spermatium (3-5) 4-5 (6-5) 4 width spermatium (1-5) 2 (2-5) 1-7 TETRASPORANGIAL CONCEPTACLE external diameter 140-210 75-100 surface diameter (110) 132 (160) VS internal diameter (78) 100 (122) VS height (66) 82 (106) 50-105 65-85 VS roof height (14) 21 (26) roof type zonal zonal zonal no. basal cell rows 1+4 ostiole simple simple simple no. tetrasporangia up toc. 8 length tetrasporangium (39) 54 (65) 46-63 40-50 width tetrasporangium (26) 39 (50) 30-42 20-30 & Tokida, 1963, Pl. IX, Fig. 4) of P. sargassi are similar to those seen in Fosliella; furthermore, judging by the fact that material from Trondheim purporting to be Pneophyllum sargassi proved to be Fosliella (fide Masaki & Tokida, 1963), it would seem that the taxa are easy to confuse. A further character which suggests a relationship to Fosliella is the apparent absence of procarps bearing two carpogonia in English material of Pneophyllum sargassi. Most records of Fosliella indicate that this genus always has single carpogonia, while some bicarpogonial procarps are present in all Pneophyllum spp. Masaki & Tokida (1963) report bicarpogonial procarps in P. sargassi, but their drawing shows only single carpogonia; on the other hand, their illustrations of Fosliella farinosa (Masaki & Tokida, 1960) show bicarpogonial procarps. In addition present material of Pneophyllum sargassi may show connections between the fusion cell and subtending cells (Fig. 85C). These features need further investigation. The final character of interest in the present context is the spore segmentation pattern. Fosliella is distinguished from Pneophyllum in the present study by having a four-celled central element to the germination disc, as opposed to an eight-celled one. My own attempts to germinate spores from P. sargassi were unsuccessful but Chihara (1974a) and Notoya (1976a) have both germinated spores and found a Corallina-type germination pattern which is different from both the Fosliella and Pneophyllum types. Notoya (1976a) found a similar pattern in species of Neogoniolithon and Pseudolithophyllum sensu Adey, which are both mastophoroid algae, and also in Lithothamnion which is in a different subfamily. This, therefore, seems to supply added negative evidence that P. sargassi may not belong to the genus Pneophyllum. However, it seems best to refer it to Pneophyllum for the present until detailed information is available to indicate its true affinities. FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 455 Acknowledgements I would like to express my thanks to Professor E. B. G. Jones, Dr R. L. Fletcher, Dr W. F. Farnham, and Mrs L. M. Irvine for their continual guidance and support during the seven years spent on the present investigation. I am also grateful to the following colleagues for discussion and advice regarding botanical, nomenclatural, and textual matters: Professor P. S. Dixon, Dr D. J. Garbary, Dr H. W. Johansen, Mme M. Lemoine, Mr J. H. Price and Dr P. C. Silva. Dr W. J. Woelkerling has helped in many ways including making slides of herbarium specimens and suggesting the use of the generic name Pneophyllum. Ihave had an interesting correspondence with Professor E. Voigt regarding fossil species. Mr R. Ross wrote the latin descriptions for which I am most grateful. Mr A. E. W. Hawton printed the photographs and Mrs P. A. Davies and Mrs G. Perks typed the text and I thank them all for their skilful help. Further invaluable technical help was given by Mr G. Bremer regarding scanning electron microscopy, and by Mrs A. Davis, Mrs M. Funnell, Mr R. Healey, Mr J. P. Hepburn, Mr K. Purdy, and Mr S. I. Honey. Many colleagues have collected material for me and