ISSN 0968-0446 Bulletin of The Natural History Museum THE NATURAL HISTORY MUSEUM PRESENTED GENERAL LIBRARY Botany Series THE NATURAL HISTORY MUSEUM VOLUME 24 NUMBER 2 24 NOVEMBER 1994 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History)), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Dr S. Blackmore Editor of Bulletin: Mrs M.J. West 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 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. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 716 Andover Hampshire SP10 1YG Telephone: (0264) 334748 Fax: (0264) 334058 Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6 months for the UK, and 9 months for the rest of the world. World List abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.) © The Natural History Museum, 1994 Botany Series ISSN 0968-0446 Vol. 24, No. 2, pp. 101-180 The Natural History Museum Cromwell Road London SW7 5BD Issued 24 November 1994 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain at The Alden Press, Oxford Bull. not. Hist. Mus. Land. (Bot.) 24(2): 101-114 Issued 24 November 1994 Observations on the benthic marine algal flora of South Georgia: a floristic and ecological analysis DAVID M. JOHN Department of 'Botany ', The Natural History Museum, Cromwell Road, London SW75BD, UK PHILIP J.A. PUGH British Antarctic Survey, Natural Environmental Research Council, High Cross, Madingley Road, Cambridge CBS OET, UK IAN TITTLEY Department of Botany, The Natural History Museum, Cromwell Road, London SW75BD, UK CONTENTS Introduction Materials and methods Results THE'NATURAt HISTORY MUSEUM 101 102 103 Distribution of shore algae . . 103 Systematic list j 4 ut'li Iob4 103 Discussion Shore ecology ••PRESENTED • Ill Ill Marine algal flora GENE R At ttB RARY 112 References . . 113 SYNOPSIS. The pattern of littoral zonation of benthic algae on rocky and boulder shores in Husvik Harbour, on the north-east coast of South Georgia, was investigated during the austral summer of 1990/91. Distribution patterns are similar to those on other shores in the sub-Antarctic region of the Southern Ocean, except for the absence in the littoral fringe of a 'Hildenbrandia' or 'Hildenbrandia-Bostrychia association'. The supralittoral fringe is devoid of macroalgae and gives way to a eulittoral zone consisting of a series of belts. An uppermost belt dominated by Porphyra followed by ones dominated or co-dominated by Nothogenia fastigiata, Iridaea cordata, and Adenocystis utricularia. The lowermost belt is either dominated by Palmaria georgica or co-dominated by this red alga and Schizoseris condensata. The upper vertical limit of crustose coralline algae and the kelps Macrocystis pyrifera and Durvillaea antarctica define the sublittoral fringe; the latter is only present on wave-exposed shores. The 103 species of algae known from South Georgia are critically evaluated, four are endemics and 12 are known only from South Georgia and Tierra del Fuego. The biogeographical affinities of the algal flora of South Georgia lies with other sub-Antarctic islands and the southern tip of mainland America. INTRODUCTION South Georgia is an isolated island in the Southern Ocean that lies between latitudes 53°56'-54°55'S and longitudes 34°45'_38°15'W. It is roughly crescent-shaped (Fig. 1) and the second largest of the circum- Antarctic islands. Its steep rocky coast, dissected by deep fjords and bays, provides suitable though very hostile habitats for benthic organisms. The algal flora of South Georgia remains little-known since biological exploration has been limited to a few bays on the more sheltered north-east coast. The first publications specifically on its marine algae are those of Reinsch (1888, 1890) and were based on material collected by the 1882-1883 German International Polar Year Expedition. The next major period of collecting on the island was during the 1901-1903 Swedish South Polar Expedition, but most of the material was lost when the expedition's ship 'Antarctica' foundered. Fortu- nately, the journals belonging to the expedition's botanist, Carl Skottsberg, survived and he re-visited the island in 1909 to make further observations and new collections. Informa- tion contained in his journals was used to write an ecological account of South Georgian algae which was subsequently published (Skottsberg, 1941). Some of the numerous expedi- tions en route to higher Antarctic latitudes have found safe anchorage at South Georgia and used the opportunity to collect algae from its shores. Accounts of the ecology of Antarctic and sub-Antarctic ©The Natural History Museum, 1994 102 D.M. JOHN, P.J.A. PUGH AND I. TITTLEY marine algae still remain almost wholly descriptive. Early information on sublittoral assemblages was obtained indi- rectly by trawling, dredging and the collection of drift plants. The advent of SCUBA diving has enabled direct observations and the hand collection of undamaged specimens of sublit- toral algae at several localities in the Southern Ocean (Neus- chul, 1968; Delepine et al., 1966; Lamb & Zimmermann, 1977). Unfortunately SCUBA diving to investigate algal ecology has yet to be carried out on South Georgia. Observa- tions on its sublittoral algal ecology date back to the early years of this century and are restricted to dredged collections (see Skottsberg, 1941). Although incomplete and unevenly distributed, there is sufficient information to enable some preliminary conclusions to be reached on the distribution patterns of littoral algae in the Southern Ocean (see Stephen- son & Stephenson, 1972; Luning, 1990; Delepine et al., 1966; Zaneveld, 1964). Few experimental studies and the absence of reliable quantitative data have meant that information is lacking on ecological interactions involving algae. The present study examines the distribution and abun- dance of shore algae on the north-east coast of South Georgia, critically reviewing all published and unpublished records, and makes some observations on the biogeographi- cal affinities of its algal flora. MATERIALS AND METHODS Five transects were studied at Husvik Harbour (36°40'W, 54°11'S), Stromness Bay on the north-east coast of South Georgia (Fig. 1). This relatively wave-sheltered bay has small rocky outcrops of volcanoclastic sandstone-shale turbidite (Macdonald et al., 1987) separated by coarse sand or gravel beaches; all its shores are free from ice-scouring. Three of the transects were on rocky shores: A was north of the whaling station and below the catcher boat 'Karrakatta' on its repair slipway; B and C were on the southern side of the Harbour and on the west (B) and east side (C) of Kanin Point (Fig. 1). The other two transects were on boulder shores at Brain Island Point (D) on the northern side of Husvik Harbour and between the Husvik 'villa' and Kanin Point (E) on the southern shore. The benthic algal communities on shores in Husvik Har- bour were monitored at intervals from October 1990 to mid-January 1991. It was considered that by the end of the period the cover abundance of shore algae was close to its seasonal maximum. The distribution and cover of macroalgae were determined between 25 January and 15 February 1991 along the five shore transects. Each transect was surveyed from low water to the top of the supralittoral fringe using an Abney level mounted on a wooden block to give a base to horizontal line-of-sight distances of 20 cm. Slope distance between stations along a transect was measured with a tape. From these data both horizontal and vertical distances between sampling stations were calculated and shore profiles drawn (see Fig. 2). Vertical heights were corrected to Chart Datum (C.D.) using Admiralty Tide Tables (Admiralty, 1990, 1991). The tidal range is c. 1.2 m and the cumulative measurement error was +0.1 m at high water mark. Fig. 1 Map of Husvik Harbour on the north-east coast of South Georgia showing the position of the five investigated transects (A-E). Insert of South Georgia shows an arrow indicating the position of Husvik Harbour. W.S., whaling station. BENTHIC MARINE ALGAL FLORA OF S. GEORGIA 103 Horizontal distance on shore (m) Fig. 2 Shore profiles of the five transects. The algal assemblages were photographed in situ and all material collected was preserved in 5% seawater-formalin. The relative cover abundance of the dominant macroalgae at each sample station was calculated to within ±3% and expressed as a percentage of total rock covered. All algal specimens are deposited in the Herbarium at The Natural History Museum, London. RESULTS Distribution of shore algae Despite the small tidal amplitude (c. 1.2 m) marine influences extend vertically by several metres where the coast is exposed to heavy swells and severe wave action. Only in rocky bays and inlets is the sea sufficiently calm to enable a detailed analysis of the distribution of shore organisms. In Husvik Harbour, the upper limit of the littoral zone is about 1.5-1.7 m above C.D. on rocky shores (Figs 3-5) and 0.75-1.2 m on the boulder shores (Figs 6-7), based on the vertical limit of the membranaceous red alga Porphyra. Total algal cover shows considerable variation (Fig. 8) in response to several factors well-known to influence algal abundance and distribu- tion, i.e. shore topography (see Fig. 2), shore aspect, nature of the substratum, and type and degree of exposure to water motion. Rocky shore transect A is partially shaded, faces due south and is positioned along a surge channel. Transects B and C run across rocky shores that are in close proximity to each other (see Fig. 1), the former more wave-sheltered due to the presence of a rock spur. The other two transects (D, E) run over gently sloping boulder shores and most algae are restricted to the sides of the boulders. Porphyra accounts for the high total algal cover on the upper part of the eulittoral zone in transects B and E. The algal vegetation of the rocky shores (Figs 3-5) com- prises 12 to 13 species. The upper eulittoral was dominated by Porphyra whose cover-abundance was particularly high on gently sloping shores (B, C; see Fig. 2). Associated with Porphyra were Nothogenia fastigiata and Iridaea cordata; these latter two red algae were found commonly in protected niches. They were not present higher than l.lm above C.D. and tended to be confined to the lower part of the Porphyra belt. The lower eulittoral zone was dominated by the red alga Palmaria georgica, accompanied by lesser amounts of Por- phyra sp., Schizoseris condensata and Adenocystis utricularis; the latter grew as isolated clumps or single plants confined to sheltered niches. In the lower eulittoral zone the filamentous red alga Ceramium rubrum was frequently intermingled with clumps of Schizoseris. Small Macrocystis and encrusting calcareous red algae ('lithothamnia') were observed around Chart Datum and below. Only 7-9 algal species were found on the boulder shores D and E (Figs 6, 7), with the former unusual in having a sparse cover of Porphyra and a lower eulittoral zone dominated by colonial diatoms. The brownish layer of diatoms covered the sides of many boulders, occupying the position of the Palmaria/ Schizoseris association on rocky shores. The upper eulittoral of shore E was dominated by Porphyra, confined to the sides of boulders, and the lower eulittoral zone had a very sparse covering of a mixture of algae. Systematic list A list of the marine benthic algae from South Georgia based on a critical evaluation of published records, and an examina- tion of material collected by one of us (PJAP) in January and 104 D.M. JOHN, P.J.A. PUGH AND I. TITTLEY Enteromorpha bulbosa green Porphyra sp. Acrosiphonia pacifica Desmarestia menziesii Macrocystis pyrifera Adenocystis utricularis Porphyra spp. Nothogenia fastigiata Iridaea sp. Iridaea cordata Schizoseris condensata Palm aria georgica Fig. 3 The vertical distribution and percentage cover abundance of seaweeds on the rocky shore transect A. February 1991 and specimens in the Herbarium of The Natural History Museum, London. Nomenclatural revisions and re-examination of material has resulted in some of the species listed here published under different names. Given under each entry is the name(s) by which the species has been reported in the list area (synonyms, misdeterminations). For further information on many of the entries, see Papenfuss (1964). Qualifying notes accompany some of the entries especially where there is an element of doubt attached to the records. Chlorophyta Acrosiphonia pacifica (Montagne) J. Agardh Reported as Cladophora arcta, C. pacifica, Spongomorpha arcta and S. pacifica. Cladophora incompta (Hooker f. & Harvey) Hooker f. & Harvey BENTHIC MARINE ALGAL FLORA OF S. GEORGIA Level on shore (m) 105 filamentous diatoms Enteromorpha bulbosa green Porphyra sp. Desmarestia sp. Desmarestia menziesii Adenocystis utricularis Porphyra spp. Nothogenia fastigiata Iridaea sp. Iridaea cordata Schizoseris condensata Pa I mar ia georgica Fig. 4 The vertical distribution and percentage cover abundance of seaweeds on the rocky shore transect B. Enteromorpha bulbosa (Suhr) Montagne Reported as E. novae -hollcmdiae . All plants collected during the Swedish 1907-1909 expedition and attributed to this species by Hylmo (1919) may equally well be a form of E. intestinalis (see Ricker, 1987: 36). The principal features distinguishing E. bulbosa from E. intestinalis are the smaller dimen- sions of its cells and their thicker walls (Lamb & Zimmermann, 1977), or the absence of any branching of the thallus (Ricker, 1987). Enteromorpha gunniana J. Agardh Closely related to E. bulbosa, with a plant from Macquarie Island attributed to E. gunniana by Ricker (1987: 36) considered to fall 106 D.M. JOHN, P.J.A. PUGH AND I. TITTLEY Level on shore (m) Enteromorpha bulbosa green Porphyra sp. green filamentous alga Acrosiphonia pacifica Desmarestia menziesii Adenocystis utricularis Porphyra spp. Nothogenia fastigiata Iridaea sp. Iridaea cordata Schizoseris condensata Ceramium rubrum Palmaria georgica Fig. 5 The vertical distribution and percentage cover abundance of seaweeds on the rocky shore transect C. within the form range of E. bulbosa. Doubt attaches to the species identification of this alga from South Georgia. Entonema subcorticale Reinsch This taxon and the following are little-known epiphytes, possibly identical to Entocladia (see Ricker, 1987: 24). Entonema tenuissimum Reinsch ?Prasiola crispa (Lightfoot) Meneghini subsp. antarctica (Kuetzing) Knebel Recorded as P. antarctica. Doubtful record as no text entry and yet listed in Reinsch's index (Reinsch, 1890); this is the likely source of Hylmo's (1919) secondary citation. BENTHIC MARINE ALGAL FLORA OF S. GEORGIA 107 Level on shore (m) o -r o p bo Enteromorpha bulbosa green Porphyra sp. Adenocystis utricularis Porphyra spp. Nothogenia fastigiata Iridaea sp. Fig. 6 The vertical distribution and percentage cover abundance of seaweeds on the boulder shore transect D. filamentous diatoms Prasiola tllifbrmis Reinsch var. minuta Reinsch Status of this taxon requires investigation. Ulothrix sp. Reported as Hormiscia parasitica. According to Papenfuss (1964: 1), this record is 'probably represen- tative of one of the species of Ulothrix that has been reported from Antarctica'. Ulva lactuca Linnaeus var. macrogyna Reinsch Status of this taxon requires investigation. Phaeophyta Adenocystis utricularis (Bory de Saint- Vincent) Skottsberg Recorded as Chroa sacculiformis . Ascoseira mirabilis Skottsberg Reported as Lessonia fuscescens var. linearis. Caepidium antarcticum J. Agardh Cladothele decaisnei Hooker f . & Harvey Recorded as Stictyosiphon decaisnei. Corycus lanceolatus (Kuetzing) Skottsberg Recorded as C. prolifer. Desmarestia antarctica Moe & Silva South Georgia is its northernmost limit of distribution. Its minute gametophytic stage is endophytic on Curdiae recovitzae (see Moe & Silva, 1989), a red alga also at the limit of its range. Desmarestia ligulata (Lightfoot) Lamouroux Recorded as D. firma. Desmarestia menziesii J. Agardh Recorded as D. aculeata var. compressa, D. compressa and D. harveyana. Desmarestia pteridoides Reinsch Status of this plant remains uncertain, see remarks in Skottsberg (1907: 20). Desmarestia willii Reinsch This southern hemisphere species is closely related to D. viridis known only from the northern hemisphere. According to Ricker (1987: 126), the southern hemisphere species shows wide variation in key characters and so is less distinct than Reinsch (1888: 191) indicated when justifying its creation. Durvillaea antarctica (Chamisso) Hariot Ectocarpus constanciae Hariot Ricker (1987: 67) separates it from E. siliculosus upon its greater cell diameters, tapering of apical branches, absence of hook-shaped laterals, presence of fewer corticating rhizoids covering lower axes, helical coiling of ribbon-like plastids, and more elongate and solitary plurangia. He accepts that the two species may form part of the form range continuum of E. siliculosus. 108 D.M. JOHN, P.J.A. PUGH AND I. TITTLEY Level on shore (m) Enteromorpha bulbosa green Porphyra sp. Acrosiphonia pacifica Adenocystis uthcularis Porphyra spp. Nothogenia fastigiata Iridaea cordata Schizoseris condensata Palmaria georgica Fig. 7 The vertical distribution and percentage cover abundance of seaweeds on the boulder shore transect E. Ectocarpus exiguus Skottsberg Recorded as E. humilis. Ectocarpus siliculosus (Dillwyn) Lyngbye Recorded as E. confervoides and E. fasiculatus . Elachista meridionalis Skottsberg Geminocarpus austrogeorgiae Skottsberg Geminocarpus geminatus (Hooker f. & Harvey) Skottsberg Recorded as Ectocarpus geminatus. Halopteris funicularis (Montagne) Sauvageau Recorded in table (Skottsberg, 1941: 76) showing Antarctic distribu- tion of algae. No indication given as to source and not mentioned in Papenfuss's (1964) catalogue of Antarctic and sub-Antarctic algae. Halopteris obovata (Hooker f. & Harvey) Sauvageau Himantothallus grandifolius (A. & E.S. Gepp) Zinova Recorded as Laminaria saccharina var. angustata, Himantothallus spiralis and Phyllogigas grandifolius. Lessonia fuscescens Bory Recorded as L. flavicans. It seems very likely that this large brown alga, characteristic of many sub-Antarctic shores, grows attached on South Georgia despite Skottsberg's (1921) statement (p. 47) that 'Drifted pieces have been observed along the coast of S. Georgia where according to my impression this plant does not grow'. Macrocystis pyrifera (Linnaeus) C. Agardh Recorded as M. pyrifera var. longibullata. Melastictis desmarestiae Reinsch Status of this monotypic genus is uncertain. Myrionema densum Skottsberg Myrionema incommodum Skottsberg Adenocystis the host alga. Myrionema inconspicuum Reinsch Very similar to M. densum and M. corunnae; Skottsberg (1921) suggests M. inconspicuum and M. densum to be conspecific. Myrionema macrocarpum Skottsberg Record from a table showing Antarctic distribution of algae (Skotts- berg, 1907). No indication given as to source and Papenfuss (1964) does not mention it from South Georgia. Myrionema? paradoxum Reinsch Reinsch's description and illustration of this species and M. incon- spicuum (Reinsch, 1890) tire equivocal, see comments in Ricker (1987: 83). BENTHIC MARINE ALGAL FLORA OF S. GEORGIA 109 (B) (E) E o> I w o 1 0) 100 100 100 100 0 20 40 60 80 100 Area of rock covered by algae (%) Fig. 8 Total percentage cover abundance of seaweeds at different levels on each of the five shore transects. Petalonia fascia (O.F. Mueller) Kuntze Recorded as Phyllitis fascia and Ilea fascia. Petroderma maculiforme (Wollny) Kuckuck Phaearthron austrogeorgica (Skottsberg) Pedersen Recorded as Xanthosiphonia austrogeorgica. Pilayella littoralis (Linnaeus) Kjellman Recorded as Pylaiella opposita. Scytosiphon lomentaria (Lyngbye) Link Scytothamnus fasciculatus (Hooker f. & Harvey) Cotton Recorded as 5. australis. Stegastrum porphyrae Reinsch Status of this monotypic genus is uncertain. Syringoderma australe Levring Utriculidium durvillaei (Bory) Skottsberg Morphologically very similar to Adenocystis utricularis. Differs in the cortical cells possessing a single (not several) plastid and only plurilocular sporangia known. Some authors have suggested that they are alternate phases of a single species. Rhodophyta Ahnfeltia plicata (Hudson) Fries Anisocladella serratodentata (Skottsberg) Skottsberg Recorded as Delesseria erratodentata. Antithamnion ptilota (Hooker f. & Harvey) Gibson Ricker (1987: 240) examined the type (Callithamnion ptilota Hooker f. & Harvey, 1845) and concluded that it 'may be identical to D[asyptilon\. pellucidum, as both species have similar apical develop- ment and branching patterns'. He left unresolved the question of conspecificity since no reproductive structures were present on the type. If they should prove to be conspecific then the correct epithet for the species will be Dasyptilon ptilota. Ballia callitricha (C. Agardh) Kuetzing Bostrychia vaga Hooker f. & Harvey ?Calliblepharis ciliata (Hudson) Kuetzing Recorded as Rhodymenia ciliata var. ligulata. The identity of this plant remains uncertain. Callithamnion montagnei Hooker A new record for South Georgia and deposited in the Herbarium in the Department of Botany, The Natural History Museum, London. It was discovered as one of three red algae (Ceramium rubrum, Plocamium secundatum) entangled with or growing epiphytically on a specimen of Phycodrys quercifolia collected from Possession Bay in January 1867 by Dr R.O. Cunningham. ?Callonema olivaceum Reinsch The identity of this taxon remains uncertain (Papenfuss, 1964). ?Callophyllis cristata (C. Agardh) Kuetzing Recorded as Euthora cristata and Rhodymenia cristata. A questionable determination according to Skottsberg (1941). Callophyllis linguata Kylin Callophyllis variegata (Bory de Saint Vincent) Kuetzing Recorded as Kallymenia multiflda and Callymenia multifida. Ceramium diaphanum (Lightfoot) Roth Ceramium involutum Kuetzing Ceramium rubrum (Hudson) C. Agardh ?Choreocolax rhodymeniae Reinsch Parasitic alga on Palmaria. The identity of this species remains uncertain. 110 D.M. JOHN, P.J.A. PUGH AND I. TITTLEY Cladodonta lyallii (Hooker f. & Harvey) Skottsberg A new record for South Georgia based on three specimens in the Herbarium in the Department of Botany, The Natural History Museum, London. Material collected in January 1867 by Dr R.O. Cunningham from Possession Bay during the extra-tropical South America survey of H.M.S. Nassau. Clathromorphum obtectulum (Foslie) Adey Recorded as Lithophyllum aequabile, L. discoideum f. aequabilis and Antarcticophyllum aequabile. Colacodasya inconspicua (Reinsch) Schmitz Recorded as Polysiphonia inconspicua and Merenia inconspicua. Parasitic on Heterosiphonia. Curdiea recovitzae Hariot ex Wildemann Recorded as Kallymenia reniformis f. carnosa and Callymenia reni- formis. Delesseria salicifolia Reinsch Indistinguishable on vegetative features from D. lancifolia, but separated on form of the cystocarp and specialized sporophylls containing tetrasporangia. Delisea pulchra (Greville) Montagne Recorded as Bonnemaisonia prolifera. Falklandiella harveyi (Hooker f.) Kylin Recorded as Dasyptylon harveyi, Plumaria harveyi and Euptilota harveyi. For discussion of distinctions between this genus and others including Dasyptilon, see Moe & Silva (1979). Georgiella confluens (Reinsch) Kylin Recorded as Ptilota confluens, Euptilota confluens and Plumariopsis eatoni. This genus and Plumariopsis are endemic to the southern hemisphere and are morphologically similar. Moe & Silva (1983) tabulate a number of vegetative and reproductive features that distinguish them. Heterosiphonia berkeleyi Montagne Recorded as Merenia microcladioides and Heterosiphonia merenia. Hydrolithon discoideum (Foslie) Mendozoa & Cabioch Recorded as Lithophyllum discoideum, Pseudolithophyllum discoi- deum and Spongites discoidea. ?Hymenocladiopsis crustigena Moe Recorded as Gracilaria prolifera. Doubt concerns the exact status of the South Georgian material. Moe (1986) has investigated the type of Gracilaria prolifera and states that it is 'certainly placed incorrectly as to genus .. Although nothing precludes its assignment to Hymenocladiopsis, I hesitate to propose a new combination without seeing tetrasporangial material or material in which the presence or absence of gland cells can be determined with certainty. It is possible that the plants from the Antarctic Peninsula on which the new genera is based are conspecific with Gracilaria prolifera'' . Should this prove to be the case then the epithet 'prolifera' would have priority. Iridaea cordata (Turner) Bory de Saint- Vincent Recorded as /. micans and /. cordata f. ligulata (type locality of trivial growth form). Leister (1977) regards several earlier described species as conspecific with /. cordata. Iridaea obovata Kuetzing Recorded as Iridaea macrodonta and Rhodoglossum macrodontum. Microrhinus carnosus (Reinsch) Skottsberg Recorded as Delesseria carnosa and Chauvinia carnosa. Mesophyllum schmitzii (Hariot) Mendoza Recorded as Lithothamnion schmitzii. The original record on which this report is based has not been traced (see Ricker, 1987: 175). ?Myriogramme livida (Hooker f. & Harvey) Kylin Skottsberg (1941: 76) tabulated the distribution of Antarctic and sub- Antarctic algae. No indication was given as to source and Skottsberg's record was not mentioned by Papenfuss (1964) in his catalogue of Antarctic and sub- Antarctic algae. Myriogramme smithii (Hooker f . & Harvey) Kylin Recorded as Nitophyllum fuscorubrum and N. smithii. Nereoginkgo adiantifolia Kylin Neuroglossum ligulatum (Reinsch) Skottsberg Recorded as Delesseria ligulata and Choreocolax delesseriae. Nothogenia fastigiata (Bory de Saint- Vincent) Parkinson Recorded as Chondrus crispus var. pigmaeus and Chaetangium fastigiatum. Palmaria decipiens (Reinsch) R.W. Ricker Recorded as Rhodymenia decipiens, R. palmata sensu Reinsch, R. palmetta var. multiloba, Leptosarca alcicornis and L. decipiens. According to Ricker (1987: 221), it is morphologically very similar to the type species of Palmaria (P. palmata) but distinguished by the readiness of its tissues to soften in 4% formalin-seawater. Palmaria georgica (Reinsch) R.W. Ricker Recorded as Rhodymenia georgica, R. palmatiformis and R. palmati- formis var. austrogeorgica. The overlap in thallus morphology of the two Palmaria species has led Ricker (1987: 224) to suggest that P. georgica may be 'merely a diminutive form or an ecotype of P. decipiens'. P. georgica separates from P. decipiens by forming tufts composed of many fronds, its bushier form, presence of cortical hairs, and occupying a different habitat. Pantoneura plocamioides Kylin Phycodrys austrogeorgica Skottsberg Phycodrys quercifolia (Bory) Skottsberg Recorded as Delesseria quercifolia. 1 Mi v I lop nora antarctica A. & E.S. Gepp Recorded as Ahnfeltia plicata and Phyllophora ahnfeltioides . Phyllophora appendiculata Skottsberg According to Skottsberg (1953: 542), this species might be identical with Gymnogongus turqueti Hariot (a Phyllophora, see Ricker, 1987: 202). Picconiella plumosa (Kylin) De Toni Recorded as Dasya? pectinata and Pteronia plumosa. ?Plectoderma minus Reinsch Uncertainty surrounds the taxonomic status of this taxon. The genus was established by Reinsch (1874) for two species of simple, crustose coralline algae. According to Chamberlain (1983: 308), this species 'was probably Melobesia membranacea judging by the cell shape and general disc pattern . . . No specimens have been found of either species so further identification is not possible'. Plocamium cartilagineum (Linnaeus) Dixon Recorded as P. coccineum. Plocamium hookeri Harvey Plocamium secundatum (Kuetzing) Kuetzing Polycoryne radiata Skottsberg Parasitic on Schizoseris dichotoma. BENTHIC MARINE ALGAL FLORA OF S. GEORGIA Polysiphonia anisogona Hooker f. & Harvey Porphyra endiviifolium (A. & E.S. Gepp) Chamberlain Recorded as Monostroma endiviifolium. Chamberlain (1963: 152) believed Hylmo's (1919: 6) record from South Georgia was based on a misidentification. New collections from the island now confirm its presence. Porphyra umbilicalis (Linnaeus) Kuetzing Recorded as P. laciniata and Wildemannia laciniata. Pseudolaingia larsenii (Skottsberg) Levring Recorded as Delesseria larsenii. Pterothamnion simile (Hooker f. & Harvey) Naegeli Recorded as Callithamnion pinastroides var. ramulosum, Antitham- nion ramulosum and A. simile. Sarcodia montagneana (Hooker f. & Harvey) J. Agardh Schizoseris condensata (Reinsch) R.W. Ricker Recorded as Delesseria condensata, Nitophyllum condensata, N. multinerve pro parte, N. a/fine, Schizoseris laciniata and Delesseria laciniata. See Ricker (1987: 285, 286) for discussion of nomenclature and synonymy. Schizoseris dichotoma (Hooker f. & Harvey) Kylin Recorded as Delesseria polydactyla, Myriogramme multinervis, Nito- phyllum polydactylum and Schizoseris polydactyla. Synarthrophyton patena (Hooker f. & Harvey) Townsend Recorded as Lithothamnium antarcticum. Rejected records Cystosphaera jacquinotii (Montagne) Skottsberg South Georgia: Neuschul (1968). Despite mentioning this taxon as occurring at this locality in the text (p. 10), it is not indicated on his distribution map nor the source of the record given (see Lamb & Zimmerman, 1977: 174). Plumariopsis eatoni (Dickie) De Toni The report of this taxon from South Georgia by Kylin & Skottsberg (1919) is probably a misdetermination for the closely related Geor- giella confluens (see above). Moe & Silva (1983) believe this species does not occur in Antarctica despite earlier reports. DISCUSSION Shore ecology The upper part of the eulittoral zone of rocky shores within Husvik Harbour is dominated by belts of Porphyra or mixed belts of Porphyra/ Nothogenia. Similar belts were described by Skottsberg (1941) at this shore level at May Cove (=Mai- viken), Cumberland Bay where an ' Adenocystis- Chlorophyceae association' was observed on the lower shore. Skottsberg makes no reference to belts of Schizoseris or Palmaria, but simply mentions that these two red algae were largely confined to tide pools containing a reasonably diverse algal community. He reported a 'Rhodymenia [= Palmaria]- Lithophyllum association' in rock-lined pools and a 'Rhody- menia [= Palmaria] association' in stony pools, while brackish-water pools contained Adenocystis, Scytosiphon, and Utriculidium. In the present study green algae were frequent in the littoral zone although the two most abundant taxa (Enteromorpha, Acrosiphonia) did not form a distinctive 111 belt. Only on boulder shores were green algae found to be relatively abundant. On one of the rocky shores a coating of a green filamentous alga was observed in the upper eulittoral zone (probably Ulothrix; not collected). No mention is made by Skottsberg (1941) of any green algae in his study of a stony beach at Boiler Bay (=Grytviken), Cumberland Bay where algal diversity was considerably less compared to the boulder shores studied at Husvik Harbour where Adenocystis and Nothogenia ('Adenocystis-Chaetangium [ = Nothogenia] asso- ciation') dominated. Unlike Husvik Harbour, the shores in Grytviken are affected by small icebergs ('growlers') pro- duced by calving glaciers of which seven occur in the Cumber- land Bay complex. Ice-scour and abrasion are known to be major physical factors affecting sessile shore organisms in high latitudes (see Keats et al., 1985). Our observations on the distribution of shore algae show close agreement with many of those made by Skottsberg (1941) in the early years of this century. He was of the opinion that his general observations enabled him to have a 'fair idea of the composition of the vegetation [algae]' of South Georgia. Of the 21 stations described by Skottsberg, all were relatively sheltered and confined to north-easterly shores. If he had been as familiar with South Georgia as claimed then it is surprising that he overlooked the bull kelp Durvillaea antarctica, first reported by Will in 1890 and yet going unmentioned by Reinsch (1888, 1890) who determined his algal material. Will's report of Durvillaea growing on rocky headlands on South Georgia was confirmed by Hay (1988, based on personal observations by Knowles) who states it forms (p. 426) 'a distinctive band in the low intertidal zone on the outer coast between Cumberland Bay and Royal Bay'. In 1991 one of us (PJAP) found a large bed of Durvillaea growing in the sublittoral fringe at Tonsberg Point (see Fig. 1). A large proportion of the tidal drift within Husvik Harbour comprises Durvillaea, much of it probably originates from this Point. On wave-exposed coasts Durvil- laea antarctica characterizes the sublittoral fringe as reported on other sub-Antarctic shores (see Stephenson & Stephen- son, 1972). The general distribution pattern of shore algae described on South Georgia during the short austral summer resembles those of sub-Antarctic and Antarctic rocky shores so far investigated (Knox, 1960; Kenny & Haysom, 1962; Price & Redfearn, 1968; Smith & Simpson, 1985; Delepine et al., 1966; Stephenson & Stephenson, 1972, among others). Absent from the littoral fringe are macroscopic growths of algae and the upper eulittoral subzone is defined by the genus Porphyra, below which appears Nothogenia. Simpson (1976) used the upper limit of Porphyra to define the upper limit of the eulittoral zone on Macquarie Island. Most algae on the lower shore seem confined to pools with the exception of Adenocystis, Iridaea, Schizoseris and Palmaria. As men- tioned above, these algae are present in the lower eulittoral zone and are common at this level on other shores of the Southern Ocean (see Stephenson & Stephenson, 1972). Cal- careous coralline algae or 'lithothamnia', often characteristic of the lower eulittoral subzone on lower latitude shores, form evident crusts in the sublittoral fringe on South Georgia. These crusts correspond to the "Lithophyllum-Lithothamnion association' of Antarctic shores, an association typically con- fined to rock pools and whose upper limit indicates the mean low tide level on sloping rocky shores (see Delepine et al., 1966). The presence of a 'Hildenbrandia' or "Hildenbrandia- Bostrychia association' on several sub-Antarctic shores has 112 not been observed on South Georgia. The sublittoral fringe on South Georgia is defined by the upper distributional limit of two kelps: juveniles of Macrocys- tis pyrifera on sheltered to moderately wave-exposed shores, and mature plants of Durvillaea antarctica on wave-exposed headlands. These brown algae characterize this fringe on sub-Antarctic and cold temperate shores, both reaching the southernmost limit of their distribution in South Georgia and Tierra del Fuego. The presence of Macrocystis on South Georgia led Skottsberg (1941) to remark (p. 36) that 'from the very first the visitor gets the impression that, regarded from an algological viewpoint. South Georgia must be included in the Subantarctic zone1. This statement was made at a time when Skottsberg had rejected Durvillaea antarctica as indigenous to the island. The high diversity and biomass of subtidal algal vegetation markedly contrasts with the low diversity and comparative barrenness of the littoral zone. On rocky bottoms the large kelps are the canopy dominants. In the entrance to sheltered inlets such as Grytviken (= Boiler Bay), a "Macrocystis- Desmarestia-Pteronia-Plocamium association' was reported by Skottsberg (194 1) as growing on stones down to a depth of 25 m. He noted that some of the dominant lower littoral algae (e.g. Palmaria sp. [as Rhodymenia palmatiformis]) grew to a depth as great as 30 m and described a number of associations which, albeit sampled indirectly by trawling, seemed to vary according to depth and substratum type. Marine algal flora The South Georgian algal flora is surprisingly diverse (103 species, 75 genera) considering the few shores (mostly wave- sheltered) investigated on its north-eastern coast during just a few austral summers. Undoubtedly species records for the island would increase if wave-exposed shores were visited and more subtidal collecting was undertaken. Skottsberg (1964), in recalling his two summers on South Georgia, states (p. 149) 'my journal from 1902-03 contains rough drawings of unknown Rhodophyceae lost in the shipwreck and never collected a second time'. Of the 103 species of marine algae known from South Georgia, nine are green algae, 35 brown algae, and two of the 47 red algae are unpublished records (Cladodonta lyallii, Callithamnion montagnei). The island is the type locality for 32 species (four genera), and yet South Georgia has just four endemic species and one endemic genus (Tables 