ISSN 0968-0446 Bulletin of The Natural History Museum THE NATURAL HISTORY MUSEUM PRESENTED GENERAL LIBRARY Botany Series THE NATURAL HISTORY MUSEUM VOLUME 28 NUMBER 1 25 JUNE 1998 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. *vl * * The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Dr S. Blackmore Editor of Bulletin: Ms M.J. Short •'•" 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. 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(Bot.) 28(1): 1-16 Issued 25 June 1998 Morphology and ecology of seedlings, fruits and seeds of Panama: Vochysiaceae THE NATURAL HISTORY NANCY C. GARWOOD ^ Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS 18 JUN 1998 pnr^cu*rgn> GENERAL LIBRARY Introduction Methods Vochysiaceae Vochysia ferruginea Mart Vochysia guatemalensis Donn. Sm Vochysia sp Key to known seedlings of Vochysia in Panama and Central America . Discussion ... Fruit Seed Embryo Seedling ................................................................................................................................................................................. 10 Phylogeny .............................................................................................................................................................................. 12 Ecology .................................................................................................................................................................................. 14 References ... ... 15 SYNOPSIS Seedlings, saplings, fruits and seeds of Vochysia ferruginea (Vochysiaceae) are described, and ecological data on these stages summarized, as part of the ongoing Seedling Flora Project centred on Barro Colorado Island, Panama. Seedlings of two other species reported from Panama are also described. The morphology of these stages is compared to that of other taxa in the family. The potential use of seed and seedling characters in phylogenetic studies within the family is discussed. Special attention is drawn to unusual two-parted cotyledonary stipules, ontogenetic changes in leaf hair morphology, and complex ptyxis and vernation of the embryonic cotyledons. 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;Garwood, 1994, 1995, in prep.). This is the account of Vochysia ferruginea Mart. (Vochysiaceae). It is the most abundant Vochysia in Panama, common in the younger forests of Barro Colorado Island and Central Panama; four other species are known, but only from a few collections (Robyns, 1967a, b\ Croat, 1978; D'Arcy, 1987). Ranging from Central America through the Amazon basin, V. ferruginea is also the most widespread species in the genus, in contrast to most species which have rela- tively limited distributions (Stafleu, 1948). Because of its fast growth rate and valuable timber, V. ferruginea has been identified as a potentially important forestry species for use in plantations or managed secondary forests in the neotropics (Finegan, 1992; Condit et al., 1993; Butterfield & Fisher, 1994). In this account, I describe the seedlings, saplings, fruits and seeds of V. ferruginea, and the seedlings of two other species known from Panama, then compare these stages to those of other species in the family to identify characters of potential phylogenetic importance. Fruit and seed characters have been important in delimiting the © The Natural History Museum, 1998 tribes and genera of Vochysiaceae; subgeneric classifications and keys to species, however, have been based almost solely on floral and vegetative characters (Warming, 1875; Stafleu, 1948-1954; Keay & Stafleu, 1953). There have been few comparative studies of the gross morphology or anatomy of fruits and seeds (Corner, 1976; Boesewinkel & Venturelli, 1987). Seedlings of few species have been described in detail (delAmo, 1979; Ricardi et al., 1987; Flores, 19930, b). METHODS Standard methods for germinating seeds, producing voucher speci- mens for each seed collection, growing and harvesting seedlings, collecting unvouchered seedlings (wildlings) from the forest, and manipulating the descriptive database were developed for the Seed- ling Flora Project (Garwood, 1994, 1995) and were used for this account. The detailed descriptions follow the standard format devel- oped for the Project. Descriptive terms have been standardized for use across all families in the Project, but generally follow common use. See de Vogel (1980) for definitions of seedling-specific terms and Corner (1976) and Boesewinkel & Bouman (1984) for seed- specific terms. To facilitate identifying isolated seeds, the base of the N.C. GARWOOD seed is arbitrarily defined as the part nearest the hilum, rather than using a reference point within the fruit. Leaf nodes of seedlings are numbered sequentially from the first leaf-bearing node above the cotyledons. The following less conventional terms are used: for length of pubescence or size of other surface features - minute (< 0. 1 mm), short (0. 1 -0.5 mm), long (0.5- 1 .0 mm), very long (> 1 .0 mm); for relative thickness of the seed coat or pericarp layers compared to total width of the structure - very thick (> 25% width), thick (10- 25%), thin (1-10%), and very thin (< 1%). VOCHYSIACEAE HABIT AND DISTRIBUTION. Trees or shrubs, rarely subshrubs. Neotropical except for one very small African genus; 6-8 genera and c. 200 species, 3 genera and 8 species from Panama, 1 genus and species known from Barro Colorado Island. TAXONOMIC REFERENCES. Stafleu ( 1 948), Keay & Stafleu ( 1 953), Robyns (1967a, b), Cronquist (1981), D'Arcy (1987), Kawasaki (in press). Vochysia Aubl. HABIT AND DISTRIBUTION. Trees, shrubs or rarely subshrubs. Neotropical; c. 100 species, 5 species in Panama, 1 species known from Barro Colorado Island. Vochysia ferruginea Mart, in Mart. & Zucc., Nov. Gen. sp. pi. 1: 151,1.92(1824). Fig. 1. HABIT AND DISTRIBUTION. Trees, 6-35 m tall, of moist to wet lowland and premontane forests, usually in secondary forest. Nica- ragua to Peru and Brazil. COLLECTIONS. Panama. Barro Colorado Island: Garwood 1 630A (F, BM, PMA, adult vouchers), Garwood 1630B-F (seedlings), Garwood 2005 (seedlings), Foster s.n. (seedlings with fruit voucher); 50 seedlings to 37 cm tall examined. Fruits Infructescences terminal and upper-axillary thyrses, of 1- to 5-flow- ered helicoid cymose branches, usually with < 1 fruit maturing per branch. Rachis densely short-pubescent, with hairs 2-branched, ± sessile to short-stalked, appressed to ascending, straight to sinuous; lateral branches (including pedicels) 8-15 mm long, slightly to moderately pubescent. Receptacles undeveloped. Stipes absent. Fruits dry capsules, from superior ovaries. Capsules ± woody, loculicidally dehiscent; completely septate; 17-26 x 8-12 x 8-12 mm; straight; oblong to obovate in outline, 3-lobed to 3-cleft in t.s., not com- pressed; margins entire; base rounded to abruptly truncate; apex rounded to slightly retuse, with stylar scar to 1 mm wide; surface dark brown to black-brown, dull, slightly to prominently warty, often irregularly clear- to white-scaly, glabrous. Seeds 1-3, 1 per locule, intermediary, separated by mesocarp and endocarp, com- pressed laterally; long axis of seed radially parallel to long axis of fruit. Placentae axile. Funicle absent. Valves 3, dehiscing from apex, slightly spreading apart then each folding back, remaining firmly attached at base. Fruit wall thick, 0.4-1.5 mm wide, thickest cen- trally, 3-layered. Exocarp very thin, = 0.05 mm wide, ± hard-papery, dark brown to black-brown. Mesocarp thick, 0.2-1.3 mm wide, thickest centrally between locules, hard-spongy to ± woody, with 2 large longitudinal resin-filled canals near axis between each pair of locules and several smaller chambers along locules (resin drying red-brown and glassy), light to medium brown. Endocarp thin, ~ 0. 1 mm wide, crustaceous, cream to light brown; inner surface smooth, minutely transversely fibrous, glabrous, glossy, medium brown-orange. When fruit unripe, exocarp grey-green. Seeds Seeds 20-25 x 5-6 x 1-2 mm; hilar-anatropous; 1 -winged; trans- versely half-obovate in outline, compressed and elliptic in t.s. in seminiferous area, planar in wing; margins constricted distally in seminiferous area on hilar side, constricted below wing on opposite side; base widely truncate; apex widely rounded; outer surface ± smooth to slightly rough, minutely rough under hairs at higher magnification, densely long- to very long-pubescent, with hairs sinuous, appressed, light to medium brown-orange; glossy, medium brown to brown-orange, dark brown under hairs; not exfoliating. Wing lateral, fibrous, ~ 15-17 x 5-6 mm, ± half-elliptic in outline, planar, asymmetrically rounded at apex, entire along margins, with curved ridge slightly raised, ± flattened, extending from base to 3/4 length of wing and forming thickened straight margin; formed from very long, interwoven, testal hairs. Seminiferous area lateral, ~ 10- 1 1 x 4-5 mm, ± elliptic in outline, compressed, darker than wings, with short fringe of interwoven hairs on side opposite wing. Aril absent. Sarcotesta absent. Hilum basal, along straight side, exposed, flush, light brown, lighter than testa, linear, 7-10 mm long. Micro- pyle subbasal to marginal (visible internally at tip of radicle, possibly continuing through thickened ridge of wing and terminating along straight side past hilum), concealed externally. Lens absent. Storage reserves absent. Embryo large, = 10-11 mm long, filling seminifer- ous area, straight, ± oblong or ellipsoid, compressed; axis well-developed, with long 4-winged hypocotyl separated from very short radicle by annular ridge at collet, half-exposed; cotyledons well-developed, thin, with compressed obliquely supervolute ptyxis and convolute (sensu imbricate) vernation, with wider side of each cotyledon angled down and partially enclosing hypocotyl, with shape obscured by folding and base hidden within folded blade; plumule rudimentary. Vascular bundle half-lateral on hilar side, unbranched, thin, with tip exposed at end of hilum and fragments of the funicular bundle often exposed in hilum; passing through short raphe, terminating marginally at tip of seminiferous area. Seed coat very thin, ± papery, undifferentiated at maturity. Testa very thin, ± thicker under hilum and around micropyle, ± papery, 1 -layered, medium to dark brown, densely long-pubescent (see above); inner surface irregular and minutely rough, glabrous, glossy, dark brown. Tegmen absent. Fig. 1 Vochysia ferruginea Mart. Fruit: A, undehisced fruit; B, dehisced capsule; C, t.s. Seed: D, seed, lateral view; E, embryo in seminiferous region, with seed coat partially removed; F, embryonic cotyledon and radicle, with one side of cotyledon unfolded and second cotyledon removed to show primary veins, radicle, and angle of folding; G, t.s. through seminiferous region at level marked by arrow in E-F, view toward radicle and wing. Seedling: H, seedling; J, cotyledon; K, cotyledonary stipules, hypocotyl, and petiole ridges; L-M, cotyledonary stipules after epicotyl expands; N, first leaf; O, stipules at first node. Sapling: P, young sapling; Q, branched sapling; R, leaf at node 15; S, stipules at about node 20. Scale bars: black or black and white, 1 mm units; white only, 1 cm units. Symbols: ac, annulate collet; ab, abaxial surface of cotyledon; ad, adaxial surface of cotyledon; ^-c,, cotyledons 1 and 2; en, endocarp; ex, exocarp; m, midvein of cotyledon; ms, mesocarp; r, radicle; re, resin canals; sd, seed; so, outline of seed (dotted line); ts, testa; f,-f4, folds of cotyledon. SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE N.C. GARWOOD Seedlings and saplings Seedlings phanero-epigeal with photosynthetic cotyledons. DEVELOPMENT. Radicle emerges from hilar side of seed near wing. Hypocotyl elongates into loop, then straightens, often carry- ing seed upward. Cotyledons emerge from seed and fully expand before first leaves are initiated. Shoot rest period is short during the cotyledon stage, about 1 month in sun and 3 months in shade. Seed reserves are exhausted by the end of this period, as albino seedlings die at the cotyledon stage without initiating any leaves. Growth is continuous, with a pair of leaves initiated about every month in sun and every 2-3 months in shade, although shade-produced leaves are smaller. ROOTS. Primary- root 30-50 x 0.6-0.8 mm, circular in t.s., slightly sinuous, cream to white, becoming medium orange-brown; narrow annular ridge encircles axis at collet, becoming indistinct with age; surface texture and colour between hypocotyl and root change conspicuously and abruptly at collet. Secondary roots to 0.2 mm wide, sinuous, very sparsely scattered along primary root. Root hairs appear abruptly and densely on primary root just below collet. Sapling root system: taproot thick, long-tapering; secondary roots ± thin and common along primary root; tertiary and higher order roots very fine and numerous. STEMS. Hypocotyl 15-67 x 0.25-0.50 mm, ± square in t.s., be- coming circular in t.s. and hollow with age; 4-ridged to 4-winged, each wing to 0.5 mm tall, continuing at apex into lower margin of cotyledon petiole and terminating at base abruptly above collet; glabrous; medium yellow-green to green, becoming dark brown or red-brown. Epicotyl 3-12 x 0.3-0.5 mm, ± square in t.s., becoming circular with age, 4-winged, each 0.3-1.0 mm tall and terminating apically in stipules, glabrous or sparsely short-pubescent, medium green to yellow-green. Hairs short, T-shaped (ratio of arm lengths = 1 : 1 ), ± sessile, appressed, straight, and light brown to orange-brown. Cataphylls absent. Internodes 1-10, 2-15 mm long; later internodes to 34 mm; similar to epicotyl except later internodes becoming densely short- to long-pubescent, with hairs long, unequally 2-branched (arm length ratio to 8:1), short-stalked, ascending, and curved or sinuous or coiled. Older stems dark brown, with wings often red-brown, hollow, often sparsely pubescent with age; wood cream-coloured. Lenticels appearing on stems > 2 mm wide, sparse on stems 2—4 mm wide; mostly 0.5 mm long, ± round to elliptic, light brown, lighter on dark stems. Buds: Cotyledonary buds hidden in axils and inconspicuous above cotyledon scars, rarely protruding to == 0.5 mm in older seedlings, moderately short-pubescent. Lateral buds usually slightly protruding out of axils, to 1 mm long at later nodes, moderately pubescent at early nodes to densely pubescent at later nodes, with accessory bud above sylleptic branches at later nodes. Terminal buds naked, partially covered by stipules, moder- ately short-pubescent at early nodes to densely long-pubescent at later nodes, with hairs light brown to orange-brown. COTYLEDONS. Two, opposite, foliaceous, simple, petiolate, stipu- late, emergent from seed, spreading apart, persistent at least until node 3 matures, falling sometime before node 9 matures (between 9 and 20 months). Blades 10-15 x 13-23 mm; straight, flat, depressed obovate, with right half (viewed from the lower abaxial side) usually slightly larger, 2-lobed to 2-cleft; apex of lobes obtuse to broadly obtuse; base emarginate or cordate, often asymmetrical with right lobe slightly larger, or half-rounded/half-cordate; margins entire. Blades glabrous; green above, lighter below. Venation palmate, 3-nerved; acro-brochidodromous, basal and perfect. Primary veins 3; moderate-sized; impressed to slightly raised above, moderately raised below; central primary vein forked, with straight stem '/2-3/4 distance to apex and branches slightly recurved; lateral primary veins basal, = 3/4 length of blade, straight to curved, diverging at narrowly acute angle, looped, joining branches of central vein to form large ± triangular to obovate intercostal area, the pair of intercostal areas usually spanning about 60-70% of the width of the cotyledons. Secondary veins moderate-sized basally along lateral primaries becoming fine apically; 16-24, 2-4 along midvein, 6-10 along lateral primaries, 6-10 along branches of midvein; mostly marginal, alternate along midvein; 'A>-3/4 length of blade (mostly 'A>- '/4 along branches of midvein), longest basally; mostly curved or arched along lateral primaries, hooked along branches of midvein, curved to zigzag along midvein, rarely scythe-shaped; diverging at narrowly acute angles (< 45°) basally on lateral primaries to perpen- dicular angles along branches of midvein; looped marginally, sometimes percurrent in intercostal area, mostly obtusely joined; slightly raised above, slightly raised below. Inter-secondary veins absent. Tertiary veins reticulate within intercostal area, looped in marginal area. Petioles 0.5-1 .5 mm long, transversely oblong in t.s., straight to slightly curved, 2-ridged above, with ridges continuing into lateral primary veins, 4-winged along margin, with lower wings continuing into hypocotyl wings and upper wings terminating in fleshy knobs near centre of hypocotyl, glabrous. Stipules usually fused and interpetiolar, sometimes separate and paired; composed of two structures, both variable in shape and development. The lower part usually a ± conical projection between the petioles at the base of the upper marginal ridge of each petiole, sometimes with two or three separate projections or these absent, mostly 0.125-0.25 mm long, usually perpendicular to the stem, sometimes gland-like apically. The upper part usually a flattened interpetiolar scale-like outgrowth just above the petioles, initially folded over the terminal bud, becoming erect and visible as the epicotyl expands, 0.25-0.5 mm tall and 0.5-0.75 mm wide, ± transversely oblong to depressed ovate, usually 2- to 5-lobed or -cleft, but sometimes divided into two separate parts or unlobed, the apex of lobes gland-like when young; persistent, papery, green becoming light brown. LEAVES. Opposite, decussate on trunk axis, distichous on branches; simple; short-petiolate, stipulate. Leaf blades 17-32 x 3-9 mm at first node; 20-67 x 5-20 mm at nodes 2-10; to 142 x 49 mm at later nodes; 65-125 x 25-37 mm on branches; straight; narrowly elliptic at nodes 1-3, narrowly elliptic or narrowly obovate at later nodes, elliptic or narrowly elliptic or narrowly obovate or linear-obovate on branches; apex narrowly obtuse or acuminate at nodes 1-10, acumi- nate to long-acuminate at later nodes and on branches, mucronate at most nodes; base mostly attenuate at early nodes, sometimes nar- rowly cuneate at later nodes, mostly acute on branches; margins entire, sometimes undulate. Blades green above, lighter below; flat between secondary veins above when fresh; papery; densely minutely pusticulate below (from internal crystals), but weakly so at nodes 1- 2; densely minutely pusticulate above, with pustules usually white on larger leaves; slightly to moderately short- to long-pubescent on blade and veins below at early nodes becoming densely long- to very long-pubescent only on the midvein at later nodes; glabrous on blade above, slightly to moderately ± short-pubescent on midvein at early nodes to densely long-pubescent at later nodes; slightly to moder- ately short- to long-pubescent along margin, often also densely minute-pubescent when young at later nodes; pubescence light brown to orange-brown. Hairs on midvein above simple, ± erect, straight to slightly curved, mostly 0.5-0.75 mm long on first node, to 1-2 mm long on later nodes; those on blade and veins below T-shaped (ratio of arm lengths ~ 1:1), ± sessile, appressed and straight at early nodes, mostly 0.25-0.75 mm long, becoming pro- gressively more unequally 2-branched (ratio of arm lengths mostly SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE < 4:1 on blades but to 8:1 on midvein), short-stalked, ascending, curved or sinuous or coiled at later nodes; those on margins T-shaped or 2-branched (as above) and, at later nodes, also minute, simple, ± erect and sinuous. Young leaves conduplicate, spreading open before half-grown, densely pubescent, light green, rigid, erect to horizon- tal. Venation pinnate; brochidodromous. Midvein moderate-sized; straight; flush to slightly raised above, impressed at later nodes and when fresh, prominently raised below. Secondary veins ± fine; 14— 17 on leaves at first node, 15-30 at nodes 2-10, to 30 at later nodes and on branches; alternate; mostly '/6-'/3 length of blade, longest centrally; mostly hooked or arched, sometimes also slightly zigzag; diverging at moderately acute angles (45"-65°); looped, sometimes diffusely looped basally on larger leaves, obtusely joined; slightly raised above, slightly raised below. Inter-secondary veins infre- quent. Tertiary veins reticulate within intercostal area, looped in marginal area. Petioles 0.5^.0 mm long, < 2 mm long at nodes 1- 1 0; ± circular in t.s., straight to slightly curved; shallowly channelled, with erect narrow margins above; sparsely to moderately pubescent. Stipules paired; 0.5-1 .0 mm long at nodes 1-6, to 3 mm long at later nodes and on branches; simple, triangular to narrowly triangular; persistent, ± papery; densely pubescent, pale against dark older stems; the inner margin and base continuing to and perhaps above the petiole; the outer margin and base usually terminating abruptly, but sometimes forming a low interpetiolar ridge with the stipule of the opposite leaf as the stem thickens at early but not later nodes. ARCHITECTURE. Trunk axis monopodial to at least 37 cm tall (based on one individual). Initial axis orthotropic. Branches plagiotropic but ± ascending, with leaves distichous through twist- ing of internodes; sylleptic; alternate and perpendicular at two consecutive nodes (semi-decussate); first produced 3 1-34 cm above the collet (at about nodes 17-18). First branch internode 60-75 mm long, much longer than superadjacent trunk internode (10-30 mm). First branch leaf half- to full-size of later branch leaves. Single or paired sylleptic secondary branches were produced on the primary branches 24-29 cm from the trunk. Leaves held horizontal. Several factors indicate that growth of the trunk axis above the branches stopped or slowed considerably while the branches were develop- ing, so that overall growth of the trunk was rhythmic (see Fig. 1Q): the trunk axis was very short (about 4 cm long) compared to the length of the branches (28-38 cm long); it was non-woody and very thin compared to the woody internodes on the trunk below the branches and on the lower parts of the branches; and it had matured only one pair of leaves compared to 7-8 pairs on the branches . At the time of harvest, the trunk axis had re-initiated growth, although the primary and secondary branches were still elongating. Rhythmic growth, if substantiated, would suggest Massart's model. Ecology PHENOLOGY. In Panama, Vochysiaferruginea usually flowers from late March to early July, and sometimes to a lesser extent in September-October, and its fruits mature August-October (Croat, 1978). Ripe fruits were dehiscing and releasing mature seeds in October (Garwood 1630A). Elsewhere in Central America, it prima- rily flowers in April-June and October-December and fruits July-September and December-January (Croat, 1978;Flores, 19936; Moreira & Arnaez, 1994). DISPERSAL. When ripe, valves of the dark brown capsules open and fold back, exposing the glossy bright orange-brown endocarp and releasing the seeds. Seeds are wind-dispersed. VIABILITY. Seeds remain viable for 2-3 months when stored at ambient temperature (24-26°C) and moisture levels if well-aerated (Flores, 1993ft). Flores (1993ft) considers the seeds orthodox, al- though the initial moisture content of the seeds (23%) is rather high. GERMINATION. Seeds germinate rapidly (13 days) at moderate percentages (39-50%) in both sun and shade in the growing house (this study). Other studies also report rapid germination (8-14 days) in the forest or growing house, but higher germination percentages (80-95%) (Flores, 1993ft; Moreira & Arnaez, 1994). ESTABLISHMENT AND GROWTH. The earliest stages of natural seed- ling establishment and growth have not been studied, although Flores (1993ft) noted that germinating seeds and young seedlings suffered high predation from ants and other herbivores. Boucher et al. (1994) reported exceptional seedling densities and sapling growth after Hurricane Joan in Nicaragua in October 1989, which killed all adults in the population. Four months after the hurricane, there were 94 500 seedlings and saplings (0.1-1.5 m tall) per hectare, but few young trees (3.2-10 cm DBH). The extremely open canopy, created by the hurricane through defoliating and felling trees, favoured survivorship and growth of these juveniles, such that young trees were abundant four years later and were growing on average at about 1 .4 cm DBH/year. Under the less open canopy of Barro Colorado Island, not recently if ever disturbed by hurricanes, similar-sized individuals (1-9.9 cm DBH) grew on average only 0.45 cm DBH/ year, but larger individuals (> 10 cm DBH) grew more rapidly, 1.3- 1.7 cm DBH/year (Condit et al., 1993). The population dynamics and appropriate management strategies for this species may differ greatly in hurricane-prone areas such as Nicaragua compared to hurricane-free areas such as Panama. Where hurricanes are common in the Caribbean, they occur mostly August- October during the period Vochysia ferruginea is maturing and dispersing its seeds. A hurricane occurring late in the season might blow the wind-dispersed seeds over large areas of disturbed forest, expanding the population. An earlier hurricane might kill adult trees before seeds could be matured and dispersed, preventing regenera- tion from seed and eliminating the local population, unless most regeneration comes from established shade-tolerant juveniles. It is not yet known whether seedling recruitment after disturbance is primarily from recently germinated seedlings from the current year's seed crop or older shade-tolerant juveniles surviving from previous years. Vochysia guatemalensis Donn. Sm. in Bot. Gaz. 12: 131, pi. 22 (1887). Vochysia hondurensis Sprague in Bull. Misc. Inform., Kew 1922: 183(1922). Fig. 2A-D. HABIT AND DISTRIBUTION. Trees, 30-55 m tall. Veracruz, Mexico, to Panama. COLLECTIONS. Belize. Cayo District, Chiquibul: Palmar, Garwood 403 1 (BM, seedlings with seed voucher); New Maria, Monro 867 to 871, Monro, Howe & King M4162X, M7231X, M7340X (seed- lings) and M6002X (sapling); 28 seedlings to 17 cm tall examined, sapling height about 2 m. TAXONOMIC NOTE. The seeds associated with Garwood 4031 unmistakenly identify the seedlings to genus. If one accepts the synonymy of Vochysia guatemalensis and V. hondurensis (Standley & Steyermark, 1949; Gaos, 1978; Flores, 1993a), then only one species has been reported from Belize (Stafleu, 1948; Standley & Steyermark, 1949). That the mature leaves and stems are nearly glabrous supports the identification of these collections as V. guatemalensis. N.C. GARWOOD Fig. 2 Vochysia guatemalensis Donn. Sm. Seedling: A, cotyledon; B, first leaf; C-D, cotyledonary stipules. Vochysia sp. Seedling: E-F, cotyledons; G, cotyledonary stipules; E, Hampshire & Whitefoord 269; F-G, Garwood 135. Scale bars: black, 1 mm; white, 1 cm. Seedlings and saplings Seedlings phanero-epigeal with photosynthetic cotyledons. ROOTS. Primary root 30-70 x 0.8-1 .2 mm, circular in t.s., slightly sinuous, cream-coloured; narrow annular ridge encircles axis at collet, becoming indistinct with age. Secondary roots sinuous, scattered along primary root. STEMS. Hypocotyl 50-85 x 1 mm, square to rectangular in t.s., becoming circular with age; 4-ridged or 4-winged, each wing < 0.25 mm tall, continuing at apex into lower margin of cotyledon petiole; glabrous; green. Epicotyl 5-22 x 1 mm, ± square in t.s. when young, becoming circular with age; 4-winged, each wing< 0.25 mm tall and terminating apically in a stipule; glabrous; green. Cataphylls absent. Internodes 1-16, 3-5 mm long; 4-winged, each wing about 0.25 mm tall and terminating apically in a stipule; glabrous; green. Older stems light brown to grey-brown, hollow. Buds: Cotyledonary buds hidden in axils, sparsely pubescent. Lateral buds protruding out of axils, < 0.5 mm long at nodes 1-15, sparsely short-pubescent at first node, becoming densely pubescent at later nodes. Terminal buds naked, sparsely pubescent at first node, becoming moderately pu- bescent at later nodes. Hairs on all buds clear to white, mostly < 0.25 mm long. COTYLEDONS. Two, opposite, foliaceous, simple, petiolate, stipu- late, emergent from seed, spreading apart, persistent until at least leaves at nodes 2-3 mature. Blades 15-26 x 25-37 mm; straight, flat, ± depressed obovate, with right half (seen from lower or abaxial side) usually larger at both base and apex; apex truncate to emarginate, often obliquely so, rarely ± rounded; base cordate, rounded, to ± truncate, often obliquely so; margins entire. Venation palmate, 3- nerved; acro-brochidodromous, basal and perfect. Primary veins 3, ± fine, slightly raised above, moderately raised below; central primary vein forked, with straight stem '/2 to 3A distance to apex and branches slightly recurved; lateral primary veins basal, about 3A length of blade, mostly curved, diverging at narrowly acute angle, looped, joining branches of central vein and forming ± small obovate or triangular intercostal area or rarely branching before joining the central vein and forming a ± indistinct intercostal area, the pair of intercostal areas, when distinct, usually spanning about 30-50% of the width of the cotyledon. Secondary veins moderate-sized basally along lateral primaries becoming fine apically; 12-16, 0-1 along midvein, 8-10 along lateral primaries, 4-5 along branches of midvein; mostly marginal; to 2/3 length of blade, longest basally; mostly arched or hooked; diverging at narrowly acute angles basally on lateral primaries to perpendicular angles along branches of midvein; looped marginally, acutely to obtusely joined; flush above, moder- ately raised below. Petioles 2-3 mm long, transversely oblong in t.s., 2-ridged above, with ridges continuing into lateral primary veins; 4- ridged along margins, the lower ridges continuing down into the ridges or wings on the hypocotyl and the upper ridges terminating at the stipules; glabrous. Stipules separate and paired or sometimes fused and interpetiolar, composed of two structures, both variable in shape and development. The lower part a pair of ± conical projec- tions between the petioles near the base of the upper marginal ridge of each petiole, one or both sometimes absent, mostly 0.25-0.75 mm long, usually perpendicular to the stem, sometimes gland-like apically. The upper part usually a pair of flattened scale-like outgrowths between and just above the petioles, initially folded over SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE the terminal bud but becoming erect and visible as the epicotyl expands, each ± ovate to rhombic, 0.5-0.75 mm tall and wide, the apex often 3-lobed and gland-like, the pair sometimes fused to- gether, then 0.5-0.75 x 0.75-1.5 mm tall and wide and the apex multilobed or irregular. LEAVES. Opposite, decussate on seedling trunk axis, sometimes 3- whorled on sapling trunk axis; simple, short-petiolate; stipulate. Leaf