I would particularly like to record my thanks to the late Dr H. Blackler and to Dr G. T. Boalch, Mme J. Cabioch, Dr A. T. Critchley, Dr M. de Valéra, Dr P. W. G. Gray, Dr M. D. Guiry, Mr N. A. Jephson, Professor T. Masaki, Mr O. W. Morton, and Dr W. Prud’homme van Reine. I am grateful to the Directors of the following herbaria for allowing me to examine the specimens in their care: BM, C, CHE, CN, CO, DBN, E, GLAS, L, PC, and USNC. Mr P. W. James and Mr J. R. Laundon have provided the editorial expertise for which I am much indebted. I am grateful to the trustees of both the Special Fund of Bedford College, University of London, and the Botanical Research Fund for financial assistance and to the Science Research Council for Studentship number B78/101389 for the year 1978/79. Finally, but crucially, without the tolerant and encouraging help of my husband, my daughters, and my parents this work would not have been possible. References Adams, J. 1908. A synopsis of Irish algae, freshwater and marine. Proc. r. Ir. Acad. 27B: 1-60. Adey, W. H. 1970. A revision of the Foslie crustose coralline herbarium. K. nor. Vidensk. Selsk. Skr. 1970 (1): 146. Adey, W. H. & Adey, P. J. 1973. Studies on the biosystematics and ecology of the epilithic crustose Corallinaceae of the British Isles. Br. phycol. J. 8: 343-407. Adey, W. H. & Johansen, H. W. 1972. Morphology and taxonomy of Corallinaceae with special reference to Clathromorphum, Mesophyllum and Neopolyporolithon n.gen. (Rhodophyceae, Cryptonemiales). Phycologia 11: 159-180. 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An asterisk (*) denotes a figure; if this occurs after reference to a group of pages (eg 340—343*) the figure, or figures, occur within the group, not necessarily on the page with the asterisk. Alcyonidium 298 Amphiroa 298, 342 verruculosa 342 Amphiroideae 331 Bornetia 408, 410 secundiflora 304—6, 308, 361, 405, 410 Bryopsis balbisiana 306, 404* Calliblepharis ciliata 297, 410 Carpolithon 299, 316, 436, 445 Chaetolithon 315 Chaetomorpha vasta 386 Chondrus 355 crispus 336, 349, 382, 396, 399, 401, 410, 442, 444 Choreonema 315, 316, 318 thuretii 293, 338 Cladophora 387, 389 rupestris 338, 386, 390 Cladostephus spongiosus 410 Conferva vasta 385 Corallina 293, 297, 298, 312, 389, 395 membranacea 297, 298, 341, 344 officinalis 338, 394, 395 virgata 344 Corallina-type spore germination 454 Corallinaceae 293, 330, 331* Corallinoideae 331 Cryptonemiales 330, 332 Cystoseira foeniculacea 410 nodicaulis 338, 349, 401 Dermatolithon 293, 297—300, 344, 354, 359 corallinae 338 lejolisii 316, 359 litorale 324 pustulatum 300, 336, 338, 385 f.laminariae 338 sp.297, 300, 338, 359 Desmarestia aculeata 410 Dictyota dichotoma 338 Dumontia-type spore germination 319 Epilithon 299, 308, 344 membranaceum 308, 339, 344 van heurckii 387 Euheteroderma 299 Eulithothamnion 299, 315 Eumelobesia 299, 310, 312, 315-17, 340, 341, 436 Fosliella 293, 295, 297-301, 312, 319-335, 340-343* , 354, 387, 452, 454 chamaedoris 354 cruciata 341, 342 farinosa 297, 300, 306, 319*, 322*-324, 327, 328, 334-336*, 338-342, 343-351*, 357, 367, 382, 383, 385, 389, 395, 413, 429, 437, 439, 445, 447, 452, 454 f.callithamnioides 333, 341, 342, 345*, 351-352 var.chalicodictya 341, 342 var.solmsiana 351 inexspectata 293 lejolisii 339, 359, 360 limitata 339, 376 minutula 339, 385 f.lacunosa 394, 395 paschalis 323, 341, 342 tenuis 339, 413, 437, 439, 442, 443* valida 429, 430* 462 Y. M. CHAMBERLAIN Fosliella — cont. zonalis 437 f.myriocarpa 411, 413 Fucus serratus 338, 351, 382, 383, 401 vesiculosus 376, 377 Furcellaria 389 lumbricalis 338, 386, 390, 395, 396, 399, 401, 410 Galaxaura 297 Gastroclonium