1,2). The taxonomic status of these endemics and endemic infraspecific taxa (Iridaea cordata f. ligulata, Prasiola filiformis var. minuta, Viva lactuca var. macrogyna) is questionable. Twelve algal species are known only from South Georgia and Tierra del Fuego (Kuhnemann, 1972), an island lying about 2150 km due west of it. About 22% of the South Georgian algae also occur in the Northern Hemisphere, while Entonema tenuissi- mum, Calliblepharis ciliata, Callonema olivaceum and Callo- phyllis cristata are only known in the Southern Hemisphere from this island; all four records are regarded as doubtful. Almost half of its species are known from the Antarctic, the remainder reach their southernmost limit within the sub- Antarctic region. About 47% of the South Georgian marine algae are known from sub- Antarctic islands at similar or more northerly latitudes and mainland South America (see Tables 1,2). The remainder are confined to more southerly latitudes with a few reaching their northernmost limit at South Georgia (e.g. Desmarestia antarctica). D.M. JOHN, P.J.A. PUGH AND I. TITTLEY Table 1 An analysis of the marine benthic algae of South Georgia. Distribution A s-A SG SG+TF NH SG only only type loc. Acrosiphonia pacifica + Adenocystis utricularis + Ahnfeltia plicata + Anisocladella serratodentata Antithamnion ptilota Ascoseira mirabilis + Ballia callitricha + Bostrychia vaga Caepidium antarcticum 1 Calliblepharis ciliata Callithamnion montagnei 1 Callonema olivaceum 1 Callophyllis cristata Callophyllis linguata Callophyllis variegata + Ceramium diaphanum Ceramium involutum + Ceramium rubrum IChoreocolax rhodymeniae Cladodonta lyallii Cladophora incompta Cladothele decaisnei Clathromorphum obtectulum Colacodasya inconspicua Corycus lanceolatus Curdiea recovitzae + Delesseria salicifolia Delisea pulchra + Desmarestia antarctica + Desmarestia ligulata Desmarestia menziesii + Desmarestia pteridoides Desmarestia willii + Durvillaea antarctica + Ectocarpus constanciae Ectocarpus exiguus Ectocarpus siliculosus Elachista meridionalis Enteromorpha bulbosa + Enteromorpha gunniana + Entonema subcorticale Entonema tenuissimum Falklandiella harveyi Geminocarpus austrogeorgiae + Geminocarpus geminatus + Georgiella confluens + Halopteris funicularis Halopteris obovata Heterosiphonia berkeleyi + Himantothallus grandifolius + Hydrolithon discoideum 1 Hymenocladiopsis crustigena + Iridaea cordata (form) + Iridaea obovata + Lessonia fuscescens Macrocystis pyrifera Melastictis desmarestiae Mesophyllum schmitzii + Microrhinus carnosus IMyriogramme livida Myriogramme smithii + Myrionema densum Myrionema incommodum + Myrionema inconspicuum BENTHIC MARINE ALGAL FLORA OF S. GEORGIA Table 1 com. Distribution 113 A s-A SG only SG+TF only NH SG type loc. Myrionema macrocarpum Myrionema paradoxum Nereoginkgo adiantifolia Neuroglossum ligulatum Nothogenia fastigiata Palmaria decipiens Palmaria georgica Pantoneura plocamioides Petalonia fascia Petroderma maculiforme Phaearthron austrogeorgica Phycodrys austrogeorgica Phycodrys quercifolia Phyllophora antarctica Phyllophora appendiculata Picconiella plumosa Pilayella littoralis 1 Plectoderma minus Plocamium cartilagineum Plocamium hookeri Plocamium secundatum Polycoryne radiata Polyslphonia anisogona Porphyra endiviifolium Porphyra umbilicalis IPrasiola crispa (subsp. antarctica) Prasiola filiformis (var. minuta) Pseudolaingia larsenii Pterothamnion simile Sarcodia montagneana Schizoseris condensata Schizoseris dichotoma Scytosiphon lomentaria Scytothamnus fasciculatus Stegastrum porphyrae Synarthrophyton patena Syringoderma australe iflva lactuca (var. macrogyna) Utriculidium durvillaei Key to abbreviations: A, Antarctic (latitudes <55° S); s-A, sub- Antarctic; SG, South Georgia; TF, Tierra del Fuego; NH, Northern Hemisphere. A recent ordination analysis of the algal floras of the Southern Oceans by John et al. (1994) supports the findings of Lawson (1988: fig. 7) and Ricker (1987) who contend that the sub-Antarctic region represents a single circumpolar province. They observed no sharp discontinuity between the algal floras of the Antarctic and sub-Antarctic despite the principal surface water currents running counter to one another in the two regions. The close similarity between different sub- Antarctic islands is remarkable considering the remoteness and often vast distances separating many of them. It is speculated that much of the sub-Antarctic flora results from the long distance dispersal by the West Wind Drift of plants originating on South American shores. In conclusion, further investigations are required to deter- mine just how typical are the distribution patterns of algae in Table 2 Summary of an analysis of the marine benthic algae of South Georgia. Distribution A s-A SG SG+TF NH SG only only type loc. Chlorophyta 4 5 1 0 4 1 Phaeophyta 14 21 2 6 7 13 Rhodophyta 29 28 1 6 11 18 TOTAL 47 54 4 12 22 32 Key to abbreviations: A, Antarctic (latitudes <55° S); s-A, sub-Antarctic region; SG, South Georgia; TF, Ticrra del Fuego; NH, Northern Hemisphere. Husvik Harbour compared to other South Georgian shores. Advances in our knowledge of the seaweeds of South Geor- gia and the Southern Ocean in general continue to be hampered by lack of material (especially collected by SCUBA diving), paucity of taxonomic research on new or historically-important collections, absence of seasonal obser- vation on shore algae and few experimental studies designed to provide information on dynamics including plant-animal interactions. ACKNOWLEDGEMENTS. We are grateful to the British Antarctic Survey for providing research and logistic facilities, and particular thanks are due to the members of the 1990-1991 summer season expedition to Husvik, South Georgia (especially Mr H. MacAlister) for their help to PJAP with collection of material. We are indebted to Dr N. Jarman, formerly of the South African Antarctic Programme, for advice on collecting material given during his visit to Husvik on the yacht 'Diel'. Thanks go to Mrs Linda Irvine for assistance with the nomenclature of coralline algae, Mr J.H. Price and Dr Richard Moe for help with the literature, and Mrs Marian West for her editorial expertise. Finally, special thanks go to Dr G.W. Lawson for provid- ing information on the distribution of algae in the Southern Ocean. REFERENCES Admiralty Tide Tables. 1990. 1991. Volume 3: The Atlantic and Indian Oceans (including tidal stream predictions). London. Chamberlain, Y.M. 1963. The identity of Monostroma endiviifolium A. and E.S. 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Phaeophyccen. In O. Nordenskjold (Ed.), Wissenschaftliche Ergebnisse der schwedischen Siidpolar- Expedition 1901-1903 4 1(6): 172 pp., 10 pis, 1 map. Stockholm. 1921. Botanische Ergebnisse der schwedischen Expedition nach Patago- nien und dem Feuerlandc 1907-1909. VIII. Marine algae. 1. Phaeophyceae. K. svenska VetenskAkad. Handl. 61(11): 1-56. 1941 . Communities of marine algae in subantarctic and antarctic water. K. svenska VetenskAkad. Handl. Ill, 19(4): 1-92. Skottsberg, C. 1953. On two collections of Antarctic marine algae. Ark. Bot. II, 2: 531-566. 1960. Remarks on the plant geography of the southern cold temperate zone. Proc. R. Soc. B, 152: 447-457. 1964. Antarctic phycology. In R. Carrick, M. Holdgatc & J. Prevost (Eds), Biologic antarctique: 147-154. Hermann & Cie. Smith, J.M.B. & Simpson, R.D. 1985. Biotic zonation on rocky shores of Heard Island. Polar biol. 4: 89-94. Stephenson, T.A. & Stephenson, A. 1972. Life between tidemarks on rocky shores. San Francisco. Will, H. 1890. Vegetationsverhaltnisse Sudgeorgiens. In G. Neumayer (Ed.), Die deutschen Expeditionen und ihr Ergebnisse (1882-1883). 2. Bes- chreibende Naturwissenshaften. . . . : 366-499, pis 1-19. Berlin. Zaneveld, J.S. 1964. The benthic algal vegetation of Antarctica. Bull. U.S. Antarct. Pro'). Off. 5(10): 66-69. Bull. not. Hist. Mus. Lond. (Bot.)24(2): 115-159 Issued 24 November 1994 Studies in Pseudocyphellaria (Lichens) Palaeotropical species (excluding Australia) DAVID J. GALLOWAY Department of Botany, The Natural History Museum, Cromwell Road, London SW75BD SYNOPSIS. Twenty-nine species of Pseudocyphellaria are recorded from the palaeotropics (from Africa to the eastern Pacific but excluding Australia) viz., P. argyracea, P. aurata, P. beccarii, P. carpoloma, P. clathrata, P. crocata, P. crocatoides, P. desfontainii, P. dissimilis, P. dozyana, P. gilva, P. godeffroyii, P. haywardiorum, P. homalosticta, P. insculpta, P. intricata, P. maculata, P. multifida, P. neglecta, P. pickeringii, P. poculifera, P. prolificans, P. punctillaris, P. reineckeana, P. rigida, P. semilanata, P. stenophylla, P. sulphurea and P. trichophora. Details of their anatomy, chemistry, morphology and distribution are presented together with a key. The following new combinbations are proposed: P. beccarii (Kremp.) D.J. Galloway and P. trichophora (Vain.) D.J. Galloway. INTRODUCTION Species of Pseudocyphellaria are conspicuous, leafy, foli- ose lichens best developed and with richest biodiversity in rainforest, shrubland and successional vegetation, or subal- pine and alpine grassland habitats of the Southern Hemi- sphere cool temperate zone, with major areas of speciation being New Zealand (Galloway, 1988) and southern South America (Galloway, 1992). In tropical regions Pseudocyphellaria is most commonly found in montane or mossy cloudforest between 1600 and 3600 m (see dicussion on altitudinal zonation in Sipman (1993)), but several widespread species such as P. aurata, P. crocata and P. intricata occur at lower altitudes and in lowland and coastal sites as well. The contribution of species of Pseudocyphel- laria to the 'Lobarion' alliance in south-east Asian forests is discussed in Wolseley (1991). During a study of Australian species of Pseudocyphel- laria (in preparation) very many collections from neigh- bouring areas in the Pacific Basin were examined, so that it is now possible to present a preliminary revision of palaeo- tropical species. Neotropical taxa in Pseudocyphellaria are discussed in accounts of Ecuadorean (Galloway & Arvids- son, 1990) and Brazilian (Galloway, 1993) collections, and in catalogues of Central American (Imshaug, 19560), Mexi- can (Imshaug, 1956Z?) and West Indian (Imshaug, 1957) lichens. Earlier accounts discussing or listing palaeotropi- cal taxa referable to Pseudocyphellaria include: Montagne (1856), Krempelhuber (1875), Zahlbruckner (1908, 1943), Vainio (1913, 1924), Magnusson (1940), Zahlbrucker & Mattick (1956), Szatala (1956), Joshi & Awasthi (1982), Hawksworth & Shaw (1984), Streimann (1986) and Swins- cow & Krog (1988). The context of palaeotropics used in the present account refers to all land in the tropical-subtropical zone outside of the neotropical region, viz., from Africa to the eastern Pacific bordering North, Central and South America and lying roughly between the tropics of Cancer and Capricorn at latitudes 35° north and south of the equator. Species of Pseudocyphellaria discussed in this revision are * Part III In Bibliotheca Lichenologica 46: 1-275 (1992). generally conspicuous lichens, some often reaching a great size and being among the largest and most rapidly growing of foliose lichens. They grow on twigs, bark, soil, or rock, often over or intermingled with bryophytes or other lichens in a wide variety of habitats. Since all taxa contain cyanobacteria either as a primary photobiont or as internal cephalodia they are efficient nitrogen fixers and important contributors to rainforest nitrogen budgets, a role which is of importance in the maintenance of rainforest biodiversity (Galloway, 1994). Detailed accounts of anatomical, morphological and chemical characters useful in species delimitation in the genus are given in Galloway (1988, 1992) and are not repeated here. The importance and possible role of triterpenoids in Pseudocyphellaria are discussed by Galloway (1991) and Wilkins (1993). The undoubted importance of species of Pseudocyphellaria in a variety of tropical ecosystems makes a modern account of this genus a vital necessity. However, in offering this present revision as a contribution to tropical lichenology I must admit to its being almost entirely a herbarium study. I have col- lected only briefly from Peninsular Malaysia in the palaeotro- pics and consequently have not had the advantage of assessing variation in the field. Accordingly, I have taken a rather broad view of the limits of taxa and in widespread species, such as P. argyracea, P. crocata, P. gilva, P. intricata and P. sulphurea for example, I accept a wide morphological variation which seems acceptable based on the variation of these taxa in temperate habitats. It is hoped that the present revision will form a working baseline to the genus in the tropics and encourage lichenologists to undertake closer regional studies of it. Type and other material was obtained from or examined in the following herbaria: AK, B, BM, BR, BSIP, CBG, COLO, E, G, GB, H, H-ACH, H-NYL, KEP, KLU, L, LD, LG, M, MEL, NY, PC, PC-LENORMAND, PC-THURET, S, TNS, TUR-VAINIO, UKMB, UPS, UPS-THUNBERG, UPSV, US, W, WU and from the following private herbaria: Dr A. Aptroot (Baarn), Dr L. Arvidsson (Goteborg), Prof. G. Degelius (Askim) [Prof. Degelius's lichens are now at UPS], Dr P. Diederich (Luxembourg), Dr K. Kalb (Neu- markt), and Prof. C.W. Smith (Honolulu). ©The Natural History Museum. 1994 116 D.J. GALLOWAY SYSTEMATIC TREATMENT Key to palaeotropical species of Pseudocyphellaria 1 Medulla white 2 Medulla yellow 26 2 Photobiont green 3 Photobiont blue-green 9 ate 7. P. crocatoides Phyllidiate 19. P. neglecta 25 Upper surface deeply faveolate 17. P. maculata Upper surface undulate 12. P. gilva 26 Sorediate or isidiate/phyllidiate 27 Not sorediate , isidiate or phyllidiate 5 . P. clathrata 27 Sorediate 28 Isidiate/phyllidiate 20. P. pickeringii 3 Pseudocyphellae white 4 28 Soralia linear, confluent, labriform 2. P. aurata Pseudocyphellae yellow 4. P. carpoloma Soralia derived from small, marginal, crowded isidia 2 1 . P. poculifera 4 Pseudocyphellae present on upper surface 5 Pseudocyphellae not present on upper surface 6 The species 5 Isidiate-phyllidiate 15. P. homalosticta Without isidia or phyllidia 24. P. reineckeana 1. Pseudocyphellaria argyracea (Delise) Vain, in Hedwigia 6 Without isidia or lobules 7 3?. 35 (189g) Sficta argyracea Delise in Mem. Soc. linn. Normandie 2: 91 pi. 7, fig. 30 (1825). Stictina argyracea I Lobes broad, punctate-impressed 28. P. sulphurea (Delise) Nyl., Syn. meth. lich. 1(2): 334 (1860). Cyanisticta Lobes narrow 27. P. stenophylla argyracea (Delise) Gyeln. in Reprium Spec. nov. Regni itjfiHo veg. 29: 2 (1931). Type: He de la Reunion. Sur des troncs 8 Lobes broad, rounded, phyllidiate 18. P. multiiida , . Lobes narrow, punctate-impressed 22. P. prolificans ou les rochers mosses des hautes regiones, Wory de St-Vincent, ex Herb. Bory (PC-THURET-lectotype (Gal- 9 Pseudocyphellae white 10 loway & James, 1986: 429)). Pseudocyphellae yellow 21 10 With soredia or isidia 11 Sticta argyracea var. sorediifera Delise in Mem. Soc. linn. Without soredia or isidia 16 Normandie 2: 92 pi. 7, fig. 31 (1825). Stictina argyracea var. II Sorediate 12 sorediifera (Delise) Nyl., Syn. meth. lich. 1(2): 334 (1860). Isidiate/phyllidiate "". ]""""!"!'.!!!!!"!""'.! 15 Pseudocyphellaria argyracea var. sorediifera (Delise) Malme in Bih. K. svenska VetenskAkad. Handl. 25(3/6): 12 Upper surface smooth or punctate-impressed; 2 hopanes 24 (1899). Type: Madagascar, without specific locality, Present 13 collector or date (PC-LENORMAND-lectotype (Gallo- Upper surface faveolate or punctate-impressed; nopane tnol „ jarnes igof,. A^rm Sticta boryana Delise in Mem. Soc. linn. Normandie 2: 102 13 Upper surface plane or undulate; pseudocyphellae on lower pi. 8, fig. 37 (1825). Pseudocyphellaria boryana (Delise) surface rare or absent 16. P. intricata D.J. Galloway in Lichenologist 17: 303 (1985). Type: He Upper surface punctate-impressed; pseudocyphellae on lower de Bourbon [Reunion], Plaine de Chicots, sur le bois mort surface prominent 14. P. haywardiorum des for£ts montagneuses, Bory de St-Vincent s.n. (PC- 14 Isidiate 14 THURET-holotype). Phyllidiate 9. P. insculpta Sticta rigidula Delise in Mem. Soc. linn. Normandie 2: 97 pi. 8, fig. 34 (1825). Stictina argyracea f. rigidula (Delise) Nyl. 15 Isidia associated with pseudocyphellae 1 . P. argyracea \, . , ,. „• f D TTT -> oo^ ... ,n „ . . ... in Hue, Nouv. Archs Mas. Hist. nat. Paris 111, 2: 295 Isidia not associated with pseudocyphellae .... 10. P. dissimilis .... ,^ ,- \ -» in_ ^ (1890). Sticta argyracea f. rigidula (Delise) Zahlbr., Cat. 16 Pseudocyphellae present on upper surface 17 Uch. univ. 3: 371 (1925). Cyanisticta rigidula (Delise) Pseudocyphellae absent on upper surface 3. P. beccarii QW. Dodge in Beih. nov. Hedwigia 12: 178 (1964). Type: 17 Isidia absent 18 ^e de Bourbon [Reunion], sur 1'ecorce, Bory de Isidia or marginal lobules present 23. P. punctillaris St-Vincent s.n. (PC-THURET-lectotype (Galloway & James, 1986: 432)). 18 Upper surface plane 19 Sticta f lav escens Delise in Mem. Soc. linn. Normandie 2: 117 Upper surface scrobiculate 25. P. rigida pl ^ fig 4? (lg25) ^^ argyracea var flavescens 19 Upper surface smooth, not areolate-scabrid 20 (Delise) Nyl. in Hue, Nouv. Archs Mus. Hist. nat. Paris Upper surface areolate-scabrid 13. P. godeffroyii HI, 2: 295 (1890). Sticta argyracea var. flavescens (Delise) ~0 t Zahlbr., Cat. lich. univ. 3: 372 (1925). Cyanisticta flave- 20 Margins of lobes tomentose-hairy 29. P. tnchophora . \ ' ,/,..,;;.,-- Margins of lobes glabrous 26. P. semilanata scens (Delise)CW. Dodge in Beih.nov. Hedwigia 12: 173 (1964). Type: He de Bourbon [Reunion], sur Tecorce, 21 Sorediate or isidiate/phyllidiate 22 gory fa St-Vincent (PC-THURET-holotype). Without soredia or isidia 25 Sticta aspem Laurer in Unnaea 2: 41 (1827) . Sticta argyracea 22 Isidiate/phyllidiate 23 var. aspera (Laurer) Kremp. in Verh. zool. -hot. Ges. Wien Sorediate 6. P. crocata 18: 316 (1868). Stictina argyracea var. aspera (Laurer) Mull. Arg. in Revue mycol. 9: 138 (1887). Cyanisticta 23 Phyllidiate or with marginal or laminal proliferations ... 2 Q (Laurer) c w Dod in ^ ^ Hedwigia 12: 170 (1964). Type: Mauritius, Sieber 40 (L 910,215-1683- 24 With laminal and marginal proliferations, not truly phyllidi- lectotype (Galloway & James, 1986: 430)). STUDIES IN PSEUDOCYPHELLAR1A (IV) Stictina argyracea f. insidiata Nyl. in Cromb., J. Linn. Soc. (Bot.) 15: 435 (1876). Sticta argyracea f. insidiata (Nyl.) Zahlbr., Cat. lich. univ. 3: 371 (1925). Type: Ins. Rod- riguez, I.E. Balfour 2279 (H-NYL 34058-holotype; BM-isotype). (The holotype material in Nylander's her- barium is a small scrap taken from a larger collection in Crombie's herbarium (BM) which is preserved as two separate specimens, only one of which is numbered 2279. All three specimens are labelled 'f. isidiata Nyl.' and not insidiata as appears in the protologue.) Cyanisticta javanica Gyeln. in Reprium Spec. nov. Regni veg. 29: 297 (1931). Type: Java, Prov. Preanger, in monte ignivomo Papandayan, 1750 m, Schiffner 3309 (L 956.124 594-isotype). Pseudocyphellaria horridula H. Magn. in H. Magn. & Zahlbr., Ark. Bot. 31A: 82 (1943). Cyanisticta horridula (H. Magn.) Szatala in Annls hist. -not. Mus. natn. hung. 7: 41 (1956), comb, inval. Type: Hawaii ad truncos muscosos in paludosis ad Waimea, 4000', J.F. Rock 6 (W-isotype). Pseudocyphellaria horridula var. excrescens H. Magn. in H. Magn. & Zahlbr., Ark. Bot. 31A: 83 (1943). Type: Hawaii. Maui, lao Valley, Faurie 566 (W-not seen). Pseudocyphellaria argyracea is a characteristic, laminally pseudocyphellate, pseudoisidiate, white-medulla species which is widespread in the palaeotropics from the Indian Ocean islands to the Pacific basin as far east as the Galapagos Islands (Weber, 1986) and as far south as New Zealand (Galloway, 1988) and southern Chile (Galloway, 1992). Typi- fication of this species is discussed by Galloway & James (1986) and a detailed account of the morphology and anatomy is given in Galloway (1988: 64-68). CHEMISTRY. Methyl gyrophorate, gyrophoric acid, 7p-acetoxyhopane-22-ol, hopane-7(3, 22-diol (tr.), hopane- 15a, 22-diol. 117 OBSERVATIONS. Pseudocyphellaria argyracea is slatey grey- blue to blue-black, often tinged red-brown when wet, pale grey to grey-brown when dry; it has a white medulla, a cyanobacterial photobiont and white pseudocyphellae on both upper and lower surfaces. It is characterized by numer- ous, evenly spaced, laminal, white pseudocyphellae which at maturity become pseudoisidiate with corticate, fingerlike pseudoisidia, concolorous with the thallus, developed in clusters, some of which may become abraded and appear granular-sorediate. New isidia often develop from older abraded structures. Superficially P. argyracea resembles some broad-lobed forms of P. intricata but differs in the pseudoisidiate clusters associated with the laminal pseudocyphellae. The taxon P. boryana (Galloway, 1985ft; Galloway & James, 1986: 432-3) has narrower, more dis- sected, ± dichotomously branching lobes and distinctive, marginal proliferations, the laminal pseudocyphellae of which are not sorediate or associated with isidia. However, it seems only to be an extreme form of P. argyracea and is therefore placed in synonymy with this species. DISTRIBUTION AND ECOLOGY. Pseudocyphellaria argyracea is a widespread palaeotropical taxon (Fig. 1), extending from East Africa (Swinscow & Krog, 1988) to southern South America (Galloway, 1992) northwards to India (Awasthi, 1965, 1988), China (Wei, 1991) and Japan (Yoshimura, 1974), and southwards to New Zealand (Galloway, 1988). On bark of saplings and small trees and on mossy trees, rotting logs in humid, shady montane and cloud forest, often in crowns of trees, from 700 to 3650 m. Also at lower elevations in coastal sites. SPECIMENS EXAMINED. Africa. Tanzania: Tanga. Usambara Mountains, Amani, Santesson 23370 (UPS); ibid, Brunntha- ler (W, WU); Nazumbei, Brunnthaler (WU); Ukaguru Mts, road from Mandege Forest Station to Rubeho, Pocs, Harris Fig. 1 Distribution of Pseudocyphellaria argyracea in the palaeotropics. 118 D.J. GALLOWAY & Mwanjabe 6588 (BM); Uluguru Mts, Mwere Valley, Pocs, Farkas, Geissler, Iversen, Steiner & Temu 86158 (BM). Uganda: Masaka, Nanuzinna swamp, Lye L 640 (BM); Kigezi, Kinkizi, Swinscow 3U 56/5 (BM). Kenya: Mt Kenya, 200-2100 m, Swinscow K48.34A, K 51/3 (BM). South Africa: Transvaal. Drakensberge, Werdermann & Oberdieck 1849 (B). Comoro I: Anjouan, MTingui Peak, Benson 183 (BM). Madagascar: ?Andrangoloaka, ?Likora (W); sine loco, Herb. Persoon (L); Roxburgh (BM); sine loco, Lam en Meeuse 5966 (L). Mauritius: sine loco, Robillard (W); [ISO- TYPE] Sieber 40 (W); McGregor 1819 (BM); Vacquois, Ayres (BM). Reunion: sine loco, Richard (H-NYL 34064); Cirque de Salazie, K. & A. Kalb 26560, 26561, 26564 (Herb. Kalb); Cirque de Cilaos, de Sloover 17.463 (LG). Seychelles: Silhouette, Gardiner 1905 (BM). Rodriguez Is: Balfour 2279 (BM). Sri Lanka: Nuwara Eliya, Meltzer s.n. (Herb. Aptroot); Rampodde, Almquist (H-NYL 34065); v.Beusekom 290 (Herb. Aptroot); Habgalla, Thwaites (BM); above Pattipola, Horton Plains, van Steenis 19924c (L). Thailand: Nakhon Sawan, Touw 8237 (Herb. Aptroot). Malaysia: Pahang. Eraser's Hill, Dransfield 481 (BM); Eras- er's Hill, Burkill 2084 (L); Eraser's Hill, Galloway (BM, KEP, KUL). Indonesia. Sumatra: sine loco, Korthals (L). Java: Tjibodas, Koernich 2a (Herb. Aptroot); Arvidsson & Nilsson (GB); Mt Kawi, Mt Panderman, Groenhart 1931, 1936, 2632 (L); Mt Gede, Schiffner 3289b (L); sine loco, Junghuhn (L); Mt Ardjuno, Groenhart 29, 668, 1531, 1988, 7332 (L); Mt Lawu, Clason 986 (L); Mt Wilis, Groenhart 1537, 1838 (L); Mt Pangerango, Schiffner (WU). Flores: sine loco, Verheijen 5201 (Herb. Aptroot). Sulawesi: sine loco, De Vriese (L). Philippines: Luzon. Pampanga. Mt Pinatubo, Elmer 22270 (B, BM). Mindanao. Butuan, Weber 1352 (US); Rizal, 1911, Ramos 13634 (BM); sine loco, Camming 2156 (BM). Papua New Guinea: Eastern Highlands. Chimbu. Mt Wilhelm, Weber & McVean (Herb. Aptroot L54979, COLO); Aptroot 18235, 18333, 18651, 32786 (Herb. Aptroot); Kashi- wadani 10882, 10924, 11188, 11199, 11410, 11418 (TNS); Wade (COLO); McVean 66182 (CBG); track to Mt Wilhelm, Sipman 21923 (B); Pindaunde Valley, Sipman 15905, 15906, 22098 (B); Goroka. Mt Gahavisuki Provincial Park, Aptroot 32420 (Herb. Aptroot); Streimann 18215 (CBG); Daulo Pass, Streimann 18110 (CBG). Morobe. Lake Wamba, Koponen 33406 (Herb. Aptroot); Mt Kaindi, Weber & McVean (COLO); Streimann & Bellamy 17665 (CBG); Herzog Moun- tains, Streimann & Umba 11015 (CBG); Gumi Divide, Stre- imann 22761 (CBG); Ekuti Divide, Streimann 20147 (CBG). Southern Highlands. Margarima, Streimann 24393 (CBG); Munie Logging Area, Streimann 23674 (CBG); Onim For- estry Station, Streimann 24649 (CBG); laro River, Streimann 23979 (CBG). Milne Bay. Woodlark Island, Kumei 57, 71 (CBG). Central. Mt Albert-Edward, Kashiwadani 12002 (TNS). Western Highlands. Tumbang Village, Streimann 21351, 21371 (CBG). Solomon Islands: Guadalcanal Island. Mt Popomansiu, Hill 9314, 9523, 9563, 9690, 9704, 9853 (BM). Bougainville. Lake Luralu, Kajewsky 1930 (BM). New Caledonia: Riviere Bleue, Hill 11689 (BM); Riviere Blanche, Hill 11699 (BM). Fiji: Viti Levu. Nandarivatu, Green (BM). Tahiti: sine loco, Vieillard & Panchon (H-NYL 34064); Aorai, v.Balgooy (Herb. Aptroot). Hawaiian Islands: Hawaii. Waimea, Rock [Lichenes Sandwicenses No. 6] (W). Kauai. Kaholnamano, Rock [Lichenes Sandwicenses No. 11] (W). Oahu. Central Waianae Mts, Waianae Kai Forest Reserve, Honua Stream, Smith 1611 (Herb. Smith); Honou- liuli Forest Reserve, Puu Kaua, Smith 4126 (Herb. Smith); trail to Puu Kalena, Smith 1549a (Herb. Smith); Koolau Mts, Koolauloa District, Kahana Valley, Vitt 14691 (H). Galapa- gos Islands: Isla Pinzou. Sipman L106 (COLO). 2. Pseudocyphellaria aurata (Ach.) Vain, in Ada Soc. Fauna Flora fenn. 7: 183 (1890). Sticta aurata Ach., Methodus: 277 (1803). Type: ? England, Devon, without specific locality, ex Herb. Hudson-label incomplete (H-ACH Fig. 2 Distribution of Pseudocyphellaria aurata in the palaeotropics. STUDIES IN PSEUDOCYPHELLARIA (IV) 1534-holotype). For additional synonymy see Galloway (1988). Pseudocyphellaria aurata is bright lettuce-green tinged yellow-gold when wet, pale green-grey, often tinged or becoming reddish on storage when dry; it is a characteristic, yellow-medulla species which is widespread in tropical regions of the world, for example in Ecuador (Galloway & Arvidsson, 1990: 116-118) and is also found in drier, warmer, coastal areas in cool temperate regions, particularly in the Southern Hemisphere where it is known from New Zealand (Galloway, 1985a, 1988), eastern Australia and Chile (Gallo- way, 1992). Palaeotropical material closely approximates in both anatomy and morphology the description given in Galloway (1988: 68-69). CHEMISTRY. Pulvinic acid, pulvinic dilactone, calycin, 3p-acetoxyfern-9( 1 1 )-en- 12-one , 3p-acetoxyfern-9( 1 1 )-en- 123-cl, fern-9(ll)-ene-3p, 12p-diol, 3p-acetoxyfern-9(ll)-en- 19p-ol, 3p-hydroxyfern-9(ll)-en-12-one, lupeol acetate (Wilkins & Elix, 1990). OBSERVATIONS. Pseudocyphellaria aurata is a cosmopolitan species having a yellow medulla, a green photobiont, yellow pseudocyphellae on the lower surface, and prominent, mar- ginal, labriform, ± linear yellow soralia, that often erode back the lower surface and contain coarse, granular yellow soredia. Apothecia rather rare, submarginal, distinctly pedi- cellate, exciple concolorous with thallus, margins ragged, yellow-sorediate. Spores brown, broadly fusiform-ellipsoid, 3-septate (25-)30-32 x 6-7 |xm. DISTRIBUTION AND ECOLOGY. Pseudocyphellaria aurata occurs on trees, shrubs and rocks in both open, sunny sites as well as in the forest canopy and on branches and twigs in moderate shade in montane forests. From sea level to 2300 m. Widespread in the tropics (Fig. 2) and in cool temperate regions (Galloway, 1988, 1992; Galloway & Arvidsson, 1990). SPECIMENS EXAMINED. Africa. Tanzania: Arusha National Park. Mt Meru, Renvoize 2493c (BM). Kenya: Mt Marsabit, Lye (BM). Uganda: Lake Mulehe, Swinscow (BM). Malawi: Zomba Plateau, Jellicoe (BM). South Africa: Cape Peninsula. Hout Bay, Maas Geesteranus 14659 (Herb. Aptroot); near Knysna, Werdermann & Oberdieck 913b (B); Cape of Good Hope, Ecklon (B); Table Mt, Eaton (BM). Natal. Sim (BM). Transvaal. Kowyns Pass near Graskop, Sipman 19.926 (B); Long Tom Pass, Sipman 20.094 (B); Drakensberge, Werder- mann & Oberdieck 1855 (B). Madagascar: Imarina, Cowan (BM). Reunion: sine coll. (BM); southern slope of Piton des Nieges, near Cilaos, Arvidsson & Nilsson 2536 (GB); Cirque de Cilaos, de Sloover 17.471, 17.473, 17.631, 17.780 (LG). Rodriguez I: Balfour (BM). Malaysia: Pahang. Fraser's Hill, Dransfield 517 (BM); Fraser's Hill, Galloway (BM, KEP, KLU); Cameron Highlands, Tanak Rata, Degelius As-547, As-550 (UPS). Indonesia. Sumatra: Bukittinggi, Hensen (Herb. Aptroot). Java: Tjibodas, Koernich 2b (Herb. Aptroot). Philippines: Luzon. Benguet. Mt Santo Tomas, Aptroot 20449 (Herb. Aptroot); Sipman 21751 (B); Baguio, Luneta Hill, Degelius As-905 (UPS). New Guinea: Morobe. Edie Creek Road, Streimann NGF 39103 (CBG); v.Royen NGF 16288 (Herb. Aptroot); Kwama River Valley, Koponen 33232 (Herb. Aptroot); Upper Watut River, Streimann 17072 (CBG); Herzog Mountains, Streimann & Umba 10988, 11137 (CBG); Honzeukngon village, Aptroot 17773 (Herb. 119 Aptroot); Pouyu Village, Streimann & Tamba 12575, 12693 (CBG); Upper Nawata Band, Streimann 33973 (CBG); Manki Trig, Streimann & Bellamy 12942 (CBG); Mt Susu, Streimann 34182 (CBG); Kauli Lake, Streimann 34099 (CBG); Mt Kaindi, Streimann 33418 (CBG). Madang. Finis- terre Range. Teptep Village, Aptroot 32286 (Herb. Aptroot). Eastern Highlands. Lapegu, Streimann 18275, 18396, 18427, 18443 (CBG); Mt Michael, Streimann 18786 (CBG); Goroka. Mt Gahavisuki Nature Reserve, Aptroot 18848 (Herb. Aptroot). Western Highlands. Kagamuga, Streimann 21701 (CBG). New Caledonia: sine loco, Compton 716 (BM). Norfolk Island: Mt Pitt Reserve, Duncombe Road, Streimann 34595 (CBG); Broken Pine, Elix 18315 & Streimann (BM). Kermadec I.: Cheeseman (BM). Hawaiian Islands: Oahu. Waianae Mts, Honouliuli Forest Reserve. Puu Kaua, Smith 4126 (Herb. Smith); Waianae Kai Forest Reserve, Honua Stream, Smith 1611 (Herb. Smith). Fiji: Viti Levu. Nadari- vatu. Nadala, Degener 31807b (B); District Commisioners House, Degener 31814ad (B); Mba, Smith 5965 (BM). Samoa: sine loco, Powell (BM). Galapagos Islands: Isla Charles. Trail from Black Beach to highlands, Weber & Lanier (COLO); Weber 328, 427 (COLO). Isla Duncan. Summit, Cavagnaro (COLO). Isla Isabella. Volcan Cerro Azul, Sipman L-67 (COLO); Weber & Lanier (COLO). Isla San Cristobal. West of El Junco, Lanier (COLO). 3. Pseudocyphellaria beccarii (Kremp.) D.J. Galloway, comb. nov. Fig. 3. Basionym: Sticta beccarii Kremp. in Nuovo G. hot. ital. 7: 11 (1875). Stictina beccarii (Kremp.) Mull. Arg. in Flora, Jena 65: 301 (1882). Type: Sarawak, O. Beccari, Lichenes Bornenses, No. 121, 1866 (M-lectotype, selected here; BM, M-isolectotypes). Stictina fragillima var. subpunctulata Nyl. in Leight., Trans. Linn. Soc. Lond. 27: 164 (1869). Stictina subpunctulata (Nyl.) Stizenb. in Flora, Jena 81: 138 (1895). Sticta sub- punctulata (Nyl.) Hue in Nouv. Archs Mus. Hist. not. Paris IV, 3: 54 (1901). Pseudocyphellaria subpunctulata (Nyl.) Vain, in Philipp. J. Sci. sect. C, Bot. 8: 119 (1913). Cyanisticta subpunctulata (Nyl.) Szatala in Annls hist. -not. Mus. natn. hung. 7: 41 (1956), comb, inval. Type: Ceylon, Central Province, G.H.K. Thwaites C.L. 22 (BM- lectotype, selected here). Stictina junghuhniana Mull. Arg. in Flora, Jena 65: 300 (1882). Pseudocyphellaria junghuhniana (Mull. Arg.) D.D. Awasthi in Beih. nov. Hedwigia 17: 104 (1965). Type: In Insula Java, Junghuhn (L-910,215-1406- lectotype, selected here). Stictina junghuhniana var. laevis Mull. Arg. in Flora, Jena 65: 300 (1882). Stictina subpunctulata var. laevis (Mull. Arg.) Stizenb. in Flora, Jena 81: 128 (1895). Sticta subpunctulata var. laevis (Mull. Arg.) Zahlbr., Cat. lich. univ. 3: 399 (1925). Pseudocyphellaria junghuhniana var. laevis (Mull. Arg.) D.D. Awasthi in Beih. nov. Hedwigia 17: 104 (1965). Type: Ceyloniae, in montanis cum forma genuina speciei, altit. circ. 6000-pedali, Nieter (G-not seen). Thallus irregularly spreading in entangled clones, 8-15(-25) cm diam., loosely attached centrally, apices free, ascending. Lobes linear-elongate, 3-8(-15) mm wide, l-5(-10) cm long, ± dichotomously to irregularly branching, contiguous or discrete at margins, ± imbricate centrally, apices divergent. 120 D.J. GALLOWAY Fig. 3 Pseudocyphellaria beccarii. Lectotype Stictina fragillima var. subpunctulata (BM). Scale in mm. rounded, truncate or sharply to bluntly furcate. Margins entire, noticeably thickened, ridged below and often con- spicuously pseudocyphellate, very rarely with occasional small lobules developed. Upper surface dark grey-black or blue-black, suffused red-brown at apices, pale glaucous buff or greyish when dry, conspicuously deeply to shallowly punctate-depressed or dimpled, irregular to undulate or wrinkled, not faveolate, coriaceous, matt or shining in parts, maculate, without isidia, phyllidia, pseudocyphellae or sore- dia. Medulla white. Photobiont cyanobacterial. Lower surface pale buff or whitish at margins, red-brown to black centrally or sometimes uniformly pale brown from margins to centre, irregularly wrinkled to ± bullate, tomentum rather variable, from scattered thin patches centrally to densely and uni- formly developed from margins to centre, pale buff or whitish to red-brown or blackened. Pseudocyphellae white, widely scattered to common and ± crowded, round to irregular, O.l-l(-l.S) mm diam., conical-verrucose, margins distinctly raised and sharply defined, concolorous with lower cortex, decorticate area flat to ± concave, projecting above thin tomentum or usually sunk in thick tomentum, often ± conspicuous at margins. Pycnidia mainly marginal, crowded in lines, rarely scat- tered on upper surface, ostiole hemispherical brown-black, 0.1 mm diam. Apothecia marginal or submarginal, rarely laminal, sparse to moderately frequent, rounded to subirregular, 2-5 mm diam., sessile to subpedicellate, exciple coarsely verucose- scabrid, obscuring disc when young, pale buff to brown, disc STUDIES IN PSEUDOCYPHELLARIA (IV) 121 to dark red-brown, to ± blackened, consistently darker than margin, concave at first, plane to subconvex at maturity, matt, smooth, epruinose. Epithedum pale yellow-brown, 12-15 (Jtm thick. Hymenium colourless, 100-115 |xm tall. Ascospores pale brown, 3-septate, ellipsoid, apices pointed, 33.5-42.5 H4.5) x (6.5-) 8.5-11 jxm. CHEMISTRY. Tenuiorin, methyl gyrophorate, gyrophoric acid, 7p-acetoxyhopane-22-ol, hopane-7pi, 22-diol (tr.), hopane-15a, 22-diol. OBSERVATIONS. Pseudocyphellaria beccarii is characterized by a white medulla; a cyanobacterial photobiont; a dimpled, punctate-impressed upper surface; ± dichotomously branch- ing lobes with entire margins; scattered, flecklike white pseudocyphellae on the lower surface; and a two-hopane chemistry with tenuiorin, methyl gyrophorate and gyrophoric acid as accessory substances. It is distinguished from P. insculpta in the absence of marginal isidia and phyllidia, from P. sulphurea in having a cyanobacterial photobiont, and from P. semilanata which has scattered white pseudocyphellae on the upper surface. DISTRIBUTION AND ECOLOGY. Widespread in the palaeotro- pics (Fig. 4), from Madagascar eastwards to Fiji and Samoa but not known from Hawaii. Also in north-eastern Australia. An epiphyte of montane rainforest and mossy cloudforest on trees and shrubs, 1500-2800 m. SPECIMENS EXAMINED. Madagascar: sine loco, Thompson (M). Sri Lanka: sine loco, Beccari 12 [Crittogame di Ceylan No. 12] (M); Nuwara Eliya, IBlallu 74 (W). Burma: sine loco, Lobb (BM). Malays!: Sabah. Mt Kinabalu, Sipman & Tan 30960, 31084, 31377 (B); Richards (BM); Clarke 86 (BM). Indonesia. Java: Mt Ardjuno, Groenhart 1516, 1852, 1993, 1998, 2042, 4615, 7245, 7327, 7329 (L); Mt Kawi, Groenhart 1829, 1951, 1963, 7255 (L); Mt Lawu, Clason 982 (L); Mt Wilis, Groenhart 1538 (L); Batu, Roomaker 1985 (L); Mt Gede, Groenhart 7271 (L); Mt Megamendung, Schiffner 1159, 3351 (L, WU). Kalimantan: sine loco, Beccari (M). Philippines. Luzon: Benguet, Mt Pulog, Jacobs B47 (Herb. Aptroot); Curran, Merritt & Zschokke (US); Pauai, McGregor (E, US); Pampanga. Mt Pinatubo, Elmer 22270 (B, BM); Laguna. Mt Banajao, Merrill 7525 (US); Mt Malinao, Edano 37208 (L). Mindoro: Alag River, Merrill 5497 (US). Papua New Guinea: Eastern Highlands. Chimbu. Mt Wilhelm, Sipman 21959, 22137 (B); Borgmann 776, 896 (B); Kashiwadani 10845, 10917, 10965, 11080 (TNS); McVean 66149 (CBG); Lake Aunde, Aptroot 18466 (Herb. Aptroot); Pindaunde Valley, Weber & McVean (B, COLO); Goroka. Daulo Pass, Streimann 17989 (CBG). Morobe. Cromwell Mountains, Siwea, Koponen 30545 (Herb. Aptroot); Huon Peninsula, Mt Rawlinson, Hoogland 9315 (Herb. Aptroot, COLO); Mt Missim, Bellamy 203 (CBG); Ekuti Divide, Streimann 20121, 20168, 20187, 20212, 34148 (CBG); Wagau-Malolo Track, Streimann 19577 (CBG); Upper Watut River, Streimann 23137 (CBG); near Honzeu- kngon village, Aptroot 17998, 18041 (Herb. Aptroot); Raw- linson Range, Strong Clemens 12490 (COLO); Aiuwa-Bakia Track, Streimann & Tamba 12281 (CBG); Yinimba, Stre- imann 19710 (CBG); Mannasat, Hoogland 9466 (COLO). Central. Mt Albert-Edward, Kashiwadani 11748, 11770, 11816, 11982 (TNS). Western Highlands. Nebilyer River, Streimann 20599 (CBG); Mur Mur Pass, Streimann 21196, 22404 (CBG). Southern Highlands. Munie Logging Area, Streimann 23248, 23309, 23613, 23615 (CBG); Onim Forestry Station, Streimann 24639 (CBG); Enga. Mt Hagen-Wabag Road, Streimann 21256 (CBG). Solomon Islands: Guadalca- nal Island. Mt Popomansiu, Hill 9290, 9560, 9572, 9632, 9694, 9706 (BM); Mt Gallego, Hill 8172 (BM). Kolomban- gara Island. Ridge west of Kolombangara River, Hill 10686 (BM). Society Islands: Raiatea. Tetooroa, 200 m, Moore L26 (Herb. Aptroot). Fiji: Viti Levu. Mt Tomanivi [Victoria], Smith 5205a (BM); Mt Victoria, Selling (S); Lam 6831 (L); Fig. 4 Distribution of Pseudocyphellaria beccarii in the palaeotropics. 