blades 30-69 x 1 1-19 mm at first node; 30-82 x 13-17 mm at nodes 2-15, to 195 x 52 mm on saplings; straight; narrowly obovate to narrowly elliptic at nodes 1-15 and on sapling; apex acuminate or long-acuminate, rarely acute at first node; bases cuneate to narrowly acute; margins entire. Blades green, papery at early nodes becoming subcoriaceous on larger saplings; sometimes moderately pusticulate above and below on later leaves, the pustules pale above; the blade nearly glabrous when fully expanded, the midvein below often sparsely short-pubescent, but moderately to densely pubescent on the blade and midvein below and along margins when expanding leaves of seedlings are < 10 mm long. Hairs simple, basifixed, < 0.25 mm long, clear to white. Venation pinnate, brochidodromous. Midvein moderate-sized; straight; flush to slightly raised above, prominently raised below. Secondary veins thin, 14-22 at first node, to 22 at later nodes; alternate; mostly '/4 to V3 length of blade, longest centrally; mostly hooked or arched, diverging at a moderately acute angle; looped, obtusely joined; slightly raised above and below. Intersecondary veins infrequent. Tertiary veins reticulate in inter- costal area, looped in marginal area. Petioles 1 mm long at first node, to 2 mm long at nodes 2-15, to 7 mm on saplings; shallowly channelled, with erect narrow margins above. Stipules paired; 1 .0- 1.2 mm long at first node, to 1.7 mm long at nodes 2-15; simple, narrowly triangular; the inner margin and base continuing above the petiole and apparently terminating near the lateral bud; the outer margin and base usually terminating abruptly, but sometimes form- ing a low interpetiolar ridge with the stipule of the opposite leaf as the stem thickens at some early nodes; persistent; often sparsely short-pubescent toward the apex. Vochysia sp. Fig. 2E-G. COLLECTIONS. Panama. Colon Province: Santa Rita Ridge, Garwood 135 (seedlings with seed fragment); Chiriqui Province: Fortuna, Hampshire & Whitefoord 269 (BM, seedlings); 15 seed- lings to 9 cm tall examined. TAXONOMIC NOTE. Specific identification is not yet possible, as Garwood 135 had only a seed fragment attached and Hampshire & Whitefoord 269 lacked seeds altogether. In addition to Vochysia ferruginea, V. allenii Standl. & L.O. Williams, V. jefensis A. Robyns, and V. speciosa Warm, have been reported from eastern Panama and V. hondurensis (- V. guatemalensis) from western Panama (D'Arcy, 1987). As immature leaves of both collections have T-shaped hairs > 0.25 mm long on the lower surfaces (see below), it is not V. guatemalensis, which has simple basifixed hairs < 0.25 mm long (see above). The T-shaped hairs on the first leaves ally it with V. ferruginea, but the cotyledons are about 1.5-2 times larger. Thus, the Chiriqui collection is a new species record for western Panama. As the cotyledons from the Chiriqui collection are consistently larger than those from the Colon collection (Fig. 3), two species might be represented. Given their similarity and the paucity of material, they are described together below, with the few differences noted. Seedlings Seedlings phanero-epigeal with photosynthetic cotyledons. ROOTS. Primary root 45-70 mm long, 1 mm wide, circular in t.s., slightly sinuous, medium brown. STEMS. Hypocotyl 50-86 x = 1 mm long and wide, ± square to rectangular in t.s., 4-ridged to 4-winged, each wing < 0.25 mm tall and continuing at apex into lower margin of petiole; glabrous. Epicotyl (immature) 4-ridged or 4-winged, each wing < 0.25 mm tall; glabrous. Buds: Cotyledonary buds hidden in axils (Garwood 1 35) or 0.75 mm tall and densely pubescent (Hampshire & Whitefoord 269). COTYLEDONS. Two, opposite, foliaceous, simple, petiolate, stipu- late, emergent from seed, spreading apart, persistent. Blades 14-21 x 22-40 mm (14-17 x 25-32 mm in Garwood 135, 19-21 x 34-^0 mm in Hampshire & Whitefoord 269); straight, flat, depressed obovate, with right side (viewed from lower or abaxial surface) usually larger than or sometimes equal to left side; apex truncate, emarginate, or cordate, sometimes obliquely so; base rounded, truncate or emarginate, sometimes unequal; margins entire. Blades green, glabrous. Venation palmate, 3-nerved; acro-brochidodromous, basal and perfect. Primary veins 3, moderate-sized; central primary vein forked, with straight stem2/3 to3A distance to apex and branches recurved; lateral primary veins basal, about 3/4 length of blade, mostly curved, diverging at narrowly acute angle, looped, joining branches of central vein to form ± small obovate or triangular intercostal area, the pair of intercostal areas usually spanning about 40-50% of the width of the cotyledon. Secondary veins moderate- sized basally becoming fine apically; 10-16, 0-2 along midvein, 6-9 along lateral primaries, 3-5 along branches of midvein; mostly marginal; '/3 to nearly full length of blade, longest basally; mostly arched to hooked, diverging at narrowly acute angles basally to perpendicularly apically; looped marginally, acutely to perpendicu- larly joined; flush above, moderately raised below. Tertiary veins reticulate within intercostal areas, looped in marginal areas. Petioles about 2 mm long, transversely oblong in t.s., 2-ridged above, with ridges continuing into the lateral primary veins; 4-ridged along margin, with lower ridges continuing into hypocotyl ridges and upper ridges terminating in the stipules; glabrous. Stipules usually fused and interpetiolar, sometimes separate and paired; composed of two structures, both variable in shape and development. The lower part a pair of ± conical projections between the petioles near the base of the upper marginal ridge of each petiole, sometimes absent or bifid, mostly 0.25-0.5 mm long, usually perpendicular to the stem, sometimes gland-like apically. The upper part usually a flattened scale-like outgrowth between and just above the petioles, initially folded over the terminal bud but becoming erect and visible as the epicotyl expands, ± transversely oblong to depressed ovate, 0.5- 0.75 x 0.75-1.0 mm tall, 3- to 5-lobed and gland-like at the apex, sometimes divided. LEAVES (only immature leaves at first node present). Opposite, decussate on trunk axis; simple, short-petiolate. Blades moderately pubescent below and along margins when expanding blades are < 5 mm long, sparsely pubescent when half-expanded. Hairs 0.25- 0.625 mm long, T-shaped, sessile, pale orange-browa Venation pinnate, brochidodromous. Key to known seedlings of Vochysia in Panama and Central America 1 . T-shaped hairs present on lower surface of young leaves at first and early nodes 2. Cotyledons 13-23 mm wide and 10-15 mm long V. ferruginea N.C. GARWOOD 2. Cotyledons 22-40 mm wide and 14-21 mm long V. sp. Simple basifixed hairs present on lower surface of young leaves at first and early nodes or hairs absent; cotyledons 25-37 mm wide and 1 5-26 mm long V. guatemalensis DISCUSSION Vochysiaceae is a moderate-size, primarily neotropical family di- vided into two tribes. Vochysieae includes Vochysia (100 species), Callisthene (12 species), Salvertia (1 species), and Qualea sensu lato (77 species, including Ruizterania); Erismeae includes Erisma (16 species) and Erismadelphus (2 species), the only African taxa (Stafleu, 1948-1954; Keay & Stafleu, 1953; Marcano-Berti, 1969; Kawasaki, in press). The monotypic Euphronia (= Lightia), some- times included in Vochysiaceae (Lleras, 1976; Cronquist, 1981) or segregated as the Euphroniaceae (Marcano-Berti, 1989, 1990), is not closely related to Vochysiaceae according to recent molecular studies (Litt et al., 1995) and will not be further discussed. Fruit There are two basic types of fruit in Vochysiaceae (Warming, 1 875; Stafleu, 1948- 1954; Keay & Stafleu, 1953). The dehiscent trilocular capsules of Vochysieae, derived from superior ovaries, contain winged seeds. The indehiscent unilocular fruits of Erismeae, derived from inferior to semi-inferior ovaries, have persistent enlarged winged calyx lobes. Within Vochysieae, fruits of Callisthene have a thick persistent central column and a fragile exfoliating exocarp not found in the other genera. Several of the 12 ovules per locule mature into seeds in Qualea (Stafleu, 1953); one or more of the few ovules per locule mature in Callisthene, (Stafleu, 1952). In Vochysia and Salvertia, however, only one of the two ovules in each locule usually develops into a mature seed (Warming, 1875; Stafleu 1948; Standley &Steyermark, 1949;Robyns, 1967a;Gaos, 1978;Moreira&Arnaez, 1994; this study), although two mature according to Flores (1993a, b). Fruits of Vochysia ferruginea are typical of the genus (Stafleu, 1948), being 3-locular loculicidal capsules, oblong or obovate in outline and 3-angled. Variation in size, shape, and surface texture of fruits in Vochysia is not well-documented: fruits of many species are still unknown (Warming, 1875; Stafleu, 1948; but see van Roosmalen, 1985). The gross morphology of fruits has been little studied. Stafleu ( 1 948) described the pericarp of the genus as consisting of a black or dark bluish exocarp 'strongly adhering to the shining golden-yellow endocarp' , an observation repeated by later authors (Robyns, 1 967a; van Roosmalen, 1985), but Flores (1993a, b) noted the presence of a well-developed mesocarp in both V. ferruginea and V? guatemalensis. In V. ferruginea, there is a brown to black exocarp, a light to medium brown spongy to woody mesocarp, thickest near the centre of the fruit, and a pale crustaceous endocarp of even thickness around each locule (Fig. 1C). The mesocarp is penetrated by longitudinal resin canals adjacent to the locules (Fig. 1C) and is also well-vascularized (Flores 1993a, b). In mature fruit, the endocarp is as well-developed around empty locules as around those filled with mature seeds, indicating that development of each locule is not dependent on the growth of fertilized ovules within it. The locules complete develop- ment before the seeds and determine the shape of the seed wing (Boesewinkel & Venturelli, 1987). Seed Seeds are winged in the Vochysieae. The wings are large and unilateral in Vochysia, Qualea, and Salvertia, but small and circum- ferential in Callisthene, and are formed from compacted testal hairs in Vochysia and Salvertia, but from extensions of the testa in Qualea and Callisthene (Warming, 1875; Stafleu, 1948-1953; Boesewinkel & Venturelli, 1987). The seeds of V. ferruginea are typical of the genus (Warming, 1875). The wing is on the anti-hilar side of the seed, and the micropyle extends along the seminiferous area and terminates near the hilum, as in V. pygmaea Bong, and Salvertia (Boesewinkel & Venturelli, 1987). In contrast, the wing of Qualea is on the hilar side of the seed, such that the micropyle extends along the anti-hilar margin of the wing and opens at the apex of the wing away from the hilum (Boesewinkel & Venturelli, 1987). The final shape of the wing in Vochysia is determined by the shape of the locule: the unicellular hairs grow upward from the testa to fill and take their shape from the empty locule (Boesewinkel & Venturelli, 1987). Hence, much of the variation in size and shape of seeds among species in Vochysia probably arises from differences in the size and shape of the mature locules. There has been little work on seed anatomy of Vochysiaceae (Corner, 1976; Boesewinkel & Venturelli, 1987). Stafleu (1948, 1 953) described the testa of Vochysia as brown, thin, and chartaceous, and the tegmen as very thin, paper white, and adhering to the testa, and the testa of Qualea as chartaceous. The mature seed coat of V. pygmaea, however, consists only of the outer layers of the testa, because the inner layers of the testa are crushed and the tegmen resorbed during development (Boesewinkel & Venturelli, 1987). The seed coat also appeared as one undifferentiated layer at maturity in V. ferruginea (this study), but Flores (1993a, b) described a dark exotesta distinct from the dead cell layers and air spaces of the mesotesta and endotesta and remnants of the tegmen in V. ferruginea and V. guatemalensis. In contrast, the mature seed coat of Qualea densiflora Warm, and Callisthene retains both the inner layers of the testa and tegmen (Boesewinkel & Venturelli, 1987). The seed coat of Erisma and Erismadelphus is thick and contains a large branched vascular bundle (Boesewinkel & Venturelli, 1987). The vascular bundle of V. ferruginea, hitherto undescribed in Vochysieae, is unbranched and terminates at the tip of the seminiferous area. The hilum of V. ferruginea (7-10 mm long) is longer than in V. pygmaea (3-4 mm long), and stretches over a greater proportion of the length of the winged seed (35-40% versus about 20%). Due to the long hilum, the seed type of V. ferruginea is described as hilar-anatropous. Endosperm is absent in mature seeds of Vochysiaceae (Bentham & Hooker, 1867; Baillon, 1874; Warming, 1875; Petersen, 1896; Stafleu, 1948-1953; Standley & Steyermark, 1949; Robyns, 1967a; Gaos, 1978; Boesewinkel & Venturelli, 1987; Flores, 1993o, b). None was seen in Vochysia ferruginea. Occasional reports that endosperm is rarely well-developed (Cronquist, 1981; Niembro Rocas, 1989) do not appear well-founded. A thin, whitish aril, adhering to the testa, was reported in V. guatemalensis (Gaos, 1978) and in the family (Niembro Rocas, 1989): this appears to be a mistranslation of Stafleu's (1948) description of the tegmen (see above). Embryo In all genera, the embryo of the mature seed is straight and the plumule and radicle poorly developed (Warming, 1875; Stafleu, 1948-1954). In Vochysia ferruginea, the embryonic axis is well- developed; the hypocotyl is relatively long compared to the poorly developed radicle, compressed, 4-winged, and annulate at the collet. The winged hypocotyl and annulate collet persist into the seedling stage, confirming the identification of the structures in the seed. In Vochysieae, the embryonic cotyledons are generally thin, folia- ceous, and folded in various ways (see below), whereas, in Erismeae, SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE they are thick, fleshy, and not or scarcely folded (Baillon, 1874; Warming, 1875; Boesewinkel & Venturelli, 1987; Kawasaki, in press). The cotyledons of Erismadelphm are plano-convex and slightly convolute at the margins (Boesewinkel & Venturelli, 1987), while those of Erisma are narrowly elongate and semi-cylindrical (Baillon, 1874), plano-convex (Boesewinkel & Venturelli, 1987), or flat (Stafleu, 1954). The embryonic cotyledons have been described as unequal in Vochysia (Warming, 1875; Stafleu, 1948) and Salvertia (Stafleu, 1 948), although no comparable information was given for Qualea or Callisthene. Unequal could mean either that each cotyledon is asymmetrical or that the two cotyledons are heterocotylar, differing in shape or size. Cotyledons are asymmetric in two species of Vochysia and one species of Qualea but symmetric in one species of Vochysia and Salvertia (Table 1 ), based on the illustrations of the unfolded embryo in Warming ( 1 875). In V.ferruginea, each cotyle- don is usually asymmetric, with the right side (viewed from the lower abaxial surface) larger, at least basally. The base of the smaller side tends to be rounded, and that of the larger side cordate, as in the expanded cotyledon of the seedling (Fig. U). The embryonic coty- ledons of V. ferruginea, when unfolded, and the fully expanded cotyledons of the seedling (Fig. 1 H, P), are isocotylar. The larger side of one cotyledon opposes the smaller side of the other on the embryo and seedling, hence, the two cotyledons are identical in shape, not mirror-images. Most monographic and floristic works indicate that the embry- onic cotyledons of Vochysiaceae are folded, but give insufficient detail to characterize the complex pattern of folding. It is usually unclear which component of folding is described: ptyxis, the folding of an individual organ such as a leaf in the bud or cotyledon in the seed, or vernation, the relationship of the immature organs to each other (Bell, 1991). Some terms, such as convolute, are used for both ptyxis and vernation but have different meanings (Bell, 1991). Definitions of terms vary considerably among authors. In Vochysia ferruginea, each cotyledon is folded four times around the longitudinal axis of the embryo (Fig. 1E-G). The folds (f,-f4) are parallel to the radicle but not parallel to the midvein of the cotyledon (Fig. IF). The margin of the larger side of the cotyledon (the right side as viewed from the lower abaxial surface) is on the exterior of the embryo, while the margin of the smaller side is in the interior. The folded smaller side is completely en- closed within the folded larger side. The inner margin is narrowly bent back with the adaxial surfaces appressed at the first fold (f,). Because of the oblique folding of the cotyledon, the base of the larger side of each asymmetric cotyledon is folded around the hypocotyl, nearly obscuring it (Fig. IF). Except for the com- pressed nature of the embryo and distinct lines of folding, ptyxis is essentially supervolute (often called convolute) but oblique. It is not conduplicate, although the cotyledon appears to be folded around the midvein in cross-sections through the middle of the embryo (Fig. 1G). Vernation is essentially convolute sensu Bell (1991), although unusually compressed. Each cotyledon partially overlaps the other, with inner margins opposite near the centre of the embryo and outer margins opposite on the exterior (Fig. 1G). Confusingly, vernation is not convolute sensu Radford et al. (1974), because one rolled or supervolute cotyledon is not com- pletely enclosed within a second rolled cotyledon. Convolute vernation and supervolute ptyxis are also found in Vochysia pygmaea, V. tucanorum Mart., and Salvertia convallari- dorea A. St.-Hil., as seen in transverse sections of the embryo (Warming, 1875: figs 16-11, 17-11; Boesewinkel & Venturelli, 1987: fig. 10B). The embryos are less compressed and angular than in V. ferruginea, and the inner margins of the cotyledons overlap rather than being folded back individually. Embryos of V. guatemalensis, V. obscura Warm., V. tucanorum, and 5. convallaridorea (Warming, 1875: figs 13, 1611, 1711; Flores, 1993a: fig. 9) are externally similar to that of V. ferruginea, and the position of the outer cotyledon margin is also consistent with convolute vernation. Ptyxis is prob- ably oblique is most species because the hypocotyl is hidden, as in V. ferruginea, but may differ in V. obscura, given that the hypocotyl is well-exposed in spite of the deeply lobed cotyledon base. Although folding appears somewhat angular in section because the seed is compressed, cotyledons are not plicate in the sense of fan-like. Thus, there is one basic pattern of vernation and ptyxis for cotyledons of Vochysia and Salvertia, with minor variations, whether cotyledons are asymmetric or not. Embryonic cotyledons of the genus Vochysia have previously been described as convolute or rolled (Bentham & Hooker, 1867; Petersen, 1896; Flores, \993a, b), spirally convolute (Baillon, 1874), obliquely spirally convolute (Warming, 1875), spirally folded (Boesewinkel & Venturelli, 1987), and spirolobal (Stafleu, 1948). The cotyledons of the species discussed above are not spirolobal, a term now used for bent embryos with incumbent once-folded coty- ledons (Duke, 1969;Radfordetal., 1974). The remaining descriptions are not inconsistent with convolute vernation and/or obliquely supervolute (= convolute) ptyxis, but do not completely characterize ptyxis and vernation. In Qualea, embryonic cotyledons have been described as convo- lute or rolled (Petersen, 1896; Boesewinkel & Venturelli, 1987) or convolute and plicate (Warming, 1875; Stafleu, 1953). The cotyle- dons of Q. grandiflora Mart. (Warming, 1875: fig. 8-II) are more complexly folded than in Vochysia, although the complete pattern cannot be resolved from the embryo and its cross-section. Supervolute ptyxis and convolute vernation, as in Vochysia, can be eliminated, however, and folding may be irregular but is not fan-like. The exposed margins of the cotyledon are very uneven, almost lobed. In contrast, the exposed cotyledon margins of Q. multiflora Mart. (Warming, 1875: fig. 7-II) are unlobed and parallel, similar to Vochysia except that they are close together on the exterior of the embryo. There appears to have been some folding around the longitudinal axis through the radicle, but the specific pattern is again unknown. In both species of Qualea, the folded cotyledons nearly obscure the embryonic axis. Thus, ptyxis and vernation appear very different in these two Qualea species compared to the homogeneous pattern observed in Vochysia. Embryonic cotyledons in Callisthene have been described as plicate and/or convolute (Warming, 1875; Petersen, 1896; Stafleu, 1952; Boesewinkel & Venturelli, 1987). In C. major Mart. (Warm- ing, 1875: fig. 3-II), the embryo is straight and compressed. Each cotyledon is irregularly folded, appearing more wrinkled (corru- gate) than plicate on the surface; each margin is involute, folding back with the adaxial surfaces touching. The folded cotyledons are pushed together for most of their length and width, but the folds are only rarely synchronous between the two. The involute margin of the first cotyledon is exposed at the edge of the embryo and overlaps the involute but hidden margin of the second, whereas the exposed involute margin of the second cotyledon abuts, rather than overlaps, the hidden margin of the first. No standard terms describe this pattern of folding, but corrugate-involute ptyxis and semi-convolute or valvate-convolute vernation might be used. In contrast to other genera in Vochysieae, the embryonic axis is fully exposed and the embryo appears almost spathulate, except for the wrinkled slightly overlapping cotyledons. Embryos of Vochysia, Qualea, and Callisthene are rich in lipids and poor in starch, those of Erisma are poor in lipids and starch, and those of Erismadelphus are rich in lipids and starch (Boesewinkel & 10 Venturelli, 1987). The seeds of V. hondurensis are rich in protein, as well as lipids (Blanche et al., 1991). Seedling There is little published information on seedlings and saplings of Vochysiaceae. Only Vochysia duquei Pilg., V. guatemalensis, and V. ferruginea (del Amo, 1979; Ricardi et al., 1987; Flores, 19930. b; this study) have been described in detail. Limited data are available for ten additional species and some information can be inferred from the embryos of five species (Tables 1-2). In Erismeae, cotyledons of Erismadelphus are cryptocotylar, hypogeal, and food-storing. Em- bryonic cotyledons ofErisma are also fleshy and food-storing, but it is not known whether these are cryptocotylar or phanerocotylar, or epigeal or hypogeal, after germination. No other information on seedlings is available for Erismeae. In Vochysieae, cotyledons are all phanerocotylar, epigeal, and foliaceous (Table 1 ). Where known, they persist after the first leaves are produced, and presumably continue to photosynthesize. In Vochysia thyrsoidea Pohl, they are the major photosynthetic organ 20- 10- 15 20 Length (mm) Fig. 3 Relationship of cotyledon length and width in Vochysiaceae. Vochysia species: circles, V. ferruginea (Panama: Garwood 1630; Garwood 2005; Foster s.n.; Costa Rica: Flores, 19936); squares, V. guatemalensis (Belize: Garwood 4031; Monro 868-871, Monro et al. M4162X; del Amo, 1979; Costa Rica: Flores, 1993a); diamonds, V. sp. (Panama: Hampshire & Whitefoord 269); octagons, V. sp. (Panama: Garwood 135); inverted triangles, V duquei (Venezuela: Ricardi et al., 1987); open triangle, V. surinamensis (Guyana: Polak, 1992); triangle with black circle, V. sp. (French Guiana: Oldeman & Fundter, 1986); triangle with white circle, V thyrsoidea (Brazil: Rizzini, 1965). Other genera (stars): C, Callisthene major (Brazil: Labouriau et al., 1964); Q, Qualea grandiflora (Brazil: Thomazini et al., 1973). Shading of symbols indicates country of origin: black, Panama; cross-hatching, Belize; horizontal hatching, Costa Rica; vertical hatching, Mexico; white. South America. Dashed lines represent width to length ratios of 2.0 (upper line) and 1.3 (lower line). Data are measurements of all seedlings in each collection, values cited in references, or my estimates from illustrations in references where values are not given. Overlapping points have been off-set by 0.5-1 .0 mm to improve readability. N.C. GARWOOD for many months (Rizzini, 1965). Cotyledons range from about 1 to 3 cm long and 1 to 5 cm wide (Fig. 3). They are smallest in Callisthene major and largest in an unidentified species of Vochysia (Fig. 3). Within Vochysia, those of V. ferruginea are the smallest (Fig. 3). Cotyledons are mostly 1.3 to 2.0 times wider than long in Vochysia, Salvertia, and Callisthene, but only 0.8 to 1.3 times wider than long in Qualea, and broadest above the middle in Vochysia and Salvertia. They are broadest around the middle in Callisthene and Qualea (Table 1; Fig. 3). They are asymmetric or symmetric in Vochysia (Fig. U and 2 A, E-F), as in the embryos, but this may be variable within some species or collections (Table l).The apex of Q. grandiflora was short-acuminate, that of C. major ± obcordate, and those of Vochysia truncate, rounded, retuse or rarely obcordate (Fig. U and 2A, E-F; Table 1 ). Venation can only be assessed in Vochysia, but the pattern is unusual. The midvein forks toward the apex, whether the apex is rounded or lobed, and two strong lateral prima- ries diverge from the base: in most species, the lateral primaries join with the recurved branches of the midvein to form triangular to obovate intercostal areas (Fig. U and 2A, E-F; Table 1). Smaller veins form one or more concentric series of loops around the central intercostal areas. The cotyledonary stipules in the three species of Vochysia de- scribed in this study are unusual two-parted structures (Fig. 1K-M; Fig. 2C-D, G). The lower part usually consists of one to three narrow projections between the petioles, usually perpendicular to the hypocotyl, situated at the base of the upper (adaxial) surface of the petiole where the upper ridge or wing of the petiole terminates. The upper part consists of a flattened interpetiolar scale-like outgrowth just above the petioles, initially folded over the terminal bud but becoming erect and visible as the epicotyl expands, completely or partially divided or torn into two to five parts but rarely unlobed. The apex of the lower and/or upper parts often appears gland-like. Neither part appears to be fused over the petioles, and the axillary buds are often visible above the cotyledonary petiole. Although there was some intraspecific variation, the stipules of V. ferruginea usually had one lower projection and a lobed upper part and those of V. guatemalensis two lower projections and two separate upper parts. The reduced stipules of V. guatemalensis have also been described as 2-3 trichomes of a glandular nature (Flores, 1993a), but the thick 3-lobed pair of structures illustrated (fig. 7e) appear rather large and fleshy for trichomes and may be a different form of the variable structures described in this study. Two-parted cotyledonary stipules similar to those of Vochysia have not been seen in seedlings of more than 600 dicotyledonous species in 88 families surveyed from Panama (Garwood, in prep.), although the fleshy cotyledonary stipules of Lafoensia (Lythraceae) are similar to the lower stipular parts of Vochysia. Leaves are opposite at the early nodes in Vochysia, Qualea, and Callisthene (Table 2). In Vochysia, the pair of leaves at the first node is decussate with respect to the cotyledons and leaves at the second node (Table 2). In Q. grandiflora, the first pair of fully developed leaves is parallel to the cotyledons but decussate to the second pair of leaves (Table 2); a pair of 'prophylls' between the cotyledon and first pair of leaves (Thomazini et al., 1973) might be vestiges of the 'missing' pair of decussate leaves or homologous to the interpetiolar cotyledonary stipules of Vochysia. Leaves remain opposite into the adult stage in V. ferruginea, but become 3- then 4-whorled in V. guatemalensis sometime in the sapling stage. The initial axis of C. major appears to be plagiotropic, with opposite distichous leaves smaller than the cotyledons, but this will need confirmation because it is difficult to interpret details in the published photograph of the seedlings (Labouriau et al., 1964: fig. 26). Stem characters are known only from Vochysia. The collet, the SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE 11 Table 1 Cotyledon characters in seedlings and embryos of Vochysiaceae. Genus Species Source Emer- gence Posi- tion Devel- Persistence Width:Length Ratio Shape Base Apex Symmetry Venation Stipules opment node months Vochysia V. densiflora Spruce 18* - - F - 1.7 2 5-7 8 A V duquei Pilg. 15 P E Fr 2 >1 1.6-1.9 2-3,6 4-6 3-5 S 7,2 + V.ferruginea 6 P E F 3-8 >9 1.3-1.9 2 3-6 6-7 A 1-3 f+f,g Mart. 10 - E - - - _ _ _ _ _ _ 4b P E F 2 1.6 2 6 5 8^4 7 f V. guatemalensis 6 P E F 3 1.2-1.8 2 3-6 3-5 A 1-3 s+s,g Donn. Sm. 10 - E - - _ _ _ _ _ _ _ _ 4a P E F / 1.2-1.3 2 6 4 A 7,4 s,g 1,5 P E F 6 1.5 2 4-5 4 A 1,2 V. obscura Warm. 18* - - F - - 1.4 2 7 4-5 S V surinamensis Stafleu 14 P E n 6 2.7 1 — 3-4 — — _ V. thyrsoidea Pohl 13 - E - = 12 - - - - _ _ _ 16 P E Fr 1 >12 = 7-2 7 2 4 _ _ _ V. tucanorum Mart. 18* - - F - - 7.6 2 6-7 3^t A V. sp. (Brazil) 2 P E F - - - 1 _ - — _ _ V. sp. (Fr. Guiana) 11 P E F - 2.7 2 4 5 5 2,5 V. sp. (Panama) 6 P E F 1 1.4-2.1 2 3-5 4-6 S,A 1-3 s+f,g Salvertia S. convallaridorea 18* - - F _ - 1.6 2 2-3 4-5 S A. St.-Hil. Qualea Q. grandiflora 12 P E F 2 >6 - - - - _ _ _ Mart. 3 P E F _ 4 - - — _ _ _ _ 17 P E F 2 >6 1.1 5 3^f 7 Al P Q. multiflora Mart. 18* - - F - 0.8-1.3 4 3 9 A Callisthene C. major Mart. 8 P E Fl 1-3? 1.2-1.5 3 - 5-6 _ _ _ 18* - - F - 1.2-1.6 - - - - - - Erisma E. spp. 7* - - R - - - - - - Erismadelphus E. exsul Mildbr. 