ovatum 338, 399, 410 Gelidium 389 pusillum 352 Gigartina stellata 338 Gracilaria verrucosa 338, 410 Guerinea 301, 310, 354, 395 callithamnioides 352, 392 Haliptilon 293 Halimeda 297 Halopitys 355, 372 incurvus 305, 336, 370, 373, 376, 401, 416, 444, 445, 447, 449, 452 Hapalidium 298-302, 353, 354, 387, 413 callithamnioides 302, 310, 352, 388*, 392, 394*, 395, 437 coccineum 302, 388*, 437 confervicolum 385-387 confervoides 340, 385, 387, 437, 439 phyllactidium 300, 385, 387, 388* , 437, 439 roseolum 300, 306, 308, 402, 404* roseum 306, 308, 437 zonale 302, 303*, 388*, 435, 437-439* Heteroderma 299, 301, 310, 312, 315-318, 320, 340, 353, 354, 413, 436, 437 inaequilaterata 385, 395 lejolisii 359-361, 366 minutula 385 nicholsii 312 sargassi 445, 447 f.parvula 445, 447, 452 subtilissima 312 zonale (is) 413, 437 Heterodermia 310 Hildenbrandia 301 Jania 293, 395 rubens 338 Laminaria digitata 382, 385, 401 hyperborea 410 ochroleuca 410 saccharina 382, 385 sp. 338, 351, 413 Laurencia 452 intermedia 447 pinnatifida 338, 401, 402, 410 Lepidomorphum 299, 316 Leptophytum 293 Lithocystis 299-301, 354 allmanni 339, 385, 387 Lithophylloideae 331 Lithophyllum 293, 298, 299, 354 macrocarpum 385 sargassi 445 zonale 436 Lithoporella 318 Lithothamnion (um) 293, 298, 299, 354, 454 membranaceum 340, 343 Littorina 439 Lomentaria clavellosa 301 Mastophoroideae 293, 328, 330-332* Melobesia 293, 297-319, 344, 354, 387 callithamnioides 315-317, 323, 339, 342, 351, 352, 392 chamaedoris 312 coccinea 315 confervicola 339, 385 confervoides 315, 339, 340 corticiforme (is) 304, 306, 307* farinosa sensu Kiitz. 305, 315, 343, 367, 369, 370 farinosa 298, 300, 304—317* , 327, 328, 339, 340, 343, 344, 413 f.macrocarpa 342 f.solmsiana 351, 413 var. borealis 343 var.callithamnioides 351 fosliei 385, 387, 389 granulata 343, 344 inaequilaterata 316, 385, 395 lejolisii 304—308* , 314—317*, 328, 339, 340, 359-361, 366, 367, 369*, 370, 373, 402, 405, 413 f.limitata 317, 361, 376, 377 limitata 323, 376, 377 marginata f.sargassi 445, 447 membranacea 297—312*, 315, 319, 336, 338, 359, 376, 389, 390*, 445, 447, 452 membranacea sensu Aresch. 360 membranacea sensu P. Crouan & H. Crouan 343 membranacea sensu Lloyd 360 microspora 395, 397* minuta 343, 344 minutula 317, 339, 340, 387, 389, 392, 394*, 395 f.lacunosa 317, 339, 387, 392, 394, 395, 413 f.typica 385, 387 myriocarpa 302, 316, 339, 410, 412*, 413, 419, 421, 437, 439 orbiculata 298 paschalis 342 pruinosa 356, 359 pustulata 298, 306, 383 yer FOSLIELLA AND PNEOPHYLLUM IN THE BRITISH ISLES 463 Melobesia — cont. rosea 295, 305, 306, 308, 309*, 402, 404*, 405 roseola 308 sargassi 445, 447 solmsiana 351, 352 trichostoma 437, 439, 442, 443 verrucata 298 zonalis 339, 340, 343, 387, 410, 411, 413, 419, 421, 429, 436, 437, 439 f.myriocarpa 410, 413 Melobesioideae 323, 331 Mesophyllum lichenoides 338 Metagoniolithoideae 331 Naccaria-type spore germination 319 Neogoniolithon 342, 354, 454 Palmaria 355, 372, 376, 383 palmata 306, 338, 351, 370, 376, 382, 385, 399, 410, 452 Paracentrotus lividus 352 Patella 439 Phyllactidium 299, 300, 386 confervicola (um) 300, 385, 386* , 390* Phyllophora crispa 338, 399 pseudoceranoides 338, 399 Phymatolithon 293 Plectoderma 300, 308 majus 308 minus 308 Pliostroma 299, 410, 413, 436, 437, 445 Plocamium 387, 389 coccineum 390 Pneophyllum 293, 298-301, 304, 308, 310, 312, 318-334, 352-—355* , 378, 405, 412, 424, 449, 452, 454 concollum 304, 305, 308, 315, 324, 327, 328, 334—336*, 338, 343, 359, 361, 367, 370, 376, 383, 395, 402—410*, 412, 419 confervicolum 295, 312, 333, 338-340, 342, 354, 385-392*, 395, 412, 413, 439 f.minutulum 302, 312, 333, 345*, 352, 392-395* , 437 fragile 294, 