122 D.J. GALLOWAY Mt Tomanivi [Victoria], Smith 5205a (US); Ra, ridge from Mt Namama toward Mt Tomanivi, Smith 5712 (BM). Samoa: Upolu. Mountains near Tiave, Schultz-Motel 4350 (B). 4. Pseudocyphellaria carpoloma (Delise) Vain, in Hedwigia 37: 34 (1898). Sticta carpoloma Delise in Mem. Soc. linn. Normandie 2: 159 pi. 19, right hand figure (1825). Stictina carpoloma (Delise) Nyl., Syn. meth. lich. 1(2): 339 (1860). Saccardoa carpoloma (Delise) Trevis., Lichenotheca ven- eta exs. no. 75 (1869). Cyanisticta carpoloma (Delise) Gyeln. in Reprium Spec. nov. Regni veg. 29: 2 (1931). Type: New Zealand, 'Sur les vieux arbres a la Nouvelle Zelande', Bay of Islands, 1824, ?/?.P. Lesson (PC- LENORMAND-holotype). For additional synonymy see Galloway (1988: 80-85). Pseudocyphellaria carpoloma is bright lettuce-green to olive- green when wet, pale olivaceous-greenish when dry; it is a characteristic dichotomously branching, white-medulla, green-photobiont species with yellow pseudocyphellae below, which is widespread in New Zealand (Galloway, 1988) but has not been previously correctly identified from tropical regions. It is discussed in detail in Galloway (1988: 80-85). Earlier tropical records of P. carpoloma such as Magnusson (1940, 1956), Magnusson & Zahlbruckner (1943), Szatala (1956) refer to specimens of P. gilva, a cyanobacterial spe- cies. CHEMISTRY. Methyl evernate, tenuiorin, methyl gyrophor- ate, evernic acid (tr.), gyrophoric acid (tr.), hopane-7(3, 22-diol, hopane-6a, 7(3, 22-triol, 7fi-acetoxyhopane-6a, 22-diol (tr.), 6p-acetoxyhopane-7p, 22-triol (tr.), norstictic (tr.), stictic, cryptostictic, and constictic acids, pulvinic dilac- tone, pulvinic acid and calycin. OBSERVATIONS. Pseudocyphellaria carpoloma is character- ized by dichotomously branching lobes with entire margins and a distinctive, faveolate upper surface. It has a white medulla, a green photobiont and prominent yellow pseudocyphellae on the lower surface and projecting from the margins. Spores grey-brown, oval-ellipsoid, thickened 1-septate to 3-septate at maturity, (20-)22-25(-27) x 7-11 um. DISTRIBUTION AND ECOLOGY. In the palaeotropics known so far only from Papua New Guinea and Norfolk Island from humid forest at 900 and 2500 m (Fig. 5). It is probably more widespread in the palaeotropics and is need of further collec- tion. SPECIMENS EXAMINED. Papua New Guinea: Madang. Finis- terre Range. Teptep Village, Aptroot 31927 (Herb. Aptroot). Norfolk Island: Mt Bates, Henderson (E). 5. Pseudocyphellaria clathrata (De Not.) Malme in Ark. Bot. 26A(14): 9 (1934). Sticta clathrata De Not., Osserv. Sticta: 19 (1851). Crocodia clathrata (De Not.) Trevis., Lichenotheca veneta exs. no. 75 (1869). Type: Brazil, in sylvis insulae S. Sebastiano, 1839, Casaretto (BM- lectotype (Galloway & Arvidsson, 1990: 119)). For addi- tional synonymy see Galloway & Arvidsson (1990: 119). Pseudocyphellaria clathrata is bright lettuce-green with a yellow-gold tinge when wet, grey-green often suffused red- dish when dry or on storage; it is a characteristic rosette- forming to irregularly spreading, yellow-medulla species which is widespread in tropical regions (Galloway & Arvids- son, 1990; Galloway, 1993). Palaeotropical material exam- ined agrees in all respects with the anatomical and morphological details given in Galloway & Arvidsson (1990: 121-126). CHEMISTRY. Similar to that of P. arvidsonii and P. aurata (Galloway & Arvidsson, 1990) containing calycin, pulvinic Fig. 5 Distribution of Pseudocyphellaria carpoloma in the palaeotropics. STUDIES IN PSEUDOCYPHELLARIA (IV) lactone, pulvinic acid and a mixture of unidentified fernene triterpenoids. OBSERVATIONS. Pseudocyphellaria clathrata is a widely dis- tributed pantropical species having rather large, broadly rounded to subdichotomously or irregularly branching lobes with entire, non-sorediate, non-phyllidiate margins and is characterized by a yellow medulla, a green photobiont, yellow pseudocyphellae on the lower surface, a glabrous to partly pubescent or tomentose upper surface which is ± distinctly punctate-impressed, and distinctly pedicellate mar- ginal to submarginal apothecia. Apothecia distinctly pedicel- late, areolate-scabrid to white-tomentose, concolorous with thallus. Spores fusiform-ellipsoid, pointed at one or both ends, 3-septate at maturity, pale red-brown, (15.5-)18-20.5(-22.5) x 3.5-4.5 jjun. DISTRIBUTION AND ECOLOGY. Widespread in tropical regions of the world (Swinscow & Krog, 1988; Galloway & Arvidsson, 1990; Galloway, 1993) and recently collected in northern India by Dr K.P. Singh (Fig. 6). In humid montane forest, in canopy branches and main branches of shrubs and trees, rarely on rocks, 400-1600 m. SPECIMENS EXAMINED. Africa. Uganda: Kabale, Burnet 230 (BM); West Mengo, Lye L 196 (BM). Zimbabwe: sine loco, Sim (BM). Tanzania: Ngorongoro Crater, Pocs & Chuwa 89027/Z (BM); Usumbara Mountains, Amani, Moberg 1491b (UPS). Kenya: Ngong Hills, Meyink (BM); Mt Kenya, Swin- scow (BM); Kakamega District, Santesson 21764 (UPS); Kajiado District, Ngong Hills, Moberg 1413 (UPS). South Africa: Cape Province. Table Mountain, Sipman 20.186 (B); Disa Gorge, Table Mt, Pillans (BM). Angola: Golungo Alto, Welwitsch (BM). Madagascar: sine loco, Sykora, 1894 (WU); Imerina, Andrangoloaka, 1880, Hidebrandt 2156 (WU); Imerina, Wills (BM). Reunion: sine loco, Lepervanche Mezieres (M); southern slopes of Piton des Nieges, near 123 Cilaos, Arvidsson & Nilsson 2538 (GB); Cirque de Cilaos, de Sloover 17/472 (LG). Java: Mt Ardjuno, Groenhart 9769 (L). Tjibodas, Arvidsson & Nilsson 2494 (GB). Philippines: Luzon. Benguet, Bangio, Elmer 8991 (BM). Papua New Guinea: Eastern Highlands. Goroka, Lapegu, Streimann 18272, 18289 (CBG). Morobe. Mt Kaindi, Kashiwadani 10516 (TNS); Kauli Lake, Streimann 34079, 34092 (CBG); Yinimba, Streimann 19177 (CBG). Western Highlands. Baiyer River Sanctuary, Streimann 21105 (CBG); Kagamu- nga, Streimann 21301 (CBG); Minj, Streimann 21504 (CBG). New Caledonia: He des Pins. Tribu de Gadji, Hill 12099 (BM). Norfolk Island: sine loco, F. Bauer (W). 6. Pseudocyphellaria crocata (L.) Vain, in Hedwigia 37: 34 (1898). Lichen crocatus L., Mant. pi.: 310 (1771). Type: India, without specific location, Konig (LINN 1273.137- holotype). For additional synonymy see Galloway (1988: 113) and Galloway & Arvidsson (1990: 126). Sticta xanthosticta Pers. in Gaudich., Voy. Uranie: 201 (1827). Type: [Hawaii] in insulis Sandwicensibus, Gaud- ichaud (L 910.187.685-lectotype, selected here). Sticta crocata f. sandwicensis Zahlbr. in Rech., Denkschr. Akad. Wiss. Wien 88: 29 (1911). Cyanisticta sandwicensis (Zahlbr.) Gyeln. in Reprium Spec. nov. Regni veg. 29: 6 (1931). Type: Hawaii. Vulkan Kilauea, auf arten von Metrosideros , April 1905, N. Rechinger 2544 (W-lectotype, selected here). Pseudocyphellaria hawaiiensis H. Magn. in Acta Horti goth- oburg. 14: 21 (1940). Cyanisticta hawaiiensis (H. Magn.) Rasanen in Suomal. elain-ja kasvit. Seur. van. kasvit. Julk. 20(3): 17 (1944). Type: Hawaii. Molokai, between Upper Mountain Camp and Pepeopae, 9 July 1938, O. Selling 5842 (UPS-isotype). Cyanisticta hawaiiensis var. scrobiculata Rasanen in Suomal. elain-ja kasvit. Seur. van. kasvit. Julk. 20(3): 17 (1944). Fig. 6 Distribution of Pseudocyphellaria clathrata in the palaeotropics. 124 D.J. GALLOWAY Type: Tahiti. Sine loco, Vieillard (H-not seen). Cyanisticta hawaiiensis var. xanthocardia Rasanen in Suomal. elain-ja kasvit. Seur. van. kasvit. Julk. 20(3): 17 (1944). Type: Tahiti. Sine loco, Vieillard (H-not seen). Pseudocyphellaria crocata is dark slate-blue to blue-black or glaucous brownish often suffused red-brown when wet, pale olivaceous grey or blue grey to red-brown when dry; it is widespread in both tropical and temperate regions of the world and shows both throughout and within its range a considerable plasticity of form which has led to an extensive synonymy. Palaeotropical material is also very variable but within the range of anatomy and morphology recorded by Galloway (1988: 115-118). Hawaiian material recorded as P. hawaiiensis (Magnusson, 1940; Magnusson & Zahlbruckner, 1943) tends to have narrower, ± dichotomously branching lobes with the soralia often restricted to small, scattered, marginal clumps, but in a range of recently collected material (Prof. C.W. Smith, pers. comm.) a complete sequence from broad-lobed, laminally and marginally sorediate forms to narrow-lobed, ± dichotomously branching, sparsely margin- ally sorediate forms was observed, all of which fall within the species range of variation. CHEMISTRY. Tenuiorin, methyl gyrophorate, gyrophoric acid (tr.), hopane-6a, 7(3, 22-triol (major), 6a-acetoxyhopane-7p, 22-diol (tr.), 7(3-acetoxyhopane-6a, 22-diol (tr.), hopane-7fi, 22-diol (tr.), ± physciosporin (tr.) norstictic (tr.), stictic, cryptostictic, constictic and ± salazinic acids, pulvinic acid and pulvinic dilactone (Elix et al., 1992). OBSERVATIONS. Pseudocyphellaria crocata is characterized by a white medulla; a cyanobacterial photobiont; yellow laminal and marginal soralia on the faveolate to plane upper surface, and yellow pseudocyphellae on the lower surface. Apothecia are rather variable in occurrence varying from moderately common to rare or absent. Spores are broadly ellipsoid, smoky olive-brown to dark brown, thickened 1-septate to irregularly 3-septate, straight or slightly curved, 22.5-27(-29.5) x 7-9 u,m. It has a characteristic chemistry including hopane-6ot, 7(3, 22-triol, stictic acid metabolites, tenuiorin and methyl gyrophorate and yellow pigments. It is distinguished mainly by its soredia from other members of the P. cracato-group; P. gilva has neither soredia nor isidia or phyllidia; P. crocatoides has characteristic marginal lobules but no soredia or true isidia; P. neglecta has marginal and laminal phyllidia which may erode and become pseudosoredi- ate; while P. desfontainii has terete to subsquamiform isidia which never become sorediate. DISTRIBUTION AND ECOLOGY. Widely distributed in the palaeotropics and neotropics and in cool temperate zones of both Northern and Southern Hemispheres (Fig. 7). One of the most widely distributed species in the genus. It occurs in a wide variety of habitats from sea level to 4200 m (in Papua New Guinea) as an epiphyte of trees, shrubs, on rotting logs and on the forest floor, on both shaded and sunny rocks, and on soil in alpine grasslands. In the palaeotropics it occurs most commonly in humid, shaded woodlands, montane for- est, cloudforest and alpine grasslands. SPECIMENS EXAMINED. Africa. Tanzania: Kiliminjaro, Bigger 1966 (BM). Kenya: Mt Marsabit, Lye L660 (BM); Aberdare Mts, Swinscow (BM). Uganda: Usumbara Mts, Hoist 2665 (BM); Sasa River above Bulambuli, Lye L 501 (BM). South Africa: Cape of Good Hope. Table Mountain, Wedermann & Oberdieck 48 (B); ?Tafelberg, Wilms (B). Transvaal. Hou- bosdorp, Sipman 19.786 (B); Kowyns Pass near Graskop, Sipman 19.936 (B); Long Tom Pass, Sipman 20.093 (B). Madagascar: sine loco, Baron (BM). Reunion: Cirque de Cilaos, auf der Strasse zwischen Cilaos und Ilet a Cordes auf den Col du Taibit, K. & A. Kalb 26567 (Herb. Kalb); Piton de la Grande Montee, pres des sources Reihlac, de Sloover Fig. 7 Distribution of Pseudocyphellaria crocata in the palaeotropics. STUDIES IN PSEUDOCYPHELLARIA (IV) 17.257 (LG); SW du Piton Mare-a-Boue, de Sloover 17.299 (LG); Cirque de Cilaos, de Sloover 17.538, 17.577, 127.927 (LG). Mauritius: Pouce Mt, Ayres (BM); Curepipe, sine coll. (BM). Sri Lanka: Central Province, Thwaites C.L. 24 (BM). Malaysia: Pahang. Eraser's Hill, Burkill 2099 (L); Eraser's Hill, Galloway (KEP); Cameron Highlands, Tanak Tara, Degelius As-567 (UPS). Sabah. Mt Kinabalu, Sipman & Tan 31105, 31079, 31024 (B). Sarawak. Gunong Mulu National Park, Argent & Coppins 5116 (BM). Indonesia. Sulawesi: sine loco, Herb. Lugd. Batav. (L). Java: Tjibodas, Koernich 6a (Herb. Aptroot); Sipman & Zainal 30094 (B); Kawi Moun- tains. Mt Panderman, Groenhart 1934 (Herb. Aptroot); sine loco, Jelinek (B); Idgen Plateau, Kebun Djampit, Pos 7379 (B); sine loco, Junghuhn (L); Mt Pangerango, Schiffner 3288 (W); Cibodas Botanical Garden, Arvidsson & Nilsson 2488 (GB). Philippines: Luzon. Benguet, Mt Santo Tomas, Aptroot 20447, 20448, 20452, 20453 (Herb. Aptroot); Mt Pulog, Merrill 6458 (BM). Mindanao. Mt Apo, Copeland 1093 (BM). West Irian: Eipomek-Tal, Hiepko & Schultze- Motel 2019, 2277 (B); Carstensz Mts, Hope (COLO). Papua New Guinea: Morobe. Saruwaged, Sipman 24336, 24386, 24429, 24462 (B); Kaisinik, Kashiwadani 10448, 10686, 10745, 10760 (TNS); Mt Missin, Kashiwadani 10428 (TNS); Rawlinson Range, Strong Clemens 12444 (COLO); Araulu Logging Area, Streimann 13593, 13620 (CBG); Koke Village, Streimann & Tamba 11658, 11752 (CBG); Honzeukngon village, Aptroot 17826, 17928, 17930, 18019 (Herb. Aptroot); Ekuti Divide, Streimann 20164, 20355, 20362 (CBG); Herzog Mountains, Streimann & Umba 11113 (CBG); Spreader Divide, Streimann & Tamba 11895, 12073, 12208, 12211 (CBG); Slata Creek, Streimann 14045 (CBG); Herzog Moun- tains, Streimann & Umba 11134 (CBG); Yinimba, Streimann 19712 (CBG); head of Black Cat Creek, Streimann 25644 (CBG); Gumi Divide, Streimann 22764, 25729 (CBG); Madang. Finisterre Range, Teptep Village, Aptroot 31931, 32285, 32289, 32290, 32291 (Herb. Aptroot). Eastern High- lands. Chimbu. Mt Wilhelm, Borgmann 732b, 919 (B); Aptroot 18282, 18396, 18528, 18655, 32828, 32834 (Herb. Aptroot); Kashiwadani 10880, 10921, 10967, 10998, 11011, 11051, 11062, 11125, 11144, 11312, 11329, 11354 (TNS); McVean 66123, 66234, 66254 (CBG); Pindaunde Valley, Aptroot 31354, 32742, 33112 (Herb. Aptroot); Toromam- buno, Walker 8315 (CBG); Lake Piunde, Sipman 21985, 22132 (B); Goroka. Mt Zapaliga, 2650 m, Iserentant 9546 (Herb. Aptroot); Mt Gahavisuki Provincial Park, Sipman 22185 (B); near Mopei Village, Streimann 18844 (CBG); Daulo Pass, Streimann 18021, 18029, 18077 (CBG). Western Highlands. Yobobos, Hoogland & Schodde 7639 (B); Kubor Range. Nona River, Vink 16473 (Herb. Aptroot); Milep Area, Vinas 7644 (CBG); Jimi-Waghi Divide, Streimann 22317 (CBG); Kum Magnei Mtn, Streimann 20646-7 (CBG); Nebilyer River, Streimann 20597 (CBG); Kagamuga, Stre- imann 20429, 21751 (CBG); Central. Mt Albert-Edward, Kashiwadani 11501, 11758, 11936, 12000 (TNS); 2 km N. of the Waiotape Airstrip, Kashiwadani 11657, 12045, 12242 (TNS); Ehu Creek, Streimann & Naoni 16615 (CBG); Mt Victoria area, V.Roy en 10957 (CBG). Southern Highlands. Mt Giluwe, Streimann 24219 (CBG); Onim Forestry Station, Streimann 23592, 24562, 24614, 24631, 24635, 24638, 24640 (CBG); Enga. Mape Creek, Streimann 22112 (CBG); Gulf. Hepataewa, Streimann 33845 (CBG). Solomon Islands: Guadalcanal Island. Mt Gallego, Hill 8381 (BM); Mt Popomansiu, Hill 9379, 9458, 9625, 9725 (BM). New Cale- donia: Noumea. Mt Koghi, Hill 11509 (BM). Sarramea. Col 125 d'Amieu, Hill 11884, 11958 (BM). Norfolk Island: Selwyn Pine Road, Streimann 34663, 34661 (CBG); Mt Pitt, Stre- imann 34817 (CBG); track from Red Road to Mt Bates, Streimann 34444 (CBG). Fiji: Viti Levu. Nadarivatu. District Commisioners House, Degener 31814ae,u (B); Nandarivatu, Green (BM); Nadala, Degener 31807 (B); Mt Nairosa, Smith 4100, 4420 (BM). Samoa: Upolu. Rechinger (W); Tutuila, Reinecke 62a (BM). Tahiti: Fautaua Valley, Setchell & Parks 5442 (BM). Hawaiian Islands: Hawaii. Mauna Loa, Rock [Zahlbruckner: Lich. Rar. Exsic. 171] (BM, B, W); Kipuka Ki, Degener 31426 (B); Weber & Bujakiewicz (B); Waimea, [ad truncos muscosos in paludosis] Szatala [Lichenes Sandwi- censes] (B); Glenwood, Faurie 938 (BM); Saddle Road, Kipuka, Smith 4953 (Herb. Smith) Kauai. Sine loco, [on trees] Heller (B); Hanapepe River, Heller 2630a (BM); Mt Gay summer house, Faurie 297 (BM); E. of Kalalau Look- out, Wedin 3722 (UPS). Maui. Haleakala. Puu Uianiau, 7000 ft, Degener 22242 (B); Haleakala National Park, Kalua awa, Medieros (Herb. Smith): Oahu. Mt Kaala summit, Degener 30064 (B); Puu Hapapa near Kolekole Pass, Doty (B); Waianae Range, Kaala Natural Area Reserve, Wedin 3698 (UPS); Honouliuli Forest Reserve, Smith 4125 (Herb. Smith). Galapagos Islands: Isla Sant Cruz. Academy Bay, Weber (COLO); Horneman 4/64 (COLO). 7. Pseudocyphellaria crocatoides D.J. Galloway in Gr aphis Scr. 5: 8 (1993). Type: Fiji. Taveuni, Mt Utugatau, near summit, tree trunks in rainforest, c. 1140 m, 22 April 1970, G. Degelius P-236 (UPS-holotype; BM-isotype). Fig. 8. Thallus in irregular rosettes or loosely spreading, 4-7(-9) cm diam., loosely attached centrally, margins free and ± ascend- ing. Lobes narrow, 2-5 mm wide, rarely to 10 mm wide, irregularly to complexly branched, ± imbricate centrally, ± discrete at margins. Margins sinuous or ragged, dentate- incised to ± richly phyllidiate, slightly thickened below and occasionally also ridged above, occasionally with protruding yellow pseudocyphellae. Upper surface dark slate-blue to blue-black when wet, pale glaucous greyish when dry, undu- late, matt, smooth, wrinkled to subfaveolate especially at lobe apices, very fragile, friable when dry, pliable when wet. Phyllidia common and conspicuous, mainly marginal, very variable, ± dorsiventral, simple to coralloid-branched, 0.2-1 mm tall (to 3 mm long), fringing lobes with clustered, finger-like proliferations. Maculae occasional to frequent, white or pale yellowish, ± distinctly reticulate, following shallow ridges and in faveolae (x 10 lens). Isidia, pseudocyphellae and soredia absent. Medulla white. Photo- biont cyanobacterial. Lower surface pale yellowish white to buff-brown at margins darkening centrally, tomentum pale whitish to grey or buff, to brown-black centrally, rather sparse at margins, thick and entangled centrally. Pseudocyphellae yellow, low-conical, scattered, ± rounded, 0.1 mm diam. or less, margins only slightly raised, decorti- cate area flat to convex. Pycnidia rather sparse, solitary, scattered, at margins and along laminal ridges, ostiole red-brown, punctate-depressed, 0.1 mm diam. or less. Apothecia very rare, solitary, marginal, sessile, constricted at base, rounded, cupuliform, 1-1.5 mm diam., exciple pale buff or brownish, ± translucent when wet, coarsely corrugate-scabrid, with a conspicuous, irregularly dentate margin, disc red-brown, slightly roughened and with a thin 126 D.J. GALLOWAY imiimiiimiiimiimimimiiiii Fig. 8 Pseudocyphellaria crocatoides. Isotype (BM). Scale in mm. white pruina. Epithecium 10-14 u,m thick, red-brown. Hyme- nium colourless to pale straw, 70-80 u,m tall. Ascospores not seen. CHEMISTRY. Pulvinic acid, pulvinic dilactone, 6a-acetoxyhopane-7(3-22-diol (tr.), ?p-acetoxyhopane-6a, 2-diol (tr.), hopane-7p, 22-diol (tr.), hopane-6a, 7(3, 22-triol (major), tenuiorin, methyl gyrophorate, gyrophoric acid (tr.), ± physciosporin (tr.), norstictic (tr.), stictic, cryptostic- tic, constictic acids. OBSERVATIONS. P. crocatoides is a characteristic species of the P. crocata group and has a white medulla, a cyanobacte- rial photobiont, yellow pseudocyphellae on the lower surface and a chemistry containing stictic acid metabolites, hopane- 6a, 7p, 22-triol as the dominant triterpenoid, and the pig- ments calycin, pulvinic acid and pulvinic dilactone (Galloway & Kemp, 1993). It is distinguished by the distinctive marginal (rarely laminal) lobulate proliferations, and a smooth upper surface without isidia or soredia, characters which separate it from P. crocata which is yellow-sorediate; from P. dozy ana which is white-sorediate; from P. desfontainii which has terete to squamiform isidia; from P. neglecta which has phyllidia which erode to become pseudosorediate; and from P. gilva which has entire margins and is without soredia, isidia, phyllidia or lobulate proliferations. DISTRIBUTION AND ECOLOGY. Known from Fiji, Papua New Guinea (see below) and also eastern Australia (Fig. 9). Still very poorly collected. Palaeotropical collections so far seen are from humid, shaded, montane rainforest, 840-3000 m. SPECIMENS EXAMINED. Fiji: Taveuni. Mt Utuigatau, Degelius P-243 (UPS); Nandarivatu, Green (BM). Papua New Guinea: Morobe. Mt Kaindi, Streimann 17622 (CBG). Eastern High- lands. Mt Wilhelm, Kashiwadani 10866 (TNS). 8. Pseudocyphellaria desfontainii (Delise) Vain., Result. Voy. Belgica, Lich.: 29 (1903). Sticta desfontainii Delise in Mem. Soc. linn. Normandie 2: 60 pi. 4, fig. 12 (1825). Stictina carpoloma f. desfontainii (Delise) Nyl. in Hue, Nouv. Archs Mus. Hist. nat. Paris III, 2: 297 (1890). Sticta carpoloma f. desfontainii (Delise) Zahlbr., Cat. lich. univ. 3: 374 (1925). Cyanisticta desfontainii (Delise) Rasanen in Suomal. elain-ja kasvit. Seur. van. kasvit. Julk. 2(1): 42 (1932), non C.W. Dodge (Beih. nov. Hedwigia 12: 173 STUDIES IN PSEUDOCYPHELLARIA (IV) 127 Fig. 9 Distribution of Pseudocyphellaria crocatoldes. (1964)). Type: He de Bourbon, Bory de St-Vincent (PC- THURET-lectotype (Galloway & James, 1986: 434)). Fig. 10. Cyanisticta crocata var. tingaensis Sbarbaro in Arch. Bot. 15: 102 (1939). Type: Rarotonga, June 1929, H.E. Parks (Not seen). Pseudocyphellaria ceylonensis H. Magn. in Acta Horti goth- oburg. 14: 23 (1940). Type: Ceylon [Sri Lanka], Central Province, G.H.K. Thwaites26 (UPS-isotype). Cyanisticta mougeotiana ssp. dentata Rasanen in Suomal. elaln-ja kasvit. Seur. van. Julk. 20(3): 16 (1944). Type: New Caledonia, ad corticem arboris, 1863, E. Vieillard (H-not seen). Thallus rosette-forming to irregularly spreading, 4-9(-ll) cm diam., closely attached centrally, margins ± free. Lobes 2-8(-15) mm wide, (0.5)l-3(-6) cm long, subdichotomously branching to complex-imbricate, discrete, contiguous or sub- imbricate at margins, complex-imbricate centrally. Margins entire in parts (especially at lobe apices) to indented, ragged, incised, crenulate, slightly thickened and ridged below, sparsely to densely isidiate. Upper surface dark glaucous blue to blue-black, suffused red-brown towards apices when wet, glaucous grey or pale bluish to red-brown when dry, undu- late, shallowly pitted or punctate-impressed to deeply faveo- late, ridges sharply defined to smoothly rounded, matt to slightly shining in parts, flabby when wet, brittle and rather fragile when dry, isidia easily broken off leaving yellow scars, without soredia, phyllidia or pseudocyphellae. Maculae com- mon, minute, white or yellowish, effigurate to ± reticulate, best seen when wet (use x 10 lens). Isidia sparsely to densely developed, often crowded at margins also on interconnecting ridges or in faveolae, solitary to crowded in groups, terete, simple, rarely squamiform or coralloid, 0.5-1.5 mm tall and 0.1-0.2 mm diam., concolorous with upper surface, eroding at apices and appearing pseudosorediate, breaking off and leaving yellow scars resembling pseudocyphellae. Medulla white, often suffused yellow in upper parts in some collec- tions. Photobiont cyanobacterial. Lower surface pale yellow- white or buff at margins darkening to red-brown or brown- black centrally, wrinkled-undulate, tomentose from margins to centre or with a narrow, glabrous, marginal zone, tomen- tum thick and woolly, white to dark brown or ± blackened. Pseudocyphellae yellow, common, scattered, rounded, minute, 0.2 mm diam. or less, rarely 0.5 mm diam., conical- verruciform, decorticate area flat to convex. Pycnidia laminal, scattered, inconspicuous, slightly swol- len, ostiole minute, red-brown to black, 0.1 mm diam. or less. Apothecia sparse (often absent) to ± frequent, marginal and laminal 0.5-3 mm diam., sessile, constricted at base to subpedicellate, shallowly to deeply cupuliform, ± deeply concave to undulate and ± plane at maturity, exciple promi- nent, persistent, pale whitish pink, ± translucent when wet, brownish or red-brown when dry, coarsely corrugate-scabrid, obscuring disc at first, rupturing and leaving an irregular dentate margin, disc red-brown to ± blackened, grey-white pruinose. Epithecium red-brown, 8-14 jim thick. Hymenium colourless to pale straw to pale or dark red-brown, 80-100 jim tall. Ascospores red-brown, ellipsoid, apices pointed, 1-septate, 23-28(-30.5) x (5.5-)6.5-8.5(-ll) n-m. CHEMISTRY. Tenuioirin, methyl gyrophorate, gyrophoric acid (tr.), hopane-6a, 7(3, 22-triol, 7(3-acetoxyhopane-6a, 22-triol (tr.), 6a-acetoxyhopane-7p, 22-diol (tr.), stictic, cryp- tostictic, and constictic acids, pulvinic acid, pulvinic dilactone and calycin. OBSERVATIONS. Pseudocyphellaria desfontainii is character- ized by a white medulla; a cyanobacterial photobiont; terete, ± fingerlike isidia on the upper surface; yellow 128 D.J. GALLOWAY Fig. 10 Pseudocyphellaria desfontainii. T.G.A. Green s.n. (BM) Scale in mm. pseudocyphellae on the lower surface; and a chemistry con- taining yellow pigments, a dominant hopane-triol and metabolites of the stictic acid aggregate. Its characterisitic isidia distinguish it from other members of the P. crocata group (see above under P. crocata and P. crocatoides) . DISTRIBUTION AND ECOLOGY. A strictly palaeotropical taxon (Fig. 11), occurring from Africa to the south-western Pacific eastwards as far as the Marquesas, but not recorded from Hawaii, or the Galapagos Islands (Weber, 1986). On living and dead trees, on rotting logs and on shrubs in montane rainforest, 800-3650 m. SPECIMENS EXAMINED. Africa. Tanzania: Usambara. Amani, Brunnthaler (W). Madagascar: sine coll. (L-ex Herb Paris); Ambohimitombo Forest, Forsyth Major 469, 554, 576, 583, 587 (BM). Mauritius: sine loco, McGregor (BM); Les Mares, Ouhamed 8 (BM). Reunion: Cirque de Salazie, K. & A. Kalb 26562 (Herb. Kalb); zwischen le Brule (S von St-Denis) und Plaine des Chicots, K.& A. Kalb 26563 (Herb. Kalb); south- ern slopes of Piton des Nieges, near Cilaos, Arvidsson & Nilsson 2539 (GB). Sri Lanka: above Pattipola, Horton Plains, van Steenis 19924d (L); Mount Pedro, Blatter 56 (W); Nuwara Eliya. Horton Plains, Farr Inn, Moberg 2585, 2598 (UPS); Hakgala Botanical Garden, Lundqvist 9003 (UPS); Hakgalla Botanical Garden, Degelius As-411 (UPS); Nuwara Eliya, Degelius As-440 (UPS). Malaysia: Sabah. Kota Belud, Mt Kinabalu, Sipman & Tan 31104 (B). Pahang. Eraser's Hill, Burkill 2073b, 2796 (L); Dransfield 515 (BM); Degelius As-611, As-620 (UPS); Galloway (KEP); Cameron High- lands, Tanak Rata, Degelius As-568, As-576 (UPS). Indone- sia. Java: Mt Ardjuno, Groenhart 1860 (L); Mt Gede, van Ooststroom 14592 (L); Cibodas Botanical Garden, Arvidsson & Nilsson 2487, 2490, 2497 (GB). Sulawesi: Lompo Batang, Zelf 250 (L). Papua New Guinea: Eastern Highlands. Chimbu. Mt Wilhelm, Borgmann 779 (B); Streimann 18554 (CBG); Kashiwadani 10955, 11044, 11186 (TNS); Goroka. Mt Gahavisuki Provincial Park, Aptroot 31036 (Herb. Aptroot); Daulo Pass, Weber & McVean (COLO). Madang. Finisterre Range. Teptep Village, Aptroot 31926, 32287 (Herb. Aptroot). Morobe. Kewieng, Koponen 34404 (Herb. Aptroot); Mt Kaindi, Streimann 17523, 17568, 17602, 17614, 17621, 17663-4 (CBG); Kashiwadani 10548 (TNS); Weber & McVean (COLO); Upper Watut River, Streimann 17078 (CBG); Yinimba, Streimann 19061, 19711 (CBG); track to STUDIES IN PSEUDOCYPHELLARIA (IV) Mt Missim, Bellamy 206 (CBG); Pouyu Village, Streimann & Tamba 12675 (CBG); logging area 15 km E. of Bulolo, Streimann & Bellamy 13166 (CBG); Gumi Divide, Streimann 22774 (CBG); Wau, Edie Creek Road, Sipman 15627 (Herb. Aptroot); head of Black Cat Creek, Streimann 25653 (CBG). Central. 2 km N. of Waiotape Airstrip, Kashiwadani 12242, 12260 (TNS); Mt Albert-Edward, Kashiwadani 11719, 11803 (TNS). Southern Highlands. Margarima-Tari Road, Stre- imann 24394 (CBG); laro River, Streimann 23950 (CBG). Western Highlands. Karpena plantation N. of Mt Hagen, Streimann 21813, 21825 (CBG); Yobobos, Hoogland & Schodde 7640 (COLO). Solomon Islands: Guadalcanal Island. Mt Popomansiu, Hill 9439, 9491, 9575, 9849, 9883 (BM). New Caledonia: Sarramea. Col d'Amieu, Hill 12021 (BM). Fiji: Viti Levu. Nggaliwana Creek Valley, Smith 5335 (L); N-Baluti trail, Selling (S); Mba, Nandarivatu, Smith 5964 (BM); Nandarivatu, Green (BM). Taveuni. Mt Utugatau, Degelius P-205, P-230 (UPS). Samoa: Upolu. Lanuto'o, Rech- inger (W). Tahiti: Aorai, v.Balgooy (Herb. Aptroot); Ono- hea Valley, Degelius P-346 (UPS); Belvedere near Papeete, Degelius P-390 (UPS); sine loco, Moseley (BM). Marquesas Islands: Nukuhiva. Tovii, Peake (BM). 9. Pseudocyphellaria insculpta (Stizenb.) D.J. Galloway in Lichenologistll: 305 (1985). Stictina insculpta Stizenb. in Flora, Jena 81: 129 (1895). Stictina impressula Mull. Arg. in Flora, Jena 71: 22 (1888). non Nyl. (Flora, Jena 57: 71 (1874), based on Stictina tomentosa var. impressula Nyl. in Annls Sci. nat. (Bot.) V, 7: 305 (1867) from South America). Sticta insculpta (Stizenb.) Zahlbr., Cat. lich. univ. 3: 388 (1925). Type: Australia. Queensland, Mt Bellenden Ker, Sayer, comm. F. v. Mueller 1887 (G 0020099-holotype). Fig. 12. Stictina diplomorpha Mull. Arg. in Flora, Jena 65: 301 (1882). 129 Pseudocyphellaria diplomorpha (Mull. Arg.) D.J. Gallo- way, Tropical lichens: their systematics, conservation & ecology: 9 (1991). Type: Ceylon [Sri Lanka], sine loco, 1876, G.H.K. Thwaites (G 001975-holotype). Stictina impressula var. sublaevis Mull. Arg. in Hedwigia 30: 48 (1891). Stictina insculpta var. sublaevis (Mull. Arg.) Stizenb. in Flora, Jena 81: 129 (1895). Sticta insculpta var. sublaevis (Mull. Arg.) Zahlbr., Cat. lich. univ. 3: 388 (1925). Type: Australia. Queensland, Bellenden Ker Dis- trict, 1889, Bailey 575 pr.p. (G 002105-holotype). Thallus orbicular to irregularly spreading in entangled clones, 5-12(-20) cm diam., loosely attached centrally, margins and apices free, ± ascending. Lobes very variable, irregularly divided, subdichotomously branching at apices to complex- imbricate centrally, l-4(-8) mm wide, 0.5-1(^4) cm long. Margins irregularly notched or incised, occasionally to densely isidiate or lobulate-phyllidiate. Upper surface vivid navy blue to blue-black when wet, olivaceous-grey suffused brownish in parts when dry, irregularly undulate, conspicu- ously dimpled, punctate-impressed, here and there minutely papillate (use x 10 lens), rather fragile, papery when dry, flabby when wet, isidiate-phyllidiate, maculate, without pseudocyphellae or soredia. Maculae frequent, minute, white, effigurate to ± reticulate imparting a delicate marbling to the upper surface. Phyllidia mainly marginal, occasionally also laminal, simple to coralloid, terete to flattened- dorsiventral, constricted at base 0.2-0. 5(-l) mm wide, 1-2 mm tall. Isidia terete, subgranular at first, becoming flattened-phyllidiate. Medulla white. Photobiont cyanobacte- rial. Lower surface pale yellow-brown or whitish to buff brown at apices, darkening centrally, glabrous in a narrow to broad marginal zone and tomentose centrally, or uniformly tomentose from margins to centre, tomentum long, silky, white to grey-black or brown-black, densely entangled to ± felted- woolly. Pseudocyphellae prominent, white, round to Fig. 11 Distribution of Pseudocyphellaria desfontainii in the palaeotropics. 130 DJ. GALLOWAY irregular, 0.1-1 mm diam., margins raised, concolorous with lower surface, decorticate area flat to convex, sunk in tomen- tum. Apothecia rare or absent to occasional, rounded, l-2(-2.5) mm diam., sessile, constricted at base to ± sub- pedicellate, exciple pale pinkish brown, translucent when wet, wrinkled-striate, with occasional to dense white, silky tomentum below, disc plane to subconcave, red-brown, matt, smooth, epruinose. Epithedum pale yellow-brown, 8-12 u,m thick. Hymenium colourless, 70-85 jxm tall. Ascospores yellow-brown to red-brown 1-3-septate, ellipsoid-fusiform, apices rounded or pointed, 28-33.5 x 6.5-8.5 |xm. CHEMISTRY. Methyl gyrophorate, gyrophoric acid, 7p-acetoxyhopane-22-ol, hopane-7|3, 22-diol (tr.), hopane- 15a, 22-diol. OBSERVATIONS. Pseudocyphellaria insculpta is characterized by a white medulla; a cyanobacterial photobiont; projecting marginal lobules or elongate phyllidia; a dimpled to punctate- impressed upper surface; and a two-hopane chemistry with methyl gyrophorate and gyrophoric acid. It is distinguished from P. prolificans and P. multifida, both of which have green photobionts; from P. beccarii which has entire margins; from P. argyracea and P. dissimilis which have terete to coralloid isidia; and from P. crocatoides which has yellow pseudocyphellae. DISTRIBUTION AND ECOLOGY. A palaeotropical species known from Sri Lanka to the south-west Pacific (Fig. 13). An epiphyte of trees and shrubs in humid montane rainforest or cloud forest, often growing amongst moss, 300-3760 m. SPECIMENS EXAMINED. Sri Lanka: Big Mount, Neitner (US); Adams Peak, Thor 391 (S); Horton Plains, World's End, Bohlin (S); Central Province, Thwaites Cll, C22 (BM). Malaysia: Sabah. Kota Belud. Mt Kinabalu, 1650-3100 m, Siptnan & Tan 31075, 31383 (B). Indonesia. Sumatra: ?Gunung Leuser National Park, Assink s.n. (Herb. Aptroot). Java: Malang, Lederer s.n. (B); Tjibodas. Mt Cede, Schiffner 3079 (L); sine loco, Zollinger (L); Mt Ardjuno, Groenhart 7328 (L). Philippines: Luzon. Benguet. Mt Santo Tomas, Sipman 21819 (B); sine loco, Loher (BM). Papua New Guinea: Eastern Highlands. Chimbu. Mt Wilhelm, Borgmann 821 (B); Aptroot 32880 (Herb. Aptroot); Kashiwadani 10843, 10861, 10865, 10912, 10952, 11045, 11450 (TNS); Bundi Gap, Fig. 12 Pseudocyphellaria insculpta. G. Thor 391 (S). Scale in mm. STUDIES IN PSEUDOCYPHELLARIA (IV) 131 Fig. 13 Distribution of Pseudocyphellaria insculpta in the palaeotropics. Aptroot 32197 (Herb. Aptroot); Pindaunde Valley, Aptroot 32741 (Herb. Aptroot). Morobe. Cromwell Mountains. Siwea, Koponen 30489 (Herb. Aptroot); track to Mt Missim, Bellamy 210a,c (CBG); Kaisinik, Kashiwadani 10738 (TNS); Mt Kaindi, Streimann 33207 (CBG). Central. Mt Albert- Edward, Kashiwadani 11807 (TNS); 2 km N. of Waiotape Airstrip, Kashiwadani 12263 (TNS). Southern Highlands. Onim Forestry Station, Streimann 24627 (CBG). Solomon Islands: Guadalcanal Island. Mt Popomansiu, Hill 9289, 9315, 9383, 9432, 9435, 9495-6, 9512, 9559-60, 9667 (BM). Kolombangara Island. South Summit, Hill 10494 (BM). Fiji: Viti Levu. Mba, immediate vicinity of Nandarivatu, Smith 5964 (US). 