9 C H R - - - - - _ _ _ E. sp. 9 C H R - - - - - Source: 1, del Amo 1979; 2, Duarte 1979; 3, Felippe & Dale 1990; 4a,b, Flores 1993a, b; 5, Gaos 1978; 6, Garwood, this study; 7, Kawasaki, in press; 8. Labouriau et al. 1964: 9, Miquel 1987; 10, Moreira & Arnaez 1994; 11, Oldeman & Fundter, 1986; 12, Paulilo et al. 1993; 13, Paviani & Haridasan 1988; 14, Polak 1992; 15, Ricardi et al. 1987; 16, Rizzini 1965; 17, Thomazini et al. 1973; 18, Warming 1875. *, all characters based only on the embryonic cotyledon in the mature seed. Emergence: P, phanerocotylar; C, cryptocotylar. Position: E, epigeal; H, hypogeal. Development: F, foliaceous; R, fleshy, presumably with stored reserves; r, somewhat fleshy or succulent, with reserves or stored water. Persistence: measured as i) the last recorded leaf node to produce leaves while the cotyledons are still present (e.g. node=3. cotyledons still present when leaves produced at node 3) or ii) the length of time [months] that cotyledons persist after germination. These measures underestimate persistence when only the youngest seedling stages have been studied. Width:Length Ratio: width divided by length. Shape: 1, obtriangular to shallowly obtriangular, including 'triangular'; 2, depressed obovate; 3, transversely oblong; 4, very widely obovate; 5, transversely widely elliptic to transversely widely oblong; 6 transversely reniform, rectangular-reniform. Base & apex: 1, short-acuminate; 2, acute; 3, rounded; 4, truncate; 5, retuse to emarginate; 6, cordate, obcordate, 2-lobed; 7, 2-cleft or -parted; 8, 3-lobed; 9, irregular. When blade is asymmetric, the base and/or apex may be unequal on the two sides (e.g. half-cordate, half-rounded) or oblique (e.g. obliquely truncate). Symmetry of blade: A, asymmetrical; S, symmetrical. Venation: 1, branches of forked central primary vein join lateral primaries forming a pair of obovate to triangular intercostal areas; 2, palmate or 3-nerved; 3, basal acro- brochidodromous; 4, basal actinodromous; 5, branches of central primary vein forked; lateral primaries each form a closed loop apically, terminating near but not joined directly to the tips of the central vein. Stipules: s, usually separate; f, usually fused and interpetiolar; f+f, f+s, and s+s, two structures (fused or separate) present at cotyledonary node (first is lower structure, second is upper structure); g, gland-like; +, present; P 'prophylls' present above cotyledons. -, Information not available. Characters in italics were inferred or estimated from illustrations in the citations, not from the text of the original author. Where authors identified the cotyledons as foliaceous, it was assumed that they were also phanero-epigeal. junction of the radicle and hypocotyl, is annulate in at least three species (Table 2; Fig. IF). The hypocotyl, epicotyl, and later internodes are usually square, with ridges or wings on each corner (Table 2). Ridges on the hypocotyl continue without interruption into the ridges on the lower margins of the cotyledonary petioles in V.ferruginea and V. guatemalensis (Figs IK, 2G; Flores, 1993a, b). In contrast, the wings or ridges on the epicotyl and later internodes terminate in small narrowly triangular to linear stipules at the top of each internode (Fig. 1O, S; Table 2), with the inner margin and base of the stipules apparently extending above the petiole but not attached to it. The winged stems do not originate directly from decurrent leaf bases, as sometimes suggested, because the attenuate base of the blade narrows into a medial pair of short thin ridges on the upper (adaxial) surface of the petiole, interrupting the lamina 12 N.C. GARWOOD Table 2 Root, stem, and leaf characters in seedlings of Vochysiaceae. Genus Species Source Collet Hypocotyl Epicotyl Internodes Phyllotaxis Leaf stipules Nodes 1-2 Later nodes Vochysia V duquei 15 a s s 0 -,v + V. ferruginea 6 a s,w s,w s,w o,d o,d +,s 4b c s - o,d o,d - V. guatemalensis 6 a s,w s,w s,w o,d o,w +,s 4a c s,w s,w o,d o,w - 1,5 c c,w s,w s,w o,d 0 +,i V. surinamensis 14 - - - S,W 0 o,v - V. thyrsoidea 13 c t - - - - V. sp. (Panama) 6 - s,w w 0 - - Qualea Q. grandiflora 17 - - - o,p - — Callisthene C. major 8 - o a? Sources and authorities for names: see Table 1. Collet (junction of hypocotyl and primary root): a, annulate ridge present; c, constriction, abrupt change, or other feature present, but nature of structure not discernable from illustration. Hypocotyl. Epicotyl, and Internode: cross-sectional shape of hypocotyl, epicotyl, and early internodes (about first ten), respectively: s, square to rectangular; c, circular; t, tuberous and circular, at least when older; w, 4 wings or ridges present. Phyllotaxis: Nodes 1-2: o, leaves at first node opposite; d, pair of leaves at first node decussate to both cotyledons and pair of leaves at second node; p, pair of leaves at first node parallel to cotyledons and decussate to pair of leaves at second node. Later nodes: o, leaves opposite and decussate (at least to node 5, sometimes reported up to node 20); d, leaves opposite and decussate on larger seedlings, saplings, and adults; w, leaves whorled on larger seedlings or saplings (and adults), with three or four leaves per node, size of juveniles at time of transition from opposite to whorled leaves unknown; v, juvenile leaves unknown, but adult leaves whorled; a, axis becoming plagiotropic with pairs of leaves distichous. Leaf stipules: +, small, narrowly triangular to linear stipules present from the first node; i, stipules intrapetiolar; s, stipules separate. -, Information not available. Characters in italics were inferred or estimated from illustrations in the citations, not from the text of the original author. between the blade and wings. It is unclear, however, whether the petiolar ridges are joined to the lateral extensions of the stipules. Striking ontogenetic changes are seen in hair morphology in Vochysia ferruginea (this study). The epicotyl and first leaves are only sparsely pubescent below, but stems and leaves at higher nodes become progressively more densely pubescent. Hairs on early internodes and lower leaf surfaces are nearly sessile, appressed, and T-shaped, with straight arms of equal length; those on later leaves and stems are short-stalked, ascending to erect, 2- branched, with the longer arm curved to coiled and about eight times longer than the shorter arm. Hairs on the adult leaves are similar, except the branches are even more unequal in length (Garwood, pers. obs.)- On seedlings to adults of V. ferruginea, hairs on the midrib above are always simple. In Vochysia sp. (Garwood 135 and Hampshire & Whitefoord 269), hairs on the epicotyl and eophylls were similar to those on V ferruginea. In contrast, hairs on the young expanding leaves of V guatemalensis were simple and basifixed, but these were usually sparse or absent on mature leaves (Flores, 1993a; this study). All parts of V duquei are glabrous (Ricardi et al., 1987). Seedlings from the Brazilian cerrado have a number of special morphological adaptations to the dry cerrado environment (Labouriau et al., 1964; Rizzini, 1965; Paviani & Haridasan, 1988). Vochysia thyrsoidea has ± fleshy succulent cotyledons, a thick watery hypocotyl which is mostly buried, minute eophylls which remain hidden between the cotyledons, greater growth of the taproot rather than shoot in the first few months, and a swollen subterranean organ which forms from the hypocotyl and upper part of the taproot in the first two years. Qualea grandiflora also forms a swollen under- ground organ. Phylogeny There have been no phylogenetic analyses of relationships within Vochysiaceae. Since the completion of Stafleu's ( 1 948- 1 954) mono- graph more than 40 years ago, the numbers of collections and species described have increased, but major systematic work has been limited to elevating one section of Qualea to Ruizterania (Marcano-Berti, 1969) and monographing Erisma (Kawasaki, in press). Fruit and seed characters have long been important in delim- iting genera within the family, but the utility of seedling characters has been untested because seedlings have not been available for study. The potential of the new seedling and seed characters de- scribed above to contribute to phylogenetic and systematic studies is discussed here. If such traits look promising, data on more species should be collected. Phanero-epigeal seedlings are reported only from Vochysieae, and crypto-hypogeal seedlings from Erismeae, following the current tribal division. In Vochysieae, shape characters of the foliaceous cotyledon fell into three groups corresponding to Qualea, Callis- thene, and Vochysia/Salvertia, although some variation occurred in all groups. Symmetry of the cotyledons varied most in Vochysia, and vernation and ptyxis of the embryonic cotyledons inQualea. Several traits generally uncommon in dicotyledon seedlings occur together in Vochysia, including unusual cotyledonary stipules and cotyledonary venation, annulate collet, and winged hypocotyls and early internodes, but these have not been assessed in many species or other genera. Supra-cotyledonary prophylls, also unusual in dicoty- ledon seedlings, were reported in one species of Qualea. The existence of considerable variation in cotyledon characters, and the presence of many unusual seedling characters in this small sample of SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE 13 about 20 species (Tables 1-2), suggests further study of other taxa would be profitable. Ontogenetic changes from the seedling to adult stages can also be important phylogenetic characters. In Vochysia, leaves of adults are either opposite or 3- or 4-verticillate, but the first seedling leaves of all species described are opposite (Table 2). In V. guatemalensis, leaves remain opposite at early seedling nodes, but gradually be- come 3- then 4-verticillate in larger saplings (Flores, 1993o). Although presence of opposite seedling leaves in all taxa would not be phylogenetically informative, the timing of the switch from opposite to verticillate leaves may provide a useful character. Adult leaves are also verticillate in Salvertia and some species of Erisma, but opposite in other taxa (Stafleu, 1948-1954). Pronounced ontogenetic changes in hair morphology were ob- served in Vochysia ferruginea, but not in V. guatemalensis. Because studies of ontogenetic changes of trichomes on leaves of seedlings and adults have helped identify groups of species within the large and complex genera of Solanum and Rhododendron (Seithe 1979, 1980; Seithe & Anderson, 1982), further study would be warranted. Unfortunately, the morphology of hairs in Vochysia is poorly known. They have been described as simple or sometimes unicellular and 2- branched (Warming, 1875; Metcalfe & Chalk, 1950); hairs have arms of equal length and short stalks in at least one species (Shanmukha Rao & Vijayalakshmi Sarma, 1992). Stafleu (1952) indicated that simple and biramose hairs occurred in Vochysieae, but did not specifically describe those of Vochysia (Stafleu, 1948), although frequent references to 'peculiar indumentum' or 'charac- teristic indumentum' suggests that their morphology is unusual. An examination of adult specimens of Vochysia at BM indicates that species with 2-branched or simple hairs are both common (Table 3). The distribution of simple and branched hairs is clearly not random with respect to the sections of Vochysia delimited by Stafleu (1948). Branched hairs are especially abundant in subsection Ferrugineae, which contains V. ferruginea. There was considerable variation among species in the morphology of the branched hairs of the adult stage, ranging from nearly sessile appressed T-shaped hairs with straight branches of equal length, to long-stalked erect 2- branched hairs with curled arms of equal or unequal length (Garwood, pers. obs.). In some species, the smaller arm was so reduced in length that the hair could easily be misinterpreted as simple, sug- gesting that at least some of the simple hairs in Vochysia might be derived from much-reduced branched hairs, as occurs in Malpighi- aceae (Anderson, 1990). The range of hair morphology from the seedling to adult stages in V. ferruginea spans nearly the entire range of branched hairs seen in the genus. Therefore, examining the ontogeny of branched hairs might reveal similarities in the early stages that would be phylogenetically informative in comparison with other species of Vochysia, or other genera. Hairs are also simple or 2-branched in Qualea (Stafleu, 1953), but stellate in Erisma (Warming, 1875; Stafleu, 1954), and simple in Callisthene (Stafleu, 1952), Salvertia (Garwood, pers. obs.), and Erismadelphus (Garwood, pers. obs.). Seed, fruit, and seedling characters also bear on the higher order relationships of the family. Vochysiaceae has usually been included in the Polygalales with Polygalaceae, Trigoniaceae, Malpighiaceae, Xanthophyllaceae, Tremandaceae, and Krameriaceae (Dahlgren, 1980;Takhtajan, 1980; Cronquist, 1981). Boesewinkel & Venturelli (1987) used anatomy of the seed coat to reassess the position of the family in Polygalales. They concluded that only Polygalaceae and Xanthophyllaceae should be retained in the Polygalales, and that the relationship of Vochysiaceae and the remaining families needed further study. Reviewing numerous floral and vegetative characters in Malpighiaceae, Anderson (1990) decided that Malpighiaceae was most similar to Vochysiaceae and Trigoniaceae and least similar to Polygalaceae and Krameriaceae. The presence of 2-branched uni- cellular hairs in Vochysiaceae, Trigoniaceae, and Malpighiaceae, and their absence in Polygalaceae, also supports this grouping (Shanmukha Rao & Vijayalakshmi Sarma, 1992). Recent molecular analyses of the angiosperms using the chloroplast gene rbcL also supported the break-up of the Polygalales, but not the grouping of Vochysiaceae with Malpighiaceae and Trigoniaceae (Chase et al., 1993). Instead, Vochysiaceae, repre- sented by Qualea, fell within the Myrtales. A more detailed study of Myrtales using the same gene confirmed the Myrtalean affinities of Vochysiaceae, represented by Vochysia, Erisma, and Qualea (Conti et al., 1996). Vochysiaceae was sister group to the Myrtaceae and these, with Heteropyxidaceae and Psiloxylacecae, formed the sister clade to that containing Melastomataceae, Memecylaceae, and other families. Work with chloroplast gene matK also supports placement of Vochysiaceae in Myrtales, but it falls between Myrtaceae (with Heteropyxidaceae and Psiloxylacecae) and Melastomataceae (with Memecylaceae) (P. Wilson, pers. comm.). Previous systematic treatments based on morphology have never included Vochysiaceae in the Myrtales (Conti et al., 1996), regard- Table 3 Distribution of simple and branched hairs in Vochysia. Section Subsection Number of species Described K,, ^'i:,!! .11 /IG/1Q\ Examined Type of hair by Matleu (iy4o) for study Simple Branched Unknown Vochysiella Decorticanthes 10 6 6 0 0 Calophylloideae 10 7 6 0 1 Ciliantha Micranthae 11 9 8 0 1 Lutescentes 23 13 6 4 3 Discolores 3 2 0 2 0 Chrysophyllae 1 1 1 0 0 Megalanthae 5 3 3 0 0 Ferrugineae 31 22 1 18 3 Pachyantha - 3 0 — — ~ Type of hair: simple, only simple unbranched hairs present on leaves, stems, and reproductive parts; branched, some or all of hairs present branched, including T-shaped hairs; unknown, specimens glabrous or hairs too small or dense to characterize. Leaves, stems, and inflorescences of one to several specimens of each species at BM monographed by Stafleu (1948) were examined under a 40x dissecting microscope. 14 ing this relationship as so remote that the presence of two of the most definitive Myrtalean characters, vestured pits in the second- ary xylem and bicollateral vascular bundles in the primary stem, was not considered significant (van Vliet & Baas, 1984). In retro- spect, many vegetative characters of Vochysiaceae are equally at home in Myrtales (sensu Dahlgren & Thorne, 1984) as in Polygalales, including opposite or verticillate leaves with entire margins, brochidodromous venation, and stipules. The highly zygomorphic flowers with one fertile stamen are, however, unu- sual in the order (Conti et al., 1996). When a thorough morphological analysis of the relationship of Vochysiaceae within Myrtales is undertaken, several seedling and seed characters de- scribed for Vochysiaceae deserve further study. Vochysiaceae share other vegetative characters with Myrtales, including the occasional presence of 2-branched unicellular or tufted hairs in Combretaceae, Lythraceae, and Myrtaceae (Metcalfe & Chalk, 1950). Cotyledonary stipules, generally un- common in dicotyledons, occur as rudimentary glandular structures in some Lythraceae, Onagraceae, and Myrtaceae, and as 2- to 3-lobed structures or glandular processes near the basal margins of the blade in other groups of Myrtaceae (Weberling, 1984). The latter type might be homologous to the unusual cotyledonary stipules of Vochysia (Flores, 1993«; this study), which are composed of an upper lobed part and lower possibly glandular part. Wings are well-developed on adult stems in some Lythraceae and Penaeaceae, and often develop auricles at the apex, which are distinct from the rudimentary stipules found closer to the petioles (Weberling, 1984). In Lythraceae, auricled wings are already present at the seedling stage in Lafoensia (Garwood, in prep.). Wings are well-developed on seedlings of Vochysia, although usu- ally reduced to ribs on adults (Stafleu, 1948; Garwood, pers. obs.). As the small stipules of Vochysia occur at the apex of the wings, they may be homologous to the auricles at the apex of the wings of these Myrtales, rather than the small, often dissected, stipules characteristic of Myrtales (Weberling, 1984). If so, Vochysia may lack true stipules. Herbarium specimens of V. ferrugineae and V. guatemalensis seedlings were carefully exam- ined for the presence of dissected or rudimentary stipules over the petioles, but none were seen (Garwood, pers. obs.). The occa- sional presence of a minute shrivelled structure, however, suggests that actively growing seedlings should be examined to verify whether minute caducous stipules are present but hidden by the small erect expanding leaves. In seedlings of Myrtaceae, auricles are lacking on the winged epicotyl and later internodes of Psidium and Calycolpus and on the winged hypocotyl of Myrcia, and wings are lacking in Eugenia, but all have rudimentary gland-like stipules (Garwood, in prep.). Seeds of Vochysiaceae and Myrtales generally lack endosperm and arils and have straight or slightly curved embryos (Cronquist, 1981; Dahlgren & Thorne, 1984). A fibrous exotegmen and an endotestal crystal layer are found in Qualea and some Lythraceae, Combretaceae, and Onagraceae, but the exotegmen lacks fibres and is crushed in Vochysia and some Melastomataceae and Myrtaceae (Corner, 1976; Dahlgren &Thorne, 1984; Boesewinkel &Venturelli, 1987). The embryonic cotyledons are variously folded in Vochysiaceae and many Myrtales, with folding frequently described as 'spiral' in Vochysia as well as Combretaceae and Oliniaceae (Dahlgren & Thorne, 1984). As brief descriptions of vernation and ptyxis in Vochysiaceae are generally incomplete or incorrect (see above), further detailed studies should be carried out throughout the order. N.C. GARWOOD Ecology Seed germination has been the most frequently studied aspect of the seed and seedling ecology of Vochysiaceae. Germination is usually rapid, occurring 1 week to 3 months after seed dispersal or sowing in species of Vochysia, Salvertia, and Qualea from both cerrado and rainforest (Rizzini, 1965; Thomazini et al., 1973; Macedo, 1977; Duarte, 1979; Sabatier, 1985; Paviani & Haridasan, 1988; de Godoy & Felippe, 1992a; Moreira & Arnaez, 1994; Flores 1993a, b; this study; M. Bustamante, pers. comm.; P.E. Oliveira, pers. comm.; P. Wilson, pers. comm.). Chemical or mechanical scarification of the seed coat, or its complete removal, may hasten germination in Q. grandiflora (Felippe, 1990; Felippe & Dale, 1990), but is not required (M. Bustamante, pers. comm.; P.E. Oliveira, pers. comm.; P. Wilson, pers. comm.). Germination is not inhibited by dark or shaded conditions in Q. retusa Spruce ex Warm., Q. grandiflora, or V.ferruginea (Macedo, 1977; Felippe, 1990; this study), although it can be somewhat depressed in direct sunlight (Flores, 19936). Recently dispersed seeds of V. maxima Ducke buried under the leaf litter either lost viability or had germinated within 2 months (Viana, 1990). Thus, there is no indication of any type of seed dormancy in the family, and seeds should form transient, rather than persistent, seed banks under natural conditions. Under artificial conditions, seeds of Qualea cordata Spreng., Vochysia guatemalensis, and V.ferruginea retained high viability for at least 2-3 months when stored at room temperature and humidity (de Godoy & Felippe, 1992a; Flores, 1993a, b), in spite of a relatively high initial moisture content in Vochysia (23-32%, fresh weight basis). Reducing the moisture content extended viability of the apparently orthodox seeds, although they were intolerant of chilling (Flores, \993a). The potential to store seed should promote the ongoing silvicultural development of the family. Seeds of all species are wind-dispersed, as indicated by the morphology of seeds of Vochysieae and indehiscent fruits of Erismeae, except Erisma calcaratum (Link) Warm, from the Ama- zonian varzea, which is water-dispersed (Stafleu, 1954). In tropical forests, seeds of Vochysia maxima are dispersed near the parent, mostly < 30 m from the trunk (Viana, 1990), seedlings of V. maxima and V. ferruginea persist in the shaded understorey (Viana, 1990; Garwood, pers. obs.), but growth is fast only in treefall gaps and larger openings (Viana, 1990; Boucher et al., 1994). In plantations, seedlings of V. guatemalensis and V.ferruginea can reach 2-3 m tall within three years (Flores 1993a, b). Thus, tall canopy Vochysia appear to be gap opportunists, regenerating from a persistent seed- ling bank following disturbances throughout the year. Regeneration of Qualea may follow a different pattern. Following a severe hurri- cane, most of the severely defoliated adults of Q. paraensis Ducke survived but there was no regeneration from seedlings or saplings; in contrast, all the equally abundant and severely defoliated adults of V. ferruginea died, but there was extensive regeneration from seedlings and saplings (Boucher et al., 1994). If this pattern is typical, species of Vochysia may be more amenable to sustainable management or plantation development than Qualea. In the drier and more open cerrado, seedlings of Qualea grandiflora, Q. cordata, and Vochysia thyrsoidea appear to grow inherently slowly, with neither extra moisture nor nutrients pro- moting rapid growth (Rizzini, 1965; de Godoy & Felippe, 19926; Paulilo et al., 1993; Felippe & Dale, 1990). Although the basis for the slow growth is not understood, many species develop special morphological adaptations to the cerrado environment, such as large subterranean storage organs and thick succulent cotyledons (Labouriau et al., 1964; Rizzini, 1965; Paviani & Haridasan, 1988). SEEDLINGS, FRUITS AND SEEDS OF PANAMA: VOCHYSIACEAE 15 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 encourage- ment and advice; S. Knapp, D.M. Williams and an anonymous reviewer for helpful comments on the manuscript; M. Beasley for bibliographic assist- ance; INRENARE for permission to work in Panama; M.L. Kawasaki, M. Bustamante, P.E. Oliveira, and P.O. Wilson for sharing work in press, unpublished data, and observations; 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 generously supported by the National Science Foundation (BSR-85-17395), the Natural Environment Research Council (GR3-695 1 ), the Smithsonian Tropical Research Institute, The Natural History Museum, and the Linnean Society of London. REFERENCES del Amo, S. 1 979. Clave para plantulas y estados juveniles de especies primaries de una selva alta perennifolia en Veracruz, Mexico. Biotica 4: 59-108. Anderson, W.R. 1990. The origin of the Malpighiaceae-the evidence from morphol- ogy. Memoirs of the New York Botanical Garden 64: 210-224. Kit illi in, H.E. 1874. Vochysiaceae. Histoire des plantes 5: 93-104. Bell, A. 1991. An illustrated guide to flowering plant morphology. Oxford. Bentham, G. & Hooker, J.D. 1867. Vochysiaceae. Genera plantarum 1(3): 975-978. 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Hair types as taxonomic characters in Solanum. In J.G. Hawkes, R.N. Lester & A.D. Skelding (Eds), The biology and taxonomy of the Solanaceae: 307- 319. London. 1980. Rhododendron hairs and taxonomy. In J.L. Luteyn & M.E. O'Brien (Eds), Contributions towards a classification of Rhododendron: 89-115. New York. & Anderson, GJ. 1982. Hair morphology and the relationships of species in Solanum sect. Basarthrum. Plant Svstematics and Evolution 139: 229-256. 16 N.C. GARWOOD Shanmukha Rao, S.R. & Vijayalakshmi Sarma. 1992. Morphology of 2-armed trichomes in relation to taxonomy: Malpighiales. Feddes Repertorium 103: 559- 565. Stafleu, F.A. 1948. A monograph of theVochysiaceae. Recueildes trauvaux botaniques neerlandais 41: 397-540. 1952. A monograph of the Vochysiaceae. II. Callisthene. Acta Botanica Neerlandica 1: 222-242. 1 953. A monograph of the Vochysiaceae. III. Qualea. Acta Botanica Neerlandica 2: 144-217. 1954. A monograph of the Vochysiaceae. IV. Erisma. Acta Botanica Neerlandica 3: 459-^80. M.I IK lk->. P.C. & Steyermark, J.A. 1949. Vochysiaceae, Flora of Guatemala: Part VI. Fieldiana: Botany 24: 2-5. Takhtajan, A.L. 1980. Outline of the classification of flowering plants (Magnoliophyta). Botanical Review 46: 225-359. Thomazini, L.I., Massa, C.S. & Piccolo, A.L.G. 1973. Qualea grandiflora Mart., considerac,5es sobre a semente, a germinagao a plantula. Revista de Agricultura, Piraciaba Brazil 48: 101-106. Viana, V.M. 1990. Seed and seedling availability as a basis for management of natural forest regeneration. In A.B. Anderson (Ed.). Alternatives to deforestation: steps toward sustainable use of the Amazon rainforest: 99-1 15. New York. van Vliet, G.J.C.M. & Baas, P. 1984. Woody anatomy and classification of the Myrtales. Annals of the Missouri Botanical Garden 71: 783-800. de Vogel, E.F. 1 980. Seedlings of dicotyledons: structure, development, types; descrip- tions of 150 woody Malesian taxa. Wageningen. Warming, E. 1875. Vochysiaceae. In K.F.P. von Martius & A.W. Eichler (Eds), Flora brasiliensis 13(2): 17-116. Weberling, F. 1984. Stipular structures. In R. Dahlgren & R.F. Thorne (Eds), The order Myrtales: circumscription, variation, and relations. Annals of the Missouri Botanical Garden 71: 641-644. Bull. not. Hist. Mus. Land. (Bot.) 28(1): 17^0 Issued 25 June 1998 A revision of the genus Mandragora (Solanaceae) STEFAN UNGRICHT* SANDRA KNAPP AND JOHN R. PRESS Department of Botany, Tne~Natural History Museum, Cromwell Road, London SW7 5BD * Present address: Waldmatt 6, CH-5242 Birr, Switzerland CONTENTS Introduction Mythological and medicinal history Taxonomic history Materials and methods Material examined Taxonomic concepts Morphometrics Cladistics Results and discussion Species delimitations using morphometric analyses Phylogeny Biogeography Taxonomic treatment Mandragora L Key to the species of Mandragora 1. Mandragora officinarum L 2. Mandragora turcomanica Mizg 3. Mandragora caulescens C.B. Clarke References Exsiccatae Taxonomic index ... 17 18 18 19 19 20 21 22 22 22 26 26 29 29 30 30 33 34 36 38 40 SYNOPSIS. The Old World genus Mandragora L. (Solanaceae) is revised for the first time across its entire geographical range. The introduction reviews the extensive mythological and medicinal as well as the taxonomic history of the genus. On morphological and phenological grounds three geographically widely disjunct species can be distinguished: the Mediterranean M. officinarum L., the narrowly local Turkmenian endemic M. turcomanica Mizg. and the Sino-Himalayan M caulescens C.B. Clarke. The generic monophyly of Mandragora L. as traditionally circumscribed is supported by cladistic analysis of morphological data. The ecological and historical phytogeography of the genus is discussed and alternative biogeographical scenarios are evaluated. Finally, a concise taxonomic treatment of the taxa is provided, based on the evidence of the preceeding analyses. INTRODUCTION The nightshade family (Solanaceae) is a cosmopolitan but predomi- nantly tropical group and is especially well represented in the Neotropics. The family is of considerable economic importance. Food plants include potato (Solanum tuberosum L.), tomato (Sola- num lycopersicum L. (=Lycopersicon esculentum Mill.)), pepper (Capsicum annuum L.) and aubergine (Solanum melongena L.j. Another species of great commercial value is tobacco (Nicotiana tabacum LJ. The family also contains ornamental plants such as garden petunias (species of Petunia Juss.) and the Chinese lantern (Phy sails alkekengi L.j. Many members of the family are poisonous, but several such as deadly nightshade (Atropa belladonna L.), henbane (Hyoscyamus niger L.) and thorn-apple (Datura stramo- nium L.) provide drugs traditionally used in medicine. The subjects of this study, the mandrakes (Mandragora L.), belong to this latter group of pharmaceutically interesting Solanaceae. The long history of mythology and medicinal use of the mandrake combined with the variable morphology and phenology have led to considerable confusion in the classification of Mandragora. The nomenclatural situation is, considering the small size of the genus, amazingly chaotic. In particular the Mediterranean Mandragora is burdened with synonyms, and some authors (Mill, in litt.) have expressed doubts about the placement of the Himalayan mandrake in Mandragora at all. This study represents the first taxonomic revision of the genus over its entire range. We address the following main questions in this study. Firstly, which of the described Man- dragora taxa are justifiable if the whole range of the morphological variation is considered? Secondly, is the genus as currently delim- ited (including the Himalayan mandrake, M. caulescens) monophyletic? Finally, does the phylogeny, combined with the distribution patterns of the taxa, support a particular historical biogeographical scenario and do the ecological requirements of the defined taxa impinge upon this? © The Natural History Museum, 1998 18 Mythological and medicinal history The origin of the myth surrounding the mandrake root is hard to trace. It is clear, however, that the mandrake was a focus of supersti- tion and belief in magic and demons for many centuries. The sometimes vaguely humanoid shape of the root and its potent, psychotropically active alkaloids are probably responsible for the long history of myths. However, the properties ascribed to the plant as a whole, and especially to its root, are highly ambiguous and the first references in Ancient Egypt and in the Bible remain controver- sial. Ornaments on the casket and throne from the tomb of Tutankhamun seem to depict mandrakes (Hepper, 1990; Roland, 199 1 ). Mandragora is not known to be native in Egypt but may have been cultivated there. The Biblical 'Dudaim' is often said to be identical with Mandragora (Moldenke & Moldenke, 1952; Fein- brun-Dothan, 1978; Fleisher & Fleisher, 1994). Dudaim is mentioned in the Song of Solomon (7: 13) and in another passage from the Old Testament (Genesis 30: 14-18), according to which it was used in relation to fertility. The 'Moly ' of Homer in his Odyssey is presumed to be the mandrake, as is the 'Baraa' of Flavius Josephus' Jewish Wars, where it is described as shining in the dark (sic) and is attributed exorcistic powers. Pliny in his Natural History and Dioscorides in De Materia Medica wrote that the mandrake was known as 'Circaeon' after Circe, the mythical sorceress who turned men into sexually supercharged swine (Marzell, 1975; Roland 1991; Fleisher & Fleisher, 1 994). The Ancient Greeks and Romans consid- ered the mandrake so potent and valuable as a narcotic and restorative plant that the collectors of its roots had to obey rigid ceremonial rules which were described by Theophrastus and Pliny (Randolph, 1905). In Medieval Europe the plant was known as 'Alraune' in Germany and 'Main de gloire' in France. Carved to emphasize the anthropo- morphic male or female character of the root, it was popular as aphrodisiac, charm against sterility and love talisman alike. But the plant was soon connected with witchcraft; for example, Joan of Arc, who was tortured and burnt at the stake in 143 1 as a heretic, was also accused of the witches' crime of possessing a mandrake (Thompson, 1934). It was believed that the plant grew under gallows and that the root owed its human shape to urine or sperm of unjustly hanged men. The gathering of the root was considered to be highly dangerous and dogs had to perform the supposedly lethal procedure of pulling out the plants, whereupon the roots would shriek so 'that living mortals hearing them run mad': Shakespeare, Romeo and Juliet Act IV (Randolph, 1905; Moldenke & Moldenke, 1952; Marzell, 1975; Roland, 1991; Talalaj et al, 1992; Fleisher & Fleisher, 1994). Perhaps the most interesting aspect of the mythology of the man- drake is convergent myths in the eastern area of the genus in the disjunct region of Central Asia, the Himalayas and China where Mandragora has a similarly long history of application and tradi- tional use. For instance in Sikkim, the roots of Mandragora caulescens, called 'Lakshmana', were used in magical rites and in folk medicine (Mehra, 1979); and in Central Asia the mandrake, called 'Khaoma-soma', was considered a holy plant (Khlopin, 1979). The mythology of the mandrake and its use in traditional medi- cine are hard to keep apart. In a wide range of applications it was used as an aphrodisiac, hypnotic, emetic and purgative, sedative and narcotic, for example as a pain-killer at childbirth as well as to relieve crucifixion agony (Duke, 1985; Talalaj et al., 1992); in fact, according to Heiser (1969), it has been questioned that Christ died on the Cross at all, suggesting that the vinegar supplied to him had contained the drug, inducing merely a deathlike trance. The second- ary metabolites of major pharmaceutical importance are chiefly S. UNGRICHT, S. KNAPP AND J.R. PRESS tropane alkaloids and their esters. Staub (1962), Hegnauer (1973), Romeike (1978), Evans (1979) and Jackson & Berry (1979) list alkaloids of Mandragora and other Solanaceae. Tropane alkaloids occur mainly in the Solanaceae, but they are not entirely restricted to this family; other tropane alkaloids of importance include cocaine, found in the family Erythroxylaceae. However, esters with tropic or related acids of alkamines derived from tropane do seem to be restricted to the Solanaceae (Romeike, 1978). The most important alkaloid components of Mandragora are: atropine, apoatropine, belladonnine, cuscohygrine, hyoscyamine, hyoscyamine-N-oxide, norhyoscyamine, scopolamine, 3oc-tigloyl- oxytropane, 3oc,6p-ditigloyloxytropane, and 6p-hydroxyhyoscy- amine. The root, stem, leaves, fruits and seeds of Mandragora all contain a different alkaloid profile. However, the root is considered to be the site of synthesis of the tropane alkaloids (Romeike, 1978) and is the most poisonous part of Mandragora, due to the high concentration therein. The total content of alkaloids in the root is around 0.4% (Kessler, 1951; Maugini, 1959; Duke, 1985), hyo- scyamine being the most important constituent (Duke, 1985). In contrast, the berries are said to be sweet and edible in moderation (Viney, 1994). The synthetic pathways leading to the alkaloids in different taxa of the Solanaceae have been proposed to be homolo- gous to each other (Tetenyi, 1987) and to be a linear or spiralling phyletic sequence. Tropane alkaloids are effective as analgesics, anaesthetics, antispasmodics, and are used to increase the circula- tion, reduce secretions and dilate pupils. Scopolamine is a hypnotic (Duke, 1985) and achieved a notorious reputation as a 'truth-drug' in police and secret service investigations (Heiser, 1969). 