295, 297, 300, 301, 352-—354*, 356-—359* , 361, 367, 373 lejolisii 304, 305, 315, 324, 328, 335, 336, 339, 340, 354, 355, 359-367*, 370, 376, 377, 385, 395, 402 limitatum 306, 323, 324, 328, 333, 335, 336, 338-340, 351, 360, 367, 370, 376-385*, 398, 413, 415, 437 lobescens 319—321* , 324, 328, 335, 336, 338, 412, 419-427*, 428, 429, 437, 439 microsporum 324, 336, 338, 382, 395—402*, 405, 445 myriocarpum 302, 323—325*, 328, 336, 338, 339, 378, 402, 405, 410-419*, 420, 421, 429, 435, 437, 439 plurivalidum 322* , 324, 328, 335, 336, 338, 354, 395, 412, 419, 420, 427—435*, 437 pruinosam 359 rosanoffii 295, 304, 305, 315, 322* , 324, 326*, 328, 333-—336* , 338, 343, 359-361, 363, 367-376* , 381, 402, 403, 410, 437, 445, 447, 452 sargassi 324, 328, 335, 338, 367, 370, 376, 395, 402, 403, 445-454* sp. 344 zonale 302, 324, 325*, 328, 329*, 335, 336, 338-340, 367, 370, 376, 381, 412, 413, 420, 428, 435-445* Polyides rotundus 399 Polysiphonia-type life history 328, 330*, 336, 419, 427 Polysiphonia elongata 338, 401, 410 Porolithon 342, 354 Posidonia 366 Pseudolithophyllum sensu Adey 354, 454 Pseudolithophyllum sensu Lemoine 354 Rhodochorton 310, 395 Rhodymenia palmata 306, 343, 351 Rhyncococcus coronopifolius 300, 357 Rissoa 414* Rissoids 338, 416, 439 Sargassum linifolium 340, 343, 344 muticum 295, 338, 383, 401, 452 serratifolium 445, 447 sp. 447, 452 Schmitziella endophloea 293 Sphaerococcus 297, 300 coronopifolius 352, 353* , 356-359 Spongites 298, 354 Udotea 297 Ulva lactuca 339 Zostera 304, 305, 339, 349, 355, 359-361, 363, 3672310; 312,315,510; 56s, 901, 004 marina 336, 361, 366, 367, 370, 373, 392 - = i oaed oe * = o> a a British Museum (Natural History) Seaweeds of the British Isles The synthesis of many years’ research carried out by the British Museum (Natural history) and the British Phycological Society, this is the second in a series of books that will be published under this title covering all the British and the majority of northern Atlantic seaweeds. Volume 1 Rhodophyta Part 2A Cryptonemiales (sensu stricto), Palmariales, Rhodymeniales. : Linda M. Irvine Following the style and format laid down in Part 1, this book deals with the species attributed to the orders Cryptonemiales, Palmariales and Rhodymeniales. Each species description incorporates notes on ecology and distribution and is supported by one or more line illustrations. Keys to aid identification are also included. When complete, this title will be the standard work of reference in its field; it will provide students and researchers with a digest of the most up-to-date and compre- hensive information available on the marine algae of Great Britain and Ireland which will be indispensable throughout the north Atlantic region and beyond. ISBN 0 565 00871 4 128 pagesillustrated £13.00. Titles to be published in Volume 11 The algae of Lightfoot’s Flora scotica. By Peter S. Dixon. A taxonomic study of the lichen genus Micarea in Europe. By B. J. Coppins. The hepatics of Sierra Leone and Ghana. By E. W. Jones and A. J. Harrington. Studies in the Corallinaceae with special reference to Fosliella and Pneophyllum in the British Isles. By Y. M. Chamberlain. Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, Dorchester a oe we BOUND 2.0 JUL 1988 aes Jy y Oy % pes * > Rete RNA tn a pean sane ‘ “a is Bitter es Sing Pere tut stay) % y y) i Breit KAS nit bite ae Aut, < . Heit iat ease gtates ; i seen (tg a * s Mo t Vy i Ke ve Maes ae ey ae Mitte ai aed Uy A bite Sooke a oe, Aan pee ii mi 7 ‘ Hi) ater ae ee — oe Mriehe) if fs fay Pics sits # brs (nat nai a eae oe By i, Pies feel yy iH cz ain ae ts 4 ene Spee ve ; trata jeay tet Sly va) rah reste, Ss Pi eae