10. Pseudocyphellaria dissimilis (Nyl.) D.J. Galloway & P. James in Lichenologist 12: 297 (1980). Stictina fragillima var. dissimilis Nyl., Syn. meth. lich. 1(2): 336 (1860). Stictina dissimilis (Nyl.) Nyl. in/. Linn. Soc. Lond. 9: 246 (1866). Sticta fragillima var. dissimilis (Nyl). Kremp., Reise Ost Freg. Novara Bot. 1: 119 (1870). Cyanisticta dissimilis (Nyl.) Rasanen in J. Jap. Bot. 16: 143 (1940). Type: Australia. Sine loco, Hampe (H-NYL 34103- lectotype (Galloway & James, 1980: 297)). Pseudocyphellaria dissimilis is dark slate-blue to blue-black when wet, pale greyish to buff when dry; it is a characteristic isidiate palaeotropical species with a white medulla and white pseudocyphellae on the lower surface which is described in detail in Galloway (1988: 122-126). CHEMISTRY. Gyrophoric acid (tr.), 7(3-acetoxyhopane-22-ol, hopane-7p, 22-diol (tr.), hopane-15a, 22-diol. OBSERVATIONS. Pseudocyphellaria dissimilis is a highly vari- able palaeotropical species having linear-elongate to shal- lowly rounded, subdichotomously to irregularly branched lobes, often ± canaliculate and with entire margins becoming isidiate or occasionally also phyllidiate. The upper surface is smooth or shallowly wrinkled, glossy or matt, minutely maculate and papillate (x 10 lens), and with laminal and marginal isidia, and/or phyllidia. Isidia are simple, ± terete at first and later may become coralloid-branched or flattened and ± phyllidiate. It has a white medulla, a cyanobacterial photobiont, and a pale to brownish often ± costate lower surface with rather sparse, short central tomentum and scattered, fleck-like pseudocyphellae. Spores are pale yellow- brown, 1-3-septate, straight or slightly curved, apices rounded or pointed, 20.5-29.5 x 7-9 |xm. It has a basic two-hopane chemistry (Wilkins, 1993) with or without traces of gyrophoric acid. Its physiology is discussed in Green et al. (1991) and Lange et al. (1993). It is distinguished from P. insculpta in lacking a punctate-impressed upper surface; from P. argyracea in lacking laminal pseudocyphellae associated with isidia; from P. desfontainii which has yellow pseudocyphellae and a different chemistry; and from P. crocatoides which has marginal and laminal proliferations, yellow pseudocyphellae and a different chemistry. DISTRIBUTION AND ECOLOGY. Apparently rather rare in the palaeotropics where it has to date been positively identified only from Papua New Guinea and Norfolk Island collections from montane rainforest. It is common in north-eastern Australia and throughout New Zealand (Galloway, 1988) and is also recorded from Juan Fernandez but not from continen- tal South America (Galloway, 1992). SPECIMENS EXAMINED. Papua New Guinea: Eastern High- lands. Goroka. Mt Zapaliga, Iserentant 9534 (Herb. Aptroot). Norfolk Island: sine loco, sine coll. (MEL). 11. Pseudocyphellaria dozyana (Mont. & Bosch) D.J. Gallo- way in Lichenologist 17: 304 (1985). Sticta dozyana Mont. & Bosch, Syll. gen. sp. crypt.: 326 (1856). Stictina dozyana (Mont. & Bosch) Nyl., Syn. meth. lich. 1(2): 335 (1860). 132 D.J. GALLOWAY Saccardoa dozyana (Mont. & Bosch) Trevis., Lichenoth- eca veneta exs. no. 75 (1869). Type: Java, without specific locality or date of collection, Junghuhn (L 910,215-1471- lectotype (Galloway & Arvidsson, 1990: 128)). NOTE. Material in PC-HUE is similar in all respects to the Leiden specimen, both are fertile having young, immature, marginal fruits and obviously represent parts of the same collection. The Paris material has a printed label 'Herbarium R.B. van den Bosch'and is further labelled in van den Bosch's hand 'Sticta intricata Del. Java Junghuhn', to which Mon- tagne has added 'Sticta Dozyana M. et V.d.B.'. Stictina mougeotiana var. albocyphellata Nyl., Syn. meth. lich. 1(2): 341. 1860. Type: Ins Borbonia [Reunion], without specific locality, collector or date (H-NYL 33995- holotype). Pseudocyphellaria dozyana belongs to the P. crocata group (white medulla, cyanobacterial photobiont and a chemistry dominated by stictic acid metabolites and hopane-6ot, 7(3, 22-triol) and has a punctate-impressed to faveolate upper surface with characteristic white marginal soralia with a pale yellow tinge to the exposed medulla beneath the white soralia. CHEMISTRY. Tenuiorin, methyl gyrophorate, hopane-6a, 7(3, 22-triol, stictic, constictic, cryptostictic acids and traces of unidentified triterpenes. OBSERVATIONS. Pseudocyphellaria dozyana is dark slate- blue to blue-black when wet, pale glaucous-greyish when dry; it has a white medulla, a cyanobacterial photobiont, white pseudocyphellae on the lower surface (rarely these may be pale yellowish at margins), and conspicuous, sinuous, mar- ginal soralia containing grey- white, granular soredia often eroding faint yellowish below. Lobes are broadly elongate- laciniate and are conspicuously punctate-impressed to faveo- late. The broadly elongate-laciniate lobes and the prominent punctate-impressed to faveolate upper surface, the white marginal soralia and the mainly white pseudocyphellae distin- guish this species from P. crocata which has prominent yellow soralia and yellow pseudocyphellae, and from P. bartlettii which has broadly rounded, ± rosette-forming lobes with reticulate soralia. P. intricata has a smooth upper surface with scattered laminal and marginal soralia which never erode yellowish below, and its chemistry is different, having two hopane-diol triterpenoids and lacking the stictic acid complex of metabolites. Montagne & van den Bosch's record of Sticta granulata from Java (Montagne & van den Bosch, 1857: 437-438) evidently refers to P. dozyana (Groenhart, 1936). DISTRIBUTION AND ECOLOGY. A palaeotropical species rang- ing from the Uluguru Mountains in East Africa and Madagas- car and Reunion in the Indian Ocean to the Galapagos Islands (Weber, 1993) in the eastern Pacific (Fig. 14). Known also from Ecuador (Galloway & Arvidsson, 1990). On living and dead trees and shrubs in humid primary and secondary rainforest, 550-2000 m. SPECIMENS EXAMINED. Africa. Tanzania: Uluguru Moun- tains, Mindu Hill near Morogoro, Pocs & Ochyra 88102/AO (BM). Madagascar: near Aulananarin, Pool (BM). Reunion: Cirque de Cilaos: Aufsteig von der Strasse zwischen Cilaos und Ilet a Cordes aud den Col du Taibit, K. & A. Kalb 26565, 26566 (Herb. Kalb); Piton des Neiges, 1500 m, Arvidsson & Nilsson (GB). Indonesia. Sulawesi. Sine loco, Herb. Lugd. Batav. (L); Monado, Quindal (M). Java. Sine loco, Jung- huhn s.n. (L); Mt Merbabu, Surjanto 1608 (L); sine loco, Junghuhn (L); Mt Gede, Schiffner 1149 (L); sine loco, sine coll. (H-NYL 34070, 34072). Papua New Guinea: Eastern Highlands. Goroka. Lutheran Guesthouse, Sipman 22324 (B). Central. Near Dabamura, 40 km NE of Port Moresby, Streimann & Naoni 14957 (CBG). Morobe. Herzog Moun- tain, Streimann & Umba 11050 (CBG). Western Highlands. Fig. 14 Distribution of Pseudocyphellaria dozyana in the palaeotropics. STUDIES IN PSEUDOCYPHELLARIA (IV) 133 Baiyer River Sanctuary, Streimann 21042 (CBG); Kagamuga, Streimann 24787 (CBG). Fiji: Viti Levu. Mba, slopes of Mt Nairosa, eastern flank of Mt Evans Range, Smith 4100 (US). Galapagos Islands: Isla Santa Cruz. Table Mt, Weber 288 (COLO). Isla Santiago. Above James Bay, Pike 2732 (COLO). Isla Charles. Weber 443 (COLO); trail from Black Beach to highlands, Weber & Lamer (COLO). 12. Pseudocyphellaria gilva (Ach.) Malme in Bih. K. svenska Vetenskakad. Hand!. 25(3/6): 32 (1899). Lichen gilvus Ach., Lichenogr. suec. prod.: 157 (1799) ['1798']. Sticta gilva (Ach.) Ach., Methodus: 278 (1803). Sticta crocata var. gilva (Ach.) Ach., Syn. meth. lich.: 232 (1814). Stictina gilva (Ach.) Nyl., Syn. meth. lich. 1(2): 339 (1860). Saccardoa gilva (Ach.) Trevis., Lichenotheca Ven- eta exs. 75 (1869). Cyanisticta gilva (Ach.) Gyeln. in Reprium Spec. nov. Regni veg. 29: 5 (1931). Type: [South Africa] Cap.b.Spei, Thunberg (UPS-THUNBERG 26816- lectotype (Galloway, 1992: 130)). Cyanisticta gilva var. lanata (Pers.) Gyeln. in Reprium Spec, nov. Regni veg. 29: 5 (1931). Collema lanata Pers. in Gaudich., Voy. Uranie: 204 (1827). Type: In insulis Maclovianis [Falkland Is], Gaudichaud (?PC-not seen). Cyanisticta gilva var. pseudogilva Gylen. in Reprium Spec, nov. Regni veg. 29: 6 (1931). Type: South Africa 'Promon- torium Bonae Spei', Gueinzius (B-holotype). Cyanisticta gilva var. philippiana Gyeln. in Reprium Spec, nov. Regni veg. 29: 6 (1931). Type: Philippines. Luzon, Benguet Subprovince, May 1911, E.D. Merrill 7962 (B-holotype; B-isotype). Pseudocyphellaria lombokensis H. Magn. in Acta Horti goth- oburg. 14: 26 (1940). Type: [Java] East India. Lombok, Goenoeng Rindjani, 1925, T.A. Tengvall (?UPSV-not seen). Pseudocyphellaria gilva described originally from Table Mountain at the Cape of Good Hope in the eighteenth century (Galloway, 1992) is a palaeotropical species in the P. crocata complex of taxa, having a white medulla, a cyanobac- terial photobiont, yellow pseudocyphellae on the lower sur- face and hopane-6a, 7(3, 22-triol as the principal terpenoid present in the medulla. Spores are red-brown, 1-3-septate, fusiform-ellipsoid, apices pointed, 22-30 x 9-11 u,m. The species is described in detail in Galloway (1992: 130-135). CHEMISTRY. Tenuiorin, methyl gyrophorate, hopane-6a, 7(5, 22-triol, stictic, cryptostictic, constictic, salazinic (tr.) and norstictic (tr.) acids, calycin, pulvinic acid and pulvinic dilac- tone. OBSERVATIONS. Pseudocyphellaria gilva is livid slate-blue suffused red-brown in parts when wet, pale grey-brown, olivaceous-brown to brown-black when dry; it has a white medulla; a cyanobacterial photobiont; irregularly branching to imbricate lobes with ± subdichotomously branching api- ces, entire margins often with conspicuous, yellow pseudocyphellae; a glossy, undulate to subfaveolate upper surface, lacking soredia, isidia, maculae or phyllidia; a dark red-brown to black lower surface with conspicuous, conical- verruciform, yellow pseudocyphellae; apothecia are rare to frequent, young fruits with a characteristic red-ochre margin to the disc which may sometimes be slightly grey-pruinose. It shows a wide range of variation throughout its range (Gallo- way, 1992: 133). It is distinguished from P. crocata by lacking soredia; from P. crocatoides in the thicker, darker thallus and the absence of marginal proliferations; from P. desfontainii in the absence of isidia; from P. beccarii in having a cyanobacte- rial photobiont, yellow pseudocyphellae and a different chemistry. The palaeotropical taxon with which it has been confused (see Magnusson, 1940), P. carpoloma, has a green photobiont, much more divergent, dichotomously branching lobes and a different chemistry (Code D of Wilkins & James (1979)). DISTRIBUTION AND ECOLOGY. A palaeotropical species (Fig. 15) ranging from South Africa (the type locality is Table Mountain) through the south-west Pacific tropics to southern South America (Galloway, 1992). On trees and shrubs, amongst mosses and overgrowing rocks in humid montane forest or cloud forest, 250-3600 m. Also in eastern Australia from Queensland to Tasmania. SPECIMENS EXAMINED. Africa. South Africa: Kirstenbosch, Almborn [Lichenes africani 9] (BM, L); Table Mt, Garside (L); Sipman 20.165, 20.194 (B); Eaton (BM); McGillivray (BM); Simon's Bay, Wright (BM); Cape, Drege (BM). Mau- ritius: sine loco, Blackburn (BM). Malaysia: Sabah. Mt Kinabalu, Samsudin (UKMB). Indonesia. Java: sine loco. Horsfield (BM); Mt Ardjuno, Groenhart 26, 32, 42, 1857, 1858, 1871 (L); Mt Kawi/Mt Panderman, Groenhart 1825, 1956, 7262, 7263, 7264, (L); Mt Merbabu, Surjemto 1612 (L); Tjemorokandang, Groenhart 7261 (L). Philippines: Luzon. Benguet, Pauai, 2100 m, McGregor 8528 (B); Merrill 7962, 7972 (BM); Mt Santo Tomas, Sipman 21777d (B); Aptroot 20358 (Herb. Aptroot); Degelius As-854, As-876 (UPS). Mindanao. Mt Apo, Copeland 1089, 1092 (B, MEL). Papua New Guinea: Eastern Highlands. Chimbu. Mt Wilhelm, Weber & McVean (B, COLO); Aptroot 18211, 32828 (Herb. Aptroot); Borgmann 756, 934, 936 (B); Kashiwadani 10883, 10916, 10965, 10975, 10980, 10997, 11095, 11127, 11291, 11400, 11404, 11428, 11467 (TNS); McVean 6699, 66189 (CBG); Wade 8065 (COLO); Weber [Lich. Exs. 373] (BM); Pindaunde Valley, Sipman 15908, 21988 (B); Bundi Gap, Aptroot 32550); Kombugomambuno, Mundua 139 (CBG); 2 km N. of Chimbu Airstrip, Kashiwadani 12432 (TNS); Lake Aunde, Aptroot 18462, 18477 (Herb. Aptroot); Goroka. Mt Gahavisuki Provincial Park, 2400m, Aptroot 31029, 31140 (Herb. Aptroot); Streimann 18204 (CBG); Sipman 22193 (B); Daulo Pass, Hoffmann 89-441 (Herb. Aptroot); Stre- imann 17968, 18080, 18116 (CBG); Wopeia. Near Aiyura, Streimann 18328 (B); track to Mt Michael, Streimann 18475 (CBG); near Hogabi Village, Streimann & Bellamy 18687 (CBG); Kassam Pass, Streimann & Umba 11427-8, 11504 (CBG). Morobe. Saruwaged, Sipman 24385 (B); Mt Sarawa- ket [Saruwaged] Southern Range, Koponen 32731 (Herb. Aptroot); Hekwangi Village, Streimann 19362 (B); track to Mt Missim, Bellamy 210a,c (CBG); Upper Watut River, Streimann 17179, 17239 (CBG); Mt Kaindi, Streimann & Bellamy 17675, 19875 (CBG); Streimann 22495, 22510 (CBG); Yakwoi River, Streimann 19261 (CBG); Rawlinson Range, Strong Clemens 12444 (COLO); Pouyu Village, Stre- imann & Tamba 12712 (CBG); Ekuti Divide, Streimann 20173, 24932 (CBG); head of Black Cat Creek, Streimann 25643, 25646-7 (CBG); Logging Area 15 km W. of Bulolo, Streimann & Bellamy 13142 (CBG); near Hunzeukngon Village, Aptroot 18023 (Herb. Aptroot); Gumi Divide, Stre- imann 25062 (CBG). Western Highlands. Mt Karoma, Veld- kamp & Wiakabu (Herb. Aptroot); Baiyer River Sanctuary, 134 D.J. GALLOWAY Fig. 15 Distribution of Pseudocyphellaria gilva in the palaeotropics. Streimann 21116 (CBG); Tumbang Village, Streimann 21361 (CBG); Mur Mur Pass, Streimann 21174 (CBG). Madang. Finisterre Range, Teptep Village, Aptroot 31982, 31992, 32292, 32294 (Herb. Aptroot). Southern Highlands. Lai River, Streimann 22213, 22216 (CBG); laro River, Streimann 23827, 23837 (CBG); Munia Logging Area, Streimann 23320, 23325, 23666 (CBG); Lama Sawmill Logging Area, Stre- imann 24690 (CBG); Paunde Logging Area, Streimann 23346, 23354 (CBG). Enga. Mape Creek, Streimann 21555 (CBG). 13. Pseudocyphellaria godeffroyii (Kremp.) D.J. Galloway in Lichenologist 17: 304 (1985). Sticta (Stictina) godeffroyi Kremp. in J. Mus. Godeffroy 1(4): 99, tab. 14 fig. 10 (1874). Type: Fiji, Viti Levu, Noggara, Dr E. Graffe 67 (M-lectotype (Galloway, 1985: 304)). Fig. 16. Stictina intricata var. gymnoloma Nyl., Syn. meth. lich. 1(2): 335 (1860). Type: Fidji insulae, Milne (H-NYL 34090- lectotype (Galloway, 1985: 304)). Thallus rosette-forming, wide-spreading, 5-12(-25) cm diam., closely attached centrally, margins ± free. Lobes broadly rounded, 8-15(-20) mm diam., contiguous to over- lapping at margins, imbricate centrally. Margins entire, sinu- ous, shallowly to deeply notched, slightly thickened. Upper surface dark glaucous blue to dark malachite green-blue, suffused brownish at apices when wet, pale buff brown or red-brown to pale cinnamon brown when dry, undulate, irregularly pitted or shallowly wrinkled in places, minutely roughened to verrucose-scabrid, scabrosity best seen at lobe apices (use x 10 lens), apices minutely white-tomentose, coriaceous, tough when dry, pliable, flabby when wet, pseudocyphellate, without isidia, phyllidia or soredia. Pseudocyphellae white, round to irregular, scattered, occa- sional to frequent, 0.2-1.5 mm diam., large prominent pseudocyphellae ± ulcerose with a raised margin, decorticate area flat to concave. Medulla white. Photobiont cyanobacte- rial. Lower surface pale yellowish buff or brownish at mar- gins, darkening centrally, uniformly short velvety tomentose, tomentum very even, pale buff to dark red-brown. Pseudocyphellae white, prominent, scattered, often crowded, minute, at margins 0.1-0.3 mm diam., larger centrally, round to irregular, 1-2 mm diam., margins very shallowly raised, concolorous with lower cortex, decorticate area flat to con- cave, distinctly granular- roughened. Pycnidia prominent, solitary or crowded, marginal and laminal, raised, to 0.5 mm diam., hemispherical, ostiole red-brown, 0.1-0.2 mm diam. Apothecia sparse to frequent, often crowded at centre of thallus, rare at margins and lobe apices, sessile, constricted at base, round to irregular-deformed through mutual pressure, 1-3 mm diam., exciple prominent, persistent, pale brownish, conspicuously verrucose-scabrid forming a distinctive corrugate-scabrid margin to disc, disc concave to plane, pale to dark red-brown, shining, epruinose. Epithecium pale red- brown, 14-22 (Jim thick. Hymenium colourless to pale straw, 80-95 u-m tall. Ascospores 1-3-septate, pale yellow-brown to red-brown, fusiform-ellipsoid, apices rounded or pointed, straight or curved, 28-33.5 x (5.5-)6.5-8.5(-ll) |xm. CHEMISTRY. 7p-acetoxyhopane-22-ol, (tr.), hopane-15a, 22-diol. hopane-7(3, 22-diol OBSERVATIONS. Pseudocyphellaria godeffroyii is character- ized by a white medulla, white pseudocyphellae on both upper and lower surfaces, a cyanobacterial photobiont, and broad, rounded lobes with a distinctive-scabrid-areolate upper surface. It is distinguished from P. punctillaris which has a scabrid-areolate upper surface and isidia or lobules at the margins; from P. rigida which has a scrobiculate upper STUDIES IN PSEUDOCYPHELLAR1A (IV) 135 Fig. 16 Pseudocyphellaria godeffroyii. T.G.A. Green s.n. (BM). Scale in mm. surface (not areolate-scabrid); from P. semilanata which has a smooth, not areolate-scabrid upper surface; and from P. trichophora which has a smooth upper surface and distinctive tomentose-hairy lobe margins. DISTRIBUTION AND ECOLOGY. Apparently restricted to Fiji where it occurs on trees in open slopes and on trees and scrub in humid, montane rainforest, 700-1100 m. SPECIMENS EXAMINED. Fiji: Viti Levu. Sine loco, Milne (BM); Mba, Nandarivatu, Smith 5963 (BM); Nandarivatu, Green (BM); Naggarra, Graeffe 64, 69 (M). 14. Pseudocyphellaria haywardiorum D.J. Galloway in Bull. Br. Mus. nat. Hist. (Bot.) 17: 159 (1988). Type: New Zealand. North Island, South Auckland, Red Mercury Island, on tea tree (Leptospermum) bark, August 1971, B. W. & G.C. Hayward H 40.4 (AK 161261-holotype). Pseudocyphellaria hawardiorum is a palaeotropical species of rather limited distribution in the South Pacific. It is discussed in detail in Galloway (1988: 159-162) and in Elix et al. (1992). CHEMISTRY. 73-acetoxyhopane-22-ol, (tr.), hopane-15a, 22-diol. hopane-7p, 22-diol OBSERVATIONS. Pseudocyphellaria haywardiorum is a palaeotropical sorediate species having ± rounded to irregu- larly laciniate lobes with coarsely granular to pseudoisidiate (x 10 lens) laminal and marginal soralia, and a conspicuously punctate-impressed upper surface. Neither surface depres- sions nor soralia are arranged in a reticulate pattern. It has a white medulla, a cyanobacterial photobiont, and prominent, large, white pseudocyphellae on the lower surface well delim- ited from the densely and evenly red-brown to brown-black tomentum. Thalli are dark grey-blue to blue-black when moist, olive brown to yellow-grey when dry. Apothecia very rare, generally absent. Spores fusiform-ellipsoid, apices pointed, yellow-brown, 1 -septate (27.5-)30-32(-34) x 6-7 jim. It has a simple two-hopane chemistry. P. haywardiorum is distinguished from P. intricata by the punctate-impressed upper surface and the ± bullate lower surface with its prominent, large, raised pseudocyphellae, characters which also separate it from the isidiate species P. argyracea. It is separated from P. dozy ana by the punctate- 136 D.J. GALLOWAY impressed upper surface and the chemistry (P. dozyana is in the P. crocata group and has stictic acid metabolites and a hopane triol, and not just the two hopane-diols of P. haywar- diomtri). DISTRIBUTION AND ECOLOGY. In the region known to date only from Norfolk Island where it is rare, occurring on Araucaria heterophylla and a tree fern stem (Elix et al., 1992). Known also from northern New Zealand and eastern Australia (Queensland and New South Wales). 15. Pseudocyphellaria homalosticta Vain, in Philipp. J. Sci. Sect. C, Bot. 8: 117 (1913). Type: Phillipines. Luzon. Prov. Rizal. Ad truncos arborum, February 1911, M. Ramos, Forest Bureau 13453 (TUR-VAINIO 10317- holotype). Fig. 17. Pseudocyphellaria amphistictoides Vain, in Univ. Calif. Publs Bot. 12: 6 (1924). Type: Tahiti. Fautaua Valley, May 1922, W.A. Setchell & H.E. Parks 5443 (BM-isotype). Thallus 3-10(-14) cm diam., irregularly spreading, loosely attached centrally, margins and apices ascending. Lobes (l-)2-6(-8) mm diam., subdichotomously to intricately branched. Margins entire at apices, soon becoming isidiate- raas.vi Fig. 17 Pseudocyphellaria homalosticta. Holotype (TUR-VAINIO 10317). Scale in mm. STUDIES IN PSEUDOCYPHELLARIA (IV) phyllidiate or proliferating into long, narrow lobules, slightly thickened, ridged below, scattered white pseudocyphellae present, sometimes appearing ± sorediate. Upper surface bright lettuce green when wet, pale olive green or buff brownish when dry, brittle, fragile, easily damaged when dry, pliable when wet, maculae and soredia absent. Isidia com- mon, very variable, terete, simple at first becoming 1-3- branched to coralloid, 0.1 mm diam., to 5 mm long, becoming dorsiventral and phyllidiate, primarily marginal, rarely developing from margins of laminal pseudocyphellae. Phyllidia developing from terete isidia or intermixed and independent of them, dorsiventral, with minute pseudocyphellae below, elongate, to 5 mm long. Pseudocyphellae white, scattered, punctiform, 0.1 mm diam. or less, ± flat, occasionally with isidia developing from margins. Medulla white. Photobiont green. Lower surface pale whitish buff at margins, darkening centrally, occasion- ally ± blackened at centre, glabrous, matt or glossy from margins to centre, or with scattered, thin tomentum centrally. Pseudocyphellae white, prominent, widely scattered, conical- verruciform, rounded, 0.1-0.3 mm diam., margins not promi- nent. Pycnidia solitary, ± marginal, hemispherical, 0.1 mm diam. or less, ostiole punctate, dark red-brown. Apothecia rare, marginal, rounded, 0.5-1.5 mm diam., subconvex to plane, sessile, constricted at base, exciple minutely corrugate-scabrid, persisting as verrucose margin to disc, pale buff or pinkish, translucent when wet, disc pale to dark red-brown, smooth, epruinose. Epithecium pale yellow- brown, 8-12 |xm thick. Hymenium colourless, 70-85 jxm tall. Ascospores pale yellow-brown, 1-septate, fusiform-ellipsoid, apices rounded or pointed, 25-28 x 6.5-8 u,m. CHEMISTRY. Methyl gyrophorate (tr.), ± gyrophoric and congyrophoric acids, 7p-acetoxyhopane-22-ol, hopane-7(3, 22-diol (tr.), hopane-15a, 22-diol. 137 OBSERVATIONS. Pseudocyphellaria homalosticta has a white medulla, a green photobiont and white pseudocyphellae on both upper and lower surfaces and characteristically at the margins of lobes where they can sometimes appear ± soredi- ate. It has marginal and occasionally laminal terete isidia which may become dorsiventral flattened phyllidia. It has a basic two-hopane chemistry, with or without accessory dep- sides. It is distinguished from P. prolificans which has mar- ginal and laminal phyllidia and lobules, and a punctate- impressed upper surface which is without pseudocyphellae; from P. multifida which has a smooth upper surface without pseudocyphellae and simple to squamiform phyllidia; and from P. reineckeana which has entire margins without phyl- lidia or isidia. DISTRIBUTION AND ECOLOGY. A species endemic to the south-west Pacific where it occurs from Fiji eastwards to the Marquesas Islands (Fig. 18). It is an epiphyte of trees and shrubs in dense, montane rainforest, 900-1200 m. SPECIMENS EXAMINED. Fiji: Viti Levu. Nandarivatu, Asplund s.n. (BM, Herb. L. Arvidsson); Degener 31811 (Herb. Aptroot); Mt Nanggaranambuluta [Lomalangi], Smith 4818 (BM, L); Smith 4833 (BM); ridge between Mt Nanggaranam- buluta and Mt Namama east of Nandarivatu, Smith 5009 (L); Mt Victoria, Lam 6824 (L); Green (BM). Ovalu. Sine loco, Graff e (W). Rarotonga: Tiriora, Parks 22395 (COLO); sine loco, Parks & Parks 22363a (COLO). Marquesas Is: Ua Pu. Jones 1178 (Herb. Aptroot). Nuku Hiva. Tovii, Peake (BM). 16. Pseudocyphellaria intricata (Delise) Vain, in Hedwigia 37: 35 (1898). Sticta intricata Delise in Mem. Soc. linn. Normandie 2: 96 pi. 7 fig. 33 (1825). Stictina intricata (Delise) Nyl., Syn. meth. lich. 1(2): 334 (1860). Cyanisticta intricata (Delise) Gyeln. in Lilloa 3: 76 (1938). Type: He de Bourbon [Reunion], Bory de St-Vincent (PC- Fig. 18 Distribution of Pseudocyphellaria homalosticta. 138 D.J. GALLOWAY LENORMAND-lectotype (Galloway & James, 1986: 437 (1986)). For additional synonymy see Galloway & James (1986: 437) and Galloway (1988: 169). Sticta dolera Hue in Nouv. Archs Mus. Hist. Nat. Paris IV, 3: 98 (1901). Type: [Reunion] Ins. Bourbon, sine loco, Lepervanche-Mezieres (PC-HUE 769-lectotype, selected here). Cyanisticta philippinica Gyeln. in Reprium Spec. nov. Regni veg. 29: 298 (1931). Type: Philippines, Luzon, Prov. Benguet, Pauai, 2100 m, R.C. McGregor (D-not seen). Pseudocyphellaria intricata is a widespread cosmopolitan spe- cies having a wide range of variation and paralleling the diversity of morphology seen in P. crocata. For a detailed description of the species see Galloway (1988: 169-174). CHEMISTRY. Tenuiorin (tr.), methyl gyrophorate (tr.), 7p-acetoxyhopane-22-ol, hopane-73, 22-diol (tr.), hopane- 15a, 22-diol. OBSERVATIONS. Pseudocyphellaria intricata is dark slate- blue to blue black when wet, pale greyish to buff when dry; it has irregularly laciniate to somewhat rounded lobes, with sinuous, incised or ± entire, generally sorediate margins. It has a white medulla, a cyanobacterial photobiont, a pale buff to brown, tomentose lower surface with occasional, scattered, white pseudocyphellae and a ± shining upper surface which may have scattered, erose, white to brownish laminal soralia. It has a characteristic, basic two-hopane chemistry (Gallo- way, 1988; Wilkins, 1993). It is distinguished from P. haywar- diorum by its smooth upper surface and its lower surface which is not bullate; from P. argyracea which has terete isidia associated with the laminal pseudocyphellae; and from P. dozy ana which has a faveolate upper surface and hopane-6a, 73, 22-triol as a major metabolite. DISTRIBUTION AND ECOLOGY. Widespread throughout the tropics and also in cool temperate regions of the world (Galloway, 1988, 1992; Galloway & Arvidsson, 1990). In the palaeotropics (Fig. 19) it occurs in humid, shaded woodland and montane forest and cloud forest from 400 to 2000 m (to 3600 m in New Guinea). It is nowhere a common species. SPECIMENS EXAMINED. Africa. Kenya: Mt Kenya east side, Themwe, Swinscow (BM); 2 km west of Irangi Forest Sta- tion, Swinscow (BM). South Africa: Smith's Peak, Leighton 942 (L); Knysna, Almborn [Lichenes africani 10] (L); Werd- ermann & Oberdieck 920 (B); Table Mountain, Sipman 20.189 (B); 7-800 m, [on soil] Werdermann & Oberdieck 49, 51 (B); between Devils Peak and Table Mountain, Wilms (B). Madagascar: Amboluimiloimbo Forest, Forsyth Major 543 (BM). Reunion: Piton de la Grand Montee, pres des sources Reihlac, de Sloover 17.258 (LG). Sri Lanka: Horton Plains, World's End, Bohlin (S). Malaysia: Pahang. Fraser's Hill, Burkill 2073 (L); Dransfield 514 (BM); Cameron High- lands, Bowen 4090 (E). Sabah. Mt Kinabalu, Samsudin (UKMB). Indonesia. Java: Herb. Lugd. Batav. (L); Mt Ardjunoi, Groenhart 1855, 1982 (L); Mt Panderman, Groen- hart 1954 (L); Mt Lawu, Clason 985 (L); Mt Gede, van Ootstroom 145900 (L). Phillipines: Luzon. Benguet, Merrill 7952 (BM); Mt Santo Tomas, Aptroot 20454, 20450, 20451 (Herb. Aptroot); Sipman 21812 (B). Papua New Guinea: Morobe. Saruwaged Range, Sipman 24330, 24337, 24387 (B); Kaisinik, Kashiwadani 10743 (TNS); Wau, Mt Kaindi, Kashi- wadani 10588, 10593 (TNS); Streimann 34024 (CBG); near Honzeukngon village, Aptroot 17931-2, 18025 (Herb. Aptroot); Gumi Divide, Streimann 22760, 22769 (CBG); Koke Village, Streimann & Tamba 11730 (CBG); Manki Trig, Streimann & Bellamy 12969 (CBG). Eastern Highlands. Wopeia, Streimann 18328a (B); Chimbu. Pindaunde Valley, Aptroot 32740 (Herb. Aptroot); Weber & McVean (COLO); Lake Piunde, Sipman 22121 (B); Mt Wilhelm, Kashiwadani 10846, 10885, 10998, 11199, 11417 (TNS); Aptroot 18243 Fig. 19 Distribution of Pseudocyphellaria intricata in the palaeotropics. STUDIES IN PSEUDOCYPHELLARIA (IV) (Herb. Aptroot); McVean 66115 (CBG); Imbuka Ridge above Lake Aunde, Weber & McVean (COLO): Yagle Village, Kawagle 2 (CBG); Goroka, Daulo Pass, Aptroot 31660 (Herb. Aptroot); Mt Gahavisuki Nature Reserve, Aptroot 18802, 18842 (Herb. Aptroot); 1500 m, Streimann & Kairo 18155 (CBG). Madang. Finisterre Range. Teptep Village, Aptroot 31931, 32288, 32295 (Herb. Aptroot). Cen- tral. 2km N. of Waiotape Airstrip, Kashiwadani 11559, 11653, 12244 (TNS); Mt Albert-Edward, Kashiwadani 11776 (TNS); Varirata National Park, Streimann & Vinas 14472 (CBG). Southern Highlands. laro River, Streimann 23823 (CBG). Western Highlands. Kagamuga, Streimann 21712 (CBG). Enga. Mape Creek, Streimann 21569, 22112 (CBG). New Caledonia: Tinchialit Camp, Cheeseman (BM); sine loco, Compton (BM). Norfolk Island: Mt Pitt Reserve, Mt Bates, Streimann 34386, 34331 (CBG). Fiji: sine loco, Wilson (MEL). Samoa: Upolu. Schultz-Motel 3496 (B). Tahiti: sine loco, Jelinek 53 (W). Hawaiian Islands: Oahu. Koolau Moun- tains, ridge from Tantalus to Puu Konahuanui, Smith 130as (Herb. Smith). 17. Pseudocyphellaria maculata D.J. Galloway in Bull. Br. Mus. nat. Hist. (Bot.) 17: 187 (1988). Type: New Zealand. South Island, Nelson, Maruia River, Speargrass Flat, near Springs Junction, on twigs of wayside shrubs, 22 Septem- ber 1981, D.J. Galloway (CHR 381022-holotype; BM-isotype). Pseudocyphellaria maculata is a member of the P. crocata complex of taxa characterized by a white medulla, a cyano- bacterial photobiont and yellow pseudocyphellae on the lower surface. The species is described in detail in Galloway (1988: 187-191) and material examined from New Guinea agrees in all respects with New Zealand collections from which the species was described. CHEMISTRY. Methyl evernate (tr.), tenuiorin, methyl lecan- orate (tr.), methyl gyrophorate, evernic acid (tr.), gyrophoric acid (tr.), hopane-6a, 7(3, 22-triol, norstictic (tr.), salazinic, consalazinic, galbinic acids, pulvinic dilactone, pulvinic acid and calycin. OBSERVATIONS. Pseudocyphellaria maculata is dark slate- blue to brown black, suffused red-brown when wet, olivaceous-grey to red-brown when dry; it has a white medulla, a cyanobacterial photobiont, a conspicuously faveo- late upper surface with a well-defined reticulate pattern of white (photobiont-free) maculae best seen when the thallus is wet (x 10 lens), and yellow pseudocyphellae on the lower surface and visible at the lobe margins. It lacks isidia, soredia, phyllidia, pseudocyphellae or tomentum on the upper sur- face. Its loose, straggling habit (especially in alpine grassland habitats where it is often best-developed) is also characteris- tic. It is rarely fertile. P. maculata is distinguished from P. gilva by its thinner more fragile and papery thallus, by its markedly faveolate upper surface with characteristic sharp, reticulate ridges, and the pale to buff silkily white-tomentose lower surface; from P. crocata by the absence of soredia; from P. crocatoides by the lack of marginal proliferations; from P. desfontainii by the absence of isidia; and from P. neglecta by the absence of phyllidia. DISTRIBUTION AND ECOLOGY. An epiphyte of trees and shrubs in montane rainforest and high alpine grassland habi- tats of high humidity in New Guinea, 1200-3810 m. Also in 139 New Zealand where it occurs in rainforest and alpine grass- land habitats (Galloway, 1988). SPECIMENS EXAMINED. Irian Jaya: Carstenz Mountains, Hope (COLO). Papua New Guinea: Eastern Highlands. Chimbu, Pindaunde Valley, Stone 9903 (Herb. Aptroot); Weber & McVean (Herb. Aptroot); Aptroot 32732 (Herb. Aptroot); Mt Wilhelm, Kashiwadani 11000, 11087, 11128, 11335, 11354 (TNS); McVean 66179 (CBG); Goroka, Daulo Pass, Streimann & Kairo 18138 (CBG); Mt Gahavisuki Nature Reserve, Aptroot 18803 (Herb. Aptroot). Morobe. Mt Sarawaket [Saruwaged] Southern Range, Koponen 32164, 32640 (Herb. Aptroot); track to Mt Missim, Bellamy 211 (CBG); Wau, Mt Kaindi, Kashiwadani 10537 (TNS); Mt Missim, Kashiwadani 10412 (TNS); Slate Creek and Gumi Creek Divide, Streimann 13978 (CBG); Araulu Logging Area, Streimann 13619 (CBG). Central. 2 km N. of Waiotape Airstrip, Kashiwadani 12257 (TNS). Southern Highlands. Margarima-Tari Road, Streimann 24380 (CBG). Western Highlands. Yobobos, Hoogland & Schodde 7640 (COLO). Enga. Mape Creek, Streimann 21540 (CBG). 18. Pseudocyphellaria multifida (Nyl.) D.J. Galloway & P. James in Lichenologist 12: 301 (1980). Sticta multifida Nyl., Syn. meth. lich. 1(2): 363 (1860). Sticta dissecta Laurer in Linnaea 2: 41 (1827), non S. dissecta (Sw.) Ach. (Meth. Lich.: 279 (1803)). Crocodia multifida (Nyl.) Tre- vis., Lichenotheca veneta exs. 75 (1869). Lobaria multifida (Nyl.) Hellb. in Bih. K. svenska VetenskAkad. Handl. 21(2/13): 38 (1896). Type: Nov. Holland [Australia], Sie- ber 45 (BM-lectotype (Galloway, 1988: 199)). For addi- tional synonymy see Galloway (1988: 199-200). Pseudocyphellaria multifida is a palaeotropical taxon with a highly plastic morphology and having a white medulla and pseudocyphellae, a green algal photobiont and a basic two- hopane chemistry. It is discussed in detail in Galloway (1988: 199-204). CHEMISTRY. 7p-acetoxyhopane-22-ol, hopane-7(3, 22-diol (tr.), hopane-15a, 22-diol. OBSERVATIONS. Pseudocyphellaria multifida is bright lettuce-green when wet, pale greenish grey to buff brownish when dry; it has very variable, rather delicate lobes ranging from ± broadly rounded to more usually narrow and highly divided, entangled-imbricate, the margins ragged-incised to lobulate or richly phyllidiate. The upper surface is smooth, undulate or shallowly wrinkled, punctate-impressed, with occasional, white maculae towards margins, and occasionally to densely developed, simple, squamiform, palmate-coralloid to ± strap-like phyllidia. It has a white medulla, a green photobiont, and a pale whitish, glabrous, glossy, smooth or shallowly wrinkled lower surface, with a usually poorly developed, thin, short, velvety tomentum centrally, and with scattered, white, fleck-like pseudocyphellae most noticeable at margins. Apothecia are rare. It has a basic two-hopane chemistry. It is distinguished from P. prolif icons by its plane or undulate, not distinctively punctate-impressed upper sur- face, by the unthickened, ± naked margins of the lower surface and by the frequently ± squamiform phyllidia; from P. insculpta which has a cyanobacterial photobiont and a punctate-impressed upper surface; and from P. homalosticta which has pseudocyphellae on the upper surface which occa- sionally become ± sorediate. 140 D.J. GALLOWAY Fig. 20 Distribution of Pseudocyphellaria multifida in the palaeotropics. DISTRIBUTION AND ECOLOGY. From Sabah and Java east- wards to New Guinea (Fig. 20) and southwards to Australasia where it is common throughout New Zealand (Galloway, 1988) and in Tasmania (Kantvilas, 1988). In the palaeotropics it is an epiphyte of trees and shrubs and overgrows rotting logs in humid montane rainforest, 1000-2900 m. SPECIMENS EXAMINED. Malaysia: Sabah. Mt Kinabalu, Sam- sudin (UKMB). Indonesia. Java: East Java. Mt Tengger, Groenhart 260, 1835, 7265 (L); Mt Kawi, Groenhart 7248, 7249, 7256, 7257, 7258 (L); Mt Ardjuno, Groenhart 1534, 1867, 6254 (L); Mt Andjasmoro, Groenhart 1817, 1874, 1875, 7230 (L); West Java. Tjibodas, Mt Gede, van Ootstroom 14207 (L); ibid., Groenhart 1801 (L); Batavia. Mt Pantjar, Schiffner 2978 (WU); Preanger, Tjibodas Schiffner (WU). Papua New Guinea: Morobe. Mt Kaindi, Streimann & Bel- lamy 17670 (CBG); Kauwara River, Kairo 671 (CBG); Koke Village, Streimann & Tamba 11665 (CBG); Kaurau, Kairo 379 (CBG); Spreader Divide, Streimann & Tamba 11951 (CBG); Honzeukngon village, Aptroot 17861 (Herb. Aptroot). Eastern Highlands. Chimbu. Mt Wilhelm, Kashi- wadani 11190 (TNS); Goroka. Mt Gahavisuki Nature Reserve, Aptroot 18778, 18804, 18829 (Herb. Aptroot). Southern Highlands. Onim Forestry Station, Streimann 24667 (CBG). Enga. Mape Creek, Streimann 22084 (CBG). 19. Pseudocyphellaria neglecta (Mull. Arg.) H. Magn. in A eta Horti gothoburg. 