'Quishen', a Chinese herbal medicine consisting of dried roots of M. caulescens, is used in Yunnan and Xizang as a ginseng substitute for all kinds of physical weaknesses (Xiao & He, 1983). In allopathic western medicine, however, the mandrake has largely lost its importance as valued source of alkaloids, although the plant itself is still sur- rounded by myth. Taxonomic history The classification of Mandragora MEDITERRANEAN PLANTS. As mentioned above, considering the small size of the genus, there has been a fair amount of confusion in respect to the classification of Mandragora. The long taxonomic history, especially of the Mediterranean Mandragora species, led to a large number of often imprecisely defined species and subspecies or varieties. As a consequence, there is a plethora of names, and their somewhat indiscriminate use renders the meaning and reliability of identifications of specimens in herbaria, as well as in the literature, highly problematic. Here we present the taxonomic history of the Mediterranean part of the genus in periods, with examples of major taxonomic works and their treatment of the genus, in order to outline the main taxonomic trends through the centuries. Pre-Linnaean: The Ancients distinguished two species that they called the female or black mandrake (Mandragorasfcemina), and the male or white mandrake (Mandragoras mas). These plants were described by both Dioscorides and Pliny (cf. Randolph, 1905 for translations), and this division was widely adopted, for instance by Parkinson (1629) and by Jean Bauhin et al. (1651) as late as the seventeenth century. The male mandrake corresponds to a spring flowering entity; the female mandrake represents an autumn flower- ing one. Caspar Bauhin (1623) and Tournefort (1719), on the other hand, each recognized three 'species'. Both authors listed Man- dragora fructu rotundo and M. /lore subcaeruleo, purpurascente. MANDRAGORA 19 Bauhin's third species was M.fructu pyri while Tournefort's was M. flore subcaeruleo, foliis minoribus, fructu globoso). Linnaeus: In hisHortus cliff ortianus, Linnaeus ( 1 738) considered these variants as mere forms of a single species. In the first edition of Species plantarum (1753), he accordingly recognized only one species that he named Mandragora officinarum. Later, however, he changed his mind with respect to the distinctness of this taxon and placed it within his genus Atropa, calling it Atropa mandragora (Linnaeus, 1759, 1762). Post-Linnaean: Linnaeus's idea of a single species did not last very long. The nineteenth and the first half of the twentieth centuries were characterized by an increasingly elaborate division of the genus (Table 1 ). Heldreich ( 1 886) additionally described an infertile hybrid between two vernal entities: Mandragora vernalis Bertol. and M. haussknechtii Heldr., called M. x hybrida Haussk. & Heldr. Post-1950: In recent decades, however, there has been a trend towards reversing this ever greater subdivision of the genus. In fact, since the 1950s there have been a number of proposals to re-adopt the two traditional, pre-Linnaean taxa only (Tercinet, 1950; Hawkes, 19726; Jackson & Berry, 1979). This viewpoint was adopted in Flora Europaea (Hawkes, 1972a), using the names Mandragora officinarum L. for a vernal entity and M. autumnalis Bertol. for an autumnal one. However, M. officinarum L. might be considered an ambiguous name since it was formerly used for an autumnal entity as well (Greuter & Rechinger, 1967; see Table 1). Hawkes (19726) rejected this reservation on the grounds of the priority, the wide use and the generic type status of the Linnaean epithet officinarum. Knapp in Jarvis et al. (1993) finally lectotypified M. officinarum L. with a specimen in the Bursar herbarium, to which no phenological data are attached. SlNO-HiMALAYAN PLANTS. In contrast to the Mediterranean plants, the classification of the disjunct Central and East Asian part of the genus was never as contentious and did not arouse much comment, in part reflecting its far younger taxonomic history. Mandragora caulescens was described by C.B. Clarke (1883) in Hooker's The flora of British India (Hooker fil. and Thomson labelled the speci- mens 'Scopolia humilis ' but this name was never published). Anisodus caulescens (C.B. Clarke) Diels and A. mariae Pascher, as well as Mairella yunnanensis H. Lev., were placed in synonymy with Mandragora caulescens in the Flora of China (Zhang et al., 1994). Grubov (1970) described a new species from Xizang, China which he named Mandragora tibetica, and Kuang Ko-zen & Lu An-ming (1978) described a new species found in Qinghai and Xizang which they named Mandragora chinghaiensis. In the Flora of China (Zhang et al., 1994) the latter two taxa were provisionally put into synonymy with M. caulescens until a more detailed revision could Table 1 Post-Linnaean classifications of the increasingly subdivided Mediterranean taxa of Mandragora. Classification type Vernal species Autumnal species 1 VERNAL, 1 AUTUMNAL TAXON Sprengel (1825) M. vernalis Bertol. 1 VERNAL, 2 AUTUMNAL TAXA Bertoloni ( 1 835), M. vernalis Bertol. Dunal (1852) 2 VERNAL, 2 AUTUMNAL TAXA Heldreich ( 1 886) M. vernalis Bertol. M. haussknechtii Heldr. 3 VERNAL, 2 AUTUMNAL TAXA Vierhapper (1915) M. mas Garsault M. hispanica Vierh. M. haussknechtii Heldr. M. autumnalis Bertol. M. officinarum L. M. microcarpa Bertol. M. autumnalis Spreng. M. microcarpa Bertol. M. autumnalis Bertol. M. foemina Garsault establish their distinctiveness. Mandragora shebbeard C.E.C. Fischer, described from a single specimen from Xizang (Fischer, 1934), was transferred to the monotypic solanaceous genus Przewalskia by Grubov (1970). One attempt to subdivide Man- dragora caulescens was made by Grierson & Long (1978) who split it into four subspecies (subsp. caulescens, subsp. flavida Grierson & Long, subsp. purpurascens Grierson & Long and subsp. brevi- calyx Grierson & Long) differentiated on the basis of corolla colour and various morphometric characters, especially of flower parts. TURKMENIAN PLANTS. Finally, Mandragora turcomanica Mizg. was described in 1942 from a small area (the Shevlan 'pocket', Mizgireva, 1955) near Kara Kala in the southwestern Kopet Dag in Turkmenistan. Only a few collections of this restricted endemic exist, and these are all to be found in herbaria of the former Soviet Union. Its status as a distinct taxon has never really been questioned, but only because so few specimens exist for comparison. Mizgireva ( 1 955) worked for 1 2 years on the detailed anatomy and morphology of this species and her revision remains the best source for data on characters and their variation. The position of Mandragora within the Solanaceae More than 200 years after the original outline by Jussieu (1789), the higher level classification of Solanaceae is far from being settled, and the placement of Mandragora in the various proposed classifi- cations is just as controversial. In the following review, the most widely followed classifications of the Solanaceae are presented together with the positions of Mandragora and the other genera used in our cladistic analysis: Anisodus Link, Atropa L., Lycium L. and Nicotiana L. In his artificial sexual system, Linnaeus (1753) placed Man- dragora within his group Pentandria Monogyna, together with Atropa, Lycium and Nicotiana. Jussieu (1789) defined the 'order' Solaneae with Mandragora in group II (fructus baccatus) alongside Atropa and Lycium. Nicotiana on the other hand was placed in group I (fructus capsularis). In the nineteenth century, classifications be- came more elaborate and tribes and subtribes were established in order to approximate a more 'natural' classification. In Table 2, the three most widely adopted classifications of the Solanaceae in this period (Dunal, 1852;Bentham, 1 876; Wettstein, 1895) are presented together with the more recent one of Baehni (1946). The last few decades have seen the analysis of new types of data, such as DNA sequences (Olmstead & Sweere, 1994) and secondary plant metabolites (Tetenyi, 1987), as well as application of cladistic methodology to morphological data (Hoare & Knapp, 1997). This has led to new hypotheses concerning relationships within the Solanaceae, as well as the placement of Mandragora. However, opinion still differs widely on whether Mandragora should be placed within the tribe Hyoscyameae (Hoare & Knapp, 1997; = Atropeae sensu Tetenyi, 1987) or in an isolated, basal monogeneric tribe Mandragoreae (Olmstead & Sweere, 1994; Hunziker, 1995; Olmstead et al., in press). MATERIALS AND METHODS Material examined This revision is based exclusively on herbarium specimens. The material examined includes the Mandragora specimens in the her- baria of The Natural History Museum of London (BM) and Kew (K) together with specimens borrowed from the following herbaria: 20 S. UNGRICHT, S. KNAPP AND J.R. PRESS Table 2 Tribal classification of the Solanaceae and the position of Mandragora and the outgroup genera Anisodus, Atropa, Lycium and Nicotiana. Author Classification Genera of this study Dunal(1852) Nolaneae Nolanineae Grabowskieae Solaneae Triguereae Solaneae Solanineae Atropineae Mandragora Atropa Lycineae Lycium Datureae Hyoscyameae Anisodus (Scopolia) Nicotianeae Nicotiana Retzieae Fabianeae Metternichieae Cestreae Bentham(1876) Solaneae Atropeae Mandragora Atropa Lycium Hyoscyameae Anisodus (Scopolia) Cestrineae Nicotiana Salpiglossidae Wettstein(1895) Nicandreae Solaneae Lyciinae Atropa Lycium Hyoscyaminae Anisodus (Scopolia) Solaninae Mandragorinae Mandragora Daturae Cestreae Cestrinae Goetzeinae Nicotianinae Nicotiana Salpiglossideae Baehni(1946) Solaneae Solaninae Sarachinae Margaranthinae Physalidinae lochrominae Goetzeinae Discopodiinae Atropeae Atropinae Mandragora Atropa Lycium Markeinae Hyoscyaminae Anisodus (Scopolia) Parabouchetiinae Anthocercideae Nicotianeae Nicotianinae Nicotiana Daturinae Nicandrinae Salpiglossideae Berkeley (UC, JEPS), Edinburgh (E), Kunming (KUN), Palermo (PAL) and Reading (RNG). The collections of the Museum National d'Histoire Naturelle in Paris (P) were examined on site during a research visit. Regrettably it was not possible to obtain any speci- mens, including the types, of Mandragora turcomanica and M. tibetica from St. Petersburg (LE) or M. chinghaiensis from Xining (HNWP). Material with the same collection number or specimens collected at the same locality and at the same date were unified and treated as one sample. Unlocalized material or specimens with illegible, doutbful or very imprecise collection locality and collector data were excluded, cultivated material being the exception. Only collec- tions with precise locality data are listed in the taxonomic account of the species. A complete listing of the specimens examined can be found in the Exsiccatae. Taxonomic concepts The adopted taxonomic methodology in this study of herbarium material is necessarily largely pattern-oriented. If process-based terms like 'population' or 'hybrid' are used, these properties are only informally applied, i.e. inferred rather than rigidly defined, let alone experimentally circumscribed. The adopted species concept is operational, i.e. the ability to distinguish between sets of specimens serves as the criterion for the delimitation of the terminal taxa (species). Cladistic analysis results in an estimate of these relationship between the species and the delimitation of higher level taxa, i.e. monophyletic groups and their hierarchical relationships. The requirement for a species description is thus at least one distinguishing qualitative, or discontinuous quantitative, diagnostic character state (monothetic species) or a unique set of character states (polythetic species) (cf. Nelson & MANDRAGORA Platnick, 1 981; Nixon & Wheeler, 1990). Morphological and pheno- logical variation plus ecological and geographical information are the data that were collected from herbarium specimens and their labels respectively. Morphometrics Specimen sampling The inherent weakness of the traditional herbarium record of a taxon for taxonomic studies is well known (Du Rietz, 1930; Anderson & Turrill, 1935; Anderson, 1941). For species that are well represented in herbaria, one can obtain a more or less accurate record of the general range of variation. Seldom or never, however, is it possible to calculate frequencies of the different variants and the resolution of the classification is limited by the lack of a random sample. Plant variation also occurs on different hierarchical levels (cf. Bateman, 1989), which can become confused and overlap. In Man- dragora the confounding of within-group and between-group variation is most pronounced in the Sino-Himalayan complex, due to the secondary growth of the plants during the late flowering and the fruiting period (ontogenetic variation, see Fig. 9, also Hoare & Knapp, 1997: fig. 5d). Combining all samples to a pooled matrix easily leads to a confusion of the two sources of variation which can seriously compromise interpretation of many applications of morphometric analyses (Gibson et al, 1984). Avoiding within- group variation, for example by calculating the mean or only considering the largest or smallest example of a particular character is not a solution, but merely reduces the total amount of variation encountered in a taxon, leading ultimately to a simplistic taxon delimitation. In the taxonomic process, characters and their variabil- ity necessarily have priority over all other considerations since taxa are attributed to characters — not the other way around (characters all attributed to individuals). It follows that the basal unit of investiga- tion, the specimen, must be evaluated in respect to its variability. Therefore, maximal as well as minimal values of the flowering part measurements of each specimen were assessed, rather than solely mean or maximal values. Character sampling A genus with a long taxonomic history is unlikely to yield any 'new' gross morphological characters which distinguish species. This is especially true when working with inadequate herbarium material (i.e. incomplete with much missing data and/or very old and often in poor condition). The emphasis in this study was thus much more on evaluating the traditional and obvious characters in respect to their ability consistently to delimit proposed taxa. The analysis was therefore carried out using the following data. GEOGRAPHY. The entire distributional range of Mandragora was divided into 20 geographic areas. These are (1) Cultivated speci- mens, typically from botanic gardens north of the Alps, (2) North Africa, (3) Iberia, (4) northern Italy and Dalmatia, (5) peninsular Italy with Sardinia and Sicily, (6) mainland Greece, (7) Aegean islands, (8) Asia Minor, (9) Cyprus, (10) Levant, (11) Kopet Dag, (12) Nepal, (13) Sikkim, (14) Bhutan, (15) Arunchal Pradesch (Assam), (16) north Myanmar (Burma), (17) Yunnan, (18) Sichuan, (19) Qinghai and (20) Xizang. PHENOLOGY. Flowering and fruiting dates were recorded, result- ing in the cumulative reproductive period. MORPHOLOGY. Corolla length, calyx length, fruit size, fruiting calyx length, leaf length, leaf length/width ratio. The corolla colour 21 of herbarium material is not reliably determinable and was thus only recorded if it was stated on the labels. The shape of the berry (globose or ovoid) was excluded as a character since it is not accessible through pressed herbarium material and is only rarely stated on the labels. Berry shape in the Solanaceae is quite variable and this variation is occasionally a developmental sequence (Bohs, 1994). Data analysis Univariate and multivariate analyses were applied as a descriptive, explorative tool to study the joint relationships of the morphological, phenological and geographical variables of the specimens at hand. As mentioned above, since the examined specimens are not a random sample, the result of inferential data analysis with signifi- cance testing would be invalid. A cluster analysis or a cladistic analysis of the data at population level, on the other hand, was rejected on the grounds that these methods assume hierarchical relationships, an assumption that is not justified. Phenotypic charac- ters may be distributed clinally or in a reticulate manner, but are unlikely to be strictly hierarchical (Crisp & Weston, 1993). If hierarchical structure exists, it would suggest that the populations have already undergone speciation and have to be treated as terminal taxa in the phylogenetic analysis. The explorative data analysis was performed on SYSTAT for the Macintosh Version 5.2 (Wilkinson, 1992). UNIVARIATE ANALYSES. Phenological data are circular (Fisher, 1993). The reproductive period was thus analysed in a polar coordi- nate system. In order to dampen peak values, which might merely reflect a particularly high collecting effort, the collection proportion of each month was converted to its logarithm. MULTIVARIATE ANALYSES. Ordination techniques reduce a data matrix of distances (similarities) among the samples to one or a few dimensions. These types of techniques seem to be most appropriate under a nonhierarchical model of infraspecific variation. Principal component analysis (PCA) is the method most commonly applied in multivariate systematics studies (James & McCulloch, 1990). It aims to show the eventual existence of distinguishable groups with discontinuous variation without a priori postulates of groups such as populations. Since data sets with missing data cannot be processed using PCA, only the most commonly represented characters of the herbarium specimens (see above and Table 3) were analysed. Fortu- nately, these are also the characters that are most commonly applied to differentiate between the traditionally recognized Mediterranean taxa. In constrast to many other analyses of specimen-based data, within-collection variation was accounted for by recording and analysing both minimal and maximal values of the flowering part lengths. Only fully developed and open flowers were considered and measured. The length/width ratio of the leaves was not included in the PCA analysis on the grounds that ratios are often not linearly related to the length measurements (Miles & Ricklefs, 1 984). All the variables have been standardized (normalized) by the standard Table 3 Variables used in the principal component analysis of the Mediterranean complex. Length measurements Variable 1 Corolla length max. mm Variable 2 Corolla length min. mm Variable 3 Calyx length max. mm Variable 4 Calyx length min. mm Variable 5 Leaf length max. cm Phenology Variable 6 Flowering time month 22 deviation to obtain a variance of one and a mean of zero by performing the analysis on a correlation matrix. Due to a lack of available specimens from Asia of the taxa Mandragora caulescens subsp. brevicalyx, M. chinghaiensis, M. tibetica and M. turcomanica, the discussion of variability among these taxa (and its implications on their delimitation) has to remain qualitative. Cladistics Cladistic analysis is a parsimony-based search for hierarchical arrangements or patterns of terminal units (in most cases species). The premises are accordingly that there is a hierarchical structure of the taxa and therefore of the characters used to define them, and that the level of homoplasy in the chosen characters is low enough not to infer convergent groups. As a strictly numerical procedure, there is no need for additional models or assumptions relating to biological processes, such as reproduction or descent (Davis & Nixon, 1992). The resulting cladogram therefore represents a synapomorphy scheme, merely suggesting closeness of relationship due to recency of shared common ancestry. It might be congruent with the actual phylogenetic tree, featuring the ancestors and descendants; how- ever, there are many possible phylogenetic trees compatible with one cladogram (Eldredge, 1979). The cladogram was rooted by means of outgroup comparison. A phylogeny of the tribe Hyoscyameae based mainly on morpho- logical characters was recently published by Hoare & Knapp (1997) suggesting that Mandragora, as traditionally defined, is mono- phyletic. To test the monophyly of Mandragora as a whole, we included in this analysis material of species representing two other genera of the Hyoscyameae, Anisodus luridus Link and Atropa belladonna (see Table 4). The three recognized terminal taxa of Mandragora represent the ingroup (M. caulescens, M. officinarum, Table 4 Taxa used in the cladistic analysis. Nicotiana glauca Graham Lycium chinense Mill. Anisodus luridus Link Atropa belladonna L. Mandragora officinarum L. M. turcomanica Mizg. M. caulescens C.B. Clarke S. UNGRICHT, S. KNAPP AND J.R. PRESS Table 6 Data matrix used in the HENNIG86 analysis Taxon Character 1 I 1234567890 1 Nicotiana glauca Lycium chinense Anisodus luridus Atropa belladonna Mandragora officinarum M. turcomanica M. caulescens 00200010000 0020000001 0 11201 111110 10201 10001 1 11011 10001 0 11011 10101 ? 11111 10111 1 M. turcomanica). The outgroup, Nicotiana glauca Graham, was chosen based upon previous cladistic analyses of chloroplast DNA data sets (Olmstead & Palmer, 1992; Olmstead & Sweere, 1994; Olmstead et al., in press). Lycium chinense Mill, was included as an additional taxon (but not defined as an outgroup, see Tables 2 and 4). Although acknowledging that there is no such thing as an a priori 'perfect cladistic character' (Thiele, 1993), the suggestion of Chappill (1989) and Stevens (1991) not to include purely morphometric, quantitative data in the cladistic analysis was followed. This limita- tion is feasible in view of the minimal size of the ingroup and the already established analysis of an extensive character set including quantitative character states (Hoare & Knapp, 1997). The characters used in the present work (Table 5) and their states are largely self- explanatory. The data matrix used in the analysis is presented in Table 6. Cladistic analyses were undertaken withHENNlG86 (Farris, 1988) using the ie option (implicit enumeration) with all characters unordered, thus avoiding any character state change assumptions. The generated cladogram statistics include length (L), the number of character state changes on the tree, the ensemble consistency index (CI) which provides a measure of character fit on the cladogram (amount of homoplasy), and the ensemble retention index (RI) which expresses the fraction of similarities on the cladogram inter- preted as synapomorphy (Farris, 1989). Both of these indices have an optimal value of 1 . RESULTS AND DISCUSSION Table 5 Character set used in the cladistic analysis. Habit 1 . Woody plant (shrub or tree) 0; herb 1 Root 2. Tap-root: enlarged tap-root absent 0; present 1 Stem 3. Axis: condensed 0; first condensed, later elongated 1 ; elongated 2 Leaves 4. At anthesis: leaves clustered in a rosette 0; leaves alternate 1 Inflorescence 5. Type: numerous flowers in panicle 0; l(-3) axillary flowers 1 Corolla 6. Shape: tubular-infundibuliform 0; campanulate-urceolate 1 Fruit 7. Type: berry 0; capsule 16 Fruiting calyx 8. Not enlarged in fruit 0; markedly enlarged in fruit 1 9. Without prominent ribs 0; with prominent ribs 1 Seeds 10. Embryo: not curved 0; curved circularly or in a spiral 1 1 1 . Testal cells deep 0; shallow 1 Species delimitation using morphometric analyses For the Mediterranean taxa a number of specific hypotheses were tested using the above described statistical analyses. Univariate analysis of reproductive period. — Are there two distinct vernal and autumnal taxa ? By adopting the two species Mandragora vernalis and M. autumn- alis, Sprengel ( 1 825) argued against Linnaeus's unified Mandragora officinarum concept. Even in later, more elaborate classifications the division into these two major groups remained important (Table 1) since it was obvious that two groups with widely differing flowering periods would hardly be able to interbreed due to the seasonal isolation. So, is there evidence in the compiled Mediterranean data set that there are two phenologically distinct species? Figure 1 shows the distribution of the collecting dates (in flower or in fruit) as an indicator of the reproductive period. It is evident that there is actually only one, very extensive period of reproductive activity from autumn to spring. In fact only during the hottest summer months does the cycle break. If conditions are MANDRAGORA 23 Mar Jun Feb Jan Dec Sep Nov Fig. 1 Relative distribution of the vegetative period of the Mediterranean complex (Mandragora officinaruiri) in the course of the year (logarithimized monthly fractions: for analysis see text). favourable, the same individual can flower twice a year, in autumn and then again in spring, as can be seen from annotated herbarium material, especially from cultivated material in botanical gardens. The key factor is probably sufficient rainfall; for instance, sudden and widespread flowering after exceptionally heavy autumn rains was recorded from Tunisia (label data from Tomkinson 72). A more detailed discussion pertaining to the influence of the climate and the summer dormancy is given in the section on ecological biogeo- graphy below. Multivariate analysis of morphometric characters. - Are there distinct clusters within the Mediterranean mandrakes that are recognizable from gross morphology? The different eigenvector values of the PCA (Table 8) indicate the relative contribution of the variables. The first three axes (see Table 7) explain 92.2% of the variation if length measurements are consid- ered only (Fig. 2a), and 87. 1% of the variation if the analysis was run including the phenological variable of flowering time (Fig. 2b). In both cases it is obvious that no clear-cut clusters can be distin- guished. Cryptic species may exist within the Mediterranean region but are likely to be revealed only if extensive, field-based studies over the entire range, as well as transplant experiments, are under- taken (Clausen et al, 1940). The information currently available does not justify recognition of more than a single species of Mediter- ranean mandrake. Other characters traditionally applied in keys but not analysed quantitatively here include the following: Corolla colour. Hawkes (1972b) considered the elaborate split- ting of the vernal and autumnal groups as unjustified and recognized only the white-flowered vernal plants from northern Italy and Dal- matia as distinct from the remaining Mediterranean group. Restricted to only a small part of the total Mediterranean range, the white- flowered plant has often been considered as the most distinctive group (Heldreich, 1886; Vierhapper, 1915). The diagnostic value of this character which has been extensively used in past and current Table 7 Principal components (PC) of the Mediterranean complex. A: Length measurements only. B: Length measurements and flowering period combined. Component Eigenvalue % of variance explained A PC 1 3.248 64.97 PC 2 0.977 19.54 PC 3 0.383 7.66 B PC 1 3.328 55.47 PC2 1.410 23.50 PC 3 0.490 8.17 Table 8 Eigenvectors of the three most important axes (component loadings). Variable PC 1 PC 2 PC 3 A 1 Corolla length max. 0.885 0.023 0.131 2 Corolla length min. 0.875 0.011 0.435 3 Calyx length max. 0.898 0.114 -0.391 4 Calyx length min. 0.924 0.065 -0.147 5 Leaf length max. -0.197 0.979 0.047 B 1 Corolla length max. 0.882 0.067 -0.160 2 Corolla length min. 0.851 0.189 0.042 3 Calyx length max. 0.897 0.123 0.066 4 Calyx length min. 0.905 0.156 0.175 5 Leaf length max. -0.181 0.865 -0.453 6 Flowering time -0.413 0.763 0.471 keys is minimal, however, if one considers that flowers are very variable in colour even within individuals (Jury et al. 12238 from Morocco, Jury & Jury 13111 from Spain). Fruiting calyx length: The relative length of the accrescent fruit- ing calyx to the berry (longer: M. autumnalis or shorter: M. officinarum) has also been proposed as a diagnostic character (Hawkes, 19726). Again, it seems that within-group variation is just as high, considering for instance a specimen housed in P of a cultivated plant which features two berries that are clearly shorter than the calyx, but three others which are longer. Leaf shape: This is perhaps the most variable character of all. One can get at least an idea of the variability if the length to width ratios of mature leaves are compared. The variability within an individual ranges from 1.7:1-4:1 (in Reverchon s.n.) to 4.6:1-10.5:1 (in Welwitsch 159). The total variability within the whole set of exam- ined specimens was found to be between 1.5:1-10.5:1. Any distinction between Mandragora officinarum and M. autumnalis or any other splitting of the Mediterranean Mandragora on morphological or phenological grounds would be intentionally giving formal status to divergent ends of clines. Thus overlapping intermediates would be difficult if not impossible to assign to either of the taxa in any other way than arbitrarily. Extensive overlap and putative hybrids must be considered if aiming at expressing the whole range of variability. This confirms Meikle's (1985) doubts concerning the validity of the different proposed Mediterranean species. Although certain plants of particular populations are quite different in appearance from some plants of other populations, when the total pattern of variation over the entire range is considered, no diagnosable taxa within the Mediterranean complex of Mandragora are discernable. Further morphological support for the unification of the Mediterranean Mandragora comes from the anatomy of the tap roots which has been shown to be identical in vernal and autumnal taxa and in their chemical composition which is also similar (Jackson & Berry, 1979). For Mandragora turcomanica, a statistically valid morphometric analysis is not feasible with the available literature data only. The 24 S. UNGRICHT, S. KNAPP AND J.R. PRESS /.• •/ • •• I •2 f 1 •2-10 1 FACTOR(2) •1 0 1 FACTORO) -3 I T I T J I I I -2 -1 0 1 2 3 -4 -3 -2 -1 0 1 2 3 FACTORC2) FACTOR(3) Fig. 2 Principal component analysis of the Mediterranean complex (Mandragora officinarum). Top pair: a. Bottom pair b. Left: PC 1,2. Right: PC 2,3. range of the Turkmenian plants is so small that variability at popula- tion level is considered equivalent with the variability over the whole range. Mizgireva ( 1 955) meticulously documented teratogenic forms of flowers and fruits of M. turcomanica, but did not cite field collected vouchers for these forms. She also documented huge variability in the shape and size of tap roots, and described the growth of the plant in detail, from seed germination to fruiting. Mandragora turcomanica is accepted as a distinct species on the grounds of the evidence given in Mizgireva (1942, 1955, 1978) and Lincevskij (1955). The whole plant is much larger than M. officinarum. The lower leaves (± 90 x 60 cm) are especially large resulting in a rosette with a diameter of 150-180 cm. Mandragora turcomanica blooms in both the autumn and the spring (Mizgireva, 1955, see taxonomic treatment), with different individuals in the population exhibiting one or the other flowering time. Geographi- cally the species is isolated, occuring only in a narrowly restricted range, and within that range it is only found in localized regions (Mizgireva, 1955). It is also widely disjunct, being some 1500 km from the closest Mediterranean Mandragora population in the Near East and about 2500 km from the nearest Himalayan population in western Nepal. In common with the Mediterranean mandrake, the Himalayan plants make up a complex with considerable variation in size, shape and colour of the flowers and leaves. The following questions are addressed: MANDRAGORA Is the recognition of the four infraspecific taxa sensu Grierson & Long within Mandragora caulescens sensible? After checking herbarium material from almost the entire range of Mandragora caulescens (north-western Sichuan and southern Qinghai excluded), Grierson & Long (1978) were well aware of the extreme variability of the morphological characters they used to describe their four subspecies. The characters used were corolla colour, calyx and corolla lengths and overall plant size. As with Mandragora officinarum, flower colour, according to observations of the fresh material given on herbarium labels, is variable within individuals as well as within local populations (e.g. Alden et al. 1 185: 'flowers yellow to purplish-brown', Long et al. 266: 'corolla yellow, purple at base, inside and out', and Polunin 631: 'petals yellow with purple veins'). Colours probably merely reflect to what extent the yellow pigments are masked by purple ones. Grierson & Long (1978) were unable to find any diagnostic morphometric discontinuities. Their proposed classification is based on the rule that due to its type status, the holotype of M. caulescens, collected in Sikkim and representing an intermediate between the extremes of the 'typical' small, yellow-flowered plants and the large, purple- flowered ones, must retain the name M. caulescens (and must thus be given the subspecific epithet caulescens). They therefore split the complex not into two but into three subspecies representing the two extreme forms (subsp. flavida and subsp. purpurascens) and the intermediates (subsp. caulescens), and additionally described a fourth subspecies with a shorter calyx (subsp. brevicalyx) based on only two collections from Xizang. As a consequence, their classifi- cation is based on non-diagnostic, largely overlapping characters, rendering M. caulescens subsp. caulescens as a 'basket' taxon for any intermediates. Indeed, it might be hypothesized that there are at least two different species present within this complex with frequent 25 introgression of the sympatric populations. However, the many putative 'hybrids' linking them and resulting in an extensive overlap of their features make them unsuitable as terminal taxa for the cladistic analysis. Furthermore, a restrictive taxonomic concept as described earlier does not allow for the recognition of infraspecific taxa if their characters and their ranges are extensively overlapping. Are Mandragora tibetica and M. chinghaiensis distinct species or are they both conspecific with M. caulescens? As in the case of the Turkmenian mandrake, the unavailability of the types and a reasonable number of specimens renders the data somewhat meagre, so the original descriptions by Grubov ( 1 970) for Mandragora tibetica and by Kuang & Lu ( 1 978) for M. chinghaiensis must provide much of the necessary information. These descrip- tions, together with their accompanying keys, which distinguish the newly described species from M. caulescens, recognize the small, yellow-flowered plants as distinct from the larger, purple-flowered ones. Both new taxa are therefore covered by the range of variability given in Grierson & Long's ( 1978) subdivision of M. caulescens into four subspecies, which are defined only by overlapping differential characters and, as such, are not acceptable as distinct taxa. Indeed, a specimen in P, Polunin et al. 878, is annotated asM. chinghaiensis by Lu An-ming and is also an isotype of M. caulescens subsp. flavida. Some doubts must remain concerning M. tibetica and M. caulescens subsp. brevicalyx that together may represent a single distinct taxon with a very short calyx. An unusual specimen from western China (Wilson 4195 (K), no precise locality given) bears a single flower with a conspicuously narrow corolla. A note by Grierson and Long on the herbarium sheet suggests that they considered this specimen not conspecific or even congeneric with M. caulescens. Without further material, however, proposing a new taxon would be just as Fig. 3 The single most parsimonious cladogram from the HENNIG86 analysis. The characters are discussed in the text, and characters states are shown in Table 5. For characters marked on the branches of the cladogram: single lines indicate unambiguous synapomorphies, parallel lines parallelisms and crosses reversals (homoplasy). 26 S. UNGRICHT, S. KNAPP AND J.R. PRESS speculative as disregarding the flower as a mere artefact of prepara- tion. The synonymy presented in the taxonomic treatment here, especially in respect to the undercollected plants from central and northeastern Xizang, is therefore provisional and subject to confir- mation once more material of both flowering and fruiting plants becomes available. Only then will it be possible to judge whether or not there exist distinct taxa not included within the range of variation of M. caulescens. Mandragora is a genus with pronounced variability, both within and between individuals and therefore necessarily within the pro- posed polymorphic species. Furthermore, discordant variation in the applied gross morphological characters impedes a workable subdi- vision based on a whole set of differential characters, rather than just one (apparently non-existent) diagnostic character. Phylogeny The cladistic analysis resulted in a single most parsimonious cladogram of length = 16, CI = 75 and RI = 73 (Fig. 3). The genus Mandragora defined in the traditional sense is monophyletic in our analysis: the species share synapomorphies in characters 3 (a con- densed axis), 4 (leaves clustered in a rosette) and 9 (fruiting calyx with prominent ribs). If the inaperturate pollen known from M. caulescens and M. officinarum (see Hoare & Knapp, 1997 for details) is also found in M. turcomanica, this too would be a synapomorphy of the genus. Mandragora officinarum and M. turcomanica are more closely related to each other than to M. caulescens. They share reversals in characters 3 (a condensed axis) and 9 (a fruiting calyx without prominent ribs), but no unambiguous synapomorphies. The peculiar life-cycle, i.e. the mode of development of the leaves and flowers, is evidence for a shared common ancestry of the Mediterranean- Turkmenian clade. Mizgireva (1955) also suggested this in her revision of the Turkmenian species. In order to determine more clearly the relationships among the three species of Mandragora, a different, probably field-based set of characters is necessary. Here, however, we are concerned more with the delimitation of terminal taxa, especially among the Mediterranean plants, and whether or not the genus as currently defined is monophyletic. A comparison between alternative hypotheses concerning the position of Mandragora based on morphological, chemical and molecular (chloroplast DNA) characters is given in Figure 4. The most pronounced discrepancy concerning the position of Man- dragora is that the molecular data set (Fig. 4a) deviates from the morphological (Fig. 4c) and chemical ones (Fig. 4b) by attributing this genus a very independent position due to extremely divergent and autapomorphic cpDNA sequences. This is reflected in the proposal of a monogeneric tribe Mandragoreae (Olmstead et al., in press). Morphological and chemical evidence, on the other hand, suggests a placement within the tribe Hyoscyameae (Hoare & Knapp, 1997; = AtropeaesenswTetenyi, 1987). The present study is far too limited in its scope to allow any conclusions concerning higher level classification within the Solanaceae. However, it is clear that our limited data set supports the inclusion of Mandragora within the Hyoscyameae (synapomorphies in characters 1, 5 and 6) but, as mentioned above, a much larger and more field-based data set will be needed to address this problem. Mandragora is an anoma- lous genus in the family, both morphologically and in terms of molecules, but this should not prevent us from eventually identifying its true affinities and relatives given adequate data. Biogeography The biogeography of an exclusively Laurasian genus within the Solanaceae is of interest because the family is largely Gondwanan in distribution. It is useful to approach biogeography from two differ- ent angles. Ecological biogeography considers a short temporal scale in evaluating ecological factors relevant for present-day distri- butions of taxa. Historical biogeography, on the other hand, is concerned with the change of distributions and the potential causes thereof on a long temporal scale. We attempt here to summarize some of the most important data with respect to the geographical distribution of Mandragora, both now and in the past, and pinpoint major gaps in knowledge. Ecological biogeography The present approximate distribution of the genus is given in Figure 5. Apart from the herbarium material examined, the sources pre- sented in Table 9 were used. Most comprehensive are the distribution data in Heldreich (1886) and Vierhapper (1915). Mandragora does not appear to penetrate into the Arabian Peninsula, being absent ABC Fig. 4 Differing hypotheses of the relationships of Mandragora within the Solanaceae. a: Olmstead et al., in press, b: Tetenyi, 1987, c: Hoare & Knapp, 1997. MANDRAGORA 27 20 Fig. 5 Distribution of the genus Mandragom. 1) M. officinarum, 2) M. turcomanica, and 3) M. caulescens. even in the most northern parts of Saudi Arabia (Migahid, 1989) as well as from Mesopotamia (Rechinger, 1964; Mouterde, 1979). The genus was naturalized in Malta in the nineteenth century but is now presumably extinct there (Borg, 1927; Maugini, 1959). Mandra- gora does not appear in works covering France and Corsica (Fournier, 1990), Albania (Demiri, 1983), Egypt (Tackholm, 1974; Boulos, 1995) or Libya (Siddiqi, 1978) and is presumed absent from these areas. Likewise, the genus is absent from the Macaronesian archi- pelagos of the Canaries, Madeira and the Azores (not recorded either by Hansen & Sunding, 1993; or Press & Short, 1994). Mandragora turcomanica is found only in the Kopet Dag range of southern Turkmenistan. It has not yet been reported from the Iranian part of these mountains (Rechinger, 1972). In eastern Asia, Man- dragora (M. caulescens) occurs in the central and eastern Himalaya Table 9 Sources used for determining the overall distribution of the genus Mandragora. Geographical area Reference Tunisia Algeria Morocco Portugal Spain (incl. Balearic Is.) Italy (incl. Sicily, Sardinia) Former Yugoslavia Greece (incl. Aegean Is.) Turkey Cyprus Syria Lebanon Israel Jordan Turkmenistan Nepal India (Sikkim, Arunachal Pradesh) Bhutan Myanmar China Bouquet (1952), Pettier- Alapetite (1981) Quezel& Santa (1963) Jahandiez & Maire (1934) Amaral Franco (1984) Lazaro e Ibiza (1907), Bonafe Barcelo (1980), Valdesetal. (1987) Maugini (1959), Pignatti (1982), Cellinese et al. (1994) Hayek &Markgraf( 1931) Halacsy (1902), Hayek & Markgraf (1931) Baytop(1978) Meikle (1 985), Viney ( 1994) Post (1933), Feinbrun-Dothan (1978), Mouterde (1979) Post (1933), Feinbrun-Dothan (1978), Mouterde (1979) Post (1933), Feinbrun-Dothan (1978) Post (1933), Feinbrun-Dothan (1978) Pojarkova (1955), Kurbanov ( 1 994) Grierson& Long (1978) Hara (1966), Grierson & Long (1978), Deb(1979) Hara (1971), Grierson & Long (1978) Grierson & Long (1978) Grierson & Long (1978), Zhang et al. (1994). Zhang &Lu( 1994) and southwestern China, but there are no records from the western extensions of these mountain ranges (Karakorum, Pamir, Hindu Kush and Tien Shan). Apart from historical factors, dealt with below, the principal factor determining the geographical distribution of taxa on a large scale is typically climate. Although one might disagree on the relative importance of means and extreme values of various climatic variables, the prime importance of climate in general has long been accepted (Cain, 1944). A convenient way of depicting and comparing different climates is by means of standardized diagrams. The purpose of the so-called ecological climate-diagrams is to portray the climate of a locality in a readily comparable way and visually to present the most important climatic factors affecting the growth of plants (Walter et al., 1975). By choosing a scale at which 10°C correspond to 20 mm precipita- tion, a relatively wet (dark) and relatively dry (light) season can be identified on the diagrams (Fig. 6). These climate diagrams make it evident that the range of Mandragora comprises not only quantita- tively different climates but qualitatively different climate types. The amount and especially the main season of rainfall is particularly significant. The Mediterranean climate is characterized by mild, rainy winters and hot, dry summers. Although the total amount of rainfall for other locations for Mandragora officinarum may differ somewhat from that presented here (Iraklio, on the northern coast of Crete, Fig. 6), the general aspect of a winter rainfall regime remains the same all over the Mediterranean basin, even extending towards Afghanistan and the Hindu Kush. The typical vegetation consists of adapted sclerophyllous vegetation, analogous to that found in the same climate type in California, Chile, the Cape region of South Africa and southwestern Australia. The Kopet Dag is a Transcaspian mountain range surrounded by the Irano-Turanian desert. The climate there represents a continen- tal, dry version of the Mediterranean type with much lower rainfall (Fig. 6: Kisil-Arvat)) and an extended summer arid period. During this dry summer, Mandragora turcomanica exhibits a long period of xerothermic dormancy (Kurbanov, 1994). Mizgireva (1955) sug- gested that M. turcomanica is much more cold-tolerant than its mediterranean congeners. The typical vegetation in the Kopet Dag consists of broadleafed deciduous forest in the moister microhabitats and open shrubland in drier areas. The eastern part of the Mandragora range in the Sino-Himalayan 28 S. UNGRICHT, S. KNAPP AND J.R. PRESS Irakiio 27 m alt. Gangtok 1725m alt. IN so M 4* 20 0 Kisil-Arvat 101 m alt. 100 m M 40 20 0 30 20 10 0 Gyantse 3996 m alt. A Fig. 6 Climate diagrams of four representative weather stations in three different climate regimes. Given are the rainfall and temperature curves, resulting in wet (dark shading) and dry (light shading) seasons. Based on Walter et al. 1975. region is characterized by a completely different rainfall regime altogether. The influence of the monsoon renders not the winter but the summer the season of the highest rainfall. The amount of precipitation during the monsoon period, however, does vary enor- mously. The Himalayan chain functions as a rain screen resulting in high rainfall on the southern slope of the mountain range (Fig. 6: Gangtok) as well as in the southwestern Chinese mountains of Yunnan and Sichuan, and much lower rainfall in the inner regions of Xizang (Fig. 6: Gyantse). Even at high altitudes, winters are very mild in the high rainfall areas (see climate-diagram of Gangtok, about 1700 m above sea level) and snow is scarce. Generally speaking, the eastern Himalaya and southeastern Xizang, Sichuan and Yunnan enjoy a much more humid and milder climate than the western Himalaya and central Xizang. The locality data of the examined specimens suggest that the distributional boundary of the Himalayan mandrake towards the central Xizang highland lies within the 200-400 mm annual precipitation strip north of the main ranges. Due to the immense altitudinal gradient, the vegetation is strikingly zonal and ranges from subtropical forests to alpine scrub and meadows. The upper forest limit lies between about 3600 and 4200 m, the upper tree limit between 4200 and 4500 m. The belt in between is characterized mainly by a succession of various Rhodo- dendron species. This is the typical habitat of Mandragora caule- scens. The permanent snow line lies at about 4800 m in the south and 5800 m in the north towards central Xizang (Walter, 1962). Other factors affecting plant distribution, albeit on a smaller and less important scale, include soil type and competition. Man dragora officinarum especially grows in ruderal habitats, on clayey and nitrogen-rich soils. So far, it is not clear whether M. caulescens occurs on acidic (granitic) or base-rich (limestone) soils in the subalpine and alpine zone of the Himalaya. It cannot be ruled out that there might be two different, vicariant ecological groups as is often the case in the Alps, for example in Rhododendron (Walter, 1966). The type of climate and soil typically only indirectly deter- mine the occurrence of a species in a certain locality, in that they affect the fitness of that species in relation to others (Walter, 1977). Both Mandragora officinarum and M. turcomanica seem more or less restricted to low-competition localities, representing typical r- species (good colonizers but poor competitors). Historical biogeography The atemporal cladogram of a group together with historical evi- dence, namely paleoclimatic and paleogeographical data allow for the formulation of historical scenarios. A scenario is a model of the basic features (including space and time) of the evolutionary history of a group which can serve to stimulate directions of further investi- gation. However, the essentially speculative character of such a model renders it more like a tool than a serious scientific statement (Eldredge, 1979) and ad hoc assumptions are inevitable (Humphries & Parenti, 1986). In attempting to construct scenarios for the historical biogeography of any group, plate movement, eustasy (worldwide changes of sea level) and climate changes are consid- ered the major factors to be considered (Hallam, 1981). In the case of the Solanaceae, virtually no fossil record is available (PaleoNet Listserver, http://www.ucmp.berkeley.edu/Paleonet/), thus histori- cal hypotheses concerning Mandragora are inevitably founded on extant taxa only. The Mediterranean region presents a very complex tectonic jig- saw puzzle, hard to resolve adequately (Hallam, 1981). It seems clear, however, that during most of the Tertiary, summer was the rainy season and only before the Pleistocene was there a shift towards the current pattern of rainfall in winter. The plants had to adapt and reduce activity during the dry summer months (Walter, 1977). The northern Mediterranean localities for Mandragora in northern Italy and on the Dalmatian coast are very patchy, resulting in a highly fragmented range there, and the precise status of these populations is uncertain. We have been unable to discover any modern collections from these regions (Cellinese et al., 1994) and some, or all, of the populations may no longer exist. Vierhapper (1915) maintained that these plants are Tertiary relics, the scattered remnants of a formerly larger, continuous range, that have been unable to adapt to the changing climate and still exhibit the vernal phenology type. Relict status based on Tertiary processes, however, is normally attributed to groups of far wider disjunct ranges than is the case here. Hawkes ( 1972a) based his definition of M. officinarum L. on these northern Italian and Dalmatian populations only, using corolla colour and phenology. On the evidence of our multivariate analyses, Hawkes's position is untenable, since the plants fall within the range of variation exhibited by the rest of the Mediterranean plants. It could be argued that the different corolla colour (white) of the northern Italian and Dalmatian plants, coupled with their dis- crete distribution is sufficient to recognize them at least at subspecific rank. We have already pointed out that the corolla colour is only an MANDRAGORA 29 apparent distinction, with similar colours occurring in populations outside northern Italy and Dalmatia. The somewhat isolated distri- bution alone is insufficient reason to recognize the Italian and Dalmatian plants as a separate taxon. The close relationship between the Mediterranean and the Turkmenian mandrake as suggested by the cladistic analysis is reflected in a linked history of the regions particularly during the Tertiary. It can be proposed that in this period, after the Tethys Sea receded and the middle Asian mountains became exposed during their simultaneous uplift, a vicariance event of the Mediterranean and Turkmenian ancestors took place. Both extant species show a comparable adaptation to the aridization and change from a summer to a winter rainfall regime. In other words, Mandragora turcomanica can be seen as an example of a plant that has been evolving in isolation in Turkmenistan since the Tertiary from ancient Tethyan stock (Kurbanov, 1994). It is of some interest to note that there is a highly congruent pattern within Atropa where the disjunct distribu- tion of the Turkmenian A tropa komarovii Blin. & Schal., consists of a very narrow range south of the river Sumbar only a few miles away from the Mandragora turcomanica population north of the river (Kurbanov, 1994). The Sino-Himalayan area is of particular interest because of its position at the transition between the Palearctic and the Oriental floristic realms. The period of the collision of the Indian sub- continent with Asia is contentious but probably occurred in the Paleogene approximately 50 million years ago (Scotese et al., 1988) with the penetration of the Eurasian border continuing during the early Neogene (c. 20 millions years ago) (Dercourt et al., 1986). The connection between Tethys and the Mediterranean, on the other hand, probably did not close until the mid-Miocene (c. 15 million years ago) (Hallam, 1994). Climate changes during the Paleogene were severe. Temperatures increased up to the early Eocene (c. 55 million years ago) followed by a global cooling which marks the passage from a warm domain of the Mesozoic and early Palaeogene to the cool domain thereafter (Hallam, 1994). According to Rud- diman et al. (1989) much of the uplift in the Xizang plateau region occurred only in the last 5 to 10 million years (Plio-Pleistocene uplift). It must be assumed that the glaciation of the high altitude regions of the Sino-Himalayan area during the Pleistocene (Ice- Ages) was more or less complete, posing the question whether refugia further south and at lower altitudes might have served as speciation centres. Schuster (1976) argued that the Indian plate served as a vehicle for the dissemination of Gonwanan taxa into Laurasia. But at the same time he expected that due to the migration through several climatic zones the Indian plate probably arrived with a rather depleted and altered flora and therefore only remnants of the original Gondwanan flora are recognizable in India today. If angiosperms were indeed on the Indian plate in early Cretaceous time, they could probably not be assigned to presently recognizable families. On the basis of a proposed late Triassic (c. 210 million years ago) start of the Indian plate migration, serious doubt is cast on the possibility that early Solanaceae could have been 'on board'. The main objection is that the plate drift simply occurred too early, before modern groups were in existence. However, other data (Smith et al., 1994) suggest that India did not break free from Gondwana until the late Jurassic to early Cretaceous (130-140 million years) or even near the Creta- ceous-Tertiary boundary only 60-65 million years ago (Hallam, 1994). Lu & Zhang (1986) proposed southwestern China as a later, secondary speciation centre of the whole tribe Hyoscyameae on the grounds of its high species-richness. A different biogeographical scenario proposes a migration from Gondwana to the Mediterranean basin. Migration routes that are closed today were wide open in the Mesozoic. Indeed, massive and relatively unimpeded migrations from Gondwana to Laurasia might have been feasible until the late Cretaceous (Schuster, 1976). Con- trary to the first scenario, the mandrake ancestors would therefore have spread eastwards from the Protomediterranean to the develop- ing eastern Asian mountain ranges where the massive uplift possibly resulted in a vicariance event. The Mediterranean populations on the other hand would have had to adapt to the increasing aridization in the western part of the continent during the late Tertiary (Weinert, 1972) and thus slowly changed their mode of development from a summer flowering to a autumn- winter-spring flowering rhythm. Proskuryakova & Belyanina (1985) claim that the slow succession of the development phases, particularly of the germination, the long growth and flowering period and the large size of the plant speak for the great age of the Mediterranean group. They draw the conclusion that the Mediterranean section must be older, representing a Terti- ary relic, than the apparently highly adapted Himalayan mandrake. The Himalayan mandrake, in fact, might have arisen from now- extinct ancestors as late as the Quaternary glaciations. To conclude, the massive timespan covered in the Tertiary, the many fluctuations in climate and transgressions of the seas and the current very inaccurate timing of even the major events render definitive statements or falsifications concerning the above scenar- ios and the history of Mandragora hardly possible. In this context the distinction between a cladogram (what is more closely related?) and the actual phylogenetic tree (what is the ancestor?) must be stressed. The cladogram (Fig. 3) suggesting a more recent common ancestry of Mandragora officinarum and M. turcomanica is con- gruent with only one of the many possible trees equally resulting in this particular cladogram, which therefore represents a far less informative (more general) mode of expression. Thus, it does not allow for statements concerning the ancestor and does not imply that Mandragora caulescens is the older taxon. Nevertheless, it can be proposed that the major events affecting speciation and present distribution of Mandragora in the Mediterranean-Turkmenian area were probably the receding of the Tethys and the aridization of the climate in the Tertiary. In the Sino-Himalayan area, on the other hand, the collision of the Indian plate and the subsequent uplift of the Himalayan range, as well as the later glaciation periods during the Quaternary, are probably most important. TAXONOMIC TREATMENT Mandragora L., Sp. pi. 1: 181 (1753). Type species: M. officinarum L. Perennial herbs. Stout tap-root, often branched, sometimes vaguely anthropomorphic. Stem very short or with secondary growth during flowering and fruiting, sometimes branched. Leaves simple, nor- mally entire, occasionally toothed. Inflorescence of typically solitary axillary, pedicellate flowers. Floral envelope actinomorphic. Calyx 5(6)-lobed, persistent, enlarged in fruit. Corolla 5(6)-lobed. Sta- mens 5(6), included, inserted in lower half of corolla-tube; filaments filiform, pubescent at base; anthers dorsifixed, dehiscing longitudi- nally. Ovary 2-locular; ovules numerous; style elongate; stigma capitate or somewhat bilobed. Fruit a fleshy, many seeded berry, globose or ovoid, borne beneath the leaves on nodding pedicel or lying on the ground. Seeds compressed; embryo curved. FLORAL FORMULA. K(5) C(5) A5 G(2) DIVERSITY AND DISTRIBUTION. Three disjunct species: Mediterra- nean region, central Asia, Sino-Himalayan region. 