14: 30 (1940). Stictina neglecta Mull. Arg. in Flora, Jena 70: 58 (1887). Type: Australia, New England, sine collectoribus nomine (G002121- holotype). For additional synonymy see Galloway (1988: 207; 1992: 183). Pseudocyphellaria neglecta is a characteristic phyllidiate spe- cies in the P. crocata complex and is discussed in detail in Galloway (1988: 207-210). CHEMISTRY. Calycin, pulvinic dilactone (tr.), pulvinic acid, tenuiorin, methyl gyrophorate, stictic, constictic, norstictic (tr.), cryptostictic (tr.), salazinic (tr.) acids, 6a-acetoxyhopane-7(3, 22-diol (minor), 7(3-acetoxyhopane- 6a, 22-diol (tr.), hopane-6a, 7(3, 22-triol (major), hopane-7(3, 22-diol (minor), hopane-15a, 22-diol (tr.), 7(3-acetoxyhopane-22-ol (tr.), 15ot-acetoxyhopane-22-ol (tr.), retigeranic acid (minor) and traces of unidentified triterpe- noids. OBSERVATIONS. Pseudocyphellaria neglecta is lead-grey to dark blue-black or suffused red-brown when wet, olivaceous- brown, red-brown, brownish yellow or reddish to ± black- ened in exposed habitats when dry; it has linear-elongate to broadly rounded lobes with entire to crenate-incised to densely phyllidiate margins. The upper surface is undulate, wrinkled to subfaveolate, occasionally with squamiform phyl- lidia regenerating from cracks or scattered over upper sur- face, often eroding apically and appearing sorediate, or breaking off and leaving small yellow scars like pseudocyphellae. It has a white medulla, a cyanobacterial photobiont and yellow pseudocyphellae on the lower surface. It has a complex chemistry containing pigments, depsides, depsidones and hopane-6a, 7(3, 22-triol as the major triterpe- noid. For differences between P. crocata, P. crocatoides, P. gilva and P. dozyana see above under these taxa. DISTRIBUTION AND ECOLOGY. Widespread in the South Pacific from Mt Kinbalu (Sipman, 1993) eastwards to New Guinea and Tahiti (Fig. 21). It is also known from Australia (where it is extremely common and the most widely collected species), New Zealand (Galloway, 1988) and Chile (Gallo- way, 1992) where it tends to favour rather dry sites with high light intensities. In the palaeotropics it is found in more humid situations; on roadside banks, on rocks, stumps, fallen branches and rotting logs on the forest floor and as an STUDIES IN PSEUDOCYPHELLARIA (IV) 141 Fig. 21 Distribution of Pseudocyphellaria neglecta in the palaeotropics. epiphyte of trees and shrubs in montane forest and mossy cloudforest, 700-3500 m. SPECIMENS EXAMINED. Malaysia: Sabah. Mt Kinabalu, Sip- man & Tan 30961 (B); Samsudin (UKMB). Indonesia: Sumatra. Mt Horintji, Meyer 7791 (L). Philippines: Luzon. Benguet. Mt Santo Tomas, Sipman 21750 (B). Papua New Guinea: Central. Mt Victoria area, Iswan Swamp, van Roy en 10957 (Herb. Aptroot). Madang. Finisterre Range, Teptep Village, Aptroot 30955 (Herb. Aptroot). Morobe. Mt Kaindi, Weber & McVean (Herb. Aptroot, COLO); Streimann 19908, 33171, 34072, 34078 (CBG); Herzog Mountains, Streimann & Umba 11130 (CBG); Near Honzeukngon village, Aptroot 18019 (Herb. Aptroot); Gumi Divide, Streimann 25731 (CBG); Aiuwa-Bakia Track, Streimann & Tamba 12347 (CBG); Eraulu Logging Area, Kairo 338 (CBG); Mt Missim Track, Streimann 22923 (CBG). Eastern Highlands. Goroka. Daulo Pass, Weber & McVean (COLO); Mt Gahavisuki Nature Reserve, Aptroot 18806 (Herb. Aptroot); Chimbu. Mt Wilhelm area, Aptroot 18321, 18407, 18601, 18654 (Herb. Aptroot). Southern Highlands. Onim Forestry station, Stre- imann 24559 (CBG). Enga. Mape Creek, Streimann 21535, 21554 (CBG). Solomon Islands: Guadalcanal Island. Mt Popomansiu, Hill 9558 (BM). Tahiti: Aorai, v.Balgooy 1864a (Herb. Aptroot). 20. Pseudocyphellaria pickeringii (Tuck.) D.J. Galloway in Bull. Br. Mus. nat. Hist. (Bot.) 17: 218 (1988). Sticta pickeringii Tuck., U.S. Expl. Exped. 17 (Bot.): 138 (1874). Type: New Zealand, Bay of Islands, Wilkes Expe- dition, sine collectoribus nomine (FH-holotype). Pseudocyphellaria flavicans auct., non (Hook.f. & Taylor) Vain. For additional synonymy see Galloway (1988: 218-219). medulla, isidiate, widespread palaeotropical lichen which is discussed in detail in Galloway (1988: 218-224). CHEMISTRY. Pulvinic acid, pulvinic dilactone, calycin, 2a, 3^, 22a-triacetoxystictane, 2a, 3(3-diacetoxystictane-22-ol, stictane-3p-22a-diol, 2a-acetoxystictane-3(3, 22a-diol, 3(3-ace- toxystictane-2a,22a-diol, stictane-2a, 3(3, 22a-triol, 3(3, 22a- diacetoxystictane, 2a, 3(3-diacetoxystictane-22-ol, 3(3-aceto- xystictane-22-ol, pseudocyphellarin A, isopseudocyphell- arin A, nephroarctin, phenarctin, 2'-O-methylphenarctin, l'-chlorophenarctin, 2'-O-methylisopseudocyphellarin A, 2'-O-methylpseudocyphellarin A (Elix et al., 1992). OBSERVATIONS. Pseudocyphellaria pickeringii is bright lettuce-green suffused golden-yellow when wet, pale lemon- yellow to golden yellow when dry; it is a palaeotropical species forming irregular rosettes on bark, rocks and soil. It has variable, ± rounded to complexly divided, rather ragged, incised lobes whose margins are ± isidiate-phyllidiate. The upper surface is coriaceous, smooth to faveolate in parts to ± scabrid-areolate (x 10 lens), with marginal and laminal terete isidia and flattened, ± dorsiventral phyllidia present. It has a yellow medulla, a green photobiont, and the lower surface is pale yellow to red-brown with a velvety pale tomentum and scattered, yellow, often inapparent pseudocyphellae. Apoth- ecia are sparse or absent, though occasionally frequent in some specimens, sessile to subpedicellate with a conspicuous, whitish buff, coarsely verrucose-scabrid exciple. Spores are colourless, l-3(-5)-septate, fusiform-ellipsoid, 25-29. 5(-32) x 6.5-7 (jun. It has a complex chemistry of pigments and stictane triterpenoids (Elix et al., 1992; Wilkins, 1993). It is distinct from P. aurata which has characteristic marginal, labriform yellow soralia; from P. dathrata which has entire margins without either isidia, phyllidia or soredia; and from P. poculifera which has fragile, marginal sorediate isidia. Pseudocyphellaria pickeringii is a characteristic yellow- DISTRIBUTION AND ECOLOGY. Widespread in the palaeotro- 142 D.J. GALLOWAY pics from the Philippines eastwards to Hawaii and the Mar- quesas (Fig. 22) and common in the South Pacific in New Zealand (Galloway, 1988) and Australia. On living and dead branches of trees and shrubs (often in canopy branches) in humid montane rainforest and in open habitats of high light, 200-2700 m. SPECIMENS EXAMINED. Philippines: Luzon. Benguet, Baguio, Williams 1636 (B); Merrill 7953, 7956 (BM); Ramos 13510 (BM); Mt Santo Tomas, Degelius as-822 (UPS). Papua New Guinea: Morobe. Saruwaged Range, Sipman 24428 (B); Yinimba, Streimann 19025 (CBG); Mt Kaindi, Kashiwadani 10524, 10567 (TNS); near Honzeukngon village, Aptroot 18021 (Herb. Aptroot); Gumi Divide, Streimann 22771 (CBG). Madang. Finisterre Range, Teptep Village, Aptroot 31938, 32007, 32021, 32293 (Herb. Aptroot). Eastern High- lands. Goroka, Daulo Pass, Streimann 18020 (CBG); Mt Gahavisuki, Lambley 100/84, 102/84 (BM); Kassam Pass, Streimann & Umba 11476 (CBG). Central. Mt Albert- Edward, Kashiwadani 11793 (TNS); Myola, Lambley 92/85 (BM). Western Highlands. Baiyer-Jimi Divide, McVean 68181 (CBG). New Caledonia: Sarramea. Col d'Amieu, Hill 11959, 11982 (BM); sine loco, Compton 1300 (BM). Norfolk Island: Mt Pitt Reserve, Streimann 31931, 31944 (CBG); track from Mt Bates, Streimann 34252 (CBG); Selwyn Pine Road, Hoogland 6587 (L). Samoa: Tutuila, Lutisal (B). Hawaiian Islands: Oahu. Pauoa, Heller (BM, L, US); Waianae Mountains, Mokuleia Forest Reserve, Smith 3151 (Herb. Smith); Koolau Mountains, Kahuku Forest Reserve, Smith 1660 (Herb. Smith); Honolulu, Faurie 441 (BM). Kauai. Hanapepe Valley, Heller (BM, L); Faurie 87 (BM). Maui. South slope of Haleakala, Auwahi, James & Smith 84/2 (BM); Haleakala, Faurie 591, 592 (BM). Marquesas Islands: Nukuhiva. Tovii, Peake (BM). 21. Pseudocyphellaria poculifera (Mull. Arg.) D.J. Galloway & P. James in Lichenologist 12: 301 (1980). Sticta pocu- lifera Mull. Arg. in Flora, Jena 65: 304 (1882). Type: Lord Howe Island, Mt Gower, F. v. Mueller (G 002123- holotype; BM, MEL-isotypes). This characteristic rosette-forming to irregularly spreading, yellow-medulla species is described in detail in Galloway (1988: 224-228) and in Elix et al. (1992: 71-72). CHEMISTRY. Pulvinic acid, pulvinic dilactone, calycin, 3p-acetoxyfern-9(ll)-en-12-one, fern-9(ll)en-3p, 12p-ol, 3p-hydroxyfern-9(ll)-en-12-one, 3p-acetoxyfern-9(ll)-en- 12p-ol, 3p-acetoxyfern-9(ll)-en-19p-ol and unidentified trit- erpenoids (Elix et al., 1992). OBSERVATIONS. Pseudocyphellaria poculifera is bright lettuce-green suffused yellow-gold when wet, pale green-grey when dry; it is a palaeotropical species characterized by a yellow medulla, a green photobiont, yellow pseudocyphellae on the lower surface, greenish yellow, mainly marginal (occa- sionally also laminal), densely clustered, minutely coralloid, rather delicate isidia which are ± corticate at first, but soon erode and become sorediate. It is quite commonly fertile, the apothecia being marginal or submarginal and distinctly pedi- cellate and with granular isidiate margins. Spores are pale to dark red-brown, 3-septate, broadly fusiform-ellipsoid, (18-)20-23(-25) x 5.5-7.5 u,m. It has a complex chemistry of pigments and fernene triterpenoids (Elix et al., 1992). It is readily distinguished from the related P. aurata which has characteristic linear-labriform marginal soralia; and from P. pickeringii in the nature of the isidia, the structure of the exciple, the attachment of the apothecia, and in the size and colour of the spores. DISTRIBUTION AND ECOLOGY. A palaeotropical species (Fig. 23) which is known from East Africa (where it is extremely rare) and from Peninsular Malaysia and the Philippines Fig. 22 Distribution of Pseudocyphellaria pickeringii in the palaeotropics. STUDIES IN PSEUDOCYPHELLARIA (IV) eastwards to Fiji and Norfolk Island where it is common (Elix et al., 1992) and southwards to eastern Australia and north- ern New Zealand (Galloway, 1988). It occurs on bark and twigs of a variety of phorophytes in open conditions, occa- sionally on rocks also. It has an altitudinal range from sea level to 750 m throughout most of its range; the sole East African specimen seen was from 2100 m. SPECIMENS EXAMINED. Africa. Uganda: Butandiga Bugishu, A.S.T. 2555 (BM). Malaysia: Pahang. Eraser's Hill, Gallo- way (KEP). Indonesia. Java: Near Wonosobo, Oka 4087 (L). Papua New Guinea: Morobe. Kasu Village, Kairo 563, 573 (CBG). New Caledonia: Sarramea. Col d'Amieu, Hill 11872, 11883 (BM). Norfolk Island: Cascade, Ralston (BM, COLO); Prince Phillip Drive, Streimann 36480 (CBG); Capt. Cook Monument, Streimann 32045 (CBG); Mt Pitt Reserve, track to Hollow Pine, Streimann 31994 (CBG); Mt Pitt Road, Streimann 31943, 31923 (CBG); King Fern Valley, Streimann 34552 (CBG); Mt Pitt, Streimann 34814 (CBG); track at end of Sellwyn Pine Road, Streimann 34648, 34655 (CBG); off Selwyn Pine Road, Filmy Fern Trail, Streimann 32106, 32086, 32159 (CBG); Mt Bates, Streimann 34228 (CBG); Bird Rock Track, Streimann 34899 (CBG); track from Mt Bates, Stre- imann 34300 (CBG); summit of Mt Bates, Ralston 90b (BM); east side Mt Bates, Hoogland 11.157 (BM); north side Mt Bates, Green 1424 (BM); Now-now Valley, Hoogland 11.257 (BM); 'High ground', sine loco, Milne (BM). Fiji: Viti Levu. Nandarivatu, Smith 5965 (L); sine loco, Milne (BM). 22. Pseudocyphellaria prolificans (Nyl.) Vain, in Philipp. J. Sci. sect. C, Bot. 8: 117 (1913). Sticta prolificans Nyl. in Annls Sci. Nat. Bot. IV, 15: 42 (1861). Crocodia prolifi- cans (Nyl.) Trevis., Lichenotheca veneta exs. 75 (1869). Type: New Caledonia, ad cortices sylvarum in Kanala, Vieillard 1795 (PC-holotype). Fig. 24. 143 Pseudocyphellaria multipartita Vain, in Philipp. J. Sci. Sect. C, Bot. 8: 116 (1913). Type: Philippines, Luzon, Batangas Prov. Ad truncos arborum et supra muscos, November 1907, H.M. Curran & M.L. Merritt Forest Bureau 7809 (TUR-VAINO 10291-lectotype, selected here). Thallus very variable, orbicular to irregularly spreading, often forming densely entangled clones, 5-12(-18) cm diam., loosely to closely attached centrally, margins and apices free, ± ascending. Lobes very variable, most commonly rather narrow, linear-laciniate, rather ragged, (l-)2-5(-8) mm wide, l-4(-8) cm long, to occasionally broadly rounded, 8-15 mm wide, subdichotomously branching at or near api- ces, complex-imbricate, entangled centrally, apices truncate, rounded or ± furcate, divergent. Margins entire towards apices becoming densely phyllidiate towards centre, notice- ably thickened above and below, pseudocyphellae often prominent on lower margin. Upper surface lettuce-green to olive-green when wet, undulate, irregularly to markedly dimpled, punctate-impressed, often with dense to widely scattered papillae (use x 10 lens), thin and papery to tough, coriaceous, without isidia, maculae, pseudocyphellae or sore- dia. Phyllidia common, conspicuous, mainly marginal but also laminal, occasionally terete at first, soon becoming flattened-dorsiventral, constricted at base, 0.1-0.5 mm wide, l-5(-10) mm tall, simple, branched to ± coralloid, single or densely clustered and proliferating. Medulla white. Photo- biont green. Lower surface whitish or pale yellow-brown or pinkish fawn, smooth or wrinkled, glossy at margins to chestnut or red-brown centrally, or uniformly pale from margins to centre, tomentose from margins to centre or only in older parts, tomentum very variable, thick and woolly to thin, ragged and scattered in patches, white, silky, to dark- brown, red-brown or ± blackened. Pseudocyphellae com- mon, conspicuous, white, round to irregular, 0.1-1.5 mm Fig. 23 Distribution of Pseudocyphellaria poculifera in the palaeotropics. 144 D.J. GALLOWAY HiiniiimiiHiHiiimnimmniiinmiiiiHiiiBimimiiiiii Fig. 24 Pseudocyphellariaprolificans. Holotype (PC). Scale in mm. diam., margins raised, projecting from tomentum, decorti- cate area flat. Pycnidia common, scattered on upper surface, or concen- trated in groups or lines at lobe margins, hemispherical, 0.1 mm diam., ostiole raised, red-brown. Apothecia rather rare, often absent, when present promi- nent, marginal or submarginal, rounded 2-4(-5.5) mm diam., subpedicellate, pedicel short, 1-1.5 mm wide, pale whitish buff, exciple pale whitish buff to yellow-brown, translucent when wet, massive, corrugate-scabrid, obscuring disc when young, persisting at maturity as a thick, prominent margin, disc ± deeply cupuliform at first, subconcave to plane at maturity, smooth, shining, pale to dark chestnut-brown to red-brown, epruinose. Epithecium pale yellow-brown, to 14 |xm thick. Hymenium colourless, 110-125 |xm tall. Ascospores yellow-brown, simple to 1-septate, fusiform- ellipsoid, apices rounded or pointed, 28-33.5 x 6.5-8 jxm. CHEMISTRY. Tenuiorin, methyl gyrophorate, gyrophoric and congyrophoric acids, 7p-acetoxyhopane-22-ol, hopane-7p, 22-diol (tr.), hopane-15a, 22-diol. OBSERVATIONS. Pseudocyphellaria prolificans is a wide- spread palaeotropical species having a white medulla; a dimpled, punctate-impressed upper surface, often also con- spicuously papillate (use x 10 lens); conspicuously thickened margins below; scattered to densely clustered and proliferat- ing marginal and laminal phyllidia; a white medulla; a green STUDIES IN PSEUDOCYPHELLARIA (IV) photobiont; and a two-hopane chemistry with tenuiorin, and gyrophoric acid and derivatives. It is distinguished from P. sulphured by the marginal and laminal phyllidia; from P. insculpta by the green photobiont; from P. multifida by the dimpled, punctate-impressed upper surface, the proliferating phyllidia and thicker margins; and from P. homalosticta and P. reineckeana in lacking pseudocyphellae on the upper surface. DISTRIBUTION AND ECOLOGY. Widely distributed in the palaeotropics (Fig. 25) from Sri Lanka eastwards to Fiji and Samoa in the south-western Pacific. Also known from north- eastern Australia. On twigs, branches and trunks of trees and shrubs, and on rotting logs in humid montane rainforest and cloudforest, 500-3600 m. SPECIMENS EXAMINED. Sri Lanka: Kandy, Moon (BM). Malaysia: Sabah. Mesilau River, Hale 29225 (TNS); Mt Kinabalu, Samsudin (UKMB). Indonesia. Sumatra: Mt Sago near Pajakumbuh, Meijer B 8273 (L). Java: Tjibodas. Mt Cede, v. Ooststroom 14457, 14465, 14597 (Herb. Aptroot); sine loco, Koernich s.n. (Herb. Aptroot); Mt Kwai, Mt Panderman, Groenhart 2639, 2640 (L); Mt Gegerbentang, [on tree] Groenhart 2232 (L); Tjibeureum Falls, Schiffner 1575 (L); sine loco, Junghuhn (L); Mt Pangerango, Schiffner 1155, 2970, 3079 (L, BM, WU); sine loco, Blume (L). Philippines: Luzon. Benguet. Mt Santo Tomas, Aptroot 20321, 20392 (Herb. Aptroot). Mindanao. Mt Batangan, Warburg 14214c (B). Palawan. Brookes Point. Mt Mantalin- gahan, Sipman & Tan 29978 (B). Irian Jaya: Bamler s.n. (B); Vogelkop Penin. Nettoti Range. Mt Nettoti, v. Roy en & Sleumer 7476 (Herb. Aptroot). Papua New Guinea: Eastern Highlands. Chimbu. Lake Aunde, v.Balgooy 316 (Herb. Aptroot); Mt Wilhelm, Weber & McVean (Herb. Aptroot); Aptroot 31580 (Herb. Aptroot); Bundi Gap, Aptroot 32195 (Herb. Aptroot); Kotdame, Mundua 214, 220 (CBG); Goroka. Mt Gahavisuki Provincial Park, Aptroot 31036 145 (Herb. Aptroot); track to Mt Michael, Streimann 18502, 18541, 18809, 18825 (CBG). Southern Highlands. Mt Giluwe, Lambley (BM); Onim Forestry Station, Streimann 24761 (CBG); Margarima-Tari Road, Streimann 24369 (CBG). Morobe. Wau, Edy Creek Road, Sipman 15619 (Herb. Aptroot); Mt Missim, Bellamy 1530 (B); Yinimba, Streimann 19135 (CBG); track to Mt Missim, Bellamy 202, 202a (CBG); Streimann 18772 (CBG); Ekuti Divide, Rau 702 (CBG); Streimann 20126, 20136 (CBG); Mt Kauwara, Kairo 687-8 (CBG); Kaisinik, Kashiwadani 10739 (TNS); Araulu Logging Area, Streimann 13552 (CBG); Slate Creek & Gumi Creek Divide, Streimann 13868 (CBG); Wagau-Mulolo Track, Stre- imann 19615 (CBG); Spreader Divide, Streimann 11841, 11900 (CBG); Bulolo-Watutu Divide, Streimann 25033 (CBG); Honzeukngon village, Aptroot 18024 (Herb. Aptroot). Milne Bay. Mt Moiba, Pullen 7744 (Herb. Aptroot). Madang. Finisterre Range, Teptep Village, Aptroot 31959, 31997 (Herb. Aptroot). Central. Near Myola, Lambley (BM); Mt Albert-Edward, Kashiwadani 11523, 11723, 11821, 12011 (TNS); 2 km N. of Waiotape Airstrip, Kashiwadani 12100 (TNS). Western Highlands. Milep Area, Vinas 7647 (CBG). Solomon Islands: Guadalcanal Island. Mt Popomansiu, Hill 9283, 9288, 9513, 9561, 9670, 9687, 9709, 9827, 9841, 9856 (BM); Mt Gallego, Hill 8173, 8363 (BM); southern slopes of Mt Makarakomburu, Glenny 2048, 2055, 2125 (BSIP). Kolombangara Island. Ridge west of Kolom- bangara River, Hill 10544, 10573 (BM). New Caledonia: [ISOLECTOTYPE] sine loco, Vieillard 1795 (B); Riviere Blanche, Hill 11696 (BM). Kermadecs: [Raoul] Sunday Island. Sine loco, Milne (NY). Fiji: Viti Levu. Sine loco, Degener (GZU); ridge between Mt Nanggaranambuluta [Lomalangi] and Mt Namama, Smith 5004 (BM, L, US); western slopes of Mt Nanggaranambuluta, east of Nandari- vatu, Smith 4818 (US); Mt Evans Range, Smith 4280 (BM, US); Mt Victoria, Green (BM); Lam 6823 (L). Ngau. Herald Bay, Smith 7828 (US). Ovalu. Sine loco, Graffe (W). Samoa: Fig. 25 Distribution of Pseudocyphellaria prolificans in the palaeotropics. 146 D.J. GALLOWAY Upolu. See Lanonmea, Hochsentimer! (B). Rarotonga: Con- nells Gully, Parks 22171 (COLO). 23. Pseudocyphellaria punctillaris (Miill. Arg.) D.J. Gallo- way in Graphis Scr. 5: 9 (1993). Stictina punctillaris Miill. Arg. in Hedwigia 30: 48 (1891). Stictina fragillima f. punctillaris (Miill. Arg.) Stizenb. in Flora, Jena 81: 129 (1895). Sticta fragillima f. punctillaris (Miill. Arg.) Zahlbr., Cat. lich. univ. 3: 382 (1925). Type: Australia. Queensland, near Mt Bellenden Ker, 'Whelman Pools. Austral, orient', 1889, P.M. Bailey 567 (G 002544- holotype). Fig. 26. Thallus rosette-forming, 5-10(-15) cm diam., closely attached from margins to centre. Lobes broadly rounded, 5-8(-12) mm wide, contiguous or imbricate at margins, crowded-imbricate centrally. Margins entire, sinuous, broadly rounded at apices, becoming markedly lobulate- phyllidiate centrally, thickened and sometimes inrolled below. Upper surface dark malachite green to glaucous-blue when wet, pale yellowish fawn to brownish or pale red- brown, darker at apices when dry, undulate to wrinkled, conspicuously verrucose-scabrid (use x 10 lens), thick, coria- ceous, without isidia, maculae or soredia. Phyllidia marginal, rarely regenerating from cracks on upper surface, 0.2-1.5 mm diam., lobulate, distinctly jointed at base, simple to subcoralloid. Pseudocyphellae white, numerous, often crowded, conspicuous, 0.1-0.5 mm diam., round to elongate, conical-verruciform, margins raised, decorticate area flat. Medulla white. Photobiont cyanobacterial. Lower surface pale yellowish white to buff at margins, cinnamon-brown to red-brown centrally, uniformly tomentose from margins to centre, tomentum thick, woolly, pale at margins, brownish or greyish centrally, often obscuring pseudocyphellae. Pseudocyphellae white, scattered, round to irregular, 0.1-1 mm diam., most noticeable at margins, decorticate area flat or concave, granular, margins not raised. Pycnidia common, prominent, swollen, solitary or crowded, marginal and laminal 0.2-1 mm diam., ostiole punctate, red-brown. Apothecia sparse to frequent, submarginal and laminal, central rarely at lobe apices, sessile, constricted at base, round to deformed through mutual pressure, 1-3 mm diam., shallowly concave to plane, exciple prominent, coarsely Fig. 26 Pseudocyphellaria punctillaris. J. K. Bartlett 32147 (AK, BM). Scale in mm. STUDIES IN PSEUDOCYPHELLARIA (IV) verrucose-scabrid, obscuring disc when young, persisting as a coarse, dentate-verrucose margin to disc at maturity, pale brownish, disc shining, slightly roughened, pale to dark red-brown, epruinose. Epithecium red-brown, 8-14 ^m thick. Hymenium colourless, 110-115 u,m tall. Ascospores 1-septate, contents vacuolate, pale red-brown or yellow- brown, broad-ellipsoid, apices pointed or rounded, 25-28 x 8.5-11 (Jim. CHEMISTRY. Tenuiorin, methyl gyrophorate, gyrophoric acid, ?p-acetoxyhopane-22-ol, hopane-7p, 22-diol (tr.), hopane-15a, 22-diol. OBSERVATIONS. P. punctlllarls has rather shortly ± subdi- chotomously branching lobes with a white medulla, a scabrid- areolate upper surface, a cyanobacterial photobiont and white pseudocyphellae on both upper and lower surfaces. It has a simple two-hopane chemistry with tenuiorin and gyro- phoric acid present as accessories. Known from eastern Australia and the Philippines (Galloway & Kemp, 1993). It is distinguished from P. godeffroyii by the marginal lobulate phyllidia ; and from P. rigida (which has a scrobiculate upper surface), P. semilanata and P. trichophora by its scabrid- areolate upper surface and marginal phyllidia. DISTRIBUTION AND ECOLOGY. A characteristic Pacific spe- cies (Fig. 27) known from Indonesia and New Guinea where it seems to be most commonly collected, also from the Philippines, Samoa and in eastern Australia (Galloway & Kemp, 1993), and a single record from Hawaii. It is an epiphyte of rainforest trees and shrubs with an altitudinal range of 100 to 3650 m. Still rather poorly collected in the region. SPECIMENS EXAMINED. Indonesia. Java: sine loco, sine coll. (L). Philippines: Luzon. Mt Makiling, Degelius As-704, As-706 (UPS). Papua New Guinea: Eastern Highlands. Chimbu. Pindaunde Valley, Weber & McVean (Herb. 147 Aptroot); Felsspitze, Ledermann (B); Mt Wilhelm, McVean 66179 (CBG); top of Kassam Pass, Streimann 17915 (CBG). Morobe. Ekuti Divide, Streimann 22615 (CBG). Milne Bay. Woodlark Island, Kumei 34, 41, 95 (CBG); Soma 3 (CBG). Samoa: Savai. Lake Mafane, Bartlett 32147 (AK, BM). Hawaiian Islands: Hawaii. Waimea, Rock [Sandwicenses No. 6] (B). 24. Pseudocyphellaria reineckeana (Mull. Arg.) D.J. Gallo- way in Lichenologist 17: 305 (1985). Stictina reineckeana Mull. Arg. in Reinecke, Bot. Jb. 23: 295 (1896). Sticta reineckeana (Mull. Arg.) Zahlbr. in Rech., Denkschr. Akad. Wiss. Wien 81: 262 (1907). Type: Samoa, sine loco, 1895, Reinecke (G 002145-lectotype (Galloway, 1985: 305)). Fig. 28. Thallus irregularly spreading, 2-4(-8) cm diam., loosely attached centrally, margins ± free. Lobes narrow to medium, (l-)2-4(-8) mm wide, 5-15(-25) mm long, dichotomously to irregularly branching, divergent, discrete at apices, complex- imbricate centrally, apices pointed or smoothly rounded. Margins entire, only very slightly thickened below. Upper surface bright lettuce green or olive-green suffused brownish when wet, pale glaucous-green to pale buff to dark green- brown when dry, undulate, smooth, to irregularly wrinkled, not faveolate or punctate-impressed, matt or shining, rather brittle, friable when dry, without isidia, maculae, phyllidia or soredia. Pseudocyphellae white, scattered, rather sparse, minute, 0.1 mm diam. or less, fleck-like, punctiform, margins not noticeably raised. Medulla white. Photobiont green. Lower surface pale whitish buff to brownish, darkening centrally, smooth or shallowly wrinkled, glossy at margins, thinly to thickly tomentose centrally, tomentum ragged, whitish to pale buff, often inapparent, to thick, black or brown-black, woolly-entangled and obscuring lower surface Fig. 27 Distribution of Pseudocyphellaria punctillaris in the palaeotropics. 148 D.J. GALLOWAY Fig. 28 Pseudocyphellaria reineckeana. Lectotype Sticta reineckeana (G 002145). Scale in mm. and pseudocyphellae. Pseudocyphellae white, minute, fleck- like, 0.1 mm diam. or less, widely scattered, margins not noticeably raised. Pycnidia marginal, in groups or lines, minute, punctiform, 0.1 mm diam. or less, ostiole brown-black. Apothecia rare (often absent) to occasional, marginal or submarginal, sessile, constricted at base, rounded, 0.2-2 mm diam., exciple prominent, coarsely verrucose-scabrid, obscur- ing disc when young, persisting as verrucose-areolate margin at maturity, pinkish brown, translucent when wet, disc orange to red-brown, shining, epruinose. Epithecium pale yellow-brown, 5.5-9 u,m thick. Hymenium colourless to pale straw, 85-100 u,m tall. Ascospores 1-septate, yellow-brown to red-brown, contents vacuolate, ellipsoid, apices rounded or pointed, 25-31 x 8.5-11.5 fjun. CHEMISTRY. Methyl gyrophorate, gyrophoric acid (+ to + + ), 7p-acetoxyhopane-22-ol, hopane-7p, 22-diol (tr.), hopane-15a, 22-diol. OBSERVATIONS. Pseudocyphellaria reineckeana is character- ized by rather narrow lobes, a white medulla, a green photobiont, white fleck-like pseudocyphellae on both upper and lower surfaces and rather variable tomentum on the lower surface which varies from thin and scattered or inap- parent, to thick and dark and woolly, obscuring the lower STUDIES IN PSEUDOCYPHELLARIA (IV) surface. It has a basic two-hopane chemistry with variable amounts of gyrophoric acid which give a characteristic C+ red medullary reaction. It is similar to P. homalosticta but lacks the isidia characteristic of that species; it is distinguished from several taxa which have green photobionts and a two-hopane chemistry, from P. sulphured and P. stenophylla by lacking a punctate-impressed upper surface and having pseudocyphellae on the upper surface; from P. homalosticta in lacking isidia and from P. prolificans and P. multifida in lacking marginal and laminal lobules or phyllidia. DISTRIBUTION AND ECOLOGY. An epiphyte of trees and shrubs in montane rainforest, 1000-3500 m. Apparently restricted to the south-west Pacific from Borneo to Samoa (Fig. 29). SPECIMENS EXAMINED. Malaysia: Sabah. Mt Kinabalu, Sam- sudin (UKMB). Kalimantan. Sine loco, Lobb (BM). Papua New Guinea: Southern Highlands. Lai River, Streimann 22226 (CBG). Solomon Islands: Guadalcanal Island. Mt Popomansiu, Hill 9717 (BM). Fiji: Viti Levu. Suva, Wilson (MEL); Mt Nanggaranamuluta [Lomalangi], Smith 4833 (US); Mt Victoria, Green (BM); Nandarivatu, Green (BM); sine loco, Seeman (BM). Vanua Levu. Mt Kasi, Smith 1812 (BM). Samoa: Savai'i. Reinecke 52a (B). 25. Pseudocyphellaria rigida (Mull. Arg.) D.J. Galloway in Lichenologist 17: 305 (1985). Stictina rigida Mull. Arg. in Bull. Herb. Boissier4: 89 (1896). Sticta rigida (Mull. Arg.) Zahlbr., Cat. lich. univ. 3: 398 (1925). Type: Australia. Queensland, sine loco, P.M. Bailey (G 001990-holotype). Fig. 30. Thallus rosette-forming to irregularly spreading 5-10(-15) cm diam., loosely to closely attached centrally, margins free. Lobes subdichotomously to irregularly branch- ing, 5-10(-15) mm wide, 0.5-3(-6) cm long. Margins entire, 149 sinuous, markedly thickened-ridged below with prominent, projecting white psedocyphellae. Upper surface dark navy blue to glaucous blue-grey when wet, pale glaucous grey or grey-brown when dry, undulate, irregularly wrinkled to shal- lowly faveolate, interconnecting ridges smoothly rounded, sometimes indistinct, faveolae shallow, rather papery when dry, flabby when wet, without isidia, phyllidia or soredia. Maculae white, minute, ± reticulate, best seen when wet at lobe apices (use x 10 lens), more extensive whitish or buff cyanobiont-free areas often seen. Pseudocyphellae scattered, on laminal ridges, rather sparse, white, 0.1 mm diam., mar- gins slightly raised. Medulla white. Photobiont cyanobacte- rial. Lower surface pale whitish buff or ± greyish at margins, slightly darkening centrally, wrinkled-bullate, sparsely tomentose centrally with prominent glabrous margins or tomentose from margins to centre, tomentum dark brown, woolly, entangled. Pseudocyphellae scattered, minute, white, rather sparse at margins, prominent centrally, markedly conical-verrucose, margins prominent, swollen, concolorous with lower cortex, 0.5-1 mm diam., projecting above tomen- tum. Pycnidia not seen. Apothecia marginal and laminal, rather sparsely developed in older parts of thallus, sessile to subpedicellate, strongly constricted at base, rounded, 0.5-2.5(-3) mm diam., shal- lowly concave to plane, ± undulate at maturity, exciple coarsely corrugate-scabrid, pale buff or brown, ± translucent when wet, obscuring disc at first, persisting as a ± thick verrucose-scabrid-dentate margin or sometimes ± occluded by disc, disc pale orange-brown to red-brown, shining when young, matt at maturity, smooth, epruinose. Epithecium pale yellow-brown, to 14 jim thick. Hymenium colourless, 85-100 u,m tall. Ascospores pale yellow-brown to red-brown, 1-3-septate, contents often vacuolate, ellipsoid, apices rounded or pointed, (25-)28-30.5(-33.5) x 8.5-11 u,m. Fig. 29 Distribution of Pseudocyphellaria reineckeana. 150 D.J. GALLOWAY •••••••••••••••••••••••••••••••••••••••••••••••••I Fig. 30 Pseudocyphellaria rigida. H. Kashiwadani 10920 (TNS). Scale in mm. CHEMISTRY. 7p-acetoxyhopane-22-ol, hopane-7fJ, 22-diol (tr.), hopane-15a, 22-diol. OBSERVATIONS. Pseudocyphellaria rigida is a palaeotropical species characterized by a white medulla, a cyanobacterial photobiont, white pseudocyphellae on both upper and lower surfaces, a scrobiculate-faveolate to punctate-impressed upper surface and a bullate lower surface with continuous to sparse dark tomentum. It has a basic two-hopane chemistry. The distinctive scrobiculate upper surface and rather irregu- lar lobes distinguish P. rigida from P. trichophora which has a smooth upper surface and hairy lobe margins, and from P. semilanata which has a punctate-impressed upper surface and ± dichotomously branching lobes. DISTRIBUTION AND ECOLOGY. To date known in the region only from Mt Wilhelm in Papua New Guinea, on tree bark, 3400-3650 m. Known also from north-eastern Australia. SPECIMENS EXAMINED. Papua New Guinea: Western High- lands. Mt Wilhelm en route from Kombugomanbuno to the Pindaude Lakes, Kashiwadani 10920, 10929 (TNS); van Bal- gooy 593 (Herb. Aptroot). 26. Pseudocyphellaria semilanata (Mull. Arg.) D.J. Gallo- way in Lichenologist 17: 306 (1985). Stictina semilanata Mull. Arg. in Bot. Jb. 23: 293 (1897). Sticta semilanata (Mull. Arg.) Zahlbr., Cat. lich. univ. 3: 398 (1925). Cyanisticta semilanata (Mull. Arg.) Szatala in Annls hist.- nat. Mus. natn. hung. 7: 41 (1956), comb, inval. Type: Ins. Samoa, Dr Reinecke (G 002164-holotype). Fig. 31. Cyanisticta semilanata var. epunctulata Szatala in Annls hist.- nat. Mus. natn. hung. 7: 41 (1956). Type: New Guinea, in m. Sattelberg, K. Weinland (B-holotype). Pseudocyphellaria argyracea var. reveniens Vain, in Hedwigia 38: 121 (1913). Type: Philippines, Luzon, Sorsogon, Albay, June 1908, H.M.Curran (TUR-V 10134-holotype). Thallus irregularly spreading, 8-12(-15) cm diam., loosely attached centrally, apices ± ascending. Lobes very variable, 4-10 mm wide, l-4(-6) cm long, dichotomously branching, apices divergent, furcate, pointed or rounded, discrete from margins to centre or complex-imbricate centrally, flat to ± canaliculate. Margins entire, distinctly ridged above and STUDIES IN PSEUDOCYPHELLARIA (IV) 151 Fig. 31 Pseudocyphellaria semilanata.L. Brako 4256 (NY). Scale in mm. below with often prominent tomentum from lower surface projecting at right angles. Upper surface dull slate-blue to glaucous green suffused red-brown in parts especially at margins and apices, pale olivaceous to brownish when dry, undulate, shallowly ridged to distinctly punctate-impressed, rigid, rather coriaceous when dry, pliable when wet, without isidia, phyllidia or soredia. Maculae minute, white, imparting a delicate marbling to upper surface (use x 10 lens), promi- nent, larger and ± reticulate at lobe apices. Pseudocyphellae white, scattered, minute, punctiform, 0.1-0. 2 mm diam. Medulla white. Photobiont cyanobacterial. Lower surface yellowish white or pale buff at margins, darkening to red- brown centrally, tomentose from margins to centre, tomen- tum very thick, entangled, woolly, whitish to red-brown or ± blackened. Pseudocyphellae white, prominent, scattered, round to irregular, 0.2-2 mm diam., margins swollen, promi- nent, concolorous with lower cortex, often sunk in tomen- tum. Pycnidia marginal, hemispherical, 0.1 mm diam., solitary or in groups, ostiole dark red-brown or blackened. Apothecia marginal or submarginal, rarely laminal, rare (often absent) to occasional, sessile, constricted at base, rounded, 0.5-2 mm diam., subconcave to plane, exciple prominent in young fruits, persistent to occluded at maturity, slightly roughened to coarsely scabrid-verrucose, pale brown- ish, translucent when wet, disc pale to dark red-brown, smooth, epruinose, sometimes with a small central thalline lobule of sterile tissue. Epithecium pale red-brown, 8.5-14 u.m thick. Hymenium colourless, 70-85 u.m tall. Ascospores pale yellow-brown to red-brown, 1-3-septate, fusiform-ellipsoid, (25-)30.5-33(-36) x 5.5-8 Jim. CHEMISTRY. Methyl gyrophorate (±), gyrophoric acid (± to + + ), 7p-acetoxyhopane-22-ol, hopane-7(3, 22-diol (tr.), hopane-15a, 22-diol. OBSERVATIONS. Pseudocyphellaria semilanata is character- ized by dichotomously branching lobes, a cyanobacterial photobiont, a white medulla, scattered white pseudocyphel- lae on both upper and lower surfaces, a punctate-impressed upper surface and a two-hopane chemistry. It is similar to P. beccarii but is distinguished from it by having punctiform white pseudocyphellae on the upper surface. It differs from P. godeffroyii in having a shining, dimpled upper surface 152 D.J. GALLOWAY which is never scabrid-areolate. DISTRIBUTION AND ECOLOGY. A palaeotropical species occurring in the Pacific from the Bonin Islands and Indonesia eastwards to Fiji and Samoa (Fig. 32). Epiphytic on trees and shrubs, on rotting logs and on litter in upper montane and cloud forest, and on small cut trees and fence posts; 100-3650 m. SPECIMENS EXAMINED. Bonin Islands: sine loco, Wright (US). Indonesia. Java: Laurer s.n. (B); Mt Lawu, Feekes 4969 (L); Mt Kawi, Groenhart 1828 (L); Mt Ardjuno, Groen- hart 1517, 7244, 7246, 7247 (L). Flores: sine loco, Verheijen 5202 (Herb. Aptroot). Irian Jaya: sine loco, Bander (B); Biri, Weinland (B). Papua New Guinea: Madang. Finisterre Mountains, Saidor Subdistrict, Naho-Rawa Div., Jermy 4014 (BM). Northern Distr., Tufi subdistr., Lake Ridubidubina, Hoogland 4495 (L). Eastern Highlands. Chimbu. Pindaunde Valley, Aptroot 31382 (Herb. Aptroot); Lake Aunde, 3600 m, v.Balgooy 316 (Herb. Aptroot); Mt Wilhelm, Kashi- wadani 10881, 10883, 10915, 10919, 10937, 10953-4, 10957, 11079, 11081, 11168, 11391, 11411 (TNS); Imbuka Ridge, Weber & McVean (COLO). Morobe. Saruwaged Range, Sipman 24383, 24470 (B); near Honzeukngon village, Aptroot 17851, 18018, 18020, 18022 (Herb. Aptroot); Herzog Mountain, Streimann & Umba 11032 (CBG). Southern High- lands, laro River, Streimann 23824 (CBG). Western High- lands. Nebilyer River, 2760 m, Streimann 20600 (CBG); Milne Bay. Woodlark Island, Kumei 43^4 (CBG). Solomon Islands: Guadalcanal. Mt Popomansiu, Hill 9403, 9617 (BM). Fiji: Viti Levu. Naitasiri, northern portion of Rairaimatuku Plateau between Mt Tomanivi and Nasonggo, Smith 5755 (BM, L, US). Samoa: Upolo. Lake Lanoto'o, Schultz-Motel 3425 (B); mountains east of Tiave, Schultz-Motel 4042 (B); Lanotoo, Rechinger 3100 (W). Savai'i. Lake Mafane, Bartlett 32149, 32153, 32155, 32156 (AK, BM). 27. Pseudocyphellaria stenophylla (Mull. Arg.) D.J. Gallo- way in Lichenologist 17: 306 (1985). Sticta stenophylla Mull. Arg. in Flora, Jena 65: 293 (1897). Type: New Caledonia, Mt Mu, 1886, Vieillard (G 002010-holotype). Fig. 33. Pseudocyphellaria prolificans var. angustata Rasanen in Suomal. elain-ja kasvit. Seur. van. kasvit. Julk. 20(3): 16 (1944). Type: New Caledonia, ad corticem arboris, 1863(-64), E. Vieillard (H-lectotype, selected here). Thallus irregularly spreading, often rather sparse, 2-4(-6) cm diam, loosely to closely attached centrally, margins free and ± ascending. Lobes very narrow, 0.1-0.5(-1.5) mm wide, 2-8(-15) mm long, irregularly linear-laciniate, subdichoto- mously to irregularly branching, ± free at apices, entangled centrally, plane to canaliculate (most noticeable in young, marginal parts). Margins entire, slightly thickened above, without isidia, phyllidia, pseudocyphellae or soredia. Upper surface bright lettuce green when wet, pale green-grey to pale olivaceous buff when dry, fragile, rather brittle when dry, pliable when wet, without isidia, maculae, phyllidia, pseudocyphellae or soredia. Medulla white, photobiont green. Lower surface pale whitish buff at margins to pale yellow-brown or red-brown centrally, glossy, glabrous or very sparsely and minutely tomentose in central parts, with a narrow, raised midrib in older parts. Pseudocyphellae white, rounded, punctiform, 0.1 mm or less, most common at margins, rather sparse and widely scattered centrally. Apothecia and pycnidia not seen. CHEMISTRY. 70-acetoxyhopane-22-ol, hopane-7p, (tr.), hopane-15a, 22-diol and gyrophoric acid. 22-diol OBSERVATIONS. Pseudocyphellaria stenophylla is character- ized by a white medulla; scattered, fleck-like, white pseudocyphellae on the lower surface but not present at the Fig. 32 Distribution of Pseudocyphellaria semilanata. STUDIES IN PSEUDOCYPHELLARIA (IV) 153 Illllllllllllllllllllllllllllllllll Fig. 33 Pseudocyphellaria stenophylla. Holotype (G 002010). Scale in mm. margins; a green photobiont; a naked lower surface which is ± costate centrally; very narrow lobes which are subdichoto- mously to intricately branched; and unthickened margins which are neither isidiate nor phyllidiate. It has a two-hopane chemistry with gyrophoric acid. Its very narrow lobes distin- guish it from P. sulphurea. DISTRIBUTION AND ECOLOGY. Known in the region only from the type collections in New Caledonia. Also in Australia (Queensland). SPECIMENS EXAMINED. New Caledonia: known only in the region from the type specimens (see above). 28. Pseudocyphellaria sulphurea (Schaer.) D.J. Galloway in Lichenologist 17: 306 (1985). Sticta sulphurea Schaer. in Moritzi, Syst. Verz.: 127 (1846). Type: Java, sine loco, Zollinger 1860x (L 910, 182-20-lectotype (Galloway, 1985ft: 306)). Fig. 34. Sticta richardi var. impressa Meyen & Plot, in Nova Acta Acad. Leap. Carol. 19, Suppl.: 216 (1843). Type: Manil- lae, ad truncos sylvarum, ad Meyen (L 019,211-1788- lectotype (Galloway, 1985: 306)). Sticta quercifolia Taylor in Lond. J. Bot. 6: 177 (1847). 154 D.J. GALLOWAY Pseudocyphellaria quercifolia (Taylor) Vain, in Philipp. J. Sci. sect. C, Hot. 8: 117 (1913). Type: Sri Lanka, sine collectoribus nomine, no. 40, ex Herb. Hook. (BM- holotype; H-isotype). Sticta dissimulata Nyl., Syn. meth. lich. 1(2): 362 (1860). Crocodia dissimulata (Nyl.) Trevis., Lichenotheca veneta exs. 75 (1869). Lobaria dissimulata (Nyl.) Kuntze, Revis. gen. pi 2: 876 (1891). Pseudocyphellaria dissimulata (Nyl.) Vain, in Philipp. J. Sci. Sect. C, Bot. 8: 118 (1913). Type: Java, sine loco, sine collectoribus nomine (H-NYL 33517-lectotype (Galloway, 19856: 307)). Sticta punctulata Nyl., Syn. meth. lich. 1(2): 364 (1860). Crocodia punctulata (Nyl.) Trevis., Lichenotheca veneta exs. 75 (1869). Lobaria punctulata (Nyl.) Kuntze, Revis. gen. pi. 2: 876 (1891). Type: Java, sine loco, Zollinger 1799 pr.p. (H-NYL 33481-lectotype (Galloway, 19856: 307)). Sticta demutabilis Kremp. in /. Mus. Godeffroy 1(4): 98 (1874). Pseudocyphellaria demutabilis (Kremp.) Gyeln. in Revue bryol. lichen. 6: 173 (1933). Type: Samoa, Savai, mont. veg ad arbores, E. Grdffe 106 (M-holotype; W-isotype). Sticta demutabilis f. laevis Kremp. in J. Mus. Godeffroy 1(4): 98 (1874). Pseudocyphellaria dissimulata var. laevis (Kremp.) Szatala in Annls hist.-nat. Mus. natn. hung. 7: 40 (1956), comb, inval. (Art. 33.2). Type: Samoa, Ins Upolu, E. Grdffe 101 (M-lectotype (Galloway, 19856: 307)). Sticta demutabilis f. minor Kremp. in /. Mus. Godeffroy 1(4): 98 (1874). Type: Samoa, Savai, E. Grdffe 109 (M-holotype). Sticta karstenii Mull. Arg. in Flora, Jena 64: 505 (1881). Pseudocyphellaria karstenii (Mull. Arg.) Szatala in Annls hist.-nat. Mus. natn. hung. 7: 40 (1956), comb, inval. (Art. 33.2). Type: Novae Hollandiae [Australia], North Queen- sland, Bellenden Ker Range, Karsten 2, comm. F.v. Mueller 1881 (G 002020-holotype). Sticta leucophylla Mull. Arg. in Flora, Jena 72: 506 (1889). Pseudocyphellaria leucophylla (Mull. Arg.) Szatala in Annls hist.-nat. Mus. natn. hung. 7: 39 (1956), comb, inval. (Art. 33.2). Type: New Guinea, near summit of Owen Stanley Range, Sir W. Macgregor 6 pr.p., comm. F. v. Mueller 1889 (G 002116-holotype). Pseudocyphellaria dissimulata var. hypophaea Vain, in Phil- ipp. J. Sci. Sect. C, Bot. 8: 118 (1913). Type: Philippines, Mindanao, District of Zamboanga, ± 1200 m alt., on trees, Nov.-Dec. 1911, E.D. Merrill 8351 (TUR-VAINIO 10195-lectotype, selected here). Pseudocyphellaria dissimulata var. nudior Vain, in Philipp. J. Sci. Sect. C, Bot. 8: 118 (1913). Type: Phillipines, Negros, Canlaon Volcano, 5000 ft, on trees, April 1910, E.D. Merrill 6889 (TUR-VAINIO 10191-lectotype, selected here). Pseudocyphellaria dissimulata var. curranii Vain, in Philipp. J. Sci. Sect. C, Bot. 8: 119 (1913). Type: Phillipines, Luzon, Prov. of Pampanga, Mt Arayat, ad truncum arboris, March 1910, H. M. Curran 19341 (TUR-VAINIO 10193-holotype). Thallus irregularly spreading, often in large, entangled clones, 10-20(-35) cm diam., loosely attached centrally, api- ces free, ± ascending. Lobes very variable (l-)3-8(-12) mm wide, (l-)2-8(-12) cm long, linear-elongate, ± dichoto- mously branching, contiguous or discrete, apices usually free, ± divergent, rounded, pointed, truncate or furcate, complex- entangled-imbricate centrally. Margins entire, smoothly rounded, conspicuously thickened-ridged above and below, here and there with occasional white pseudocyphellae. Upper surface bright lettuce-green when wet, occasionally suffused red-brown at apices and margins, pale grey-green, olivaceous to fawnish or red-brown when dry, shining, conspicuously dimpled, punctate-impressed to ± shallowly faveolate, smooth in parts or faveolate but generally strongly punctate- impressed, tough, coriaceous to thin and somewhat papery, without isidia, maculae, phyllidia, pseudocyphellae or sore- dia. Medulla white. Photobiont green. Lower surface pale whitish to pale yellow-brown or pinkish buff, noticeably wrinkled-ridged, thinly tomentose from margins to centre or ± glabrous, tomentum short, velvety to wispy, whitish to pale buff. Pseudocyphellae common, scattered, white, fleck-like, minute at margins, round to irregular centrally, to 0.5 mm diam., margins very slightly raised, concolorous with lower surface, decorticate area flat. Pycnidia frequent to somewhat sparse, mostly marginal in short lines or in clusters, punctate, 0.1 mm diam. or less, ostiole red-brown to black. Apothecia absent to occasional to ± frequent, marginal and submarginal, sessile, constricted at base, rounded, 0.5-2.5(-4.5) mm diam., exciple pale pinkish fawn to yellow- brown, translucent when wet, persistent, coarsely scabrid- areolate, disc subconcave to plane, smooth, matt, pale to dark red-brown, epruinose. Epithecium pale yellow-brown, 9-15 fxm thick. Hymenium colourless, 70-95 (Jtm tall. Ascospores 1-septate, yellow-brown to red-brown, fusiform- ellipsoid, (22-)25-28(-30.5) x 6.5-8(-ll) u-m. CHEMISTRY. 7p-acetoxyhopane-22-ol, hopane-7(3, 22-diol (tr.), hopane-15ot, 22-diol as constant compounds, with or without some or all of the following as accessory compounds: tenuiorin, methyl gyrophorate, gyrophoric and congyro- phoric acids (specimens with gyrophoric acid in quantity give a positive C+ pink medullary reaction). OBSERVATIONS. Pseudocyphellaria sulphurea is a widespread palaeotropical species having a white medulla, a dimpled, punctate-impressed upper surface (not or very rarely truly faveolate) with conspicuously thickened margins below, and scattered, fleck-like, white pseudocyphellae on the lower surface. In Schaerer's description of Sticta sulphurea he mentions '. . . intus sulphureis' (i.e. yellow medulla) in the account of specimen Zollinger 1860x on which he based the name. However, examination of authentic Zollinger material from Java bearing this number failed to reveal any yellow medulla and it is not clear why Schaerer mentioned sulphureis in his description. The Zollinger material which bears Schaerer's name has a uniformly white medulla, characteristic of the species, but on all other counts the material fits Schaerer's description precisely. P. sulphurea is distinguished from P. stenophylla by its wider lobes which are thickened at the margins of the lower surface; from the New Zealand endemic P. rufovirescens by the characteristic punctate-impressed, dimpled upper surface and the presence of the depside gyrophoric acid; from P. prolificans and P. multifida by lacking marginal or laminal phyllidia, isidia or proliferations; and from P. beccarii which has a similar morphology but has a cyanobacterial photo- biont. Photosymbiodemes of P. sulphurea and P. beccarii, although reported in the literature (James & Henssen, 1976) STUDIES IN PSEUDOCYPHELLARIA (IV) 155 t. C5T. WO «W. ""rs •S. Jt; . .-«••.' *•'• "Lvt' '' - ^-t>|»?, * - ** C/ Fig. 34 Pseudocyphellaria sulphured. Holotype Sf/cta quercifolia (BM). Scale in mm. have not been seen by the present author. DISTRIBUTION AND ECOLOGY. One of the most common and widespread of palaeotropical lichens (Fig. 35) growing as an epiphyte of branches and trunks of trees and shrubs in dense, humid, mossy montane forest and cloud forest, often forming large mats, also on scattered subalpine trees and shrubs in alpine grassland, 700-3600 m. SPECIMENS EXAMINED. Madagascar: Ambohinutombo for- est, Forsyth Major 457 (BM); Toshimaniko forest, Forsyth Major 96 (BM). Sri Lanka: Kandy, Moon (BM); sine loco, Macrae 131 (BM); Central Province, Thwaites C.L. 22 (BM). Thailand: Prov. Nakawng Li Thammarat, Khao Luang, van Beusekom s.n. (Herb. Aptroot). Malaysia: Pahang. Gunong Hyan, Perak, Wray (BM). Singapore. Sine coll. 6689 (BM). Sabah. Mt Kinabalu, near HQ of National Park, Ding Hou 207 (Herb. Aptroot); Mt Kinabalu, Sipman & Tan 31085, 31377a (B); 3000 m, Polak (B); Lee (COLO); Mesilau River, Hale 28113, 29194, 29256, 29290 (TNS). Indonesia. Sumatra: Mt Korinchi, 7300 ft, Robinson & Kloss (W); sine loco, Korthals (L); sine loco, Forbes (BM). Java: Salang, v. Goebel (M); Prov. Batavia, in monte Megamendong, Schiffner 3386 c (M, W); Pamaboela bei Toegoe, Kurz (M); Mt Ardjuno, trail from Sumber Brantas Estate to Mt Kembar, Groenhart 9856 (Herb. Aptroot); Mt Gede, Tjibodas, Groenhart 1807 (L); sine loco, Weiss 4517 (B); sine loco, Laurer (B); Junghuhn (B); Malang, Lederer (B); Palang, v. Goebel (W); sine loco, Reinwandt (B). Mt Gede, Palmer & Bryant 1091, 1242 (US); 156 D.J. GALLOWAY Fig. 35 Distribution of Pseudocyphellaria sulphured in the palaeotropics. Flores: sine loco, 1970, J.A.J. Verheijen 2729 (Herb. Aptroot). Philippines: Mindanao. Elmer [Kryptogamae exsic- catae editae a Mus. Hist. Nat. Vindobon. 2841] (M, B); Mt Batangan, Warburg 14214a (B); Mt Apo, Mearns s.n. (US); Davao. Mt Apo, Elmer 11535 (W); sine loco, Mearns s.n. (W); Elmer (BM, W). Luzon. Benguet, Baguio, Elmer 83; Pampanga. Mt Pinatubo, Elmer 21939 (B); Benguet. Baguio, Elmer s.n. (US); Mt Tonglon, Ramos s.n. (US); Curran s.n. (US); ?Fayobos. Mt Banohas, Elmer 7565 (W). Mindoro. Mt Halcon, Sales & Wijangco 10 (B); Merrill s.n. (US). Cama- rines. Mt Isarog, Ramos 6042 (BM). Caroline Islands: Ponape. Mt Erica, Cheatham 27 (B); Mt Tamantamansakir, Classman (W). Irian Jaya: Vogelkop Peninsula, Tamrau Range, Gunung Bagimana, van der Zon s.n. (Herb. Aptroot); Vogelkop Peninsula, Nettoti Range, Wekari River camp, van Roy en & Sleumer 8145 (Herb. Aptroot). Papua New Guinea: East Sepik. Kairuru Island, Borrell 3 (CBG). Madang. Huon Peninsula, Finisterre Range, Yupna Valley, Teptep Village, trail NNW towards Bambu, Aptroot 31919, 32000, 32009 (Herb. Aptroot). Morobe. Track to Mt Missim, Broome 89A, 208 (CBG, B); Bellamy 201, 202b, 204a, 207-8, 208a, 1392, 1453 (B); Streimann 18511 (CBG); Mt Sarawaket [Saruwaged] Southern Range, Koponen 32872 (Herb. Aptroot); Monkumbion, Hoogland 9764 (BM); Cromwell Mountains, Koponen 31200 (Herb. Aptroot); Spreader Divide, Schodde & Craven 4944 (Herb. Aptroot); Edie Creek Road, Sipman 15621 (B); Skindiwai, Kairo 391 (CBG); Ekuti Divide, Ran 697, 701, 704-5 (CBG); Kauwara River, Kairo 670, 672 (CBG); Eraulu Logging Area, Kairo 318 (CBG); Mt Missin, Kashiwadani 10405, 10410 (TNS); Mt Kaindi, Kashiwadani 10454 (TNS); Streimann 24817, 33281 (CBG); 7 km SE of Bulolo, Kashiwadani 10808 (TNS); Herzog Mountains, Streimann & Umba 10962, 11100 (CBG); Gumi Divide, Streimann 22712 (CBG). Southern Highlands. Tari, Mt Ne, Kalkman 4866 (Herb. Aptroot); Tari Gap, Lambley (BM); Lama Sawmill Logging Area, Streimann 24710 (CBG); Munia Logging Area, Streimann 23212 (CBG). Eastern Highlands. Chimbu, Imbuka Ridge, Weber & McVean (Herb. Aptroot); track to Mt Wilhelm, Sipman 21922, 21929 (B); Mt Wilhelm, Borgmann 719, 732, 805 (B); Kashiwadani 10838, 10847, 10867, 10890, 10944, 10961, 11038, 11187, 11195, 11206, 11349, 11414 (TNS); near Hogabi Village, Streimann 18615, 18662, 18694 (CBG); track to Mt Michael, Streimann 18789, 18828 (CBG); Goroka, Gahavisuka Provincial Park, Streimann & Kairo 18184, 18227 (CBG). Central District. Mt Wosa, v.Royen NGF 20269 (Herb. Aptroot); Mt Albert-Edward, Kashiwadani 11747, 11768, 11780, 11809, 11823, 11934, 12001, 12012, 12293 (TNS); 2 km N. of Waiotape Airstrip, Kashiwadani 12262 (TNS). Western Highlands. Laiagam, Yobobos, Hoogland & Schodde 7639b (B). Milne Bay. Mt Moiba, Pullen 7742 (Herb. Aptroot). New Ireland: Hans Meyer Range, Sands 1917 (BM). Solomon Islands: Kolombangara Island. Ridge west of Kolombangara River, Hill 10508, 10537, 10577, 10679, 10603, 10605, 10674 (BM); South Summit, Hill 10484-5, 10490 (BM); Poitete, Glenny 2280 (BSIP); Iriri, Glenny 2403 (BSIP). Guadalcanal Island. Mt Popomansiu, Hill 9286-7, 9330, 9376, 9436, 9443, 9445, 9477, 9478-82, 9482a, 9514-6, 9518, 9567, 9569-70, 9573-4, 9582, 9583-5, 9676, 9678, 9688, 9691, 9696, 9701-3, 9707-8, 9710, 9712, 9719, 9721, 9803, 9829-30, 9835-8, 9857-61 (BM); Mt Gal- lego, Hill 8170 (BM). Bougainville: south rim of Lake Loloru crater, 20 miles N. of Buin, Craven & Schodde 336 (Herb. Aptroot). New Caledonia: Roberts (MEL); sine loco, Comp- ton 1729 (BM). New Hebrides: Aneityum. Sine coll. (BM). Fiji: Viti Levu. N-bulti trail, Selling (S); Nadarivatu, O. & I. Degener 31812e (Herb. Aptroot; B); Mt Victoria, Green (BM); Lam 6832 (BM); Novai, Degener 31815 (B); ridge from Mt Namama to Mt Tomanivi, Smith 5712 (US); Ngau, Herald Bay, Smith 7828 (US); sine loco, Home (BM); Milne (BM). Samoa: Upolu. Near Lake Lanoto'o, Schultz-Motel 3309 (B); Mt Lanuto'o, Rechinger (B, W); Mt Fiamoe, STUDIES IN PSEUDOCYPHELLARIA (IV) Schultz-Motel 4251 (B); Viti Savai, Graeffe (BM); sine loco, Powell (BM). Savai'i. Reinecke (WU); ?Tutuila, Reinecke (WU). 29. Pseudocyphellaria trichophora (Vain.) D.J. Galloway, comb. nov. Fig. 36. Basionym: Sticta trichophora Vain, in Philipp. J. Sci. Sect. C, Bot. 8: 123 (1913). Type: Philippines. Mindanao, Camp Keithley, Lake Lanao, September-October 1907, Mary Strong Clemens 1304 (US-isotype). Thallus orbicular to irregularly spreading, 10-12(-15) cm 157 diam., loosely to closely attached centrally, free and ± ascending at margins and apices. Lobes linear-elongate (3-)5-8(-12) mm wide, 2-6(-8) cm long, attenuating at api- ces which may be blunt, rounded or shallowly furcate, ± dichotomously branching, sinuses prominent, thickened. Margins entire, conspicuously thickened-ridged above and below, with prominent white pseudocyphellae, tomentose, especially at or near apices, tomentum white, silky. Upper surface dark leaden grey or grey-blue, suffused brownish at margins when wet, pale greyish fawn when dry, tough, coriaceous, smooth or minutely and shallowly wrinkled in parts, plane to ± canaliculate, isidia, phyllidia and soredia absent. Pseudocyphellae present, minute, 0.1 mm diam. or Fig. 36 Pseudocyphellaria trichophora. Isotype (US). Scale in mm. 158 D.J. GALLOWAY less, white, punctiform, very widely scattered, inapparent. Medulla white. Photobiont cyanobacterial. Lower surface uniformly thickly tomentose from margins to centre, tomen- tum dense, woolly-entangled, pale fawnish buff at margins to dark brown or blackened centrally. Pseudocyphellae white, prominent (especially at lobe margins), rounded, 0.1-1 mm diam., conical verruciform, margins raised, sharply defined, glossy, decorticate area flat to concave. Pycnidia not seen. Apothecia very rare, marginal or submarginal, rounded, cupuliform to 2.5 mm diam., sessile, constricted at base, exciple coarsely wrinkled-scabrid, pale buff-brown, translu- cent when wet, with white silky hairs prominent below, disc concave to plane, smooth, shining, red-brown, epruinose. Epithecium red-brown, 14—20 u,m thick. Hymenium pale straw or colourless, 90-110 u,m tall. Ascospores pale red- brown, 1-3-septate, ellipsoid, apices rounded or pointed, 18-25 x 7-11 |xm (Vainio (1913: 122) gives spore dimensions as 44-50 x 4-6 REFERENCES CHEMISTRY. 7(3-acetoxyhopane-22-ol, hopane-7p, 22-diol (tr.), hopane-15a, 22-diol. OBSERVATIONS. Pseudocyphellaria trichophora has linear- elongate lobes and a coriaceous upper surface which is ± canaliculate towards margins and apices and is devoid of isidia, phyllidia and soredia, but which has rather sparse, white punctiform pseudocyphellae. It has a white medulla, a cyanobacterial photobiont, tomentose lobe margins (espe- cially at apices), a densely tomentose lower surface with conspicuous white pseudocyphellae with raised, narrow mar- gins resembling true cyphellae. It has a two-hopane chemis- try. It is distinguished from P. semilanata by the tomentose lobe margins and the nature of the pseudocyphellae, and from the New Zealand endemic P. allanii (Galloway, 1988) by the scattered pseudocyphellae on the upper surface. DISTRIBUTION AND ECOLOGY. At present known only from the Philippines and Papua New Guinea. Still very much under-collected. From humid montane rainforest at 1500 m. ADDITIONAL SPECIMEN EXAMINED. Papua New Guinea: Morobe. Aiuwa-Bakia Track, Streimann & Tamba 12290 (CBG). ACKNOWLEDGEMENTS. I am grateful to the curators and keepers of the herbaria listed in the text; to my colleagues and friends Dr A. Aptroot (Baarn), Dr P. Diederich (Luxembourg), Dr J.A. Elix (Canberra), Mr D. Glenny (Wellington), Mr P.W. Lambley (Nor- folk), Dr E. Serusiaux (Liege), Dr H.J.M. Sipman, Prof. C.W. Smith (Honolulu), and Mr H. Streimann (Canberra) for information and the loan of specimens; to Sophie Kemp for assistance with chemistry; to Anne Hume for assistance with keyboarding data; to Profs Oo Khaik Cheang and Haji Mohammed (Kuala Lumpur) for assistance and companionship in the field; and to the NRI for assistance with travel and field work in Malaysia; and Phil Hurst (The Natural History Museum, Photographic Unit) for expert photographic assis- tance. Awasthi, D.D. 1965. Catalogue of the lichens from India, Nepal, Pakistan and Ceylon. Beih. nov. Hedwigia 17: 1-137. 1988. A key to the macrolichens of India and Nepal. J. Hattori hot. Lab. 65: 207-302. Elix, J.A., Streimann, H. & Archer, A.W. 1992. The lichens of Norfolk Island 2. The genera Cladonia, Pertusaria, Pseudocyphellaria and Ramalina. Proc. Linn. Soc. N.S.W. 113: 57-76. Galloway, D.J. 1985a. Flora of New Zealand lichens. Wellington. 19856. 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Observations on the composition and distribution of the Bull. not. Hist. Mus. Lond. (Bot.) 24(2): 161-171 Issued 24 November 1994 Morphology and ecology of seedlings, fruits and seeds of Panama: Bixaceae and Cochlospermaceae NANCY C. GARWOOD Department of Botany, The Natural History Museum, Cromwell Road, London SW75BD CONTENTS Introduction 161 Methods 161 Bixaceae 162 Cochlospermaceae 166 Discussion 169 References . . 171 SYNOPSIS. Seedlings, saplings, fruits and seeds of Bixa urucurana (Bixaceae) and Cochlospermum vitifolium (Cochlospermaceae) are described, and ecological data on these stages summarized, as part of the ongoing Seedling Flora Project, centred on Barro Colorado Island, Panama. Data from these taxa and new seedling characters are used to discuss the relationship between the two families. The presence of extra-floral nectaries at leaf nodes along the stem and 'pearl bodies' on leaves, two ecologically important characters not described in standard floras, are discussed. INTRODUCTION The aim of the Seedling Flora Project, initiated in 1985, is to produce an illustrated flora of seedlings, saplings, fruits and seeds for the neotropical forest centred on Barro Colorado Island, Panama (see Garwood & Humphries, 1993). In this account, two species are described, Bixa urucurana Willd. (Bixaceae) and Cochlospermum vitifolium (Willd.) Spreng. (Cochlospermaceae), which are rare on Barro Colorado Island (BCI) but common in other parts of central Panama. Although the two families are treated separately in the Flora of Barro Colorado Island (Croat, 1978), Flora of Panama (Robyns, 19670, /?; D'Arcy, 1987), and the Flora neotropica (Poppendieck, 1981), they are closely allied and often com- bined into one family, the Bixaceae (Corner, 1976; Cron- quist, 1981). METHODS Seeds were collected in central Panama, and sown while fresh under sun and shade conditions in a screened growing house on Barro Colorado Island. Seedlings were photographed, observed for fresh characters, and harvested at irregular intervals; several individuals were grown to sapling size. Each seed collection has a unique collection number: voucher specimens and the seedlings grown from that collection are lettered sequentially according to date of collection or har- ©The Natural History Museum, 1994 vest, e.g. 1784A (adult voucher), 1784B-G (seedlings). Adult voucher specimens are deposited among the following her- baria: F, BM and PMA. All seedling, fruit and seed speci- mens described are currently at BM, where seedling voucher specimens and fruit and seed samples will be deposited; duplicates of seedlings will be distributed to other herbaria. Descriptions are based on photographs and notes of fresh specimens and on dry and preserved material from the collections described above, supplemented with general information from published floras. Taxonomic references used throughout the descriptive parts are summarized after the notes on family habit and distribution; references are cited in the fruit and seed descriptions only when they are the sole source of specific information; those used in the ecology section are cited in the text. For each species, information is stored in a detailed data- base and output in a standard format that will be used for all 700 species in the Seedling Flora Project. Descriptive terms are being standardized across all families, but generally follow common usage. See de Vogel (1980) for definitions of seedling-specific terms. The following less conventional terms are used: length of pubescence or size of other surface features, minute (< O.lmm), short (0.1-0.5 mm), long (0.5-1.0 mm), very long (> 1.0 mm). Leaf nodes are num- bered sequentially from the first leaf-bearing node above the cotyledons. A more detailed account of seed germination for all species is being prepared (Garwood, in prep.). In this account, the median time until germination is given, rather than the mean, because the distribution of germination time is highly skewed. 162 N.C. GARWOOD BIXACEAE HABIT AND DISTRIBUTION. Shrubs or trees. Neotropical, but the cultivated Bixa orellana L. is planted throughout the tropics; one genus and five species, two species in Panama and one or two species known from BCI (see below). TAXONOMIC REFERENCES. Macbride (1941), Standley & Wil- liams (1961), Robyns (1967a), Dathan & Singh (1972), Baer (1976), Corner (1976), Croat (1978), Molau (1983). Bixa L. TAXONOMIC NOTES. Five species of Bixa are present and distinct in the Amazon basin (Macbride, 1941; Baer, 1976; Molau, 1983), including the widely cultivated B. orellana L. sensu stricto and its cultivars. Typical examples of B. uru- curana Willd. and B. orellana L. occur throughout Central America, but intermediate forms have been reported (Baer, 1976) and Central American floras have combined the two taxa or ranked them as varieties of B. orellana L. sensu lato (Standley & Williams, 1961; Robyns, 1967a). See Dempsey & Garwood (1994) for further discussion of these two taxa. The BCI taxon, described as B. orellana L. by Croat (1978), is probably B. urucurana Willd. Its distribution along the shoreline of BCI (Croat, 1978), where lake levels fluctu- ate greatly throughout the year, is consistent with the habitat of B. urucurana but not B. orellana. B. urucurana is usually found along stream and river banks and in seasonally inun- dated or periodically brackish low areas, whereas B. orellana does not tolerate waterlogged soils (Baer, 1976). In addition, the only Bixa seedlings I have encountered on BCI (Garwood 2439) were growing on a sandy beach at the mouth of a small stream. One BCI specimen (Croat 12294) has been identified as B. urucurana (Baer, 1976), but I have not verified others listed in Robyns (1967a) and Croat (1978). However, it would be surprising if the commonly cultivated B. orellana has never been planted on BCI, as many ornamental and food plants have been cultivated in the laboratory clearing and older settlement sites. Bixa urucurana Willd., Enum. pi.: 565 (1809). Fig. 1. HABIT AND DISTRIBUTION. Small tree, to 10 m tall, of moist to wet lowland forest, usually along streams, rivers or other inundated areas. Brazil and Peru to Nicaragua or Guatemala. COLLECTIONS. Panama. Colon: Gamboa, along shoreline of Rio Chagres, Garwood 1830A (BM, F, PMA, adult vouch- ers), Garwood & Rand 2085 A (no voucher, same tree as Garwood 1830A); Barro Colorado Island: shoreline, Gar- woo d 2439 (seedlings); 7 seedlings to 69 cm tall examined (Garwood 1830B-D, 2085B-C, & 2439). Fruits Infructescences terminal; loosely branched panicle-like thyrse to about 22 cm long, with 1-3 fruits usually maturing per branch. Rachis to about 22 cm long; branches to about 5 cm long; pedicels 1.5-2.0 cm long with 5 large glands about 2.5 mm tall below calyx scars; all stems densely scaly. Recep- tacles undeveloped. Stipes to 1 mm long, straight, circular in t.s., scaly, remaining attached to capsule. Fruits capsules, dry (with slightly fleshy seeds); from superior ovaries, with corolla and sepal scars inconspicuous among 5 large, extra- floral glands at base; carpels 2; locules 1. Capsules papery to woody, indehiscent or tardily loculicidally dehiscent; partially septate; 15-30(45) x 15-30 x 10-25 mm, (including spines); straight; circular to transversely elliptic in outline, circular in t.s.; margins entire; base widely rounded to truncate; apex widely rounded to truncate, with a terminal cluster of tightly packed short spines; densely covered with large, stiff spines, to 5 mm tall, with enlarged bases usually touching; surface medium to dark brown to red-brown, dull, with short weak spines < 0.5 mm tall among bases of larger spines; sparsely to moderately scaly, especially around bases of larger spines; scales peltate, orange-brown to brown, ± sessile to stalked. Seeds 6-12, central, touching along 1-2 sides, slightly deformed by neighbours; long axis of seed oblique to central axis of fruit. Placentae parietal; each a 2 mm wide dark band on lower half of endocarp, with seeds in 2 rows, and dark band narrowing above seeds and continuing to apex. Funicle about 2 mm long, curved to coiled, cylindrical, persistent on placentae, flared apically into shiny, 2-lobed appendage below seed, 1.5-2 mm long, each lobe a series of undulating compressed folds. Septa marginal, extending full-length of fruit but < '/4 distance to centre, about 2 mm tall throughout its length, solid basally but perforated apically, located under placenta but hidden by endocarp. Valves 2, dehiscing from apex if tardily dehiscent, spreading apart slightly, remaining attached at base. Fruit wall thin, about 0.5 mm wide (exclud- ing larger spines), 3-layered, endocarp separating from outer fruit wall at maturity. Exocarp very thin, about 0.05 mm wide, ± crustaceous, dark brown. Mesocarp thin, about 0.25 mm wide, forming most of septa, ± cartilaginous from numerous reticulate, tough fibres, which continue into bases of larger spines; spaces between fibres filled by ± papery matrix, which is compressed, shiny and mottled with orange- brown on inner surface, medium brown with cream-coloured fibres. Endocarp very thin, *s 0.05 mm wide, dry-membran- ous, dark brown to orange-brown; inner surface rough, glabrous, dull, dark brown to orange-brown. Seeds Seeds 3.7-5.1 x 3.5-4.3 x 3.0-3.8 mm; sarcotestal; anatro- pous; widely obovate or widely obtriangular in outline, compressed except subcompressed apically, mostly ± elliptic in t.s., with both faces often concave; margins entire; base moderately acute; apex truncate; apical cap over chalazal foramen moderately raised, circular, ~ 2 mm wide, with light brown outer ring and black centre, both easily dislodged; deep medial groove extending > 3/4 length of seed, from hilum to apex along raphe; outer surface densely pusticulate when dry, papillate when moistened, rough between pustules at higher magnification, glabrous, slightly glossy when dry, medium orange-brown or red-brown when dry, papillae yellow when moistened; if sarcotesta eaten or removed, then outer surface smooth, smooth at higher magnification, gla- brous, glossy, medium brown to orange-brown, except dark black-brown at base around micropylar regions. Aril absent. The funicular appendage (see above), which Corner (1976) describes as a vestigial aril, remains firmly attached to the funicle. Sarcotesta hard when dry, fleshy when moistened; with large oil cells resin-like when dry, becoming soft and waxy when moistened, but drying firm and waxy; completely surrounding seed; initially medium brown to orange-brown 163 Fig. 1 Bixa urucurana Willd. Fruit: A, indehiscent capsule; B, forcefully opened capsule. Seed: C-D, two views; E, t.s. (through plane 'e' in F); F, medial l.s. (through plane T in E); G, oblique l.s. (through plane 'g' in E). Seedling: H, older seedling; J, cotyledon; K, first leaf. Saplings: L, young sapling; M, older sapling; N, leaf at about node 17; O, stipule at about node 24; P, stipule at node 45: R, gland at about node 40. Scale bars: black and white, 1 mm units; white only, 1 cm units. Symbols: outer to inner layers of testa (tsl-ts3) and legmen (tgl-tg3); vascular trace (v); chalazal plug (cp); endosperm (shaded); and cross-sectional planes for E, F, and G (dashed lines: e, f and g, respectively). 164 N.C. GARWOOD when dry, becoming yellow or orange-yellow when moist- ened (and remaining so upon redrying); initially ± pusticu- late when dry, becoming ± papillate when moistened (and remaining so); developing from entire testa. Hilum subbasal in medial groove, exposed, slightly recessed to flush, some- times raised around end of vascular bundle, medium orange- brown, slightly lighter than surrounding sarcotesta, black- brown if tegmen exposed, usually indistinct and ill-defined, sometimes distinct, elliptic and = 1 mm long when much of sarcotesta adheres to funicle, often with irregular patch of tegmen exposed over the micropylar region. Micropyle basal, not visible. Chalazal foramen apical, not visible until apical cap removed, conspicuous on tegmen, round, —1.25 mm wide; closed by black chalazal plug, flat-topped, ± shallowly conical, « 0.5 mm wide, on a brown, — 1.5 mm wide disk, an extremely complex structure formed from specialized chalazal tissue (hypostase) and tegmen. Endosperm thick, encasing embryo, hard, rapidly becoming soft once moist- ened, starchy (Corner, 1976), cream-coloured to white. Embryo large, to 5 mm long, central, with cotyledons in plane oblique to medial plane through raphe, straight except abruptly curved at apex of cotyledon under chalazal plug, spathulate, flattened, yellow or yellow-green; axis well- developed, completely exposed; cotyledons well-developed, flat, often slightly curved, thin, ovate in outline, cordate at base; plumule rudimentary. Vascular bundle lateral, unbranched, very thin, sometimes exposed at hilar end, passing through sarcotesta in the medial groove of the raphe, sometimes visible through sarcotesta, terminating apically in chalazal cap. Seed coat thick, mostly 0.125-0.25 mm wide, except to 1.0 mm wide over chalaza, bony, with fleshy sarcotesta, exotegmic. Testa thin when dry, thicker in medial groove, ± thick when moistened, resin-like when dry, a fleshy sarcostesta when moistened, mostly 1-layered except 3-layered at apex around chalazal foramen; outer layer very thin, crustaceous, only evident as apical cap over chalazal foramen, light brown with black centre; middle layer (see sarcotesta above) less developed beneath apical cap; inner layer thick, bony, dark-brown, only evident below apical cap, tightly appressed to tegmen and lining the chalazal foramen. Tegmen thick, bony, quickly becoming leathery during imbi- bition, impermeable on outer surface but highly permeable on inner surface, 3-layered; outer layer thick, ± bony, cream-coloured to light grey, inflexed apically forming cha- lazal foramen, with a very thin dark brown inner band that becomes thick at base around micropylar area and forms a thick dark internal ring around chalazal foramen; middle layer very thin, ± papery, light brown, except thick and spongy around chalazal plug, splitting during imbibition and partially adhering to outer and inner layers; inner layer very thin, ± papery, medium brown, except ± thick, bony and black-brown beneath chalazal plug, separating from middle layer during imbibition, becoming leathery (except under chalaza), and adhering to endosperm. Seedlings and saplings Seedlings phanero-epigeal with cotyledons haustorial during germination, then photosynthetic. DEVELOPMENT. Hypocotyl elongates into loop, the cotyle- dons emerge from seed, sometimes not fully escaping seed. Hypocotyl straightens. ROOTS. Primary root circular in t.s., curved to sinuous, light brown; surface texture and colour between hypocotyl and root change slightly and gradually around collet. Secondary roots sinuous, scattered along primary root. Root hairs not evident. Sapling root system: taproot long-tapering, very thick for much of length; secondary roots mostly very thin and common, a few moderate-sized; rootlets very fine to fine and numerous. STEMS. Hypocotyl 34-38 x 1-2 mm, circular in t.s.; smooth, drying longitudinally wrinkled; glabrous; green. Epicotyl 1-5 x 0.5-2 mm, circular in t.s., smooth, sparsely minutely scaly, green. Scales peltate, ± sessile, erect, light orange-brown. Cataphylls absent. Internodes 1-10, 3-17 mm long; later internodes to 40 mm; circular in t.s.; smooth; glands absent on internodes 1-10, present below the stipule scar and well-developed on most internodes ^ 20, secreting nectar 3-4 internodes below youngest leaf, to 1 mm tall, widely obovate, with apex rounded to obcordate, light brown-green when fresh, drying light brown on glabrous raised pad twice width of gland and darker brown than surrounding stem; sparsely scaly at internodes 1-5, moderately scaly through internode 20, densely scaly at later internodes; slightly pusticulate at early internodes, often moderately to densely pusticulate at later internodes, pustules often still filled with dark orange- brown fluid on dry specimens. Scales peltate, ± sessile, light orange-brown, — 0.05 mm in diameter at first internode to 0.10 mm in diameter at later internodes, ± circular with entire to irregular margins. Older stems dark brown-green to dark grey-brown, solid basally, often hollow apically, shal- lowly furrowed, rough and often glabrous with age as initial bark surface fragments; wood cream-coloured. Lenticels first appearing on older stems — 2-5 mm wide, common on older stems 6-8 mm wide; mostly — 0.5 mm long, raised, initially elliptic to round, widening with age, cream-coloured on darker stems. Buds: Cotyledonary buds hidden or slightly protruding to 0.2 mm out of axils, slightly protruding above cotyledon scars, glabrous. Lateral buds protruding — 0.2 mm out of axils at early nodes, to 1 mm at later nodes; glabrous at early nodes to moderately or densely scaly at later nodes; enclosed within at least 2 widely to very widely triangular scale leaves. Terminal buds covered by stipules, glabrous at nodes 1-10, gradually becoming densely scaly at later nodes. COTYLEDONS. Two, opposite, foliaceous, simple, petiolate, estipulate, emergent from seed, spreading apart, persistent until at least node 5 matures. Blades 15-18 x 8-10 mm; straight, flat, ovate; apex acute to narrowly rounded; base ± cordate; margins entire. Blades densely thick-lined below, each to 0.5 mm long, parallel to long axis of cotyledon, light