30 PHENOLOGY AND HABITAT. All three Mandragora species are geophytes with a conspicuous tap-root which is the perennating organ of these herbs, linking the annual vegetative periods. The Turkmenian mandrake with its prolonged dry season dormancy has been referred to as an ephemeroid (Proskuryakova & Belyanina, 1985), its rhythm of development more closely related to the Medi- terranean than to the Himalayan climatic cycle. Propagation is usually by seeds or vegetatively by underground buds on the tap- root. The species grow in open woodland, olive groves, grassland, fallow land, waysides, railway embankments, ruins, in rock crev- ices, grassy and stony mountain slopes and screes. The altitudinal range is 0-1200 and 3000-4900 m. ETHNOBOTANY. Once important medicinal plants with a long tra- dition in mythology and superstition. All parts of the plant contain potent narcotic alkaloids. ETYMOLOGY. Origin uncertain, possibly from the Greek |0,otv- Spcryopaa, original meaning controversial; alternatively from the Sanskrit mandros, meaning sleep and agora, meaning object or substance (Thompson, 1934). Key to the species of Mandragora 1 Young leaves arranged in a cluster on top of a stem with scales, later along the secondarily elongated, sometimes branched stem; corolla yellow to dark purple; style not exceeding stamens. Sino-Himalayan. Alt. 3000-4900 m 3. M. caulescens Leaves permanently arranged in a rosette, the stem not elongating; corolla white to pale blue or violet; style exceeding stamens 2 2 Mature lower leaves in a large rosette of 1 50- 1 80 cm; berries 40-60 mm in diameter. Turkmenian. Alt. 500-700 m 2. M. turcomanica Mature lower leaves in a rosette of less than 1 00 cm; berries less than 40 mm in diameter. Mediterranean. Alt. 0-1200 m ... 1. M. officinarum 1. Mandragora officinarum L., Sp. pi. 1: 181 (1753). Type: Herb. Burser IX: 26 (UPS-lectotype, designated by Knapp in Jarvis et al. (1993), microfiche-BM!). Figs 7, 8. = Atropa mandragora L., Syst. nal 10th ed.: 933 (1759), nom. illeg. (Art. 52.1). = Mandragora foemina Garsault, Fig. pi. med. 3: 221, t. 363 p.p. (1764). = Mandragora mas Garsault, Fig. pi. med. 3: 221, t. 363 p.p. (1764). = Mandragora acaulis Gaertn., Fruct. sem. pi. 2: 236, t. 131 (1791). = Atropa humilis Salisb., Prodr. stirp. Chap. Allerton: 132 (1796). = Atropa acaulis Stokes, Bot. mat. med. 1: 396 (1812). = Mandragora autumnalis Bertol., Elench. plant, viv.: 6 (1820). = Mandragora vernalis Bertol., Virid. Bonon. veg.: 6 (1824). = Mandragora praecox Sweet, Brit.fl. gard. 2: t. 198 (1827). = Mandragora neglecta G. Don ex Loudon, Hon. brit.: 71 (1830). = Mandragora microcarpa Bertol., Comm. Mandrag.: 12, t. 3 (1835). = Mandragora haussknechtii Heldr. in Mitt. Geogr. Ges. Jena. 4: 77, app. (1886). = Mandragora x hybrida Hausskn. & Heldr. in Mitt. Geogr. Ges. Jena. 4:77(1886). = Mandragora hispanica Vierh. in Osterr. Bot. Z. 65: 132-133 (1915). ILLUSTRATIONS. Wettstein (1895: fig. 12M-O); Feinbrun-Dothan S. UNGRICHT, S. KNAPP AND J.R. PRESS (1978: fig. 278); Pignatti (1982: figs on p. 518); Hoare & Knapp ( 1997: fig. 5c). Plant virtually stemless. Root stout, often branched, very long. Leaves arranged in a rosette, leaf shape and size very variable, but usually elliptic to obovate, max. 45 cm long, length/width ratio 1 .5- 10, variously pubescent. Flowers borne in leaf axils. Pedicels very variable in length, max. 15 cm. Calyx 6-28 mm long, lobed '/2-2/3 of the way to the base, lobes 3-15 mm, acuminate, only slightly accrescent in fruit. Corolla 12-65 mm long, campanulate, greenish white to pale blue or violet, lobed l/2 to nearly to the base, the lobes 6-60 mm. Stamens adnate to lower half of corolla; filaments 7-15 mm long; anthers 2.5^.0 mm long, yellow to brown, sometimes pale blue. Style 8-20 mm, exceeding stamens; stigma capitate. Fruit 5^0 mm in diameter, globose to ellipsoid, glossy yellow to orange when ripe, juicy, edible. Seeds 2.5-6.0 mm long, reniform, the surface reticulate, yellow to light brown. HABITAT. Open woodland, olive groves, fallow land, waysides, railway embankments, ruins, crevices. Altitudinal range 0-1200 m. VERNACULAR NAME. Mediterranean mandrake. PHENOLOGY. Flowering September to April. Fruiting November to June. CONSERVATION STATUS. Populations are scattered throughout the species range. Northern Italian populations are vulnerable (Cellinese et al., 1994) as they are apparently known only from old herbarium records and the present occurrence and abundance of these populations are unclear. DISTRIBUTION. Circum-Mediterranean. (Tunisia, Algeria, Mo- rocco, Portugal, Spain, Italy, former Yugoslavia, Greece, Cyprus, Turkey, Syria, Lebanon, Israel, Jordan). SELECTED SPECIMENS EXAMINED. TUISIA. Cap Bon, roadsides and orchards, 0-150 m, 6 October 1969, Tomkinson, M.J. 72, 72A (BM). Nabel, 60 m, 14 October 1907, Gandoger, M. 84 (K). MOROCCO. Guercif, 144 km E. of Fes near 'La Kazbah', 38 km before the town of Guercif, 34°16'N, 3°45'W, 510 m, disturbed gravelly area by road, 3 November 1994, Jury, S.L. et al. 15434 (RNG); Gharb, c. 4 km NNE of Souk-El-Arbaa-du-Rharb on road to Ksar-El-Kebir, by bridge and salt works, 34°43'N, 6°00'W, 30 m, pasture between fields at side of salt pans, 7 November 1994, Jury, S.L. et al. 15701 (RNG); Zerhoun, c. 3 km NW of Moulay Idriss and 33 km from Meknes along road to Sidi Kacem and Tanger, 34°04'N, 5°33'W, 390 m, ruins of Roman city of Volubilis, 1 November 1994, Jury, S.L et al. 15398 (RNG); Prerif Central, N. of Fes, on the road to Ouezzane, 34°22'N, 5°09'W, 210 m, roadside ruderal community, 24 October 1993, Jury, S.L et al. 12634 (RNG); Haul Ouerrha, c. 50 km N. of Fes on the road between Fes-el-Bali and Ourtzarh, 34°33'N, 5°02'W, 120 m, in olive plantation on steep N-facing slope with Zizyphus lotus, 24 October 1993, Jury, S.L et al. 12653 (RNG); Temara, 10 km S. of Rabat, 0-30 m, sandy roadside, 14-20 October 1981, Lewalle, J. 10054 (BM, RNG); 5 km W. of Oued, Beht on Rabat-Meknes road, 33°52'N, 5°58'W, 400 m, 27 September 1991, Gardner, M.F. et al. 4885 (E, RNG); Loukkos, SW of Chefchaouen, on road to Ksar-el-Kebir, 35°1'N, 5°45'W, 120 m, in cultivated fields of wheat, 21 October 1993, Jury, S.L. et al. 12542 (RNG); Loukkos, c. 1 km E. of Larache on road to Asilah, c. 200 m from Oued Loukkos, 35°12'N, 6°08'W, 5 m, at side of road by saltpans, 28 October 1994, Jury, S.L et al. 15237 (RNG), Tanga, just N. of airport, road to Cap Spartel, 35°45'N, 5°55'W, 35 m, fallow area at side of ploughed field with Urginea maritima, 16 October 1993, Jury, S.L. et al. 12217 (RNG), Tanga, E. on road to Sebta (Ceuta), Oued Dliane, 35°50'N, 5°39'W, 5 m, nitrophilous areas on fallow cultivated land on heavy clay alluvial plain, 17 October 1993, Jury, S.L. et al. 12238 (RNG); Allal-Tazi, close to the river Sedou, 1 November 1965, Talon, A. 1630 (BM). SPAIN. Baleares, Palma, between S. Sardina and S. Bernardo, 22 Septem- ber 1913, Bianor & Sabasien 1877 (BM, E, P, RNG); Province Cadiz, MANDRAGORA Fig. 7 Mandragora officinarum. a) Flowering, cultivated at RBG Kew, b) Fruiting, cultivated at RBG Kew. 32 S. UNGRICHT, S. KNAPP AND J.R. PRESS MANDRAGORA 33 Chiclana, on clayey soil by the river, 10 January 1 855, Bourgeau, E. 2304 (E, K, P);Torredonjimeno, 600 m, 10 December 1986, Espinosa, A. 870252 (K); alrededores de Cordoba, Cuesta del Espino, 2 December 1967, Galiano, E.F. & Valdes, B. 820 (E, RNG); Province Malaga, C 341 from Campillos to Ronda, 36°52'N, 5°00'W, 525 m, disturbed ground near new road cutting, 27 October 1990, Gardner, M.F. & Knees, S.G. 4769 (RNG); Province Cadiz, 23 km from Jerez de la Frontera, 36°32'N, 5°59'W, 1 30 m, ruderal area by road with abundant Silybum marianum below ruined castle of El Pedroso, 22 December 1993, Jury, S.L. & Jury, L.C. 13111 (BM, RNG); Province Ja6n, Torredonjimeno, between Porcuna andTorredonjimeno, near Pilar de Moya, 500 m, roadside on chalky soil, 3 November 1979, Ladero, M. et al. 10605 (RNG); Malaga, near Churiana, 3 December 1957, Partridge, F. 2 (BM); Campamento, 9 November 1911, Wolley-Dod, A.M. 22 (BM). ITALY. Sicily, Province Palermo, Madonie, near Polizzi, 950 m, rocky limestone slopes, 1 8 August 1964, Davis, P.M. 40134 (E); Sicily, Trapani, 200 m, steep slopes, 25 August 1964, Davis, P.M. 40182 (RNG); Sicily, Province Siracusa, Augusta, October-November 1911, Vaccari, A. 2326 (BM, K). FORMER YUGOSLAVIA. Hercegovinae, Doljnja Glavska, 6 May 1908, Reiser, C. s.n. (K); Silieski, near Ragusa, 500 m, 29 April 1909, Maly, K. s.n. (K). GREECE. Near Corinth, 30 m, waste grounds and waysides, April 1931, Atchley, S.C. 819 (K); Crete, Frangokastello, 29 March 1973, Barclay, C. 3004 (K); Crete, Canea, 1 December 1943, Bickerich, G. 15324 (BM, K); Crete, Kolokithia, December 1977, Bowen, H.J.M. 694 (RNG); Crete, Agios Nikolaos, December 1983, Bowen, H.J.M. 3088 (RNG); Peloponnese, 5 km from Pilos on the road from Kalamata, 250 m, in cultivated fields near stream, 12 November 1973, Brickell, C. & Mathew, B. 8088 (K); Crete, Canea, frequent in hills and ditches throughout Crete up to 300 m, 1 January 1940, Davis, P.H. 1100(E,K);Cyclades,Amorgos, lOApril 1940, Davis, P.M. 1438 (K); Dodecanese, Kos, Asfendion, 300-600 m, rocky slopes, 27 March 1965, Davis, P.H. 40478 (K); Crete, near Candia, Hagios-Deka, December 1913, Duffour, C. 1501 (P); Cyclades, Kea, 30 April 1961, Gathorne-Hardy, E. 14 (E); Dodecanese, Tilos, near chapel of Ayios Nicolaos c. 180 m above Livadia, 26 April 1962, Gathorne-Hardy, E. 139 (E); Crete, near Sitia, 150- 200 m, on clayey soil, 30 October 1966, Greuter, W. 7821 (E); Near Corinth, Examillia, 8 February 1931, Guiol, F. 1731 (BM); Crete, Canea, on waste- land, 10 March 1883, Reverchon, E. 117 (BM, E, K, P, PAL); Dodecanese, Kos, by track near Asclepeion, 23 March 1985, Townsend, C.C. 85/11 (K); Dodecanese, Rhodes, 9 February 1914, Vaccari, A. 1141 (K); Crete, Nomos Lasithiou, Monastiraki, roadside, 5 April 1980, Walree, A.M. 22226 (BM). CYPRUS. Kyrenia, fallow olive grove, 13 February 1949, Casey, B.C. 221 (K). TURKEY. Province Mugla, Kemer-Kestep, 50 m, edge of fields, 29 March 1956, Davis, P.H. & Polunin, O. 25469 (BM, E, K); Side, 22 February 1966, Baytop, A. et al. 8504 (E). SYRIA. Baniyas, 300 m, foot of hill, beneath tree, 11 March 1945, Norris, F.H. s.n. (BM). ISRAEL. Jerusalem, fields, 800 m, 20 February 1908, Meyers, F. 4377 (E); Mount Carmel, upper Nahal Nesher, 2 km S. of Nesher, 32°45'N, 35°31'E, 280 m, terra rossa on hard limestone, 6 April 1989, Danin, A. & Knees, S.G. 1709 (RNG); Mount Gerizim, near Nablus, fields, 700 m, 22 December 1910, Meyers, F.S. & Dinsmore, J.E. 6377 (E); District Haifa, near Waldheim (Alloney Aba), open places in Quercus ithaburensis forest, 28 March 1942, Davis, P.H. 4166 (E, K). The confusion surrounding specific epithets for the taxon we are here calling Mandragora officinarum is incredible. This morass has arisen for two basic reasons, firstly orthographic errors, and sec- ondly, the confusion over the phenological status of type specimens. Orthographic errors made by early authors persist through floristic works, adding and compounding any ambiguities that may have arisen. A name often used in synonymy (or as a valid name) of M. officinarum is Mandragora officinalis Mill., said to be validly published in 1768 (see Jackson & Berry, 1979: 508, footnote). However, in the eighth edition of The gardener's dictionary (Miller, 1768), where Miller used Linnaean epithets for the first time, Miller actually used the epithet officinarum ('MANDRAGORA (officinarum)'), notofficinalis. In the ninth edition (Miller & Martyn, 1807), the taxon was classified as Atropa mandragora, following Systema naturae (Linneaus, 1759) and the second edition of Species plantarum (Linnaeus, 1762). Mandragora officinalis was cited in synonymy, with reference made to the illustrations produced to compliment the earlier editions (Miller, 1760). These illustrations bear no specific epithets, thus the 1807 reference to the epithet officinalis is almost certainly an orthographic error for the 1768 use of Linneaus's epithet officinarum, which was not cited in synonymy in the ninth edition (Miller & Martyn, 1807). Thus the name Mandragora officinalis Mill, has persisted in the literature, but has no real nomenclatural standing. Similarly, Bertoloni (1820) attributed the epithet officinalis (see below) to Willdenow, explicitly citing 'W. En.\ Willdenow (1809), in his Enumeratio plantarum, attributed the epithet officinalis to DeCandolle (in Lamarck & DeCandolle, 1805), where it is again attributed, in error, to Miller's Gardener's dictionary. Thus it is apparent that the numerous occurences of the epithet officinalis, attributed to many authors, are traceable to a single orthographic error from Miller's 1767 edition of the Gardener's dictionary. Even if the epithet officinalis had nomenclatural standing, it would be invalid under Article 52.1 (see above), as it is homotypic with Mandragora officinarum L.: in DeCandolle (in Lamarck & DeCan- dolle, 1805) and Willdenow (1809), Atropa mandragora L. is cited in synonymy. Assumptions as to the phenological status of type specimens has also led to confusion about the correct application of epithets (e.g. Jackson & Berry, 1979). Bertoloni (1820, 1824, 1835) contributed greatly to the confusion surrounding names for the Mediterranean mandrake. In a list of plants cultivated in the Botanic Gardens at Bologna (Bertoloni, 1820), he included two species of mandrakes: Mandragora autumnalis, which he coined as a new name, and M. officinalis, which he attributed to Willdenow. In a later work (Berto- loni, 1824) he designated two different taxa: M. vernalis and M. officinarum, explicitly citing Linnaeus as the source of the latter epithet. He put M. officinalis in synonymy with M. vernalis and put M. autumnalis in synonymy with M. officinarum. In Commentarius de Mandragoris (Bertoloni, 1835), he revised the genus and recog- nized three species: M. vernalis, M. officinarum andM. microcarpa. In synonymy he left out both of the previously used epithets, autumnalis and officinalis, which he may have considered to be superfluous. The lectotype of Mandragora officinarum, selected from amongst the elements used by Linneaus in his description, bears no pheno- logical information at all. The specimen in Burser's herbarium is labelled 'Misnia, Lusatia', indicating it came from Germany. It is probable therefore that it is from a botanical garden, and since plants of Mandragora in northern European gardens usually bloom in spring, it has been assumed that this one did as well. This is not founded in fact, thus rendering much of the debate as to which name is the vernal or autumnal taxon irrelevant. 2. Mandragora turcomanica Mizg. in Trudy Turkmen. Fil. Akad. Nauk SSSR.Ashkabad2: 165 (1942). Type: Turkmenistan, west- ern Kopet Dag, Kara Kala region, southern foot of Mt Syunt, Schevlan valley, on scree slopes, 26 November 1938, O.G. Mizgireva, M. Nastacalicz & G. Nastacalicz s.n. (ASH-holotype; LE-isotype(?)). ILLUSTRATIONS. Mizgireva (1942: figs 1, 2; 1955: figs 2-7, 9, 14, 15, 18, 19, 24); Belyanina (1982: figs 1, 2). Plant virtually stemless. Leaves arranged in a rosette: lower leaves c. 90 x 60 cm, length/width ratio c. 1 .3-1 .5, broadly elliptical or ovate, 34 S. UNGRICHT, S. KNAPP AND J.R. PRESS usually with large, irregular teeth in upper half, upper leaves smaller, oblong-ovate or broadly lanceolate, usually without large teeth, on both sides sparsely hairy, the hairs mostly arranged along veins. Flowers borne l(-3) in leaf axils, occasionally the pedicels up to 2- 6-flowered. Pedicels 2-3 cm long, 7-18 cm in fruit. Calyx 15-20 mm long, lobes 10-15 mm, acuminate, accrescent in fruit. Corolla 20-25 mm long, campanulate, violet or purple, with three narrow white stripes at base, the lobes c. 15 mm. Stamens c. 10 mm long, attached to lower half of corolla; filaments 6-7 mm, densely pubes- cent at the base and in the lower Vr, anthers c. 4 mm long, pale blue. Style exceeding stamens; stigma capitate, slightly bilobed, green. Fruit 40-60 mm in diameter, glossy yellow to orange when ripe, pulpy, juicy, smelling of melons, edible. Seeds 4-5 x 6-7 mm, reniform, yellow to light brown. HABITAT. Shrubland in ravines and on stony mountain slopes, especially among Paliurus spina-christi Mill. (Rhamnaceae). Altitudinal range 500-700 m. VERNACULAR NAME. Turkmenian mandrake. PHENOLOGY. Flowering October to March. Fruiting until June. Summer dormancy until autumn rains begin. Xerothermic ephemeroid. CONSERVATION STATUS. Only small populations in a restricted area within the Syunt-Khassardagh Reserve, and in the valleys of Chokhadgh, Shevlan, Altybai, Ekechinar, Sarymsakli and Dagdanli are known (Kurbanov, 1994). According to Mizgireva (1978) the population in the Syunt-Khassardagh reserve consists of fewer than a thousand plants. DISTRIBUTION. Southwestern Kopet Dag. (Turkmenistan, ?Iran). SELECTED SPECIMENS EXAMINED. No specimens were available for examination: the data here are drawn exclusively from the literature. In the original article in which this species was described (Mizgireva, 1942) the type is cited as being in Ashkabad (ASH). Subsequently, in the Flora of the USSR (Lincevskij, 1955), the type was said to be in Leningrad (now St. Petersburg, LE) and this citation has been followed in other floristic works (Schonbeck-Temesy, 1972). Since we have been unable to obtain specimens from either herbarium, it is uncertain whether or not the holotype remains in ASH or has been transfered to LE, thus we have cited its location as in the original publication. Mizgireva's (1955) detailed study of this species documented the flowering times (Mizgireva, 1955: fig. 16) of individual plants and clearly shows that some individuals flower in autumn (October and November) while others flower in spring (February and early March). This is the same situation as that in Mandragora officinarum, and is markedly different from that found in much of the rest of the flora of Kopet Dag (Mizgireva, 1978). Mizgireva (1955) stated that the differences between her species, M. turcomanica, and the Mediter- ranean mandrakes (separated by her into M. officinarum and M. autumnalis) were largely size of plant, colour of anthers and relative sizes of calyx and corolla. It is clear from her work that these characteristics are consistent throughout the range of wild and greenhouse grown material she examined, but since she cites no specimens, nor have we been able to examine any of the known specimens of M. turcomanica, an in-depth analysis of these charac- ters relative to the range of variation found in M. officinarum awaits future study. 3. Mandragora caulescens C.B. Clarke, Fl. Brit. Ind. 4: 242 (1883). Type: India, Sikkim, Lachen, Hooker s.n. (Kl-lectotype, designated here by R. Mill). Fig. 9. = Anisodus mariae Pascher in Feddes Repert. Spec. Nov. 7: 227 (1909). = Mairella yunnanensis H. Lev., Cat. pi. Yun-Nan: 199 (1916). = Anisodus caulescens (C.B. Clarke) Diels in Feddes Repert. Spec. Nov.Beih. 12:480(1922). = Mandragora tibetica Grubov in Rast. Tsentral. Azii. 5: 108-109 (1970). = Mandragora caulescens subsp. brevicalyx Grierson & D.G. Long in Notes Roy. Bot. Card. Edinburgh 36: 144 (1978). = Mandragora caulescens subsp. flavida Grierson & D.G. Long in Notes Roy. Bot. Card. Edinburgh 36: 143 (1978). = Mandragora caulescent subsp. purpurascens Grierson & D.G. Long in Notes Roy. Bot. Card. Edinburgh 36: 143 (1978). = Mandragora chinghaiensis Kuang & A.M. Lu, Fl. reip. pop. Sin. 67: 159 (1978). ILLUSTRATIONS. Grubov (1970: pi. IV, fig. 2; pi. V, fig. 2); Kuang & Lu (1978: fig. 36, 1-6). Hoare & Knapp (1997: fig. 5d). Root stout, often branched. Stem present, max. 60 cm long. Leaves arranged in a cluster on top of the young stem above scale leaves, later along the secondary growth of the stem, max. 30 cm long, length/width ratio 1.9-5.5, oblanceolate or spathulate, glabrous to densely pubescent beneath, especially along the veins. Flowers borne in leaf axils, often nodding, pleasantly scented. Pedicels 5-10 cm long. Calyx 5-30 mm long, often nearly as long as corolla, lobed c. l/4 of the way to the base, accrescent in fruit. Corolla 5-30 mm long, campanulate to cup-shaped, yellow to purple, the venation usually darker, lobed about 1A of the way to the base, the lobes 2-10 mm. Stamens adnate in lower half of corolla; filaments 3-10 mm; anthers 1-5 mm long, white to yellow. Style 3-6 mm long, not exceeding the stamens; stigma capitate, somewhat bilobed. Ovary 2.5-5 mm in diameter, globose. Fruit 10-25 mm in diameter, globose, pale greenish white or greenish yellow. Seeds 1.5-3.0 mm long, numerous, reniform, the surface pitted, yellow to light brown. HABITAT. Grassy and stony slopes, open moorland, pastures and screes. Subalpine and alpine zone, especially among Rhododendron shrubs. Altitudinal range 3000-4900 m. VERNACULAR NAMES. Himalayan mandrake. China: qie shen. PHENOLOGY. Flowering April to September. Fruiting August to October. CONSERVATION STATUS. Locally common. Not threatened. DISTRIBUTION. Sino-Himalayan. (Nepal, India, Bhutan, Myanmar, China). SELECTED SPECIMENS EXAMINED. NEPAL. Topke Goia, 27°39'N, 87°35'E, 3810 m, in short grassland, 3 July 1971, Beer, LW. 8264 (BM); Tarakot, 28°53'N, 82°59'E, 3900 m, 25 May 1969, Dobremez, J.F. 175 (BM); Jangla Banyang, wet place, 3800 m, 5 June 1973, Einarsson, L. et al. 430 (BM); Ganesh Himalaya, Shior Khola, 4270 m, bouldery alpine meadow, Gardner, P.C. 479 (BM); Topke Gola-Tasagon, 3600-4350 m, 17 August 1977, Ohashi, H. et al. 775165 (BM); Langtang Valley, 28°14'N, 85°32'E, 4420 m, among rocks in turf, June 1949, Polunin, O. 63 1 (BM); 5 miles E. of Timure, oak wood, 41 10 m, 3 July 1949, Polunin, O. 780 (BM); pass NW of Jumla, 6 miles Pansae Dara, 3050 m, growing in loose soil recently free of snow, 30 April 1952, Polunin, O. et al. 878 (BM); Dozam Schola, near Simikot, 3510 m, grass slopes, 2 June 1952, Polunin, O. et al. 4250 (BM); near Jumla, 3810 m, open grassy slopes, 14 July 1952, Polunin, O. et al. MANDRAGORA 35 X S \U*iin I I 36 S. UNGRICHT, S. KNAPP AND J.R. PRESS 4668 (BM); Arun-Tamur watershed, Topke Gola, 3960 m, 11 May 1956, Stainton, J.D.A. 254 (BM); Balangra Pass, W. of Tibrikot, 29°06'N, 82°41'E, 3810 m, 28 May 1963, Stainton, J.D.A. 4263 (BM); Rolwaling, 27°55'N, 86°23'E, 3810 m, beneath small shrubs, 25 June 1964, Stainton, J.D.A. 4676 (BM);Khunde-Khumjung,3820m,6Junel952,Z//n/w^r/nann,A. 688 (BM). INDIA. Sikkim: Changu, 3960 m, in crevices of rocks, 28 June 1913, Cooper, R.E. 80 (E); Gnatong, 3960 m, peaty marsh, 31 August 1913, Cooper, R.E. 759 (E); Jongri, 4000-4200 m, 22 May 1960, Hara, H. et al. 443 (BM, K); Tungu, 3660-3960 m, 23 July 1849, Hooker, J.D. s.n. (K); Jongri, 4420 m, campsite towards Yakhut amongst Juniperus scrubs, 29 May 1990, Kirkpatrick, G. 53 (E); District Darjeeling, Phalut, 3350-3660 m, 30 May 1902, Lace, J.H. 2266 (E); West District, Bikbari, Choktsering Chhu valley; 27°30'53"N, 88°08'28"E, 3950 m, amongst scrub, 12 July 1992, Long, D.G. et al. 266 (E); Chulong, 4570 m, 12 September 1912, Rhomoo, L 210 (E); Gamothang, 3960 m, 12 August 1913, Rhomoo, L. 1059 (E); Giagong, 3960 m, 6 September 191 1, Ribu & Rhomoo 5492 (BM, E, K); Phalut, 3350-3660 m, 27 May 1911, Smith, W.W. 4593 (E); Jongri, 4000 m, in shelter of dwarf Rhododendron and Juniperus, 23 June 1983, Starling, B.N. et al. 92 (K); Tangshing campsite, 4000 m, open grazed areas among Rhododendron and around campsite, 24 June 1983, Starling, B.N. et al. 106 (K); summit of Sandakphu, 3660 m, small dense tufts amongst grass, 18 May 1881, Watt, G. 5373 (E, K). Arunachal Pradesh (Assam): Orka La, Bhutan frontier, 3960 m, on stony slopes, scattered, 7 June 1938, Kingdon Ward, E 13840 (BM). BHUTAN. Paco Chu, 4110m, loose sandy crevice, 15 May 1966, Bowes Lyon, S. 3 156 (BM); Phajudin Timpu, 3960 m, 5 August 1914, Cooper, R.E. 3232 (BM, E); Tare-La area, 3660 m, 19 May 1938, Gould, B.J. 109 (K); Laum Thang, 3900 m, in alpine herbage with moderate humidity, 18 May 1967, Hara, H. et al. 1 1973 (BM); Western Bhutan, 3510 m, among boulders above tree line, 17 June 1933, Ludlow, F. & Sheriff, G. 107 (BM); Byasu La, 4110m, growing among stones on open hillside, 18 May 1937, Ludlow, F. & Sherriff, G. 3073 (BM); Cheli La, 3810 m in open moorland, 4 May 1949, Ludlow, F. et al. 16139 (BM); Pangotang, 3960 m, open hillside above fir zone, 27 May 1949, Ludlow, Eel al. 18986(BM); Kantanang,Tsampa,4110 m, among shrubs on steep hillside, 3 June 1949, Ludlow, Eetal. 19040(BM); Shingbe, Me La, 3350 m, growing amongst other herbs and small shrubs, 16 May 1949, Ludlow, F. et al. 20645 (BM, E). MYANMAR (Burma). Chenochi Pass, 3600 m, 3 July 1920, Farrer, J.R. 1694 (E). CHINA. Yunnan: Diqing Prefecture, E. slope of Bai Ma Shan, 28°19'N, 99°05'E, 3895 m, among stones in Abies forest, 9 June 1993, Alden, B. et al. 1021 (E); Diqing Prefecture, Zhongdian County, above Napa Hai, N. of Zhongdian, 27°55'N, 99°34'E, 4000 m, forested ridge, snow-bed vegetation and dwarf Rhododendron scrub, 12 June 1993, Alden, B. et al. 1185 (E); western flank of the Lichiang range, 27°20'N, 3050-3350 m, open alpine pasture, June 1910, Forrest, G. 5999 (E, K); Lichiang range, 27°25'N, 3660-3960 m, July 1913, Forrest, G. 10474 (BM, E, K); western slopes of Likiang snow range, Yangtze watershed, 30 May-6 June 1922, Rock, J.F. 4211 (BM); Mo-ting shan, ME of Atuntze, eastern slopes of Yangtze- Mekong divide, 3660 m, along trail, June 1932, Rock, J.F. 22771 (E); slopes of Mt Gyi-na lo-ko, the second peak of the Yu-lung shan, 3660 m, in alpine meadows, April-May 1932, Rock, J.F. 25005 (BM, E); Chungtien, Chiren, 2200 m, grass slope, 17 May 1937, Yu, T.T. 11327 (KUN); Mekong- Salween divide, Chingpoh, 3800 m, alpine pasture lands, 14 June 1938, Yu, T.T. 19078 (E); Upper Kiukiang valley, Clulung Lungtsahmuru, 3700 m, mountain slope, open and grassy place, 9 August 1938, Yu, T.T. 19843 (E, KUN). Sichuan: Tien-chuan-hsien, 3500 m, 14 June 1936, Chu, K.L. 2791 (BM); mountains E. of Yungning, 27°48'N, 101° E, 3660 m, open stony pasture, June 1922, Forrest, G. 21407 (BM, E, K); mountains around Muli, 28°12'N, 101° E, 3660 m, July 1930, Forrest, G. 28416 (BM, E); Liuku- liangdse, 27°48'N, 3700-4200 m, 18 May 1914, Handel-Mazetti, H. von 2349 (K); Minya Konka snow range, S. of Tatsienlu, 4450 m, in meadows, July 1929, Rock, J.F. 17530 (E); Mu-li-ka-bu, 3600 m, mountain slope, grassy place, 18 May 1937, Yu, T.T. 5496 (KUN). Qinghai: Dari (Darlag) Xian, just N. of Manzhang, along the Manzhang He, between Dari and Banma, 33°17'51"N, 100°25'55"E, 4000 m, on moist rodent disturbed slope, 12 August 1993, Ho, T.N. et al. 1 172 (E); Chindu Xian, Xiwu Xiang, E. of pass on road between Xiwu and the Szechwan border, 33°09'14"N, 97°32'35"E, 4250 m. damp slope with extensive pika warrens, 1 1 Septem- ber 1996, Ho, T.N. et al. 3063 (BM). Xizang: Dotha, 3960 m, 20 June 1945, Bar & Kirat Ram 20511 (K); Mt. Demula, Yiwei County, 4100 m, 23 August 1973, Chinese Tibet Expedition 1973 1214 (KUN); District Chengtang, Dingjie County, Ladang to Xingeng, 3800-4000 m, 9 June 1975, Chinese Tibet Expedition 1975 5679 (KUN); Londre Pass, Tsarong, Mekong-Salween divide, 28°14'N, 98°40'E, 3960 m, July 1921, Forrest, G. 19630 (UC/JEPS, P, K); Tha Chu valley, 3660 m, in clumps, alpine region, in stony pastures and on grassy slopes, 11 July 1950, Kingdon Ward, F. 19613 (BM); N. of Sanga Choling, 28°41'N, 93°02'E, 4110 m, on open grassy bank, 14 May 1936, Ludlow, E & Sherriff, G. 1590 (BM); hills S. of Lhasa, 4420 m, ground left bare by melting snow, 6 June 1942, Ludlow, F. & Sherriff, G. 8676 (BM); hills S. of Lhasa, 4720 m, herb growing under boulders, 21 August 1942, Ludlow, F. & Sherriff, G. 9014 (BM); Pome, above Showa Dzong, 29°55'N, 95°25'E, 3350 m, open hillsides midst grass, 1 1 June 1947, Ludlow, F. et al. 13149 (BM, E); Sobhe La, near Tongyuk Dzong, Pome, 30°07'N, 94°54'E, 3510 m, 21 May 1947, Ludlow, F. et al. 13750 (BM, E); Province Kongbo, Nyima La, 29°38'N, 94°52'E, 3350 m, on dry ground under trees, 21 May 1947, Ludlow, F. et al. 15051 (BM, E); Lo La, Pachakshiri, 28°49'N, 93°59'E, 3810 m, in openings of Rhododen- dron forest, mostly above Abies zone, 15 May 1938, Ludlow, F. et al. 3776 (BM); Province Kongbo, south side of Lusha Chu, 29°20'N, 94°35'E, 3810 m, in Lonicera and Berberis scrub, 13 June 1938, Ludlow, F. et al. 4591 A (BM, E); valley above Tse, Tsangpo valley, 29°23'N, 94°22'E, 3350 m, in grassy banks under Rhododendron, Ludlow, F. et al. 4591 (BM). The status and identity of the previously proposed subspecies of Mandragora caulescens (Grierson & Long, 1978) are discussed in the section on the species delimitation (p. 25). It is clear from our studies that M. caulescens is extremely variable and that a detailed field-based study over the entire species range is necessary. ACKNOWLEDGEMENTS. This study was undertaken as partial fulfilment of the requirements for an M.Sc. in Advanced Methods in Taxonomy and Biodiversity at Imperial College, London and The Natural History Museum, London undertaken by S. Ungricht and supervised by S. Knapp and J.R. Press. The work has profited from discussions with many members of the staff at The Natural History Museum, particularly S. Cafferty, N.C. Garwood, CJ. Humphries, C.E. Jarvis, C. Moncrieff, N.K.B. Robson, K. Sidwell, A.R. Vickery, D.M.Williams, and C.Whitefoord. We are grateful to Z. Frankiel, K. Rottcher and Zhan Ren-bin for their translation of the crucial Russian and Chinese papers respectively. Furthermore, we are indebted to M. Gilbert and R.R. Mill for their sharing of personal observations and to N. Turland for a thoughtful and detailed review. Thanks are also due to R. Baldini in Florence and the directors and staff of the following herbaria who made their collec- tions available: E, JEPS, K, KUN, P, PAL, RNG and UC. This study has been supported by a grant from the Park Fund of the Botany Department of The Natural History Museum, London. REFERENCES \ iimi-iil Franco, J. do 1984. Nova flora de Portugal 2. Lisbon. Anderson, E. 1941. The technique and use of mass collections in plant taxonomy. Annals of the Missouri Botanical Garden 28: 287-292. & Ttirrill, W.B. 1935. Biometrical studies on herbarium material. Nature 136: 986. Baehni, C. 1946. L'ouverture du bouton chez les fleurs de Solanees. Candollea 10: 400-492. Bateman, R.M. 1989. 