brown-orange, faintly pellucid and yellow-brown against strong light; faintly pellucid thick-lined above against strong light, otherwise scarcely visible; glabrous above and below; green. Venation palmate, 3-veined; brochido-actinodromous, basal and perfect. Primary veins 3; fine to moderate-sized; moderately raised above when dry, prominently raised below; central primary vein moderate-sized, straight to sinu- ous apically; lateral primary veins basal, fine, > 3/4 length of blade, curved or slightly arched, diverging at narrowly acute angles, looped, obtusely to perpendicularly joined to second- aries of midvein. Secondary veins fine; 10-18 per cotyledon, 4-5 along midvein and 3-4 along each lateral primary vein; alternate along midvein, marginal along lateral primary veins; =s'/2 length of blade, longest centrally along midvein; arched; diverging at mostly narrowly acute angles; looped, SEEDLINGS, FRUITS AND SEEDS OF PANAMA variously joined to superadjacent secondaries; slightly raised above, slightly raised to flush below. Inter-secondary veins occasional. Tertiary veins reticulate within intercostal area, looped within marginal area. Petioles 2-3 mm long, half- circular in t.s., channelled, often warty along margins and at apex above. LEAVES. Alternate and spiral; simple; long-petiolate, stipu- late. Leaf blades 22-25 x 12-13 mm at first node; 25-40 x 14-20 mm at nodes 2-5; to 229 x 168 mm at later nodes; straight; ovate at nodes 1-5, becoming widely ovate at later nodes; apex long-acuminate at nodes 1-5, becoming acumi- nate or short-acuminate at later nodes; base cordate at nodes 1-20, usually becoming emarginate, truncate or widely rounded at later nodes; margins entire. Blades dark green above, lighter below; raised between the veins when fresh; papery; usually densely minutely light orange-brown or green-brown punctate on blade below at early nodes, becom- ing dark brown punctate at later nodes, sometimes with thick dark brown-orange lines on veins below at later nodes, each punctation usually 0.05-0.10 mm long, irregular, circular or ± elliptic in outline, usually yellow-green pellucid against strong light at early nodes to light orange-brown pellucid at later nodes, sometimes opaque and dark brown-orange at later nodes, often associated with short thin or thick pellucid canals, sometimes branched; sometimes moderately punctate above, but usually pellucid dots scarcely visible above except against strong light; glabrous to sparsely minutely scaly below on veins and blades at nodes 1-5, becoming moderately scaly at later nodes; glabrous to sparsely minutely scaly above on veins and blades at nodes 1-5, becoming mostly sparsely scaly at later nodes; glabrous along margin. Scales peltate, ± sessile, light orange-brown, mostly =£ 0.05 mm in diameter on blade, to 0.10 mm on veins below, ± circular with entire to irregular margins. Young leaves spreading open almost immediately, glabrous or sparsely scaly below at early nodes to densely scaly below at later nodes; initially densely pusticu- late above, with shiny coating above apparently secreted by pustules when leaf about l/4-3A expanded, surface becoming ± dull again when full-size, dark green-brown, becoming dark brown-green, rigid, descending, produced continually, often before previous 1-3 leaves full-size. Venation palmate, 5-veined; brochido-actinodromous, basal, perfect. Primary veins 5; moderate-sized; moderately raised above, promi- nently raised below; central primary vein straight; lateral primary veins basal, thinner than midvein, inner pair of lateral primaries >/2-3/4 length of blade, arched, diverging at moderately acute angles, looped, acutely to perpendicularly joined to secondary vein from midvein; outer pair < '/2 length of blade, arched, diverging at ± perpendicular angles, looped to diffusely looped, perpendicularly to obtusely joined to secondary vein from inner lateral primary vein. Secondary veins moderate-sized along midvein, fine along lateral veins; 4-6 along midvein and 2-3 along each inner lateral vein on leaves at first node, 4-8 along midvein and 2-4 along each inner lateral vein at nodes 2-5, 8-15 along midvein and 2-8 along each inner lateral vein at nodes ^ 12; alternate to subopposite along midvein, on outer (exmedial) side of lateral veins; == 1/3 length of blade, longest centrally along midvein; mostly arched, often hooked apically along midvein and distally along lateral primary veins; diverging at moder- ately acute angles; looped, sometimes diffusely looped cen- trally along lateral primary veins, mostly perpendicularly joined to superadjacent secondary vein; slightly raised above, 165 moderately raised below. Inter-secondary veins absent to occasional in intercostal areas along midvein. Tertiary veins looped within marginal area, ± reticulate within intercostal area at nodes 1-5, becoming straight-percurrent and oblique to midvein near margin and convex-percurrent near midvein at later nodes. Petioles 10-140 mm long, 10-20 mm long at nodes 1-5; straight, circular in t.s.; often drying channelled above, enlarged at base, pulvinate at apex, medium yellow- green to green with medium brown pulvinus; sparsely to moderately pusticulate, sparsely minutely scaly at early nodes becoming moderately scaly at later nodes, pulvini usually densely pusticulate and densely scaly at later nodes. Stipules paired; 2-5 mm long at nodes 1-5, to 19 mm long at later nodes; simple, narrowly triangular, with long acuminate apex; quickly deciduous, papery; glabrous at early nodes to densely scaly at later nodes; green becoming light-brown; leaving pair of conspicuous scars above petiole, perpendicu- lar to stem, each > 3/4 width of stem. SAP. Not evident in freshly cut leaves, bark, or roots, but slowly oozing out around central pith of cut stem, drying cream-coloured to medium orange-brown and resin-like. ODOUR. Not distinctive. ARCHITECTURE. Trunk axis monopodial. Initial axis ortho- tropic. No branches formed on plants =s 70 cm tall, but elongated lateral buds (to 4 mm long) on largest individual suggest that sylleptic branches might soon be produced. Leaves held ± horizontal to descending. Ecology PHENOLOGY. Central American B. urucurana flowers in the late rainy season September-December, with fruit develop- ing during the following dry season. Fruits from my Panama- nian collections were dark brown and dry in April and May. DISPERSAL. When ripe, the dry capsule is dark brown with stiff spines, and probably indehiscent (Baer, 1976) or at most tardily dehiscent. Seeds within the dry, indehiscent fruit are also dry and dark orange-brown to red-brown, but the sarcotesta becomes soft, papillate and bright yellow when moistened. Dispersal mode is unknown. Seeds of the genus Bixa are usually assumed to be bird-dispersed because of the bright red or red-orange sarcotesta visible in the dehisced capsule, but Baer (1976) notes there are no observations of birds taking the seeds. In the indehiscent fruits of B. uru- curana, the duller orange-brown to red-brown seeds are not visible at maturity and are unlikely to attract birds. Given its riverside habitat, suggestions of fish-dispersal (van Roos- malen, 1985) or water-dispersal (Baer, 1976) are not unrea- sonable. In addition, the spiny, dry, brown indehiscent fruits hiding pulp-covered seeds are reminiscent of Apeiba mem- branacea Spruce ex Benth. (Tiliaceae), which is mammal- dispersed. GERMINATION. Seeds of B. urucurana germinated slowly (22-38 weeks) at low percentages (5-12%) in both sun and shade in the growing house. Baer (1976) failed to germinate seeds of numerous samples of B. urucurana, although those from B. orellana sensu stricto germinated easily. Seeds col- lected from unopened fruit of the cultivated B. orellana sensu stricto have high moisture content (~ 45%, fresh weight basis [FWB]) and permeable seed coats (Goldbach, 1979), and germinate rapidly (< 1-3 weeks; Garrard, 1955; Goldbach, 166 N.C. GARWOOD 1979; Chin et al., 1984). When seeds are air-dried, they become impermeable, moisture content drops (=£ 10% FWB), and germination becomes sporadic over > 1 year. Under natural conditions, the chalazal plug must become loosened before imbibition and germination can occur, but this requirement can be circumvented if seeds are scarified by cutting off the micropylar tip of the seed coat below the radicle (Goldbach, 1979). The proportion of impermeable seeds in a collection varies from 0% (Garrard, 1955) to > 50% (Goldbach, 1979), depending on seed moisture content, period of drying, and possibly cultivar used. My collections of B. urucurana had low seed moisture contents (6-10%, FWB), suggesting that the seed coats were impermeable at time of dispersal. Seeds of B. orellana air-dried to about 10% moisture content (FWB) and stored for 12 months in the laboratory at 23°C retained high viability (95% of initial viability); those dried to about 4% moisture content had slightly lower viability (82% of initial values) over the same period (Goldbach, 1979). Baer (1976) also reports high viability after 1 year, whereas Garrard (1955) found viability was completely lost after 5 months. The latter may reflect the absence of seeds with impermeable coats in that collection. COCHLOSPERMACEAE HABIT AND DISTRIBUTION. Trees, shrubs, subshrubs or rhi- zomatous herbs. Tropical; two genera and 15 species, one genus and two species in Panama, and one species known from BCI. TAXONOMIC REFERENCES. Robyns, (1967ft), Dathan & Singh (1972), Corner (1976), Croat (1978), Poppendieck (1980, 1981). Cochlospermum Kunth HABIT AND DISTRIBUTION. Small trees, shrubs, or subshrubs. Pantropical, mostly South American; 12 species, two species in Panama, one species known from BCI. Cochlospermum vitifolium (Willd.) Spreng., Syst. 4(2): 406 (1827). Fig. 2. HABIT AND DISTRIBUTION. Small sparsely branched trees, 3-12 m tall, of dry to moist lowland forests or savannas, common in young secondary forest, and along frequently burned roadsides. Mexico to Bolivia, northern Brazil and West Indies; introduced into Paleotropics. COLLECTIONS. Panama. Colon: Gamboa, beginning of Pipe- line Road, Garwood & Lighten 1784A (F, PMA, adult vouchers); same locality, Garwood 2941 A (BM, fruit voucher); Barro Colorado Island: Garwood 3060 A (BM, seed voucher); 20 seedlings to 81 cm tall examined (Garwood 1784B-G, 2941B, 3060B). Fruits Infructescences terminal; sparsely branched panicle-like thyrses, with l(-2) fruit maturing per branch, capsules suberect to pendent. Stems woody, glabrous or pubescent; peduncles 2-5 cm long, rachis and branches about 3-20 cm long, pedicels 3-4 cm long. Receptacles undeveloped. Stipes absent. Fruits capsules, dry; from superior ovaries, with withered persistent sepals, to 28 mm long at base; carpels 5; locules 1. Capsules papery to woody, locucidally and septicid- ally dehiscent (outer and inner valves, respectively); partially 5-septate; (40-)70-90(-1 10) x 35-70 x 35-70 mm; straight; obovate or widely ovate or elliptic in outline, circular in t.s.; margins entire; base truncate to short-attenuate; apex trun- cate and centrally depressed; surface dark brown or grey, dull, densely longitudinally narrowly multi-ridged, moder- ately to densely minute- to short-pubescent; hairs simple, erect. Seeds numerous, intermediary, touching along com- pacted cotton-like hairs but seed bodies separate, compressed ± laterally; long axis of seed parallel or oblique to long axis of fruit; at maturity, free within locule. Placentae intruded parietal; each cylindrical, running ± full length of fruit, ~ 50-60 x 5-6 mm, with numerous large funicular mounds, minute ovule scars, and aborted ovules and seeds. Funicle to 0.5 mm tall, straight, widely to very widely conical, persistent on placentae. Septa marginal, extending '/2-3/4 distance to centre. Valves 10, 5 outer and 5 inner, dehiscing from apex, spreading apart, each outer valve overlapping margins of adjacent inner valves except open at apex, outer valves remaining attached at base or by thread-like extension of placentae or sometimes falling; inner valves eventually falling out. Fruit wall thin, about 1.0 mm wide, 3-layered, endocarp separating from outer fruit wall at maturity. Exocarp thin, ± woody, with tough closely spaced longitudinal fibres, dark brown or grey. Mesocarp thin on inner surface of outer valves, forming thin septa and thickened placentae, spongy to cobwebby, with conspicuous vascular fibres transverse to oblique and sinuous on valves and oblique and often branched on septa, cream to light brown. Endocarp very thin, forming 5 separate elliptic inner valves, dry-membranous, translucent, dull, light brown or orange-brown; inner surface with central longitudinal ridge and numerous ± reticulate ridges, glabrous, glossy. When fruit unripe, exocarp green. Seeds Seeds 4-5 x 3-4 x 1-2 mm; campylo-anatropous (Corner, 1976); C-shaped in outline, sometimes slightly coiled, circular in t.s.; margins entire; base eccentrically short-attenuate; apex rounded; outer surface rough except wrinkled in sinus, densely very long-pubescent except glabrous in sinus, with hairs to 14 mm long, erect, sinuous, white, fluffy and cotton- like, easily detached after dispersal; dull, mostly medium brown except red-brown or orange-brown or dark brown in sinus; exfoliating to reveal tegmen, then base rounded, outer surface smooth, minutely cellular at higher magnification, glabrous, glossy, black. Aril absent. Sarcotesta absent. Hilum basal and eccentric to subbasal, toward sinus, exposed, prominently raised on testa, not visible on tegmen after testa exfoliates, cream-coloured, lighter than testa, circular. Micropyle basal, centric, exposed but partially hidden by hilar mound, prominently raised on testa, only slightly raised on tegmen, ± conical to cylindric on testa, punctiform on tegmen. Chalazal foramen subapical in sinus, not visible on testa but conspicuous on tegmen, round, about 0.6 mm wide; filled by black chalazal plug, conical on 0.8 mm wide disk, similar in composition to tegmen. Endosperm thick, encasing embryo, ± hard, oily (Cronquist, 1981), white. Embryo moderate-sized, ~ 4-6 mm long, central, C-curved, ± spathulate when straightened, compressed; axis moderately SEEDLINGS, FRUITS AND SEEDS OF PANAMA 167 Fig. 2 Cochlospermum vilifolium (Willd.) Spreng. Fruit: A, green, undehisced capsule; B, dehisced capsule, side view; C, dehisced capsule, apical view; D, outer valve, with placenta; E, inner valve. Seed: F, complete seed with testal hairs; G, hairs removed to show surface of testa; H, testa removed to show surface of legmen; J, partial seed, testa removed to show chalazal foramen in sinus; K, l.s. of seed without hairs; L, t.s. through sinus. Seedling: M, young seedling; N, older seedling; O, cotyledon; P, first leaf. Sapling: Q, sapling; R, leaf at node 4. Scale bars: black and white, 1 mm units; white only, 1 cm units. Symbols: outer to inner layers of testa (tsl-ts2) and legmen (tgl-lg3); vascular Irace (v); hilum (h); micropyle (m); chalazal plug and foramen (cp and cf); and endosperm (shaded). 168 N.C. GARWOOD developed, exposed; cotyledons moderately developed, flat, thin, entire; plumule rudimentary. Vascular bundle lateral in sinus, unbranched, thin, exfoliating with testa; passing through raphe, terminating subapically in chalazal foramen. Seed coat very thick, bony, exotegmic. Testa very thin, ± thicker in sinus, crustaceous, loosely surrounding tegmen, often exfoliating, 2-layered except 3-layered in sinus; outer layer thin, ± crustaceous, light brown except red-brown or orange-brown in sinus, densely long-pubescent (see above); middle layer absent except ± thick in sinus, hard-spongy, cream-coloured; inner layer thin, ± crustaceous, black. Teg- men very thick, ± bony to glassy, 3-layered; outer layer very thin, waxy, ± clear; middle layer thick, bony to glassy, ± 3-banded, with cream then brown then black bands; inner layer thin except thicker under chalazal foramen, cream- coloured, membranous to spongy, with abundant red-brown compressed globules (gum cavities). Seedlings and saplings Seedlings phanero-epigeal with cotyledons haustorial during germination, then photosynthetic. DEVELOPMENT. Radicle emerges from hilar region and slightly elongates, as cotyledons absorb endosperm. Hypo- cotyl elongates into loop, then straightens, pulling cotyledons from seed. Cotyledons expand, sometimes not fully escaping seed, which is often carried upwards on tip of cotyledons. Cotyledons are usually fully expanded within 2 weeks of germination. Shoot rest period is short during the cotyledon stage, sometimes lasting < 1 week in sun. Shade-grown seedling usually die without advancing past the cotyledonary stage. Growth is continuous in sun, with a new leaf initiated about every 1-2 weeks, often before the previous leaf has reached full-size. ROOTS. Primary root ~ 30 x 1 mm, poorly developed in shade, circular in t.s., straight to sinuous, cream-coloured becoming brown; surface texture and colour between hypo- cotyl and root change slightly and gradually around collet. Secondary roots to 0.2-0.3 mm wide, sinuous, sparsely scat- tered along primary root. Root hairs appear quickly on primary root. Sapling root system: taproot with tuberous swelling (xylopodium), to 50 x 15 mm, forming 10-20 mm below collet, developing in first year, produced on some seedlings only 12 cm tall; taproot long-tapering below swell- ing, thin to moderate-sized; secondary roots thin to moderate-sized, few to common; rootlets fine, few to com- mon. STEMS. Hypocotyl 33-71 x 0.5 mm, circular in t.s.; slightly grooved apically below petioles; glabrous basally becoming densely minute-pubescent in apical '/2-V3; medium yellow- green, often medium red-brown in sun. Hairs simple, erect to ascending, curved, clear to cream-coloured. Epicotyl 9-17 x 0.5 mm, circular in t.s., densely minute-pubescent, medium yellow-green to dark green. Hairs simple, erect to ascending, curved or sinuous or coiled, clear to cream-coloured. Cata- phylls absent. Internodes 1-10, 2-17 mm long; later inter- nodes to 65 mm; circular in t.s.; densely pubescent on side above petioles but glabrous to sparsely minute-pubescent on other side, glabrous to sparsely pubescent at later nodes, with abundant pearl bodies on youngest internode; medium yellow-green to dark green. Hairs simple, erect to ascending, curved or sinuous or coiled, clear to cream-coloured. Pearl bodies minute, globular, 0.05-0.075 mm in diameter, erect, short-stalked, clear, usually drying orange-brown. Older stems medium grey-green then medium grey-brown, solid with small soft pith basally to hollow apically; wood cream to cream-brown, with ± white rays, porous. Lenticels appearing on stems > 3 mm wide, common on stems > 4 mm; «= 1/2 mm long, round, widening with age, slightly raised, dark brown, on lighter stems. Buds: Cotyledonary buds hidden in axils, usually inconspicuous above cotyledons scars, rarely protrud- ing on older stems. Lateral buds hidden in axils at early nodes to protruding out of axils at later nodes, glabrous to slightly pubescent, enclosed within at least 2 scale leaves. Terminal buds naked, covered by stipules from several nodes, moder- ately pubescent. COTYLEDONS. Two, opposite, foliaceous, simple, petiolate, estipulate, emergent from seed, spreading, persistent until nodes 2-7 mature (usually for 2-4 weeks, sometimes to 13 weeks). Blades 24-26 x 7-8 mm; straight, flat, narrowly ovate; apex narrowly acute or acute; base obtuse or emargin- ate or truncate; margins entire. Blades glabrous above and below; green above, lighter below. Venation pinnate; eucamptodromous, sometimes brochidodromous in apical third. Midvein fine; straight to slightly sinuous; impressed above except slightly raised apically when dry, moderately raised below. Secondary veins moderate-sized; 16-18; mostly alternate except opposite at 2-4 basal veins; Vj-^A length of blade, longest basally; slightly curved; diverging at narrowly acute angles; open, branching and terminating near or at margin; slightly raised to flush above, slightly raised to flush below. Inter-secondary veins occasional. Tertiary veins inconspicuous and reticulate. Petioles 4-5 mm long, straight, half-circular in t.s., moderately channelled, mostly glabrous below, densely minute-pubescent above, with abundant pearl bodies above. LEAVES. Alternate and spiral; simple; long-petiolate, stipu- late. Leaf blades 21-33 x 13-22 mm at first node; 18-67 x 12-70 mm at nodes 2-10; to 190 x 210 mm at later nodes; ovate to widely ovate at nodes 1-2, becoming widely ovate to very widely ovate at nodes 3-10, mostly widely ovate to depressed ovate at later nodes; unlobed to palmately 3-lobed or 3-cleft at first node, becoming 3-parted, with central lobe largest and 2 large basal teeth becoming increasingly lobe-like at nodes 2-20, often 5-parted at later nodes, with central lobe at least slightly larger; apex of lobes mostly acute or acumi- nate at nodes 1-10, often long-acuminate at later nodes; base retuse or emarginate or cordate at nodes 2-15, mostly cordate at later nodes; margins ± irregularly serrate at nodes 1-3, becoming ± doubly serrate at later nodes. Teeth irregularly spaced and variably sized at nodes 1-10, becoming ± regu- larly spaced at later nodes; mostly 4-15 per side at first node, to = 50 per side at nodes 1-10, and to > 100 per side at later nodes; margins mostly curved, proximal side longer than distal side; apex acute to short-acuminate; sinus angular. Blades green above, lighter below; flat between the veins when fresh; membranous; with pearl bodies common above and below on primary veins at nodes 1-10, especially basally above, and infrequent to common on primary and secondary veins at later nodes, but sometimes infrequent below; gla- brous on blade above and below, glabrous to sparsely short- pubescent along margin; glabrous to sparsely short-pubescent on primary veins below at nodes 1-10, but only at base of primary veins above; glabrous to moderately short-pubescent on primary and secondary veins below at later nodes, but SEEDLINGS, FRUITS AND SEEDS OF PANAMA 169 glabrous or sparsely short-pubescent above along primary vein and densely pubescent at base; with short pellucid canals along margin drying harder than blade, canals absent or light yellow-orange at early nodes to mostly red-orange at later nodes; similar red-orange canals often conspicuous through- out blade of young leaves at later nodes. Hairs simple, erect to ascending, straight to curved, clear to cream-coloured. Pearl bodies minute, 0.05-0.075 mm in diameter, globular, short-stalked to sessile, erect, clear to white on immature leaves, becoming red-orange to brown with age. Young leaves plicate (lobes conduplicate-induplicate), densely pubescent, light green, with red-brown petioles, rigid, erect then ± horizontal, produced continuously before or after previous leaf full-size. Venation palmate, 3- to 5-veined; actinodro- mous, basal, perfect and marginal. Primary veins 3 at nodes 1-3, 3 to 5 at later nodes; moderate-sized; impressed above, prominently raised below; central primary vein straight to slightly curved; lateral primary veins basal, diverging at moderately acute angles, straight to slightly curved, terminat- ing at apex of lobes, sometimes only slightly thicker than secondary veins at first node. Secondary veins moderate- sized; 4-8 on central lobe and 1-3 on lateral lobes on leaves at first node, 6-15 on central lobe and 2-8 on lateral lobes at nodes 2-10, to 28 per lobe at later nodes; mostly alternate; mostly l/o-lA length of blade, rarely to 2/3 at first node, longest centrally on centre lobe; slightly curved; diverging at moder- ately acute angles; open, terminating at margin in tooth or sinus; impressed above, slightly raised below. Inter- secondary veins occasional. Tertiary veins reticulate. Petioles 10-200 mm long, «£ 42 mm long at nodes 1-10; straight, circular in t.s.; shallowly channelled; glabrous below, densely minute- to short-pubescent above at early nodes becoming ± glabrous except densely pubescent apically at later nodes; with abundant pearl bodies above and below. Stipules paired; 0.3-1.8 mm long at nodes 1-10, to 5 mm long at later nodes; simple; triangular, keeled; apex acute to acuminate; decidu- ous to ± persistent, papery; short-pubescent along margin and at apex, with pearl bodies along margin; green becoming brown, with red-orange canals along margin. SAP. Resinous, drying orange-brown in stems and roots. ODOUR. Unpleasant in hypocotyl and roots. ARCHITECTURE. Koriba's model (Halle et al., 1978). Trunk axis initially monopodial and orthotropic to at least 81 cm tall; eventually sympodial, one of the 2-6 branches at each tier becoming erect (Halle et al., 1978). Branches orthotro- pic, ± verticillate (Halle et al., 1978). No major branches were produced on our seedlings, all «£ 81 cm tall. Proleptic branches, produced after apical meristem damaged, have 3-4 stipule-like scales toward base, produce several smaller leaves with fewer lobes than those formed before damage. Leaves held horizontal to descending. Ecology PHENOLOGY. Fruits mature during the late dry season, mostly after March in Panama (Croat, 1978), about 1-2(^4) months after peak flowering, while the tree is leafless (Pop- pendieck, 1981; Janzen, 1983; Bullock & Solis-Magallanes, 1990). Most seeds are shed before the rainy season begins (Poppendieck, 1980). DISPERSAL. When ripe, capsules are dry, with outer valves dark brown, spreading and alternating with yellow-brown membranous inner valves. Seeds are wind-dispersed floaters (Augspurger, 1988), being gradually liberated from the par- tially opened fruit (Poppendieck, 1981). In Costa Rica, seeds are eaten by spiny pocket mice (Janzen et al., 1990). GERMINATION. Seeds have hard, impermeable seed coats. The chalazal plug must become loosened before imbibition and germination can occur, which may occur rapidly by fires frequent in their secondary habitats or more slowly by other factors. Germination of freshly collected, untreated seeds was low in sun and shade (*£ 17%), asynchronous over a 35 week period, and bimodal: a few seeds (4-9%) germinated in the first 3 weeks in sun (9 weeks in shade), the rest (8-12%) after a gap of 10-12 weeks. In a later seed collection, a hot water pretreatment increased germination over controls in both sun and shade (50% vs 4%): germination was still asynchronous (but not bimodal) over the 23 week period and was slightly faster in sun (7 weeks) than shade (9 weeks). Ricardi et al. (1987) also report low germination (7%) in the first three weeks. Seeds stored at room temperature were > 95% viable after nearly 10 years (Garwood & Lighten, 1990), germinat- ing in 7-10 days without treatments (Garwood, unpublished data). ESTABLISHMENT. Seedlings are extremely shade intolerant and probably require large gaps to survive (Augspurger, 1984). DISCUSSION The gross morphology of seeds of Bixa urucurana Willd. is very similar to that of B. orellana L. (Dathan & Singh, 1972; Corner, 1976), including a complex chalazal plug, an embryo with cotyledons asymmetrically bent under the chalazal plug, and a sarcotesta formed by large pigment-containing cells. Seeds of the five species in the genus differ slightly externally in size and colouring of the testal pigments (Baer, 1976). The dry seeds of B. urucurana are red-brown to orange-brown (Macbride, 1941; Molau, 1983; this study); when rehydrated, seeds from my collections became bright yellow. Dry seeds of B. orellana are usually orange-red or red (Macbride, 1941; Corner, 1976; Molau, 1983); when rehydated, seeds from a commercial source (purchased at a Mexican-American gro- cery) turned bright red (Garwood, personal observation). These colour differences may reflect differences in the rela- tive amounts of bixin, a reddish oil-soluble pigment, and orellin, a yellowish water-soluble pigment, in the testa. As there were few seeds on other BM collections of B. uru- curana, I have not yet determined whether this trait is characteristic of the species. Seeds of Cochlospermum vitifolium (Willd.) Spreng. are similar in gross morphology to those described for C. religio- sum (L.) Alston (= C. gossypium DC., Poppendieck, 1980), including a complex chalazal plug, curved form with curved embryo, long testal hairs, and a thick tegmen (Dathan & Singh, 1972; Corner, 1976). Seeds of the other ten species in the genus are similar (Poppendieck, 1980). The unique and complex chalazal foramen and plug of the exotegmic seeds most strongly link the Cochlospermaceae and Bixaceae (Cor- ner, 1976); other shared characters and differences between the two families are reviewed by Poppendieck (1980) and Cronquist (1981). 170 N.C. GARWOOD Seedlings of Bixa urucurana Willd. and Cochlospermum vitifolium (Willd.) Spreng. share the following characters: cotyledons are phanero-epigeal, foliaceous, persistent, peti- olate, estipulate and glabrous; and leaves are simple, alter- nate and spiral, long-petiolate, stipulate, palmately veined, mostly ovate to widely ovate, and basally cordate at least at early nodes. Leaves of both species and cotyledons of B. urucurana contain idioblasts: these are discussed in more detail below. The two species differ in the following traits. The cotyledons of B. urucurana are ovate, palmately veined and conspicuously thick-lined (faintly pellucid) on the lower surface (see below), while those of C. vitifolium are narrowly ovate, pinnate-veined and lack lines or punctations, pellucid or not. All seedling and juvenile leaves of B. urucurana are entire and not lobed or parted, whereas the first leaves (eophylls) of C. vitifolium are irregularly serrate, usually 3-lobed or 3-cleft, and later juvenile leaves are 3- to 5-parted. Previously published short descriptions of seedlings of Cochlospermum vitifolium agree with our description, except that Duke (1969) reported the cotyledon venation as uni- nerved and Ricardi et al. (1987) did not detect stipules on the eophylls. These differences might be attributed to the fact that the primary and secondary veins are weak, but clearly visible under a strong light source, and the stipules fall off as the leaf expands, although a stipule scar is evident at the first node on my collections. A short description of the seedling of Bixa orellana L. (Duke, 1969) agrees with that given for B. urucurana, except that cotyledons are described as punctate- lineate and the illustrated cotyledons are widely ovate rather than ovate. Leaves of Bixa and Cochlospermum have branched unicel- lular secretory idioblasts and canals in the spongy mesophyll (Keating, 1970). In leaves of B. urucurana seedlings, I found irregular, circular or elliptic pellucid punctations, < 0.1 mm long, which were sometimes associated with pellucid branched or unbranched canals. Punctations and canals were scarcely visible on the blade above (except when back-lit), but were conspicuous on the blade below, light orange-brown or green-brown. When back-lit, punctations were usually translucent yellow-green at early nodes to light orange-brown at later nodes, but sometimes opaque and dark brown-orange at later nodes. On the cotyledons, the idioblasts are much larger, to 0.5 x 0.1 mm long, thick and elongate, more or less parallel to the midvein; they were scarcely visible on the upper surface (even when back-lit), but conspicuously light yellow-brown below, and translucent brown-orange when back-lit. In leaves of C. vitifolium seedlings, I found faintly pellucid canals along the margins, which were absent or light yellow at early nodes to red-orange at later nodes. Similar canals were sometimes conspicuous throughout the blade of young expanding leaves at later nodes. No canals were found in the cotyledons. Leaf pubescence differs greatly between the two genera (Keating, 1970; Poppendieck, 1980), with Bixa having multi- cellular peltate scales on the blade and veins of the leaf undersurface and Cochlospermum having unicellular simple hairs as well as multicellular globular hairs. On seedlings of B. urucurana, Dempsey & Garwood (1994) found that the peltate scales are much smaller and sparser than those on adults: they discuss the systematic significance of these onto- genetic changes. The presence of multicellular globular hairs in Cochlosper- mum vitifolium has not been noted in standard floras (Robyns, 1967fc; Croat, 1978; Poppendieck, 1981), although they have been briefly discussed in anatomical studies and monographs of the family (Keating, 1970; Poppendieck, 1980). On seedlings of C. vitifolium, I found that the multicel- lular globular hairs were 0.050-0.075 mm in diameter and most abundant on the youngest stem internode, and on stipule margins, petioles and primary and secondary veins of expanding leaves, but often persisted on mature leaves and nodes. Globular hairs on young leaves of adult specimens examined at the BM were in the same size range (Garwood, personal observation; see also Keating, 1970: fig. 4), although Poppendieck (1980: fig. 46) illustrates hairs up to 0.5 mm in diameter; these were initially abundant, especially at the base of the primary vein, but fewer were found on mature leaves. Poppendieck (1980) and Keating (1970) called these hairs vestigial, but they might be fully functional 'pearl bodies' as described by O'Dowd (1982). Pearl bodies are typically 0.5-3.0 mm long, uni- or multi-cellular, spherical or club-shaped with short stems, easily detached from the leaf, lipid-rich, colourless to opaque, lustrous and 'pearl'-like, and often produced on the youngest leaves or near extrafloral nectaries or domatia; they occur primarily in tropical taxa, particularly those from second-growth habitats (O'Dowd, 1982). Because they are rapidly removed by ants in the field as a source of food, pearl bodies are often absent or scarce on plants in their natural habitats and on herbarium specimens. Except for their small size, the globular hairs on C. vitifolium fit the morphological description of pearl bodies. Documen- tation of removal by ants or mites (given their small size) and characterization of the cell contents will be necessary to confirm their biological function. Poppendieck (1980) assumed that they contained the same clear gum or resin as the fluid-containing idioblast canals of the leaf: this has not been reported for pearl bodies (O'Dowd, 1982). Saplings of Bixa urucurana have large extra-floral nectaries below the stipule scar (Fig. 1R). I found glands were absent on the first ten leaf nodes above the cotyledons, but were well-developed and secretory above node 20. They actively secreted nectar when less than 3-4 nodes below the youngest leaf, then became inactive. Similar glands occur on adult specimens of B. urucurana and B. orellana examined at BM (Garwood, personal observation) and have been reported for all species of Bixa except B. excelsa Gleason & Krukoff (Baer, 1976). Ant attendance at similar glands on the fruit pedicel of B. orellana sensu lato increases the proportion of flowers maturing fruits (Bentley, 1977), because the species of ants present remove or deter flowering-eating insects. Ants attending the glands on the stems of saplings probably reduce leaf herbivory in a similar way. This is another vegetative trait of ecological importance not described in the standard floras (Macbride, 1941; Standley & Williams, 1961; Robyns, 1967a; Croat, 1978; Molau, 1983) or a recent field guide emphasizing vegetative characters (Gentry, 1993). The seedling characters discussed above do not provide additional evidence for a closer link between the Bixaceae and Cochlospermaceae. Shared seedling-specific traits are usually those common throughout the angiosperms, such as petiolate, estipulate, foliaceous cotyledons. The taxa often differ in less wide-spread seedling-specific traits, such as the presence of pinnate venation or linear punctations in the cotyledons. Many of the vegetative differences and similari- ties noted by Keating (1970) and Poppendieck (1980) for adult stages, such as idioblasts, secretory canals, pubescence and leaf lobing, are already manifest at the early seedling stage. SEEDLINGS, FRUITS AND SEEDS OF PANAMA ACKNOWLEDGEMENTS. I am especially grateful to Margaret Tebbs for illustrations, Simon Thornton- Wood for cross-sectional diagrams of seeds, Ruth Dempsey for composition of the figures, and Eduardo Sierra for technical assistance in Panama. I also thank C.J. Humphries for encouragement and advice, INRENARE for permis- sion to work in Panama, and the many people on BCI and at STRI who helped the Project in various ways but are too numerous to thank here individually. The Seedling Flora Project has been gener- ously supported by the National Science Foundation, the Natural Environment Research Council, the Smithsonian Tropical Research Institute, and The Natural History Museum. REFERENCES Augspurger, C.K. 1984. Light requirements of neotropical tree seedlings: a comparative study of growth and survival. J. Ecol. 72: 777-795. 