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Al-Eisawi, D. & Al-Jallad, W. 2221, M. officinarum (BM) Alden, B., Alexander, J.C.M., Long, D.G., McBeath, R.J.D., Noltie, H.J. & Watson, M.E 1021, M. caulescens (E); 1185, M. caulescens (E) Atchley, S.C. s.n., M. officinarum (BM); 315, M. officinarum (K); 819, M. officinarum (K) Atherton, G.E. 793, M. officinarum (K); 812, M. officinarum (K); 998 M. officinarum (K) Baker, E.W. 149, M. officinarum (BM) Ball, J. s.n., M. officinarum (E) Barclay, C. 3004, M. officinarum (K) Bauer, K. & Spitzenberger, F. 53, M. officinarum (BM) Baytop, A., Tanker, N. & Sezik, E. 8504, M. officinarum (E) Beer, L.W. 8264, M. caulescens (BM) Bianor & Sabasien 1877 , M. officinarum (BM, E, P, RNG) Bickerich, G. 15324, M. officinarum (BM, K) Biro, L. s.n., M. officinarum (K) Bar & Kirat Ram 20481, M. caulescens (K); 2051 1, M. caulescens (K) Bornmiiller, J. s.n., M. officinarum (K) Bourgeau, E. s.n., M. officinarum (K, P), 2304, M. officinarum (E, K, P) Bowen, H.J.M. 694, M. officinarum (RNG); 3088, M. officinarum (RNG) Bowes Lyon, S. 3156, M. caulescens (BM); 3239, M. caulescens (BM) Brickell, C. & Mathew, B. 8088, M. officinarum (K) Carr, J.W. s.n., M. officinarum (RNG) Casey, E. C. 22 1 , M. officinarum (K) Chaboisseau, C. 575, M. officinarum (BM) Chatterjee, D. 75, M. caulescens (BM) Chinese Tibet Expedition 1973 1214, M. caulescens (KUN) Chinese Tibet Expedition 1975 5679, M. caulescens (KUN) Chu, K.L. 2791, M. caulescens (BM) Clarke, C.B. 34977, M. caulescens (K); 34983 A, M. caulescens (BM); 35704, M. caulescens (K) Clarke, E.D. s.n., M. officinarum (BM) Coincy, M.A. de s.n., M. officinarum (P) Cooper, R.E. 80, M. caulescens (E); 759, M. caulescens (E); 3232, M. caulescens (BM, E) Danin, A. & Knees, S.G. 1709, M. officinarum (RNG) Davis, P.H. 1 100, M. officinarum (E, K); 1438, M. officinarum (K); 2108, M. officinarum (E, K); 4166, M. officinarum (E, K); 40134, M. officinarum (E); 40182, M. officinarum (RNG); 40478, M. officinarum (K) Davis, P.H. & Polunin, O. 25469, M. officinarum (BM, E, K) Delavay, J.M. s.n., M. caulescens (P); 263, M. caulescens (P); 1026, M. caulescens (P) Dhwoj, L 513, M. caulescens (BM, E) Dinsmore, J.E. B 377, M. officinarum (E) Dobremez, J.F. 175, M. caulescens (BM) Duff our, C. 1501, M. officinarum (P) Economides, S. 1055, M. officinarum (K) Einarsson, L, Skarby, L. & Wetterhall, B. 430, M. caulescens (BM) Espinosa, A. 870252, M. officinarum (K) Ex Herb. Postian. apud Colleg. Syriens. Protest. 952, M. officinarum (BM) Farrer, J.R. 1694, M. caulescens (E) Fay, J.M. 815, M. officinarum (K); 864, M. officinarum (K) Forbes s.n., M. officinarum (UC/JEPS) Forrest, G. 569, M. caulescens (E); 5999, M. caulescens (E, K); 10474, M. caulescens (BM, E, K); 19630, M. caulescens (UC/JEPS, P, K); 21407, M. caulescens (BM, E, K); 26951, M. caulescens (P); 28416, M. caulescens (BM, E) Galiano, E.F. & Valdes, B. 820, M. officinarum (E, RNG) Gammie, G.A. s.n., M. caulescens (BM, E) Gandoger, M. s.n., M. officinarum (K); 84, M. officinarum (K); 8400, M. officinarum (K) Gardner, M.E, Jury, S.L. & Rejdali, M. 4885, M. officinarum (E, RNG) Gardner, M.F. & Knees, S.G. 4769, M. officinarum (RNG) Gardner, PC. 479, M. caulescens (BM) Garnett, C.S. 33/5, M. officinarum (BM); 158/7, M. officinarum (BM) Gathorne-Hardy, E. 14, M. officinarum (E); 139, M. officinarum (E); 194, M. officinarum (E) Girgenti s.n., M. officinarum (BM) Gould, B.J. 109, M. caulescens (K) Greuter, W. 7821, M. officinarum (E) Guiol, F. 1731, M. officinarum (BM) Haig, E. s.n., M. officinarum (BM) Handel-Mazetti, H. von 2349, M. caulescens (K) MANDRAGORA 39 Hara, H., Kanai, H., Murata, G. Ohashi, H., Tanaka, O. & Yamazaki, T. 1 1973, M. caulescent (BM) Hara, H., Kanai, H., Murata, G., Togashi, M. & Tuyama, T. 443, M. caulescens (BM, K) Hardy, E. s.n., M. officinarum (BM) Hart, H.C. s.n., M. officinarum (BM) Hay, J.H. 1707, M. officinarum (K); H. 2484, M. officinarum (K) Helbaek, H. 74, M. officinarum (K) Heldreich, T. von s.n., M. officinarum (BM, E, K, P); 257 [1860], M. officinarum (BM); 257 [1857], M. officinarum (P); 257 [1859], M. officinarum (K); 2817 [1853], M. officinarum (P); 2817 [1860], M. officinarum (K, P) Higgins, E.K. s.n., M. officinarum (BM) Ho, T.N., Bartholomew, B. & Gilbert, M. 1 172, M. caulescens (E) Ho, T.N., Bartholomew, B., Watson, M. & Gilbert, M. 3063, M. caulescens (BM) Hooker, J.D. s.n., M. caulescens (E, K-lectotype, P) Huguenin s.n., M. officinarum (BM) /wry, S.L., Achhal, A. & Mouks, H. & Upson, T.M. 12634, M. officinarum (RNG); 12653, M. officinarum (RNG) Jury, S.L., Aitlafkih, M., Hedderson, T. & Kahouadji, A. 15237, M. officinarum (RNG) Jury, S.L., Aitlafkih, M., Hedderson, T. & Rutherford, R. W. 15398, M. officinarum (RNG); 15434, M. officinarum (RNG); 15701, M. officinarum (RNG) Jury, S.L. & Jury, L.C. 13111, M. officinarum (BM, RNG) Jury, S.L, Rejdali, M., Taleb, A. & Upson, T.M. 12542, M. officinarum (RNG) Jury, S.L, Taleb, A. & Upson, T.M. 12217, M. officinarum (RNG); 12238, M. officinarum (RNG) Jussieu, A. de s.n., M. officinarum (P) King, G. s.n., M. caulescens (E); 4368, M. caulescens (P) Kingdon Ward, F. 13840, M. caulescens (BM); 19613, M. caulescens (BM) Kirkpatrick, G. 53, M. caulescens (E) Lacaita, C.C. s.n., M. officinarum (BM); s.n., M. caulescens (BM) Lace, J.H. 2266, M. caulescens (E) Ladero, M., Lopez Guadalupe, M., Molero, J. & Perez Raya, F. 10605, M. officinarum (RNG) Lange, J. s.n., M. officinarum (K, P) Laukkonen, P. 55, M. officinarum (K) Le Testu, G. s.n., M. officinarum (BM) Lewalle, J. 10054, M. officinarum (BM, RNG) Lojacono-Pojero, M. s.n., M. officinarum (BM) Long, D.G., McBeath, R.J.D., Noltie, H.J. & Watson, M.F. 266, M. caulescens (E) Lowne, B.T. s.n., M. officinarum (BM) Ludlow, F.& Sheriff, G. 107, M. caulescens (BM); 1590, M. caulescens (BM); 3073, M. caulescens (BM); 8676, M. caulescens (BM); 9014, M. caulescens (BM) LHd/ovv, E, S/zernjft G. cfe E///O?, //.//. 13149, M. caulescens (BM, E); 13750, M. caulescens (BM, E); 15051, M. caulescens (BM, E) LwoVow, E, Sherriff, G. & Hicks, J.H. 16139, M. caulescens (BM); 18986, M. caulescens (BM); 19040, M. caulescens, (BM); 20645, M. caulescens (BM, E) Ludlow, F., Sherriff, G. & Taylor, G. 3776, M. caulescens (BM); 4591 A, M. caulescens (BM, E); 4591, M. caulescens (BM) Maty, K s.n., M. officinarum (K) Manberg, T. s.n., A/, caulescens (K) Mavzomoustakis, G.A. 8, M. officinarum (BM) Me Laren '5 collectors AC 67, M. caulescens (E, P) Meyers, E 4377, M. officinarum (E) Meyers, F.S. & Dinsmore, J.E. 6377, M. officinarum (E); B 6377, M. officinarum (K) M;7/, y.S. s.n., M. officinarum (K) Moreau, W.M. 62, M. officinarum (K) Norris, F.H. s.n., M. officinarum (BM) , H., ATanai, H., 0/z6a, //. c£ rafm/z/, K 775165, M. caulescens (BM) Olivier & Bruguiere s.n., M. officinarum (P) Orphanides, T.G. 75, M. officinarum (BM, E, K, P) Pa/ne, /A. s.n., M. officinarum (K) Pantling, R. s.n., M. caulescens (K); 46387, M. caulescens (K) Partridge, F. 2, M. officinarum (BM) Pasta/; 7. <6 Va/des, B. s.n., M. officinarum (RNG) Perraudiere, H. de la s.n., M. officinarum (P) Pette;; E s.n., M. officinarum (BM) Pichler s.n., M. officinarum (BM, E, K) Pignant s.n., M. officinarum (K) P/fard, C.-y. 1757, M. officinarum (K) Polunin, 0. 631, M. caulescens (BM); 780, M. caulescens (BM); 6616, M. officinarum (K) Po/Mrtw, O., Sytes, W/?. Psilotaceae "" Pteridaceae 60 Selaginellaceae 61 sr\ Sinopteridaceae DZ f\) Thelypteridaceae Vittariaceae "- Doubtful records References 64 Systematic index SYNOPSIS. A catalogue of the pteridophytes of Sao Tome and Principe is presented, with 156 taxa, 12 of which are endemic to the islands. Recent synonymy for the area is included. All collections examined are cited. - ' --- The pteridophytes of the neighbouring island of Bioko (Fernando INTRODUCTION _ _ _ Po) have been recently revised by Benl (1978, 1980, 1982, 1988, 1 99 1 ) who provided descriptions and keys to the taxa. The lack of an The most recent account of the pteridophytes of Sao Tome and updated checklist for Sao Tome and Principe and the difficulty in Principe (Gulf of Guinea) was produced by Alston in 1944. Since compiling the dispersed information on this group of plants have then several nomenclatural changes have taken place and new precluded its inclusion in recent floristic and biodiversity studies records have been added to the flora of the islands (e.g. Alston 1958, (Figueiredo, 1994). The incorporation of several recent collections from Sao Tome © The Natural History Museum, 1998 42 E. FIGUEIREDO and Principe at LISC, has revealed further new records and new areas of distribution for some taxa, and provided the basis for a revision of Alston's (1944) account. DIVERSITY There are 153 indigenous taxa of pteridophytes in Sao Tome and Principe occurring from sea level (e.g. Acrostichum aureum L., found in swamps and mangroves) to the highest peaks of the islands (e.g. Grammitis spp.), and belonging to 28 families and 60 genera. Three taxa were probably introduced, and became naturalized, Pityrogramma calomelanos (L.) Link (Alston, 1944), Adiantum raddianum C. Presl from the Neotropics (Alston, 1956a), and Pteris vittata L. (Alston 1958, 1959). One hundred and thirty-seven taxa occur in Sao Tome and 75 occur in Principe; 12 taxa are endemic to the islands. In Bioko, the pteridophyte richness is higher than in either Sao Tome or Principe, with 208 taxa in 31 families (Benl, 1978). However, only four taxa (1.9%) are endemic to Bioko, while eight taxa are endemic to Sao Tome (5.8%) and three are endemic to Principe (4%). This difference in the percentage of endemics is less striking than the difference found for the angiosperms. In that group, Bioko has 3.6% endemic taxa, Sao Tome has 13.4%, and Principe has 8.2% (Figueiredo, 1994). Asplenium is the genus of pteridophytes with the largest number of taxa in Sao Tome and Principe. Twenty-four taxa are represented, of which three are endemic. CONSERVATION Twenty-five taxa collected in Sao Tome and Principe in the nine- teenth century have not been collected since, even though extensive collecting took place in the 1950s and also in the 1990s. Although some of these taxa, represented in the herbarium by a single collec- tion, may be doubtful records, others were collected more than once by different collectors. The absence of these taxa amongst the collections made this century may indicate their rarity or extinction. For example, Microlepia speluncae (L.) T. Moore was collected in Sao Tome by Don (s.l.), Quintas (Ilheu das Rolas), and Welwitsch (Monte Cafe). Although Monte Cafe is a well collected area, visited by most botanists, this species has not been found during the last 100 years. Elsewhere in Africa, M. speluncae is rare and sporadic, occurring at low to medium altitudes. Sixteen taxa are represented in herbaria by a single collection, either from Sao Tome or from Principe (10 from Sao Tome and 6 from Principe), which may indicate their rarity or even extinction. When Principe alone is considered, 27 of the total of 75 taxa have been collected only once. In this case, mistaken recordings or undercollecting are also a possibility. Several species are repre- sented in herbaria by a single collection made by Barter, who visited Principe when stationed at Bioko, and therefore it is possible that there were some errors when the specimens were labelled. Principe remains poorly collected and recent collections of pteridophytes from that island were not available to confirm the presence of these taxa. Seven taxa are apparently restricted to the Pico of Sao Tome [Asplenium friesiorum C. Chr., Ctenitis cirrhosa (Schum.) Ching, Grammitis molleri (Baker) Schelpe, G. tomensis Schelpe, Huperzia ophioglossoides (Lam.) Rothm., Lycopodium clavatum L., and Xiphopteris oosora (Baker) Alston] and five taxa are confined to the Pico of Principe [Grammitis nigrocincta Alston, Hymenophyllum polyanthos var. mildbraedii (Brause ex Brause & Hieron.) Schelpe, Lomariopsis warneckei (Hieron.) Alston, Selaginella squarrosa Baker, and Triplophyllum jenseniae (C. Chr.) Holttum]. Most of these are high altitude pteridophytes which are also distributed on other African mountains. Two of these taxa, Grammitis molleri and G. tomensis, are very rare and endemic to the Pico of Sao Tome. Lycopodium clavatum is cosmopolitan but, in the tropics, it is confined to mountains. Of the twelve taxa endemic to Sao Tome and Principe, two are apparently very rare, having been collected only once: Selaginella monodii Alston and Sphaerostephanos elatus subsp. thomensis Holttum. COLLECTIONS Historical collections Exell (1944) gives a lively account of the early collectors in Sao Tome and Principe. The majority of the earliest collections of pteridophytes from Sao Tome and Principe were made by Don (in 1822), Curror (in 1839), Welwitsch (in 1853), Barter (in 1858), and Mann (in 1861). These collections are deposited at K (Curror, Barter, and Mann), BM (Don and Welwitsch), and LISU (Welwitsch). In the late nineteenth century extensive collecting was made by the Portuguese collectors Moller, Quintas, and Newton. The origi- nal set of these collections is kept at COI, but duplicates were sent to several European herbaria. The duplicates were distributed as Flora Africanae Exsiccata (F.A.E.}, a series in which different collections of the same taxon were grouped and distributed with the same number. For this reason, most F.A.E. labels refer to several localities and different collectors After examining the original sets of the Moller, Quintas, and Newton collections, Exell (1944: 383-385) tried to link originals with duplicates. However, it seems that, for the pteridophytes, in most cases the original collections were scarsely labelled, lacking the information needed to link them to the duplicates distributed. Many F.A.E. duplicates sent from COI to K had a second label referring to a single collector. The numbering on these collector labels is unique to K specimens, as the original sets at COI are mostly unnumbered. Although the second label usually refers to a single collector and locality there is no certainty as to the correspondence between the specimen and the data. The information consisting of more than one locality, found on some F.A.E. and original set labels, is worthless and misleading because it is not possible to relate it to the specimens it is attached to. This information should be omitted when those collections are cited. Together with the F.A.E. collections, the collections made by Chevalier in 1905 and Exell in 1932-33, the first sets of which are deposited, respectively, at P and BM, comprise the bulk of the material examined to produce Alston's account of the pteridophytes in Exell's Catalogue (1944). Recent collections Only a few collections were made during the twenty years that followed Exell's expedition in 1932-33. The majority of recent collections were made during the years 1954-1973, and in the 1990s. PTERIDOPHYTES OF SAO TOME AND PRINCIPE 43 The first set of the collections made by Rozeira in Sao Tome and Principe in 1954 is deposited at PO. However, several collections of pteridophytes cited in the literature (Alston, 1958) could not be located in that herbarium. A set of duplicates of Rozeira's collection is apparently deposited at BM (Exell & Rozeira, 1958) but could not be found in 1997. Some duplicates (with the name of the collector missing) exist at K. Based on Rozeira's collections, new records for the islands were published by Alston (1958) in the proceedings of the 'Conferencia Internacional dos Africanistas Ocidentais', which took place in Sao Tome in 1956. During 1956 several botanists collected in Sao Tome and Principe. As part of a Calypso expedition which took place in 1956-57, Rose made several collections in the two islands, in- cluding living material, which were deposited at P. In the same year, Monod and Thorold made important collections in Sao Tome and Principe, which are deposited at BM. These collections were studied by Alston (1958, 1959) and resulted in several new records being added to the flora of the islands. Lains e Silva also collected in Sao Tome in 1956. Although his collections have been cited in the literature, several numbers could not be found either at LISJC or LISU. In 1968-1973, Espirito Santo collected in both islands. The original sets of his collections are kept at LISJC and LISC, with some duplicates at BM and COI. Recent collections of pteridophytes from Sao Tome examined during this study were mainly made by Pinto Basto in 1990, Matos and Matos & Van Essche in 1994, and Lejoly in 1994-1995, and are deposited at LISC. Recent collections were also made by Paiva (COI), but these were not available for examination at the time of this study. CATALOGUE The catalogue includes all the taxa of pteridophytes known to occur in Sao Tome and Principe, arranged in alphabetical order, with synonyms and references to the literature for the area published since 1944. Synonyms already cited in Alston's (1944) work are omitted. All the collections examined are cited in this catalogue. Collec- tions cited in the literature are also included. Some of these specimens were not located (indicated in the text with 'not found'), and others were not available for study. Some of the collections cited but not seen by Alston ( 1 944), probably at B, were not examined during this study; these are marked with a question mark. It is possible that a future study of the unseen material may reveal misidentifications by previous authors. For each taxon, the collections are cited by alphabetical order of collectors, but collections from the same locality are grouped. Collections without locality are placed at the end of the specimens cited. These include F.A.E. and Moller collections with more than one locality on the labels. F.A.E. duplicates at K are cited under their F.A.E. number and reference is made to other labels and numbers if they exist. Most of the localities provided by the collectors consist of the name of a plantation ('ro9a') or a river. For this reason, distribution maps based on these collections would be very inaccurate. Maps with 'ro9as' and other collecting localities mentioned in the text are provided, to allow for an estimate of the distribution of the taxa (Figs 1 and 2). Data on habit, habitat, and ecology are lacking for the vast majority of the collections examined. Acrostichaceae Acrostichum L. Acrostichum aureum L., Sp. pi. 2: 1069 (1753); Alston in Exell, Cat. Vase. PL S. Tome: 85 (1944); Benl inActa Bot. Barcinon. 38: 22(1988). SAO TOME. Porto Alegre, Chevalier 14194bis (P!), 14200 (P!), salt marshes, Espirito Santo 5131 (LISC!, LISJC!); Ilheu das Rolas, F.A.E. (Quintas)69 (BM!, LISU!), Quintass.n. (COI!); PortoAlegre, Praia Jal<£, Rio Malanza, swamp, Matos 7663 (LISC!); Perseveran?a, Rose 538 (P!). PRINCIPE. Santo Antonio, salt marshes, Espirito Santo 5170 (LISC!, LISJC!), mangrove, Monod 12191 (BM!), mangrove, Rose 425 (P!); Rio Papagaio, swampy border, Quintas 1 (BM!, COI!); Praia General Fonseca, Rozeira 430 (PO!); s.l., Welwitsch 15 (BM!, LISU!). DISTRIBUTION. Pantropical. Adiantaceae Adiantum L. Adiantum mettenii Kuhn, Filic. afr.: 65 (1868) nomen; in Jahrb. Konigl. Bot. Gart. Berl 1: 338 (1881). Adiantum soboliferum sensu auct. non Wall, ex Hook., Sp.fil. 2: 13, t. 74A (1858); Alston in Exell, Cat. Vase. PL S. Tome: 82 (1944). SAO TOME. Monte Forte, 50 m, Quintas s.n. (COI!, K!); s.l., Don s.n. (BM!). DISTRIBUTION. Tropical Africa and Madagascar, extending to In- dia and Philippines. NOTE. This taxon has not been recorded in the islands since the nineteenth century. Adiantum philippense L., Sp. pi. 2: 1094 (1753); Alston in Exell, Cat. Vase. PL S. Tome: 82 (1944); Alston in Exell, Suppl. Cat. Vase. PL S. Tome: 1 (1956); Benl in Acta Bot. Barcinon. 32: 26 (1980). SAO TOME. Casal Pires, waterfall, 200 m, Espirito Santo 1 10 (BM!, COI!, LISJC!); Boa Entrada, Chevalier 14518 (P); Rio do Ouro, 1 10 m, Quintas 1249 (BM!, COI!, LISJC!); Agua Coimbra, 100 m, Quintas s.n. (COI!), Moller & Quintas s.n. (LISU!); Diogo Vaz, Rozeira 518 (PO!); Santa Catarina, between main house and Lemba, Rozeira 840 (PO!). DISTRIBUTION. Palaeotropical. Adiantum raddianum C. Presl, Tent, pterid.: 158 (1836); Alston in Exell, Suppl. Cat. Vase. PL S. Tome: 1 (1956); Alston in Exell & Rozeira in Conf. Int. Afr. Oc. 3: 79 (1958). SAO TOME. Quinta da Graca, 600 m, Espirito Santo 124 (BM!, COI!, LISJC!); Pouso Alto, 490 m, Espirito Santo 4278 (LISC!, LISJC!); Monte Cafe, Sao Carlos, secondary forest, Espirito Santo 5048 (LISC!, LISJC!); Sao Nicolau, waterfall, on wet rocks, Espirito Santo 5178 (LISC!, LISJC!); Saudade, 900 m, Exell 399 (BM!, COI!); Ponta Figo, on the way to Morro Vilela, Lains e Silva 357 (LISU!); between Pico and Monte Castro, 1200- 1800 m, mountain rainforest, Lejoly 95/13 (LISC!); Monte Cafe, 1010 m, Monod 1 1694 (BM!, COI!), Oliveira & Noronha s.n. (LISU!); NW of Pico, 1000-1200 m, Monod 12263 (BM!, COI!); Agua Ize\ common, Pinto Basto 199 (LISC!); Tras-os-Montes, Calvario, Rozeira 582 (PO!). PRINCIPE. Esperansa, near Ribeira Camarao, Rozeira 354 (PO!). DISTRIBUTION. Cultivated American species, escaped and natural- ized. 44 E. FIGUEIREDO PTERIDOPHYTES OF SAO TOME AND PRINCIPE Adiantum vogelii Mett. ex Keyserl. in Mem. Acad. Imp. Sci. Saint Petersbourg ser. 7, 22(2): 8, 3 1 ( 1 875);Alston in Exell, Cat. Vase. PI. S. Tome: 83 (1944);Tardieu-Blot inAubrev., Fl. Cameroun 3: 147 (1964); Benl in Acta Bot. Barcinon. 32: 26 (1980). PRINCIPE. Santo Antonio, Monod 12072 (BM!, COI!); Berimbau, Newton 18 (BM!, COI!, K!), Newton s.n. (K!); Cimalo, Quintas 36 (BM!, COI!); Esperan?a, between Montalegre and Maquina, Rozeira 378 (PO!); s.l., Souza s.n. (COI!). DISTRIBUTION. W. tropical Africa to Angola, Zanzibar. Aspidiaceae (=Dryopteridaceae) Ctenitis (C. Chr.) C. Chr. Ctenitis cirrhosa (Schum.) Ching in Sunyatsenia 5: 250 (1940); Benl in Acta Bot. Barcinon. 40: 41 (1991). Dryopteris crinobulbon (Hook.) C. Chr., Index filic., Suppl. 3: 84 (1934); Alston in Exell, Cat. Vase. PL S. Tome: 61 (1944). SAO TOME. Pico, 1850 m, Mann s.n. (B, K!-type of Dryopteris crinobulbon). DISTRIBUTION. Tropical and SE Africa, Madagascar, Comoros and Mascarene Is. NOTE. This taxon has not been recorded in the islands since the nineteenth century. Ctenitis lanuginosa (Willd. ex Kaulf.) Copel., Gen. fit.: 124 (1947); Benl in Acta Bot. Barcinon. 40: 41 (1991). Dryopteris lanuginosa (Willd. ex Kaulf.) C. Chr., Index filic.: 273 (1905); Alston in Exell, Cat. Vase. PL S. Tome: 62 (1944). SAO TOME. S.I., Mann s.n. (K-not found). DISTRIBUTION. Tropical and subtropical Africa, Madagascar, Comoros, Seychelles and Mascarene Is. 45 NOTE. This taxon has not been recorded in the islands since the nineteenth century. Didymochlaena Desv. Didymochlaena truncatula (Sw.) J. Sm. in J. Bot. (Hooker) 4: 196 (1841); Alston in Exell, Cat. Vase. PL S. Tome: 66 (1944); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 254 (1964); Benl in Acta Bot. Barcinon. 40: 35 (1991). SAO TOME. Rio Contador, 1270-1350 m, F.A.E. (Moller) 36 (BM!, LISU!);Trds-os-Montes, Gama s.n. (CO\!);Mollers.n. (COI!); Macambrani, 1270 m, Moller s.n. (COI!); Sela do Camelo, near Lemba, 200 m, Rozeira 3450 (PO!); Monte Cafe, 760-850 m, Welwitsch 61 (BM!, K!, LISU!). DISTRIBUTION. Pantropical. Dryopteris Adans. Dryopteris pentheri (Krasser) C. Chr., Index filic.: 284 (1905); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 447 (1959); Benl in Acta Bot. Barcinon. 40: 36 (1991). Dryopteris oligantha sensu auct. non (Desv.) C. Chr., Index filic. Suppl. 3: 93 (1934); Alston in Exell, Cat. Vase. PL S. Tome: 62 (1944). SAO TOME. Pico, Chevalier 13651 (P), 14290 (P), Moller s.n. (COI-not found, P!); between Monte Cafe and Pico, Chevalier 14580 (P); Macambrara, Ere// 131 (BM), 1 35 (BM);Tras-os-Montes, Gama s.n. (COI-not found), 800 m, Quintas 1286 (BM!, COI-not found), Rozeira 554 (K!, PO!); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain rainforest, 1200- 1800 m, Lejoly 95/9 (LISC!); between Monte Quinas and Quinas Alias, Monod 12007 (BM!); Monte Cafe, 1010 m, Monod 1 1696 (BM!); Lagoa Amelia, 1300 m, Quintas 1291 (BM!, COI-not found), Rose 199 (P!); Sao Nicolau, near waterfall, Rozeira 182 (K!, PO!); Santa Catarina, Lemba, Rozeira 856 (K!); s.l., 1850 m, Mann (K!), F.A.E. (Moller) 39 (BM!, LISU!), Moller s.n. (COI-not found), Quintas 10 (K!). DISTRIBUTION. Tropical and southern Africa. Fig. 1 Sao Tome 1 . Agua Cardoso-2B 2. Agua Coimbra-3A 3. Agua Grande-3B-4B-4A 4. Agua Ize (Ro?a)-4C 5. Agua Pinhao-3B 6. Almas-4B 7. Antonio Soares-4B 8. Bacelar-2D 9. Batepa-4B 10. Bemposta (Ro9a)-3B ll.Benfica(Roca)-3B 12. Binda(Ro?a)-lC 13. Blu-blu-4B 14. Boa Entrada (Roca)-4A 15. Bom Sucesso (Ro?a)-3B 16. Bombaim (Roca)-3C 17. Cabumbe-2C 18. Caixao Grande-4B 19. Calvario-2B 20. Cantagalo (Ro?a)-4C 21.CaoPequeno-2D 22. Cascata-2B 23. Chami90-3B 24. Colonia Acoreana (Roca)-4C 25. Cruzeiro (Roca)-3C 26. Diana (Ro?a)-4B 27. Diogo Nunes-4A 28. Diogo Vaz (Roca)-2B 29. Dona Amelia (Roca)-2B 30. Dona Augusta (Roca)-3D 3 1 . Dona Eugenia (Roca)-3D 32. Ermelinda (Roca)-2D 33. Estacao Sousa-2B 34. Famosa-4B 35. Formoso-3C 36. Guaiaquil (Ro?a)-3C 37. Guarda (Ro?a)-3A 38. Ilheu das Rolas-2E-2F 39. Java (Roca)-3B 40. Jou (Roca)-2D 41 . Lagoa Amelia-SB 42. Lemba (Roca)- IB 43. Macambrara (Roca)-3B 44. Maria Fernandes-3C 45. Milagrosa (Roca)-4B 46. Monte Cafe (Roca)-3B 47. Monte Castro-2B 48. Monte Forte (Ro9a)-2A 49. Monte Mario (Roc.a)-2E 50. Monte Quinas-2B 5 1 . Monte Rosa (Roca)-2D 52. Monte Verde-2D 53. Morro Cantagalo-3C 54. Morro Vigia -3B 55. Morro Vilela-2B 56. Nova Ceilao (Ro9a)-3B 57. Nova Moca (Ro9a)-3B 58. Novo Brasil (Ro9a)-2E 59. Novo Destine (Ro9a)-3B 60. Paga Fogo (Ro9a)-2B 61. Pedroma (Ro9a)-4B 62. Perseveran9a (Ro9a)-3D 63. Pico-2B 64. Pinheira (Ro9a)-4B 65. Ponta Figo (Ro9a)-2A 66. Porto Alegre (Ro9a)-2E 68. Praia Jale-2E 69. Praia Melao (Ro9a)-5B 70. Praia Pantufo-5B 71.Quija-2D 72. Quinas Altas -2B 73. Quinta da Gra9a (Roca)-3B 74. RibeiraMo9a-lB-2B 75. Ribeira Peixe-3D 76. Rio Abade-3C-4C 77. Rio Abade (bridge) -4C 78. Rio Campos-2C 79. Rio Caue-2D 80. Rio Contador-2A-2B 8 1 . Rio do Ouro (Ro9a)-3 A 82. Rio Malanza-2E 89. Rodia (Ro9a)-3B 90. Santa Catarina (Ro9a)-lB 9 I.Santa Irene (Ro9a)-2C 92. Santa Margarida (Ro9a)-3B 93. Santelmo (Ro9a)-3C 94. Sao Carlos (Ro9a)-3B 95. Sao Joao dos Angolares-3D 96. Sao Miguel (Ro9a)-lD 97. Sao Nicolau (Ro9a)-3B 98. Sao Pedro-3B 99. Sao Tome -4A 100. Sao Vicente-4B 101.Saudade(Ro9a)-3B 102. Sela do Camelo-2C 103. Tras-os-Montes (Ro9a)-3B 83. Rio Manuel Jorge-3B-4B-5B 104. Trindade-4B 84. Rio Manuel Jorge (waterfall )-3B 85. Rio Miranda Guedes-3C- 3D 86. Rio Paga Fogo-lB-2B 87. Rio Xufexufe-2D 88. Ro9a Nova (Ro9a)-3B 105. Vale Carmo (Ro9a)-3D 106. Vieira Machado (Roca)-3C 107. Vila Aida (Ro9a)-3D 108. Vila Conceicao (Roca)-2E 109. Vila Jose (Ro9a)-3D 110. Zagaia-2C 111. Zampalma (Ro9a)-3B. 67. Pouso Alto (Ro9a)-3B The following collecting localities could not be located: Agua Telha (Diogo Vaz), Agua Telha (near Ribeira Mo9a), Casa do Pico (1935 m alt.), Casal Pires Correia, Monte Figo, Magodinho, Morro Pempem (Tras-os-Montes), Quingombo, Santa Maria, Rio Santo Antonio. 46 E. FIGUEIREDO Fig. 2 Principe 1 . Barriga Branca- 1 C 2. Bela Vista (Ro?a)- IB 3. Berimbau-2B 4. Cajamanga- 1 B 5. Cimalo-2B 6. Dois Irmaos-2C 7. Esperanca (Ro9a)-lB 8. Fundao-lB 9. General Fonseca-2C 10. Infante D. Henrique (Rosa)- 1C H.Lapa(Ro9a)-lB 12. Mamas- 1C 13. Maquina-lB 14. Mencorne-lC 15. Montalegre (Roca)-lB 16. Morro do Leste-lB 17. Neves Ferreira (Rosa)- 1C 18. Oque Caspar (Rosa)- IB 19. 6queNazare-lB 20. Oque Pipi-lB 21.Pico-lC 22. Pico Papagaio-lB 23. Praia da Lapa-lB 24. Praia das Burras-2A 25. Precipfcio-2A 26. RibeiraBibi-lC 27. Ribeira Cambungo-lC-2C 28. Ribeira Chimboto-lC 29. Ribeira Fria (Ro?a)-2B 30. Ribeira Jambere-lB-2B 31.RioPapagaio-lB-2B 32. Santo Antonio-2B 33. Sao Joao (Rosa)-2B 34. Sundi(Ro?a)-lA 35.TerreiroVelho-2B. The following collecting localities could not be located: Porto Real, Ribeira Camarao and Rio Bambu-Porco. Lastreopsis Ching Lastreopsis aff . currori (Mett. ex Kuhn) Tindale in Victoria Natu- ralist 73: 184 (1957); Tindale in Contr. New South Wales Natl. Herb. 3: 338(1965). Aspidium nigritianum Mett. ex Kuhn, Filic. afr.: 138 (1868). Dryopteris nigritiana (Mett. ex Kuhn) Kuntze, Revis. gen. pi. 2: 813 (1891); Alston in Exell, Cat. Vase. PL S. Tome: 62 (1944). Ctenitis nigritiana (Mett. ex Kuhn) Alston in Bol. Soc. Brot., ser. 2, 30: 12(1956). PRINCIPE. Rio Bambu-Porco, 60 m, Navel 138 (LISU!), 141 B (not found); Pico, Rozeira 2607 (PO!); s.l., Barter 1906 (K-type of Aspidium nigritianum, not found). Tectaria Cav. Tectaria angelicifolia (Schum.) Copel. in Philipp. J. Sci., ser. C, Bot. 2: 410 (1907); Alston in Exell, Cat. Vase. PI. S. Tome: 66 (1944); Alston in Exell, Suppl. Cat. Vase. PL S. Tome: 1 (1956); Benl in Acta Bot. Barcinon. 40: 55 (1991). SAO TOME. Sao Vicente, 200 m, Espirito Santo 42 (BM!, COI!, LISJC!); Bombaim, Monod 11886 (BM!); between Sao Tome and Monte Cafe, Welwitsch 62 (BM!, LISU!); s.l., F.A.E. (Moller & Quintas) 43 p.p. (BM!, COI!, K also Moller 32!, LISU!), 550 m, Welwitsch s.n. (K!). PRINCIPE. S.l., Welwitsch 12b (BM!, LISU!). DISTRIBUTION. W. and Central tropical Africa Tectaria camerooniana (Hook.) Alston in / Bot. 77: 288 (1939); Alston in Exell, Cat. Vase. PL S. Tome: 67 (1944);Tardieu-Blot in Aubrev., Fl. Cameroun 3: 29 1 ( 1 964); Benl in Acta Bot. Barcinon. 40:56(1991). SAO TOME. Macambrara to Zampalma, 900 m, ground fern, Exell 293 (BM!), 294 (BM!, COI!); between Bom Sucesso and LagoaAmelia, primary forest, 1350-1420 m,Matos & Van Esschel296 (LISC!); above Monte Cafe, 1010 m, Monod 1 1693 (BM!); between Sao Pedro and LagoaAmelia, 1050 PTERIDOPHYTES OF SAO TOME AND PRINCIPE m, Monod 11711 (BM!); Calvario, 1200 m, Monod 11815 (BM!), 11817 (BM!); Lagoa Amelia, Rose 232 (P!); s.l., F.A.E. (Moller) 44 (BM!, COI!, LISJC!, LISU!, P!). DISTRIBUTION. W. and Central tropical Africa. Tectaria fernandensis (Baker) C. Chr., Index filic., Suppl. 3: 179 (1934);Alston in Exell, Cat. Vase. PL S. Tome: 67 (1944); Benl in Acta Bot. Bardnon. 40: 57 (1991). SAO TOME. Boa Entrada, Chevalier 14360 (P!), 14517 (P!); SW region, Chevalier 14591 (P!), 14617 (P!); Sao Miguel, 50 m, Espirito Santo 4742 (LISJC!); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain rainforest, 1200-1800 m, Lejoly 95/10 (LISC!); Sao Miguel, Monte Verde, secondary forest, 10-1 50 m,Lejoly95/%4 (LISC!); Pedroma,Mocquerys 1 257 126(P!);SaoJoaodosAngolares,80m,A'eH'fons.n.(COI!),50m,2M/n/a5S.n. (COI!); Agualze, common, Pinto Basto2\ 1 (LISC!); Perseveranca,/?o.se537 (P!); s.l., Don s.n. (BM!), F.A.E. (Moller & Quintas) 43 p.p. (LISJC!). PRINCIPE. Between Oque Pipi and Morro do Leste, secondary forest, 350 m, ground fern, Exell 564 (BM!, COI!), 565 (BM!, COI!); Pico Papagaio, 600 m, ground fern, Exell 684 (BM!); s.l., Barter 1907 (K!), Welwitsch 12 (BM!, Ki.LISU!). DISTRIBUTION. W. and Central tropical Africa. Triplophyllum Holttum Triplophyllum buchholzii (Kuhn) Holttum in Kew Bull. 41: 251 (1986). Tectaria buchholzii (Kuhn) Copel. in Philipp. J. Sci. 38: 138 (1929) 'buckholzii'; Alston in Exell, Cat. Vase. PL S. Tome: 67 (1944). Ctenitis buchholzii (Kuhn) Alston in Bol. Soc. Brot., ser. 2, 30: 1 1 (1956); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 448 (1959). PRINCIPE. Pico, 500-600 m, Monod 12119 (BM!), 800 m, Rozeira 886 (PO!), 91 1 (PO!), 2617 (PO!); Rio Bambu-Porco, 60 m, Navel 139B (LISU!); Infante D. Henrique, 250 m, Rozeira 2300 (PO !); Infante D. Henrique, old path to Ribeira Bibi, Rozeira 2391 (PO!); Mencorne, 200 m, Rozeira 241 1 (PO!). DISTRIBUTION. Ivory Coast to Zaire. Triplophyllum fraternum (Mett. ex Kuhn) Holttum in Kew Bull. 41:253(1986). Aspidium fraternum Mett. ex Kuhn, Filic. afr.: 132 (1868). Ctenitisfraterna(Meti.)Tardieu inNotul. Syst. (Paris) 14: 342(1953). var. elongatum (Hook.) Holttum in Kew Bull. 41: 254 (1986). Nephrodium subquinquefidum (P. Beauv.) Hook., Sp. fil. 4: 130 (1862) var. elongatum Hook., Sp.fil. 4: 130 (1862). PRINCIPE. S.l., Mam 1861 (KMectotype of Nephrodium subquinque- fidum var. elongatum). DISTRIBUTION. Endemic. NOTES. The following collections are unlocalized but may have been collected in Principe, since there are no further collections from the continent: Niger Exped., Barter s.n. (K!), s.l., Curror s.n. (K!). Based on the same type specimen, Pichi-Sermolli (1985) consid- ered this fern to be a new species (Ctenitis attenuata Pic. Serm.) and later (Pichi Sermolli, 1991) transferred it to Triplophyllum with the name T. attenuatum (Pic. Serm.) Pic. Serm. This taxon has not been recorded in the islands since the nine- teenth century. Triplophyllum jenseniae (C. Chr.) Holttum in Kew. Bull. 41: 253 (1986). 