1988. Mass allocation, moisture content, and dispersal capacity of wind-dispersed tropical diaspores. New Phytol. 108: 357-368. Baer, D.F. 1976. Systematics of the genus Bixa and geography of the cultivated annatto tree. Ph.D. thesis, University of California, Los Angeles. Bentley, B.L. 1977. The protective function of ants visiting the extrafloral nectaries of Bixa orellana L. (Bixaceae). J. Ecol. 65: 27-38. Bullock, S.H. & Solis-Magallanes, J.A. 1990. Phenology of canopy trees of a tropical deciduous forest in Mexico. Biotropica 22: 22-35. Chin, H.F., Hor, Y.L. & Mohd Lassim, M.B. 1984. Identification of recalci- trant seeds. Seed Sci. Tech. 12: 429-436. Corner, E.J.H. 1976. The seeds of dicotyledons . Cambridge. Croat, T.B. 1978. Flora of Barm Colorado Island. Stanford. Cronquist, A. 1981. An integrated system of classification of flowering plants . New York. D'Arcy, W.G. 1987. Flora of Panama. Checklist and index. Parts I and II. (Monogr. Syst. Bot. Mo. hot. Gdn 17-18). Dathan, A.S.R., & Singh, D. 1972. Development of embryo sac and seed in Bixa L. and Cochlospermum Kunth. J. Indian hot. Soc. 51: 254-266. Dempsey, R.E. & Garwood, N.C. (1994) A study of Bixa (Bixaceae), with particular reference to the leaf undersurface indumentum as a diagnostic character. Bull. not. Hist. Mus. Land. (Bot.) 24: 173-179. Duke, J.A. 1969. On tropical tree seedlings. I. Seeds, seedlings, systems, and 171 systematics. Ann. Mo. hot. Gdn 56: 125-161. Garrard, A. 1955. The germination and longevity of seeds in an equatorial climate. Gdns' Bull., Singapore 14: 534-545. Garwood, N.C. (in prep.) Survey of seed germination in Central Panama. & Humphries, C.J. 1993. Seedling diversity in the neotropics. NERC News (No. 27, October): 20-23. & Lighton, J.R.B. 1990. Physiological ecology of seed respiration of some tropical species. New Phytol. 115: 549-558. Gentry, A.H. 1993. A field guide to the families and genera of woody plants of northwest South America (Colombia, Ecuador, Peru). Washington, D.C. Goldbach, H. 1979. Germination and storage of Bixa orellana seeds. Seed Sci. Tech. 7: 399-402. Halle, F., Oldeman, R.A.A. & Tomlinson, P. B. 1978. Tropical trees and forests. An architectural analysis . Berlin. Janzen, D.H. 1983. Physiological ecology of fruits and their seeds. In O. L. Lange, P. S. Nobel, C. B. Osmond & H. Ziegler (Eds), Encyclopedia of plant physiology. New Series, 12C (Physiological plant ecology. Ill: Responses to the chemical and biological environment): 625-655. Berlin. Fellows, L.E. & Waterman, P.G. 1990. What protects Lonchocarpus (Leguminosae) seeds in a Costa Rican dry forest? Biotropica 22: 272-285. Keating, R.C. 1970. Comparative morphology of the Cochlospermaceae. II. Anatomy of the young vegetative shoot. Am. J. Bot. 57: 889-898. Macbride, J.F. 1941. Flacourtiaceae, Flora of Peru. Publs Field Mus. not. Hist. (Bot.) 13 (4): 5-52. Molau, U. 1983. Bixaceae. In G. Marling, B. Sparre & L. Andersson (Eds), Flora of Ecuador 20: 3-7. O'Dowd, D.J. 1982. Pearl bodies as ant food: an ecological role for some leaf emergences of tropical plants. Biotropical4: 40-49. Poppendieck, H-H. 1980. A monograph of the Cochlospermaceae. Bot. Jb. 101: 191-265. 1981. Cochlospermaceae. Flora neotrop. , Monogr. 27: 1-33. Ricardi, M., Hernandez, C. & Torres, F. 1987. Morfologia de pldntulas de drboles de los basques del estado Merida. Merida, Venezuela. Robyns, A. 1967a. Bixaceae. In R.E. Woodson Jnr, R.W. Schery & collabora- tors (Eds), Flora of Panama. Ann. Mo. hot. Gdn 54: 57-59. 1967 b. Cochlospermaceae. In R.E. Woodson Jnr, R.W. Schery & collaborators (Eds), Flora of Panama. Ann. Mo. hot. Gdn 54: 61-64. van Roosmalen, M.G.M. 1985. Fruits of the Guianan flora. Utrecht. Standley, P.C. & Williams, L.O. 1961. Bixaceae, Flora of Guatemala. Fieldi- ana Bot. 24(7): 65-67. de Vogel, E.F. 1980. Seedlings of dicotyledons: structure, development, types; descriptions of 150 woody Malesian taxa. Wageningen. Bull. not. Hist. Mus. Land. (Bot.)24(2): 173-179 Issued 24 November 1994 A study ofBixa (Bixaceae), with particular reference to the leaf undersurface indumentum as a diagnostic character RUTH E. DEMPSEY and NANCY C. GARWOOD Department of Botany, The Natural History Museum, Cromwell Road, London SW75BD CONTENTS Introduction 173 Materials and methods 174 Results 175 Discussion 176 Appendix 178 References . .179 SYNOPSIS. In the course of preparation of an account of Panamanian Bixa urucurana Willd. for the Seedling Flora Project, differences in leaf undersurface indumentum were found between adults and juveniles. We describe the pattern of variation of indumentum between species within the genus and between developmental stages within B. urucurana. We conclude that leaf undersurface indumentum is a useful diagnostic character for the adult stages of the genus but not the juvenile stages. INTRODUCTION Bixa is the only genus of the small woody family, Bixaceae which is widespread in tropical America and frequently cultivated throughout the tropics. The genus has often been treated as one highly variable species, Bixa orellana L. sensu lato (Eichler, 1871; Kuntze, 1891; Warburg, 1895; Standley & Williams, 1961; Robyns, 1967; Croat, 1978) or as five separate species (Macbride, 1941; Baer, 1976; Molau, 1983). Primarily fruit and indumentum characters have been used to distinguish different species. Linnaeus' (1753) protologue of B. orellana simply described the capsules as 'chestnut-like'. Willdenow (1809) described a second species, B. urucurana from Brazil, defining it by the 'white' scales densely covering the leaf underside. This species has frequently been reduced in rank to a variety of B. orellana, especially in Central America (Kuntze, 1891; Standley & Williams, 1961; Robyns, 1967). In 1831, Don published a description of B. platycarpa Ruiz & Pav. from Peru, which had flat, kidney-shaped fruits with very few bristles. Triana (1858) described B. sphaero- carpa from Nouvelle-Grenade (now Colombia), having scaly leaf undersides and spherical capsules, but this was later reduced to a synonym of B. urucurana (Pilger, 1925; Baer, 1976). Huber (1910) described B. arborea, having a kidney- shaped, dorsally compressed capsule with a wrinkled surface. Finally, B. excelsa Gleason & Krukoff was described in Gleason (1934), and is characterized by reniform capsules densely covered by glandular trichomes. Although the origi- nal descriptions of the three species restricted to the Amazon basin (B. arborea, B. excelsa, B. platycarpa) all refer to a ©The Natural History Museum, 1994 more or less kidney-shaped fruit, there are other defining features which separate them as species. Baer (1976) con- tends that they are diagnosable on fruit shape when this is more accurately defined. Other diagnostic characters include fruit ornamentation, dehiscence, life-form, and the size and density of trichomes on the undersurface of the leaf (see Table 1 for a summary of these). The primary taxonomic difficulty in the genus is separating Bixa orellana and B. urucurana. There is a high degree of variation between cultivars of B. orellana, especially in capsule shape since this has been the subject of artificial selection. Baer (1976) discusses possible intermediates or hybrids between B. orellana and B. urucurana, known pre- dominantly from Panama and Costa Rica. These apparently have trichomes of intermediate size and density, relatively short, subspherical fruits with slender spines and intermedi- ate numbers of seeds. The variation in leaf indumentum is our principal concern. The undersurfaces of leaves of all five species of Bixa have peltate scales, which are circular discs supported on short obconical stalks about 20 |o,m long. They are red-brown in colour and formed of an outer ring of cells with a central circular area which is more densely pigmented. In B. uru- curana, the scales are large, with several concentric rings of cells around the central core, irregular in outline and funneli- form. Its scales are also very densely packed, giving a tawny-brown appearance to the undersurface of the leaf. (Willdenow's (1809) original description of white scales prob- ably referred to a shiny appearance, observed due to reflec- tion of incident light at certain angles). Our study is based on herbarium material collected and 174 R.E. DEMPSEY AND N.C. GARWOOD Table 1 Summary of life form, fruit and scale characters of the five Bixa species. (Baer, 1976; Molau, 1983; Macbride, 1941) Species Fruit shape Spines Dehiscence Habit Trichome l.s. t.s. diameter density (|xm) (per mm2) B. arborea subreniform circular absent D tree 69 12 B. excelsa oblate circular stout, with glandular D tree 54 6 pubescence B. orellana oblate/cordate circular absent or slender D shrub 64 19 B. platycarpa circular flat absent D tree 63 50 B. urucurana circular/depressed oblate circular tapered I shrub 94 156 B. 'intermediates' oblate circular slender D shrub 80 59 Dehiscence: D= dehiscent, 1= indehiscent. Trees are above 30 m at maturity, shrubby species grow to about 10 m. Trichome diameter is a mean (n unknown) and density 'representative' (taken from Baer, 1976). grown in Panama for the Seedling Flora Project (see Garwood & Humphries, 1993). Accessions from this area have often been attributed to Bixa orellana L. sensu lato (Robyns, 1967; Croat, 1978). Our adult collections seem to refer to B. urucurana on the basis of both fruit and indumentum charac- ters. However, we observed fewer and smaller scales on juvenile stages of the progeny of this adult (see description in Garwood, 1994) and felt that these observations merited further investigation. We pursued three main lines of enquiry: A) Within B. urucurana, are there significant differ- ences in scale diameter and density at different positions across the lamina of the leaf or between different stages of development? How is variation in scale diameter apportioned between the central core and outer annulus of the scale? B) Do differences in scale density and diameter within juvenile individuals follow any trends associated with either leaf node or size? and C) How much variation in scale density and diameter is there between adults of the five different species of Bixa across their geographical range and is there any evidence for the existence of intermediates between B. orellana and B. urucurana'] MATERIALS AND METHODS Seeds were collected from one individual of Bixa urucurana along the Chagres River, Gamboa, Panama in May 1986 and April 1987. These seeds were germinated in a screened growing house on Barro Colorado Island (BCI), Panama. The resulting juveniles were harvested at intervals and pressed to provide herbarium specimens of various stages of development. A voucher specimen was also collected of the adult in May 1986. Two Bixa seedlings were collected from a beach on BCI, but the parental seed source was unknown. A) To determine whether significant variation occurred in Bixa urucurana in scale size and density between different positions on the lamina, and between stages of development, and to ascertain where scale diameter variation arises (whether central core or outer annulus), we took an initial sample of two leaves from each of five juveniles and one of the beach seedlings. This was also undertaken for two leaves of the adult voucher specimen. Scale density was measured as the number of scales per mm2 with a photograticule on a Leica dissecting microscope. Ten measurements were made at each of three positions across the lamina; at the edge, in the middle and adjacent to the primary vein. Scale diameter was measured using an eyepiece graticule in a compound microscope at x 125 magnification. Small squares (about 1 mm2) of leaf tissue cut from the three positions described above were placed on a glass slide and the diameter of both the dark, central portion and the entire scale were measured for ten scales at each position. The results were analyzed as follows: i) two-way (Model II) analysis of variance (ANOVA) was used to partition the variance in scale density, scale core diameter and total scale diameter into components attributable to lamina position, leaf and position x leaf interaction effects, ii) three-level nested ANOVA (Model II) was employed to partition the three scale variables into the effects of stage, individual and leaf. Variance components were calculated by the methods of Sokal & Rohlf (1981). These results and those of the following investigations were all analyzed using SYSTAT (Wilkinson, 1990). B) To determine whether variation in scale density and diameter varied with leaf node and size within an individual, two of the largest juveniles were further studied, repeating the above measurement procedures, for edges of leaves only, for all leaves available on the specimens. The leaf length and width and the node from which the leaf originated were recorded. From this data, means and standard errors were calculated for each variable at each node and the results plotted. Relationships between scale diameter, scale density, leaf node and width, were tested by a linear regression for multiple values of y per x: resulting regression mean squares were tested over the deviation mean square and the propor- tion of variance attributable to the linear regression was evaluated by the coefficient of determination (r2) (Sokal & Rohlf, 1981). The relationship between leaf-length and width was quantified using the Pearson product-moment correlation coefficient (r). C) To estimate the variation in scale size and density in the genus Bixa as a whole, a selection of specimens in the BM herbarium was chosen (see Appendix). Five edges of leaves were measured for scale density and diameter in the same manner as described above, for one leaf of each of the chosen specimens. Specimens of B. orellana from Central America, South America and the West Indies and B. urucurana from Central and South America, were chosen from those with reliable recent determinations or mature fruits. For the other three Amazonian species, all specimens with leaf undersur- faces uppermost were studied, including several type speci- mens, since little material was available. Mean scale density and mean total scale diameter were calculated for each of these specimens. STUDY OF BIXA (BIXACEAE) 175 Table 2 Results of two-way ANOVA (Model II) of effects of position (edge, middle or vein) and leaf on scale density and diameter. Two leaves on each of five juveniles and one adult were included. Scale diameter Scale density Total scale Central core Source DF MS MS MS Leaf 11 254.23 221.07 *** 91.00 3.05 3.70 ** 12.76 6165.03 311.66 94.39 Position 2 2.94 2.55 ns .16 .61 .74 ns 0.00 74.59 3.77 .21 Leaf * position 22 1.15 1.50 ns .41 .82 1.75 6.08 19.78 1.83 .41 Error 324 .77 8.31 .47 81.16 10.82 4.98 DF, degrees of freedom; MS, mean squares; F, F-statistic; P, probability: *, 0.01 < p =s 0.05; **, 0.001 < p « 0.01; ***, p « 0.001; ns, not significant; %, percent of total variance contributed by variance component of each source. Table 3 Results of three-level nested ANOVA (Model II) for stage (adult vs juvenile), individuals within each stage, and leaves within individuals, of scale diameter and density. Scale diameter Scale density Total scale Central core Source DF MS MS MS Stage 1 913.90 253.65 *** 96.11 3.01 1.41 ns 3.97 17184.20 48.45 ** 88.55 Individual 4 3.60 1.16 ns .08 2.13 3.92 ** 13.03 367.66 .68 ns 0.00 Leaf 6 3.14 3.69 ** .81 .40 .72 ns 0.00 542.11 44.34 *** 9.30 Error 108 .85 3.00 .55 83.01 12.23 2.15 DF, degrees of freedom; MS, mean squares; F, F-statistic; P, probability: *, 0.01 < p =£ 0.05; **, 0.001 < p =£ 0.01; ***, p =£ 0.001; ns, not significant; %, percent of total variance contributed by variance component of each source. RESULTS A) For scale density in Bixa urucurana, a two-way ANOVA (Table 2) indicated a significant effect of position, leaf and position x leaf interaction. However, the greatest part of the variance is accounted for by differences between leaves (94%), and a very small portion is attributable to position on the lamina or the leaf x position interaction. For total scale diameter, position and the interaction term are not signifi- cant, whilst most of the variation (91%) is again accounted for by differences between leaves. While leaf and leaf x position interaction effects were significant, most of the variation (81%) in core diameter was among replicate scales on leaves. Since position did not appear to be an important effect, further sampling and analysis was carried out on the edge of the leaves only, as this was easier and caused less damage to the specimens. The results of the three-level nested ANOVA are summa- rized in Table 3. There is significant variance in scale density and total scale diameter between adult and juvenile speci- mens of Bixa urucurana, and between leaves, but not between individuals. Most of the variation of total scale diameter (96%) and density (89%) is attributable to differ- ences between adults and juveniles. The results for central core diameter are quite different. Only the difference between individuals is significant and most of the variation (83%) occurs within replicates on leaves. Since there was so little variation in central core diameter, total scale diameter alone was used in further analyses. B) The variation of scale density, scale diameter and leaf width with node for the two largest juveniles is shown in Fig. 1. Leaf age decreases with node number: because the young- est leaves at the highest nodes are still expanding they have a small width and a very high scale density and were excluded from the analysis. For fully expanded leaves, of both indi- viduals, Garwood 1830D and Garwood 2085D, mean scale density increases with leaf node (F(1 12) = 6.601, p = 0.025; F(1 7) = 41.730, p = 0.001, respectively), although the linear regression accounts for only a proportion of the total varia- tion (r2 = 0.244 and 0.588 respectively). The increase in scale density is independent of leaf width: a linear regression of scale density on leaf width was not significant in either individual (F(1>12) = 0.794, p = 0.500; and F(K7) = 1.391, p = 0.500). We tested only leaf width, because leaf length was highly correlated with width in both individuals (r(12) = 0.892, p = 0.001; and r(7) = 0.916, p = 0.001). Scale diameter does not appear to have any significant trend associated with leaf node or size in either individual (F(1 12) = 3.966, p = 0.100 and F(1 7) = 0.633, p = 0.500 for node; F(1.12) = 0.148, p = 0.250 and F(1 7) = 0.005, p = 0.75 for width) and the proportion of variation accounted for by the linear regression was very low (r2 =£ 0.039 for node; r2 =£ 0.002 for width). The leaf dimensions of our juvenile specimens (11.5-22.6 cm long x 8.7-16.7 cm wide) exceeded the range of our adult speci- 176 R.E. DEMPSEY AND N.C. GARWOOD mens (9.4-13.6 cm long x 5.0-9.7 cm wide) and are well within the published ranges for adults of the species (5.5-27 cm long x 3-18 cm wide), (Standley & Williams, 1961; Robyns, 1967; Molau, 1983). C) Bixa orellana and B. urucurana have the least certain specific status in the genus. On a plot of mean scale diameter versus mean scale density (Fig. 2a) specimens attributable to these taxa cluster into two groups (see Appendix for list of specimens), the lower being B. orellana and the upper B. urucurana, with two exceptions. Firstly, the Panamanian Fendler 328 specimen ('F' in Fig. 2a), the BM specimen of which consists of a mature B. urucurana fruit and a separate small non-fruiting branch, clearly groups with B. orellana. Baer (1976) placed a specimen from the same collection in B. urucurana. Secondly, our juvenile specimens of B. urucurana from Panama and the unidentified Bixa sp. seedlings from BCI clearly group at the lower end of the B. orellana range. In contrast, the (adult) parent is clearly in the B. urucurana group. South American specimens of both Bixa orellana and B. urucurana group at the extremes of the ranges of both mean scale diameter and density. The Central American specimens of both taxa group towards the centre of the overall range of indumentum characters. West Indian specimens of B. orel- lana group closely with South American representatives of the same taxon. Although there are too few specimens to form a reliable cluster, the three Amazonian tree species (B. arborea, B. excelsa and B. platycarpa) also cluster with Bixa orellana (Fig. 2b). DISCUSSION There is considerable variation in the indumentum of leaf undersurfaces in the genus Bixa. There are also several different trends in this variation which need to be considered and explained before the utility of indumentum differences as a diagnostic character of species can be evaluated. Differences between positions on the same leaf were found to be much less significant than other effects (e.g. leaf, individual or age) for scale density and not significant for scale diameter. Baer (1976) stated that scale density increased near veins or at the base of the leaf, but we found little evidence for this in either adults or juveniles. Both scale density and scale diameter are radically differ- ent between adults and juvenile stages of Bixa urucurana. On average, adult scales are more than twice as large and about four times denser than those of the juveniles. There is a slight overlap, however, between the ranges of density in adults * E a & CD 1 o o Z CO 03 CO o CO 20 r 15 10 5 0 L 40 r 30 20 10 0 L 65 60 55 50 45 25 .. .00 ©00 ° o o o C) 30 35 Leaf node 40 i 45 20 15 10 5 0 L 40 30 20 10 0 65 60 55 50 45 40 25 v 30 35 Leaf node 40 Fig. 1 Relationship of scale diameter, scale density and leaf width to leaf node in juvenile specimens. (a) Juvenile Garwood 1830D. (b) Juvenile Garwood 2085D. (Open circles represent fully expanded leaves, filled triangles expanding leaves and open triangles 'borderline' leaves. Error bars for scale density and diameter represent the standard error of the mean and, where missing, the errors are smaller than the symbols. For scale diameter, the hatched line represents a non-significant linear regression. The linear regressions for scale density are significant (p =£ 0.025). Only fully-expanded leaves were included in the regression calculations.) STUDY OF BIXA (BIXACEAE) 177 E \ o o3 Q O o 0 0 G O o 00 o o 70 80 90 Diameter (pm) 70 80 90 100 110 120 Diameter (pm) Fig. 2 Scale diameter versus scale density in Bixa spp. Mean ± SE (N = 5 replicates on one leaf except for Garwood specimens marked by stars). (a) B. orellana and B. urucurana. (Filled circles represent South American collections, hatched circles West Indian collections and open circles Central American collections. Stars represent our Panamanian collections: A = Adult (N = 10 replicates from two leaves on one individual), J= Juvenile (N = 10 replicates from two leaves on each of five individuals) and S = Bixa sp. seedling (N = 10 replicates from two leaves on one individual). F = Panamanian specimen Fendler 328. (b) Five Bixa species. (Open circles and stars represent B. orellana and B. urucurana specimens from Fig. 2a; squares, B. arborea; pentagons, B. platycarpa; diamond, B. excelsa). (27-65 scales per mm2) and juveniles (2-35 scales per mm2). There is no overlap between scale diameter of the juveniles (27.6-73.6 u,m) and the adults (82.8-147.2 |xm). The difference in total scale diameter between adults and juveniles is accounted for by the outer portion or annulus of the scale: there is no significant difference in central core diameter between stages. In Bixa urucurana, the scales on adult leaves have a number of concentric rings of colourless cells around the dark central core (Baer 1976). Evidently, it is an increase in the number of these rings which increases overall scale size in the adults of this species. Although we observed these annuli in juvenile specimens, they were generally very narrow, comprising perhaps only one or two rings of cells and were occasionally completely absent, espe- cially in the very young beach-collected seedlings. The second trend within Bixa urucurana is that scale density increases with node number in juvenile specimens, whilst there is no clear trend in scale diameter with node number. However, since we found a significant difference in scale diameter between adults and juveniles, it would appear that changes in this parameter (i.e. the increase in number of rings of cells around the centre of the scale) must occur at some point between the two stages of plant development we studied. Further study of a range of intermediate, sapling stages would be required to ascertain whether there is a gradual increase in scale size with node. Scale diameter is not low due to small leaf size, since our juvenile leaves are generally larger than the adult leaves examined. Thirdly, the comparison between our specimens and those of a sample from the BM herbarium shows that two distinct clusters of taxa are formed when mean scale diameter is plotted against mean scale density (Fig. 2). The lower group with small, sparse scales includes all the specimens of Bixa orellana sensu stricto. The upper cluster of individuals with large, relatively densely packed scales, contains specimens which were originally variously identified as B. orellana sensu lato, B. urucurana, B. orellana var. urucurana or B. sphaero- carpa (see Appendix). We interpret these as B. urucurana since they form a close-knit group on the basis of indumen- tum characters and their fruit characteristics correspond, being subspherical, spiny and indehiscent. The problematic specimen, Fendler 328 ('F' on Fig. 2a) which Baer identified as B. urucurana, clearly groups with B. orellana on the basis of indumentum characters. However, there is a mature, subspherical capsule on the herbarium sheet which looks like that typical of B. urucurana. On closer inspection, it was noticed that the vegetative shoot on the herbarium sheet was non-reproductive and had never borne flowers or fruit; therefore, it must have been collected either from a com- pletely different part of the same plant or even perhaps from a different individual than the fruit. The South American specimens of the two shrubby species, Bixa orellana and B. urucurana, group at the extremes of the range of indumentum characters (Fig. 2a). Although the Central American specimens still form recognizable clusters with the conspecific specimens from South America, their indumentum characters tend to fall into the middle of the overall range for the two species. It would appear from Baer's (1976) estimates of scale density and diameter (Table 1) that some of the Central American specimens fall into his 'inter- mediate' category. However, his scale density estimate for B. urucurana, 156/mm2, is inordinately high when compared to 178 R.E. DEMPSEY AND N.C. GARWOOD our measurements: our highest mean value was 70 scales/ mm2 and our highest single value was 85 scales/mm2. Unfor- tunately, because Baer does not give his sample sizes, it is difficult to compare the measurements. The fact that speci- mens of the two Central American species are much closer to each other than are those from South America, may suggest that there is some degree of intermediacy. Perhaps if we had sampled more specimens, we would have observed more of a continuum rather than two separate clusters. Since West Indian specimens of B. orellana group with South American specimens of the same taxon, it appears that dispersal (either naturally or through cultivation) has taken place directly from South America to the West Indies, bypassing Central America. We conclude that the differences between indumentum characters in B. orellana and B. urucurana adults distinguish them as two separate species. This is reinforced by differ- ences in fruit characters (see Table 1). The three Amazonian tree species cluster with Bixa orel- lana and do not appear to be separable from it on the basis of indumentum characters alone, although it is recognized that the sample studied here is small. It is apparent that Bixa urucurana manifests a broad range of scale diameter and density as it proceeds through its development. The stages in this process apparently encom- pass all the variation observed in B. orellana and the Amazo- nian species. That four of the five Bixa species and the juveniles of B. urucurana have small sparse scales suggests that perhaps the large, densely packed scales of adult B. urucurana represent a more derived state. The demonstration of the range of variation in indumen- tum characters through development within Bixa urucurana clearly shows that the utmost caution must be exercised when using them as diagnostic characters within the genus. It is important primarily to ensure that the leaf material studied comes from a mature individual. ACKNOWLEDGEMENTS. The Seedling Flora Project has been gener- ously supported by the National Science Foundation, the Natural Environment Research Council, the Smithsonian Tropical Research Institute, and The Natural History Museum. We thank Eduardo Sierra, principal technical assistant in Panama, INRENARE for permission to work in Panama, C. Humphries for comments on the manuscript, and the many people on BCI and at STRI who helped the Project in various ways but are too numerous to thank here individually. APPENDIX Table of specimens examined. Mean (SEM) Species Collector(s) No. Country Original ID Later ID Scale density Scale diameter (No./mm2) (M-rn) B. arborea Ducke 8311 Brazil A 13 (1.2) 68 (3.7) Huber 7827 Brazil A 3 (1.0) 74 (3.3) Krukoff 8551 Brazil A A (DB 76) 9 (1.7) 57 (7.9) B. excelsa Krukoff 6831 Brazil E 4 (0.5) 49 (1.8) B. orellana Barclay 425 Ecuador O O (UM '82) 10 (1.2) 64 (2.9) Barclay 851 Colombia O O (UM '82) 6 (1.3) 56 (4.2) Bunting & Licht 1070 Nicaragua O 26 (2.2) 51 (7.4) Hahn 104 Martinique O 15 (0.5) 44 (4.5) Hall & Bockus 7849 Colombia OO 27 (1.6) 66 (3.4) Hartman 12006 Panama 0 O (RL '84) 19 (1.8) 65 (4.2) Jenman 5268 British Guiana O 7 (0.8) 66 (3.4) Mexia 6435 Peru O 0 (UM '82) 16 (2.6) 74 (5.0) Nelson & Vargas 5072 Honduras O O (RL '91) 33 (4.9) 6 (4.5) Sounders 694 Honduras O 23 (2.2) 75 (6.8) Tenorio & Miller 3313 Mexico O O (FR '83) 21 (3.1) 63 (3.4) Wagner 519 Puerto Rico O 12 (0.8) 63 (1.8) Whitefoord 3169 Belize O 18 (1.6) 64 (2.9) Whitefoord 4592 Dominica 0 16 (2.3) 64 (0.0) Williams 579 Bolivia 0 11 (2.0) 60 (5.0) Williams 649 Bolivia O 19 (3.2) 48 (3.4) Yuncker et al. 8404 Honduras O 14 (3.0) 64 (2.9) B. platycarpa Klug 3040 Peru O P (UM '82) 24 (1.8) 68 (0.3) Klug 4116 Peru O P (UM '82) 27 (3.5) 67 (6.1) Pavon s.n. Peru p P (UM '82) 27 (2.8) 81 (7.9) Ruiz & Pavon s.n. Peru p P (UM '82) 15 (1.5) 77 (0.7) B. urucurana Fendler 328 Panama - U (DB 76) 29 (2.0) 52 (3.7) Garwood 1830a Panama O U (NG '93) 42 (2.2) 117 (3.5) Garwood (J) 1830b Panama 0 U (NG '93) 8 (0.8) 42 (1.9) Garwood ( J ) 1830c Panama O U (NG '93) 10 (0.6) 48 (1.8) STUDY OF B1XA (BIXACEAE) 179 Garwood (J) 1830d Panama 0 U (NG '93) 18 (2.1) 52 (1.0) Garwood (J) 2085c Panama 0 U (NG '93) 7 (0.4) 51 (1.5) Garwood (J) 2085d Panama o U (NG '93) 9 (1.1) 51 (1.6) Garwood et al. 613 Costa Rica 0 U (RD '94) 51 (4.0) 77 (4.7) Khan et al. 875 Costa Rica 0 U (RD '94) 42 (2.4) 81 (5.4) Krukoff 1069 Brazil ou U (RD '94) 64 (1.9) 101 (0.9) Marshall & Neil 6620 Nicaragua ou U (RD '94) 37 (1.3) 88 (6.9) Philipson et al. 1445 Colombia 0 U (UM '82) 61 (1.1) 110 (10.5) Pittier 4581 Panama OP U (RD '94) 50 (2.5) 110 (5-8) Proctor 32394 Honduras ou U (RD '94) 50 (1.3) 99 (5.4) Rombouts 177 Surinam 0 U (RD '94) 54 (1.9) 96 (9.5) Schmalzel 994 Panama 0 U (RL '88) 70 (5.6) 88 (8-5) Seymour s.n. Nicaragua 0 U (RD '94) 61 (5.0) 83 (5.8) Triana s.n. Colombia S U (DB 76) 56 (6.2) 116 (11.1) B. sp. Garwood (S) 2439a Panama 7 U?(NG '93) 4 (0.4) 44 (1-7) Original ID codes: A, Bixa arborea; E, B. excelsa; O, B. orellana; OO, B. orellana var. orellana; OU, B. orellana var. urucurana; P, B. platycarpa; S, B. sphaerocarpa; U, B. urucurana. Where original determinations have been confirmed or changed, this is noted in the 'Later ID' column, giving date and identifier: DB, D. Bacr; UM, U. Molau; RL, R. Liesner; FR, F. Ramos; NG, N. Garwood; RD, R. Dempscy. Mean scale densities and diameters are calculated from 5 edge measurements on one leaf for all specimens except Garwood 1830a, b, c & d, 2089c & d, and 2439a, for which means include 10 edge measurements on each of two leaves. (J), juvenile specimen; (S), seedling; (SEM), standard error of the mean. REFERENCES Baer, D.F. 1976. Systematics of the genus Bixa and geography of the cultivated annatto tree. Ph.D. thesis, University of California, Los Angeles. Croat, T.B. 1978. Flora of Barro Colorado Island. Stanford. Don, G. 1831. A general system of gardening and botany 1. London. Eichler, A.W. 1871. Bixaceae. In K.F.P. von Martius (Ed.), Flora Brasiliensis 13(1): 422-516. Garwood, N.C. ( 1994) Morphology and ecology of seedlings, fruits and seeds of Panama: Bixaceae and Cochlospermaceae. Bull. not. Hist. Mus. Lond. (Bot.)24: 161-171. & Humphries, C.J. 1993. Seedling diversity in the neotropics. NERC News (No. 27, October): 20-23. Gleason, H.A. 1934. Plantae Krukovianae III. Phytologia 1(2): 106-111. tluher, J. 1910. Novitates florae Amazonicae. Bolm Mus. Goeldi 6: 60-90. Kuntze, O. 1891. Revisio genera plantarum. Pars 1: 44-45. Leipzig. Linnaeus, C. 1753. Species Plantarum. Stockholm. 1957 facsimile. London. Macbride, J.F. 1941. Flacourtiaccae, Flora of Peru. Publs Field Mus. not. Hist. (Bot.) 13(4): 5-52. Moiau, U. 1983. Bixaceae. In: G. Marling, B. Sparre & L. Andersson (Eds), Flora of Ecuador 20: 3-7. Pilger, R. 1925. Bixaceae. In A. Engler & K. Prantl, Die natiirlichen pflanzen- familien, 2nd ed., 21: 313-315. Robyns, A. 1967. Bixaceae. In R.E. Woodson Jnr, R.W. Schery & collabora- tors (Eds), Flora of Panama. Ann. Mo. hot. Gdn 54: 57-59. Sokal, R.R. & Rohlf, F.J. 1981. Biometry. 2nd ed. San Francisco. Standley, P.L. & Williams, L.O. 1961. Bixa, Flora of Guatemala. Fieldiana Bot. 24(7): 65-67. Triana, J. 1858. Plantes usuelles de la Nouvelle-Grenade. Bull. Soc. hot. Fr. 5: 369. Warburg, O. 1895. Bixaceae In A. Engler & K. Prantl, Die natiirlichen pflanzenfamilien, 3rd ed. 6: 307-311. Wilkinson, L. 1990. SYSTAT: The System for Statistics. SYSTAT, Inc., Evanston, IL. Willdenow, K.L. 1809. Enumeratio plantarum horti regni botanici berolensis. Berlin. Bulletin of The Natural History Museum Botany Series Earlier Botany Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown, this may also be obtained from the same address. Volume 18 The typification and identification of Calymperes No. 1 An illustrated catalogue of the type specimens in the crassilimbatum Renauld & Cardot (Musci: Greville diatom herbarium. D.M. Williams. 1988. Pp. Calymperaceae). L.T. Ellis. 1991. Pp. 193-194, 1 fig. 1-148, 74 plates. £28.00 No. 2 No. 3 No. 4 No. 5 No. 6 Volume 19 Erik Acharius and his influence on English lichenology. D.J. Galloway. 1988. Pp. 149-194, 18 figs. £8.80 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Phodophyta (Florideae) 2. Genera G.J.H. Price, D.M. John & G.W. Lawson. 1988. Pp. 195-273, 1 fig. £14.80 Some Cretaceous and Palaeogene Trinacria (diatom) species. P. A. Sims & R. Ross. 1988. Pp. 275-322, 13 plates. £9.10 A monograph of Dryopteris (Pteridophyta: Dryopteridaceae) in the Indian subcontinent. C.R. Fraser-Jenkins. 1989. 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Studies on the lichen genus Slicta (Schreber) Ach.: II. Typification of taxa from Swartz's Prodromus of 1788. D.J. Galloway. Pp. 35-48. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 4. Genera L-O. D.M. John, G.W. Lawson, J.H. Price, W.F. Prud'homme van Reine and W.J. Woelkerling. Pp. 49-90. Studies on the Cretan flora 3. Additions to the flora of Karpathos. N.J. Turland and L. Chilton. Pp. 91-100. CONTENTS 101 Observations on the benthic marine algal flora of South Georgia: a floristic and ecological analysis DM. John, P.J.A. Pugh and I. Tittley 115 Studies in Pseudocyphellaria (Lichens) IV. Palaeotropical species (excluding Australia) D.J. Galloway 161 Morphology and ecology of seedlings, fruits and seeds of Panama: Bixaceae and Cochlospermaceae N.C. Garwood 173 A study of Bixa (Bixaceae), with particular reference to the leaf undersurface indumentum as a diagnostic character RE. Dempsey and N.C. Garwood bulletin of The Natural History Museui BOTANY SERIES Vol. 24, No. 2, November 1994