47 PRINCIPE. Pico, 850 m, Rozeira 1021 (PO!). DISTRIBUTION. W. and Central tropical Africa. Triplophyllum principis Holttum in Kew Bull. 41: 246 (1986). Dryopteris variabilis (Hook.)Alston in Exell, Cat. Vase. PL S. Tome: 64 (1944) var. barteri (Hook.) Alston in Exell, Cat. Vase. PL S. Tome: 64(1944). PRINCIPE. Oque Pipi, secondary forest, 300 m, ground fem, Exell 546 (BM!, COI!); between Oque Pipi and Morro do Leste, secondary forest, 350 m, ground fern, Exell 562 (BM!, COI!); above Infante D. Henrique, second- ary forest, 300 m, ground fern, Exell 618 (BM!, COI!), 624 (BM!); Rio Bambu-Porco, 60 m, Navel 14 la (LISU!); Oque Nazare, 500 m, Newton s.n. (COI!); Infante D. Henrique, Rose 402 (P!), Rozeira 2370 (PO!); Neves Ferreira, Rozeira 2204A (PO!), 2205 (PO!); between Barriga Branca and Mamas, Rozeira 2552 (PO!); Mencorne, Rozeira 4299 (PO!); s.l., Barter 1927 (BM!, K!-holotype), Wilde et al. 371 (WAG, cited by Holttum, I.e.). DISTRIBUTION. Endemic. Triplophyllum protensum (Afzel. ex Sw.) Holttum in Kew Bull. 41: 247 (1986); Benl in Acta Bot. Bardnon. 40: 45 (1991). Aspidium subquinquefidum P. Beauv., Fl. Oware 1: 34, t. 19 (1805). Dryopteris protensa (Afzel. ex Sw.) C. Chr., Indexfilic.: 286 (1905); Alston in Exell, Cat. Vase. PL S. Tome: 63 (1944). SAO TOME. Bemposta, 600 m, Moller s.n. (BM!, COI!). PRINCIPE. SW Esperan9a, 200 m, Exell 67 1 a (BM ! ); Sao Joao, Newton s.n. (COI!); Infante D. Henrique, old path to Ribeira Bibi, Rozeira 2390A (PO!); Pico Papagaio, Welwitsch 16 (BM!, LISU!); s.l., Souza s.n. (COI!). DISTRIBUTION. Widespread in W. and Central tropical Africa. NOTE. Pichi Sermolli (1991) considered that the type of Triplophyllum protensum was a hybrid and described this taxon as a new species, T. heudelotii Pic. Serm. He considered Triplophyllum subquinquefidum (P. Beauv.) Pic. Serm. to be a distinct species. Of these two taxa only the latter was recorded for S. Tome and Principe. Triplophyllum securidiforme (Hook.) Holttum in Kew Bull. 41: 242 (1986); Benl in Acta Bot. Bardnon. 40: 43 (1991). Dryopteris securidiformis (Hook.) C. Chr., Indexfilic.: 291 (1905). D. securidiformis var. nana Bonap., Not. pterid. 14: 211 (1923); Alston in Exell, Cat. Vase. PL S. Tome: 64 (1944). Triplophyllum securidiforme var. nanum (Bonap.) Holttum in Kew Bull. 41:243(1986). SAO TOME. Macambrara, primary forest, 1050-1200 m, Exell 103 (BM!), 151 (BM!); Monte Cafe, 800 m, Moller s.n. (COI!); between Monte Quinas and Quinas Altas, Monod 12013 (BM!); Sao Joao dos Angolares, 200 m, Quintas s.n. (COI!); Ribeira Moca, Rozeira 21 19 (PO!); between Ermelinda and Cabumbe, Rozeira 2819 (PO!), 2823 (PO!); Cantagalo. 400 m, Rozeira 3152 (PO!); Sela do Camelo, 200 m, Rozeira 3451 (PO!); Formoso, Rozeira 3537 (PO!); near Rio Miranda Guedes, Rozeira 3639 (PO!); Vieira Machado, 400 m, Rozeira 3666 (PO!); s.l., F.A.E (Moller & Quintas) 40 (BM!, COI!, K also Quintas 44!, LISU!, P!). PRINCIPE. Rio Bambu-Porco, 60 m, Navel 1 39 A (LISU !), 1 4 1 C (not found); Infante D. Henrique, Rose 409 (P!), 414 (P!); Neves Ferreira, Rozeira 2201 (PO!); Ribeira Camarao, Rozeira 22 19(PO!), 2220(PO!), Rozeira 2268 (PO!); between Ribeira Jambere and Ribeira Fria, Rozeira 2237 (PO!); near Ribeira Cambungo, Rozeira 2239 (PO !); Infante D. Henrique, old path to Ribeira Bibi, Rozeira 2305 (PO!), 2326 (PO!); Infante D. Henrique, Rozeira 2374 (PO!); Mencorne, Rozeira 2408 (PO !); between Barriga Branca and Mamas, Rozeira 2562 (PO!); s.l., Barter 1929 (P!), Mann s.n. (K-not found). DISTRIBUTION. W. and Central tropical Africa. 48 Aspleniaceae Asplenium L. Asplenium aethiopicum (Burm.f.) Bech. in Candollea 6:22(1 935); Alston in Exell, Cat. Vase. PL S. Tome: 74 (1944); Benl in Acta Bot. Barcinon. 40: 24 (1991). SAO TOME. Pico, primary forest, 1987 m, epiphyte, Matos & Van Essche 7381 (LISC!); Lagoa Amelia, 1430 m, epiphyte, Matos 7495 (LISC!); between Estasao Sousa and Pico, primary forest, 1700 m, epiphyte, Matos 7554 (LISC!); Macambrara, 1270 m, Moller s.n. (COI!); Lagoa Amelia to Sao Pedro, 1200 m, Quintas 1392 (BM!, COI!); Monte Cafe, Quintas 1392 (LISJC!); s.l., F.A.E. {Moller) 31 (BM!, K!, LISU!), Moller s.n. (COI!). DISTRIBUTION. Pantropical and subtropical. Asplenium africanum Desv. in Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 322 (1811); Alston in Exell, Cat Vase. PI. S. Tome: 74 (1944); Benl in Acta Bot. Barcmon. 40:6(1991). SAO TOME. Ilheu das Rolas, Greeffs.n. (B); near Sao Tome, 8 m, Moller s.n. (COI!); Agua Ize, frequent epiphyte, Pinto Basto2\S (LISC!); s.l., Don s.n. (BM!), F.A.E. (Moller) 21 (BM!, LISU!, P!). PRINCIPE. Above Neves Ferreira, 200-350 m, epiphyte, Exell 650 (BM!, COI!); s.l., Newton (not found at COI). DISTRIBUTION. W., Central, and E. Africa. Asplenium anisophyllum Kunze in Linnaea 10: 511 (1836). Asplenium geppii Carruth., Cat. Afr. PI. 2: 269 (1901); Alston in Exell, Cat. Vase. PL S. Tome: 78 (1944). SAO TOME. Near Sao Pedro, 1280 m, F.A.E. (Moller) 26 (BM!, K also Moller 23 !, LISU!); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain forest, 1200-1800 m, Lejoly 95/18 (LISC!); NW of Pico, 1350- 1600 m, Monod 12226 (BM!); Lagoa Amelia, Quintas 1385 (BM!, COI!), Rose 206 (P!). DISTRIBUTION. W. tropical Africa. Asplenium barteri Hook., Sec. cent, ferns: t. 75 (1860); Alston in Exell, Cat. Vase. PL S. Tome: 75 (1944);Tardieu-Blot inAubrev., Fl. Cameroun 3: 192 (1964); Benl in Acta Bot. Barcmon. 40: 13 (1991). SAO TOME. Sao Miguel, Chevalier 1449 Ibis (P-not found); SW region, Chevalier 14618 (P!); Porto Alegre, Chevalier 14619bis (P!); Saudade and Pico, F.A.E. (Moller) 25 p.p. (BM!); Sao Joao dos Angolares, 80 m, Newton s.n. (BM!, COI!); Cruzeiro, 300 m, Thorold 2065 (BM!); s.l., Matos s.n. (LISC!). PRINCIPE. Esperanca, plantation, 100 m, on rocks, Exell 677 (BM!, COI!); W. of Pico, above Lapa, below 250 m, Monod 12100 (BM!); Pico, near Lapa, below 250 m, Monod 12103 (BM!); Sao Joao, 450 m, Newton 28 (BM!, COI!); Lapa (Oeste), Quintas 25 (BM!, COI!); s.l., Barter 1909 (K!), 1910 (K!). DISTRIBUTION. Widespread in tropical Africa. Asplenium biafranum Alston & F. Ballard in Hooker's Icon. pi. 34: t. 3367 (1938); Alston in Exell, Cat. Vase. PL S. Tome: 75 (1944); Tardieu-Blot in Aubrev., FL Cameroun 3: 200 (1964); Benl in Acta Bot. Barcinon. 40: 23 (1991). SAO TOME. Between Monte Cafe and Pico, Chevalier 14543 (P!); Macambrara, primary forest, 1050-1200 m, Exell 154 (BM!); Pico, primary forest, 1900-2020 m, Lejoly 94/581 (LISC!); between Lagoa Amelia and Chamico, Matos 7528 (LISC!); Santa Maria, 1350 m, Moller s.n. (COI!); E. FIGUEIREDO Lagoa Amelia, 1400 m, Monod 11747 (BM!); Calvario, 1560 m, Monod 1 1 834 (B M ! ) ; between Casa do Pico and Pico, 1 950 m, Monod 1 1 958 (BM ! ). DISTRIBUTION. Benin, Bioko, Nigeria, Cameroun. Asplenium cuneatum Lam., Encycl. 2: 309 (1786); Alston in Exell, Cat. Vase. PL S. Tome: 75 (1944); Tardieu-Blot in Aubrev., FL Cameroun 3: 219(1964). SAO TOME. Tras-os-Montes, Morro Pempem, 1200 m, Quintas 1387 (BM!, COI!). DISTRIBUTION. W. tropical Africa, tropical America, Polynesia. NOTE. This taxon has not been recorded in the islands since the nineteenth century. Asplenium currorii Hook., Sp. fil. 3: 82 (1860); Alston in Exell, Cat. Vase. PL S. Tome: 75 (1944); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 445 (1959); Tardieu-Blot in Aubrev., FL Cameroun 3: 180 (1964). SAO TOME. Macambrara, primary forest, 1050-1200 m, epiphyte, Exell 172 (BM!); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain forest, 1200-1800 m, Lejoly 95/29 (LISC!); between Bom Sucesso and Lagoa Amelia, primary forest, 1350-1420 m, Matos & Van Essche 7291 (LISC!); Moller s.n. (COI!); Lagoa Amelia, 1500 m, Monod 1 1722 (BM!); s.l., 550 m, Welwitsch 64 (BM!, LISU!). PRINCIPE. Pico Papagaio, Welwitsch 24a (BM!); s.l., Barter 1900 (K-not found), Mann 1861 (K!), Welwitsch 24 (BM!, K-not found, LISU!). DISTRIBUTION. W. tropical Africa. Asplenium dregeanum Kunze in Linnaea 10:517(1 836); Alston in Exell, Cat. Vase. PL S. Tome: 76 (1944); Alston in Exell, Suppl Cat. Vase. PL S. Tome: 1 (1956). subsp. brachypterum (Kunze ex Houlston & T. Moore) Pic. Serm. in Bull. Jard. Bot. Belg. 55: 130 (1985); Benl in Acta Bot. Barci- non. 40: 28 (1991). SAO TOME. Monte Cafe, 1200 m, epiphyte, Espirito Santo 164 (BM!, COI!, LISJC!); Macambrara, primary forest, 1050-1200 m, Exell 96 (BM!), 126 (BM!, COI!); Lagoa Amelia, 1200-1400 m, Lains e Silva 83 (LISU!), Matos 7493 (LISC!), 1400 m, Monod 1 1741 (BM!), 1400 m, Quintas 1349 (BM!, COI!), Rose 224 (P!), 240 (P!), Rozeira 154 (PO!), 1 161 (PO!); Pico, primary forest, 1900-2020 m, epiphyte, Lejoly 94/580 (LISC!), 1950 m, Moller s.n. (COI!); between Bom Sucesso and LagoaAmelia, primary forest, 1350-1400 m, epiphyte, Matos & Van Essche 7287 (LISC!); between Sao Pedro and Lagoa Amelia, 1050 m, Monod 11705 (BM!, COI!); Calvario, 1400 m, Monod 11852 (BM!); s.l., F.A.E. (Moller) 33 (BM!, COI!, K!, LISJC!, LISU!), Mann s.n. (K!), Moller s.n. (COI!). DISTRIBUTION. Zaire, Rwanda, Burundi. Asplenium emarginatum P. Beauv., Fl. Oware 2: 6, t. 61 (1808); Alston in Exell, Cat. Vase. PL S. Tome: 76 ( 1944);Alston in Exell, Suppl. Cat. Vase. PL S. Tome: 1 (1956); Tardieu-Blot inAubrev., Fl. Cameroun 3: 192 (1964); Benl in Acta Bot. Barcinon. 40: 10 (1991). SAO TOME. Sao Vicente, 200 m, common, Esptrito Santo 35 (BM!, COI!, LISC!, LISJC!); Monte Cafe, 800 m, Moller 349 (BM!, COI!); Santa Margarida, Antonio Soares, /toze/ra 1225 (PO!); s.l., Don. s.n. (BM!),800m, Quintas 8 (K!). PRINCIPE? (Beauvois, I.e.). DISTRIBUTION. Widespread in tropical Africa. Asplenium erectum Bory ex Willd., Sp. pi. 5: 328 (1810). van usambarense (Hieron.) Schelpe in Bol. Soc. Brot., ser. 2, 41: PTERIDOPHYTES OF SAO TOME AND PRINCIPE 49 207 (1967); Schelpe in Exell & Launert, Fl. Zamb., Pteridophyta: 176 (1970); Benl in Acta Bot. Barcinon. 40: 16 (1991). Asplenium quintasii Gand. in Bull. Soc. Bot. France 66: 305 (1919); Alston in Exell, Cat. Vase. PL S. Tome: 79 (1944);Tardieu-Blot in Aubrev., Fl. Cameroun 3: 198 (1964). SAO TOME. Monte Caf6, Chevalier 13673 (BM!, P!); Pico, 1850 m, Exell 345a (BM!), Newton s.n. (BM!, COI!), 1650 m, Quintas 1342 (BM!, COI!; type of Asplenium quintasii); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain forest, 1200-1800 m, Lejoly 95/19 (LISC!); between Pico and Ponta Figo, primary forest, 1600-1700 m, epiphyte, Matos & Van Essche 7389 (LISC!); Calvario, 1450 m, Monod 1 1826 (BM!). DISTRIBUTION. Widespread in tropical Africa. Asplenium eurysorum Hieron. in Engl. in Bot. Jahrb. Syst. 46: 364 (191 1); Alston in Exell, Cat. Vase. PI. S. Tome: 76 (1944). SAO TOME. Pico, Chevalier 13659 (P!), primary forest, 1987 m, epiphyte, Matos & Van Essche 7382 (LISC!); Calvario, 920 m, Espirito Santo 5070 (LISC!, LISJC!); between Bacelar and Quija, secondary forest, 160 m, Espirito Santo 5139 (LISC!, LISJC!); Macambrara, primary forest, 1050- 1200 m, ground fern, Exell 132 (BM!), 1385 m, Moller s.n. (COI!); Ribeira Peixe, Lains e Silva 301 (LISU!); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain forest, 1200-1800 m, Lejoly 95/1 (LISC!); Santa Maria, 1260 m, Moller s.n. (COI!); between Sao Pedro and Lagoa Amelia, 1050 m, Monod 11714 (BM!); between Monte Quinas and Quinas Alias, Monod 12008 (BM!); s.l., F.A.E. (Moller) 28 (BM!, LISJC!, LISU!, P!). PRINCIPE. Oque Nazare, Newton 27 (BM!, COI!, P!); Pico Papagaio, Welwitsch 14 (BM!, K!, LISU!); s.l., Barter 1896b (K!). DISTRIBUTION. Endemic. Asplenium exhaustum (H. Christ) Alston in Exell, Cat. Vase. PL S. Tome: 76, f. 2 (1944); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 445 (1959). SAO TOME. Pico, Chevalier 14587 (P!-type); Cruzeiro, 300 m, Thorold 2072 (BM!); s.l., Seabra s.n. (BM!, COI!, LISU!). DISTRIBUTION. Endemic. Asplenium formosum Willd., Sp. pi. 5: 329 (1810); Alston in Exell & Rozeira in Conf. Int. Afr. Oc. 3: 79 (1958). SAO TOME. Diogo Vaz, near Agua Telha, Rozeira 5 1 8a (PO). DISTRIBUTION. Pantropical. Asplenium friesiorumC. Chr. inNotizbl. Bot. Gart. Berlin-Dahlem 9: 181 (1924); Benl in Acta Bot. Barcinon. 40: 22 (1991). SAO TOME. Pico, 1950 m, F.A.E. (Moller) 27 (BM!, K!, LISU!, P!); between Casa do Pico and Pico, 1950 m, Monod 1 1957 (BM!). DISTRIBUTION. Widespread in tropical Africa. Asplenium hypomelas Kuhn, Filic. afr.: 104 (1868); Alston in Exell, Cat Vase. PL S. Tome: 78 (1944); Benl in Acta Bot. Barcinon. 40: 31 (1991). SAO TOME. Porto Alegre, Chevalier 13658 (P!); between Monte Cafe and Pico, Chevalier 14579 (P!); Lagoa Amelia, 1400 m, Espirito Santo 4069 (COI!, LISJC!), 1400 m, Monod 1 1735 (BM!, COI!), Rose 233 (P!), Rozeira 1 129 (PO!); Calvario, secondary forest, 920 m, Espirito Santo 5074 (LISC!, LISJC!), 1200 m,Monod 11813 (BM\),Rose 150 (P!); Macambrara, primary forest, 1050-1200 m, ground fern, Exell 120 (BM!, COI!), 1200-1350 m, ground fern, 235a (BM!, COI!);Trds-os-Montes, Gamas.n. (COI!); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain forest, 1200-1800 m, Lejoly 95/16 (LISC!); between Bom Sucesso and Lagoa Amelia, primary forest, 1350-1420 m,Matos& Van £wc/je 7295 (LISC!); Rio Contador, 1400 m, Moller s.n. (COI!); W. of Pico, 1225 m, Monod 11996 (BM!); Pico, Newton s.n. (BM!, COI!); Sao Pedro, Quintas s.n. (COI!); s.l., F.A.E. (Moller & Quintas) \\(BM\, COI!, K also Moller 10!, LISJC!, LISU!, P\),Mann s.n. (K!), Quintas s.n. (LISU!), Souza s.n. (COI!). DISTRIBUTION. Widespread in tropical Africa, extending to S. and SE Africa. Asplenium inaequilaterale Bory ex Willd., Sp. pi. 5: 322 (1810); Alston in Exell, Cat. Vase. PL S. Tome: 78 (1944); Benl in Acta Bot. Barcinon. 40: 17 (1991). SAO TOME. Ribeira Peixe, Lains e Silva 253 (LISU!), 254 (LISU!); Lagoa Amelia, 1430 m, Matos 7492 (LISC!); between Bom Sucesso and Macambrara, Matos 7605 (LISC!); Saudade, 750 m, Moller s.n. (COI!); Calvdrio, Monod 1 1856 (BM!); s.l., F.A.E. (Moller) 25 p.p. (BM!, LISJC!, LISU!), Moller 25 (BM!, COI!). DISTRIBUTION. Widespread in tropical Africa, extending to S. and SE Africa and Madagascar. Asplenium lividum Mett. ex Kuhn in Linnaea 36: 100 (1869). Asplenium protensum sensu auct. non Schrad. in Gott. Gel. Anz. [1818]: 916 (1818); Alston in Exell, Cat. Vase. PL S. Tome: 79 (1944). SAO TOME. Macambrara to Zampalma, 900 m, ground fem, Exell 296 (BM!, COI!); Agua Pinhao, 850 m, Quintas 1283 (BM!, COI!). DISTRIBUTION. E. and S. Africa, and S. America. Asplenium longicauda Hook., Sec. cent, ferns: t. 69 (1860); Alston in Exell, Cat Vase. PL S. Tome: 78 (1944); Tardieu-Blot in Aubrev., FL Cameroun 3: 1 86 ( 1 964); Benl in Acta Bot. Barcinon. 40:9(1991). PRINCIPE. Between Oque Pipi and Morro do Leste, secondary forest, 350 m, ground fern, Exell 571 (BM!, COI!); Rio Bambu-Porco, 60 m, Navel 144 (LISU!, P!); s.l., Barter 1900 (K!-syntype). DISTRIBUTION. W. tropical Africa, Madagascar. Asplenium megalura Hieron. in Brause & Hieron. in Mildbr., Wiss. Erg. Deut. Zentr.-Afr. Exped., Bot. 2: 17(1910); Benl inActaBot. Barcinon. 40: 23 (1991). var. molleri (Hieron. )Tardieu inMem. Inst. Franc. Afrique Noire 28: 190(1953). Asplenium molleri Hieron. in Engl. in Bot. Jahrb. Syst. 46: 371 (191 1); Alston in Exell, Cat. Vase. PL S. Tome: 79 (1944). SAO TOME. Lagoa Amelia, 1400 m, epiphyte, Espirito Santo 4068 (COI!, LISJC!), crater rim, 1350 m, epiphyte, Exell 202 (BM!), 1430 m, rare epiphyte, Matos 7496 (LISC!), 1400 m, Monod 1 1779 (BM!); Macambrara, primary forest, 1200 m, epiphyte, Exell 228 (BM!); Bom Sucesso, 1075 m, F.A.E. (Moller) 29 (BM!, COI!, K!-type, LISU!); between Bom Sucesso and Lagoa Amelia, secondary forest, 1 150-1300 m, Matos & Van Essche 7310 (LISC!); Calvario, 1400 m, Monod 1 1854 (BM!); W. of Pico, 870-1225 m, Monod 1 1905 (BM!); s.l., Matos s.n. (LISC!). DISTRIBUTION. The species is widespread in tropical Africa. The variety is endemic to Sao Tome. Asplenium nigritianum Hook., Sp.fil. 3: 223 (1860) et Sec. cent, ferns: t. 44 (1860); Alston in Exell, Cat. Vase. PL S. Tome: 79 (1944); Alston in Bol. Soc. Brot., ser. 2, 30: 10 (1956). SAO TOME. S.l., Mann 1 107 (K-not found). PRINCIPE. Lapa (Leste), Quintas s.n. (BM!, COI!); s.l., Barter 1895 (BM!, K-not found), (probably Lapa), Quintas B (K!). 50 DISTRIBUTION. Tropical Africa. NOTES. As shown by Alston ( 1956£), it was wrongly assumed that the type collection came from Bioko. In fact, Asplenium nigritianum does not seem to occur in Bioko. Alston (19566) referred to the type collection as Barter 225, explaining that Hooker mistook the date, 1 859, for the number and then miscopied it as 1 895 (in K specimen) or 1 898 (in BM speci- men). Since the specimen at K was not found, it was not possible to check its number. The collections of pteridophytes made by Barter which I examined range from no. 1886 to 1928, so it is likely that these figures refer to the number and not the date of collection. The type collection should be referred to as Barter 1895. The species is poorly represented in herbaria. Neither the speci- men Barter 1895 (K) nor Mann 1107 (K) could be located. In a handwritten document kept at BM, reference is made to a duplicate of the Mann collection kept at B. This taxon has not been recorded in the islands since the nine- teenth century. Asplenium paucijugum F. Ballard in Hooker's Icon. pi. 33: t. 3287 (1935); Alston in Exell, Cat. Vase. PL S. Tome: 79 (1944); Benl in Acta Bot. Barcinon. 40: 8 (1991). Asplenium variabile Hook., Sp. fil. 3: 93, t. 185 (1860) var. paucijugum (F. Ballard) Alston in Bol. Soc. Brot., ser. 2, 30: 7 (1956); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 184 (1964). SAO TOME. Monte Cafe, 800 m, F.A.E. (Moller) 24 (BM!, COI!, LISJC!, LISU!); LagoaAmelia, primary forest, 1400-1420 m, epiphyte, Matos & Van Essche 7410 (LISC!); NW of Pico, 1000-1350 m, Monod 12216 (BM!); s.l., 800 m, Moller 19 (K!), 550 m, Welwitsch 63 (BM!, LISU!). PRINCIPE. Sundi, Newton 29 (BM!, COI!). DISTRIBUTION. W., Central, and E. tropical Africa, Madagascar. Asplenium sandersonii Hook., Sp.fil. 3: 147, t.179 (1860); Alston in Exell, Cat. Vase. PL S. Tome: 80 (1944); Benl in Acta Bot. Barcinon. 40: 27 (1991). SAO TOME. Macambrara, primary forest, 1050-1200 m, epiphyte, Exell 144 (BM!, COI!); Sao Miguel, between Xufexufe and Sao Miguel, 400 m, epiphyte, Matos 7658 (LISC!); Pico, 1950 m, Moller s.n. (COI!); Bombaim, Rose 156 (P!); DonaAugusta, Dona Eugenia, epiphyte, Rozeira 1096 (PO!); LagoaAmelia, 1400 m, epiphyte, Sergio s.n. (LISU!); s.l., F.A.E. (Moller) 23 (BM!, K also Moller 21 !, LISU!), Mann (not found), Moller s.n. (COI!). DISTRIBUTION. Widespread in tropical and subtropical Africa. Asplenium unilaterale Lam., Encycl. 2: 305 (1786); Alston in Exell, Cat Vase. PL S. Tome: 80 (1944); Benl in Acta Bot. Barcinon. 40: 15 (1991). SAO TOME. Pico, 1300-1900 m, EA.E. (Moller) 30 (K-not found, LISU!); Pico, 1900 m, Moller s.n. (COI!); Rio Contador, 1300 m, Moller s.n. (COI!). PRINCIPE. S.L, Barter 1917 (K!). DISTRIBUTION. Palaeotropical. NOTE. This taxon has not been recorded in the islands since the nineteenth century. Asplenium variabile Hook., Sp.fil. 3: 93, t. 185 (1860); Alston in Exell, Cat Vase. PI. S. Tome: 80 (1944); Benl in Acta Bot. Barcinon. 40: 7 (1991). SAO TOME. 16 Grande, Sao Joao dos Angolares, 50 m, EA.E. (Quintas) 22 (K also Quintas 20!, LISU!); between Xufexufe and Sao Miguel, 400 m, epi- phyte, Matos 7657 (LISC!); Sao Joao dos Angolares, 50 m, Quintas s.n. (COI!). E. FIGUEIREDO PRINCIPE. Between Oque Pipi and Morro do Leste, secondary forest, 350 m, on rocks, Exell 567 (BM!, COI!); W. of Pico, above Lapa, below 250 m, Monod 12099 (BM!); Rio Bambu-Porco, 60 m, Navel 140 (LISU!, P!); Pico, Thorold 2085 (BM!); s.l., Mann s.n. (K-not found). DISTRIBUTION. W. tropical Africa. NOTE. Alston (1944) cited the collection Barter 1910 (BM, K). At BM, only the specimen Barter s.n. (Niger Expedition) was found. Barter 1910 (K) is Asplenium barteri. Athyriaceae Athyrium Roth Athyrium newtonii (Baker) Diels in Engl. & Prantl, Nat. Pflanzenfam. 1(4): 224 (1899); Alston in Exell, Cat. Vase. PL S. Tome: 73 (1944). SAO TOME. Pico, 1650 m, Quintas 1393 (BM!, COI!, LISJC!), 1650 m, Quintas s.n. (COI!); s.l., Newton 2 (K!-type), Newton s.n. (COI!), (probably Pico), 1650 m, Quintas 9 (K!). DISTRIBUTION. Also in Uganda and Tanzania, above 1000 m. NOTES. Newton 88 (K) cited by Alston (I.e.) is most probably a mistake for Quintas 9 (the number 88 after the collector's name is the date). This taxon has not been recorded in the islands since the nine- teenth century. Diplazium Sw. Diplazium arborescens (Bory) Sw., Syn.fiL: 92 (1806); Alston in Exell, Cat. Vase. PL S. Tome: 73 (1944). SAO TOME. Between Monte Cafe and Pico, Chevalier 14290 (P!); Pico, Chevalier s.n. (P!); Macambrara, primary forest, 1050-1200 m, Exell 128 (BM!, COI!);Tras-os-Montes, Gama s.n. (COI!); between LagoaAmelia and Chamico, primary forest, 1400 m, Matos 7526 (LISC!); Monte Cafe, Moller s.n. (COI!); LagoaAmelia, 1350-1380 m, Monod 1 1772 (BM!, COI!), Rose 234 (P!), Rozeira 1139 (K!, PO!); Cantagalo, 400 m, Rozeira 3157 (PO!); Formoso, Rozeira 3236 (PO!); Nova Ceilao, Rozeira 4905 (PO!); s.l., EA.E. (Moller) 34 (BM!, COI!, K also Moller 28!, LISJC!, LISU!, P!), Mann 621 (P!), 1200 m, Mann s.n. (K!), Moller s.n. (COI!), Rozeira 2846 (PO!). DISTRIBUTION. Also in Madagascar, Comoros, and Mascarene Is. Diplazium proliferum (Lam.) Thouars, Esquisse fl. Tristan d'Acugna: 35 (1804); Alston in Exell, Cat. Vase. PL S. Tome: 73 (1944); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 237 (1964). Callipteris prolifera (Lam.) Bory, Voy. ties Afrique 1: 283 (1804); Benl in Acta Bot. Barcinon. 38: 37 (1988). SAO TOME. Between Bom Sucesso and Lagoa Amelia, Almeida s.n. (LISC!); Boa Entrada, Chevalier 14333 (P!), 14361 (P!) 14361bis (not found); Ribeira Peixe, S. of Cabumbe, secondary forest, 350 m, Espirito Santo 3916 (COI!, LISJC!); Morro Vigia, 800 m, Espirito Santo 4253 (LISC!, LISJC!); Monte Rosa, secondary forest, 50 m, Espirito Santo 4748 (LISC!, LISJC !);Tras-os-Montes, Lains e Silva 229 (LISU!); between Monte Castro and Ponta Figo, secondary forest, 500-1070 m,Lejoly95/45 (LISC!); Dona Amelia, Santa Catarina, near Agua Cardoso, Matos 7593 (LISC!); Pedroma, Mocquerys 89/90 (P!), 131/132 (P!); Nova Moca, near Monte Cafe, 90 m, Moller s.n. (COI!); Pinheira, Moller s.n. (COI!); Rodia, 540 m, Moller s.n. (COI!); Calvario, 1200 m, Monod 11816 (BM!); waterfall near Sao Nicolau, Monod 12037 (BM!, COI!), Rozeira 571 (PO!); Cruzeiro, Rozeira 765 (PO!); between Milagrosa and Rio Abade, near Formoso, Rozeira 3188 (PO!); on the way from Santa Irene to the river, Rozeira 3319 PTERIDOPHYTES OF SAO TOME AND PRINCIPE (PO!); Sela do Camelo, near Lembd, 300-700 m, Rozeira 3459 (PO!); Monte Cafe, 760 m, Welwitsch 60 (BM!, K!, LISU!); s.l., Don 49 (BM!), F.A.E. (Moller) 35 (BM!, COI!, LISJC!, LISU!, P!). PRINCIPE. Between Oque Pipi and Morro de Leste, secondary forest, 350 m, ground fern, Exell 563 (BM!, COI!); Rio Bambu-Porco, 60 m, Navel 137 (LISU!, P!); Infante D. Henrique, Rose 413 (P!); Infante D. Henrique, near Ribeiro Chimboto, Rozeira 3 1 2 (PO ! ); Lapa, way to Pico, 300 m, Rozeira 9 1 5 (PO!); near Ribeira Camarao, Rozeira 2213 (PO!); Bela Vista, near main house, Rozeira 3916 (PO!). DISTRIBUTION. Palaeotropical. Blechnaceae Blechnum L. Blechnum attenuatum (Sw.) Mett., Fit. hort. hot. Lips.: (A, t. 3, figs 1-6 (1856); Alston in Exell, Cat. Vase. PL S. Tome: 80 (1944); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 295 (1964); Benl in Acta Bot. Barcinon. 38: 63 (1988). SAO TOME. Macambrara to Zampalma, primary forest, 800-1050 m, Exell 437 (BM!, COI!), 438 (BM!, COI!); between Casa do Pico and Pico, 1 250 m, Mono's.n.(BM!,E). DISTRIBUTION. Palaeotropical. Christella hispidula (Decne.) Holttum in Kew Bull. 31: 312 (1976); Benl in Acta Bot. Barcinon. 38: 51 (1988). Dryopteris quadrangularis (Fee) Alston in J. Bot. 75: 253 (1937); Alston in Exell, Cat. Vase. PL S. Tome: 63 (1944). SAO TOME. Agua Grande, 100 m, Moller s.n. (COI!); Saudade, 720 m, Moller s.n. (COI!); above Monte Cafe, 1010 m, Monod 11691 (BM!); between Monte Quinas and Quinas Alias, Monod 12006 (BM!). DISTRIBUTION. Pantropical. Cyclosorus Link Cyclosorus striatus (Schum.) Ching in Bull. Fan Mem. Inst. Biol. 10: 249 (1941); Benl in Acta Bot. Barcinon. 38: 55 (1988). Dryopteris striata (Schum.) C. Chr., Indexfilic.: 294 (1905); Alston in Exell, Cat. Vase. PL S. Tome: 64 (1944). SAO TOME. Lagoa Amelia, Chevalier 13653 (P). DISTRIBUTION. Tropical Africa. Metathelypteris (H. Ito) Ching Metathelypteris fragilis (Baker) Holttum in /. S. African Bot. 40: 127 (1974). Dryopteris fragilis (Baker) C. Chr., Indexfilic.: 266 (1905); Alston in Exell, Cat. Vase. PL S. Tome: 62 (1944). subsp. guineensis Benl in Nova Hedwigia 27: 150 (1976); Benl in Acta Bot. Barcinon. 38: 43 (1988). SAO TOME. Macambrara, primary forest, 1050-1200 m, Exell 133 (BM!); s.l., Newton 1 (K!). DISTRIBUTION. Bioko. NOTE. The collection cited by Holttum (I.e.) as Newton s.n. (K ex COI), 1885, is Newton 1 (K). PTERIDOPHYTES OF SAO TOME AND PRINCIPE Pneumatopteris Nakai Pneumatopteris afra (C. Chr.) Holttum in Blumea 21: 306 (1973); Benl in Acta Bot. Barcinon. 38: 57 (1988). Dryopteris oppositifolia sensu auct. non (Hook.) C. Chr., Index filic.: 281 (1905); Alston in Exell, Cat Vase. PL S. Tome: 63 (1944) p.p. quoad pi. ex Principe. Cyclosorus afer (H. Christ) Ching in Bull. Fan Mem. Inst. Biol. 10: 242 ( 1 94 1 ); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21:446(1959). SAO TOME. Sao Vicente, Espirito Santo 41 p.p. (LISC!); Sao Miguel, 50m, Espirito Santo 4744 (LISC!); between Pico and Monte Castro, 6 km S. of Ponta Figo, mountain rainforest, 1200-1800 m, Lejoly 95/30 (LISC!); Pedroma, Mocquerys 39/40 (P!); between Quinas Altas and Monte Quinas, Monod 12023 (BM!); Agua Ize, Pinto Basto 213 (LISC!); Lagoa Amelia, Rose 204 (P\), 231 (P!). PRINCIPE. Terreiro Velho, plantation, 200 m, Exell 530a (BM!, COI!); between Oque Pipi and Morro do Leste, secondary forest, 350 m, ground fern, Exell 566 (BM!, COI!); Infante D. Henrique, Rose 398 (P!); s.l., Barter 1924 (K!), Rattray s.n. (E), 300 m, Welwitsch 21 (BM!, K!, LISU!), 21bis (K!). DISTRIBUTION. Widespread in W. tropical Africa. Pneumatopteris oppositifolia (Hook.) Holttum in Blumea 21: 304 (1973); Holttum in/ S. African Bot. 40: 157 (1974); Benl inActa Bot. Barcinon. 38: 62 (1988). Dryopteris oppositifolia (Hook.) C. Chr., Index filic.: 281 (1905); Alston in Exell, Cat. Vase. PI. S. Tome: 63 (1944) p.p. quoad pi. ex Sao Tome. Cyclosorus oppositifolius (Hook.)Tardieu inNotul. Syst. (Paris) 14: 346 ( 1 953); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21:445(1959). SAO TOME. Between Lagoa Amelia and Calvario, primary forest, 150 m, ground fern, Exell 259 (BM!, COI!); Macambrara to Zampalma, 900 m, ground fern, Exell 297 (BM!); Tras-os-Montes, Gama s.n. (COI!); Ribeira Peixe, 140 m, frequent, Lains e Silva 268 (LISU!); Pico, 1500 m, Mann s.n. (K!-type); Lagoa Amelia, 1400 m, Monod 11748 (BM!); Calvario, 1200 m, Monod 11814 (BM!); between Quinas Altas and Monte Quinas, Monod 12018 (BM!); s.l., F.A.E. (Moller) 41 p.p. (BM!, COI!, LISU!, P!). DISTRIBUTION. Bioko, Annobon. Pneumatopteris venulosa (Kuntze) Holttum in Blumea 21: 315 (1973); Benl in Acta Bot. Barcinon. 38: 59 (1988). Dryopteris venulosa Kuntze, Revis. gen. pi. 2: 814 (1891); Alston in Exell, Cat. Vase. PL S. Tome: 66 (1944). Cyclosorus elatus (Mett. ex Kuhn) Alston in Bol. Soc. Brot., ser. 2, 30: 13 (1956); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 446 (1959). SAO TOME. Boa Entrada, Chevalier 13680 (P!); Macambrara, primary forest, 1050-1200 m, ground fern, Exell 242 (BM!, COI!); Lagoa Amelia, 1400 m, Monod 1 1750 (BM!); s.l., Don s.n. (BM-not found), F.A.E. (Moller) 42 p.p. (K also Moller 39!, P!). PRINCIPE. Esperan9a, Navel 145 (LISU!, P!). DISTRIBUTION. Equatorial Guinea (continent, Bioko and Annobon). Pseudophegopteris Ching Pseudophegopteris henriquesii (Baker) Holttum in Blumea 17: 15 (1969); Holttum in J. S. African Bot. 40: 128 (1974). Polypodium henriquesii Baker in Henriq. in Bol. Soc. Brot. 4: 154, t.l (1887). 63 Dryopteris henriquesii (Baker) C. Chr., Index filic.: 270 (1905); Alston in Exell, Cat. Vase. PL S. Tome: 62 (1944). Thelypteris henriquesii (Baker) Tardieu in Notul. Syst. (Paris) 14: 344(1953). Macrothelypteris henriquesii (Baker) Pic. Serm. in Webbia 23: 179 (1968). SAO TOME. Tras-os-Montes, Gama s.n. (COI!); Nova Moca, Moller s.n. (COI!), 750 m, Quintas 1464 (COI!, LISJC!, P!); Macambrard, 1250 m, Moller s.n. (COI!); Sao Nicolau, waterfall, Rozeira 162 (PO!); Trds-os- Montes, Calvario, Rozeira 562 (PO!); s.l., F.A.E. (Moller) 49 (BM!, K!-also Moller 45, type, LISU!, P!), Moller 1 (BM!). DISTRIBUTION. Endemic. Sphaerostephanos J. Sm. Sphaerostephanos elatus (Bojer) Holttum in J. 5. African Bot. 40: 167(1974). subsp. thomensis Holttum in J. S. African Bot. 40: 167 (1974). Thelypteris mauritiana (Fee) C.F. Reed in Phytologia 17: 291 (1968) subsp. thomensis (Holttum) Viane in Bull. Soc. Roy. Bot. Belg. 118:54(1985). SAO TOME. S.l., F.A.E. (Moller) 41 p.p. (COI!, P!-type). DISTRIBUTION. Endemic. NOTE. This taxon has not been recorded in the islands since the nineteenth century. Vittariaceae Anthrophyum Kaulf. Anthrophyum immersum (Bory ex Willd.) Mett. ex Hook, in Hook. & Baker, Syn.fiL: 393 (1868); Alston in Exell in Bull. Inst. Franc. Afrique Noire ser. A, 21: 442 (1959). SAO TOME. Monte Verde, secondary forest, 10-150 m, Lejoly 95/86 (LISC!); between Estacao Sousa and Pico, primary forest, 1700 m, rare epiphyte, Matos 7553 (LISC!); Cruzeiro, 300 m, Thorold 2064 (BM!). PRINCIPE. Pico, 950 m, Thorold 2081 (BM!). DISTRIBUTION. Also in Liberia and Mascarene Is. Anthrophyum mannianum Hook., Sec. cent, ferns: t. 73 (1860); Alston in Exell, Cat. Vase. PL S. Tome: 87 (1944); Benl in Acta Bot. Barcinon. 33: 6 (1982). SAO TOME. Pico, light woodland, 2000 m, epiphyte, Exell 340 (BM!); near Rio Campos, primary forest, epiphyte, Matos & Van Essche 7441 (LISC!); between Sao Pedro and Lagoa Amelia, 1250 m, Moller s.n. (COI!); Macambrara, 1 200 m, epiphyte, Moller s.n. (COI!); Sao JoaodosAngolares, 200 m, Quintas 54A (K!), Quintas s.n. (COI!); Lagoa Amelia, epiphyte, Rozeira 1097 (PO!); s.l., F.A.E. (Moller & Quintas) 60 (BM!, COI!, K!, LISJC!, LISU!), 1250 m, Moller 54 (K!). DISTRIBUTION. Widespread in tropical Africa. Vittaria Sm. Vittaria guineensis Desv. in Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 325 (1811); Alston in Exell, Cat. Vase. PL S. Tome: 86 (1944);Tardieu-Blot inAubrev., Fl. Cameroun 3: 124 (1964); Benl in Acta Bot. Barcinon. 33: 7 (1982). SAO TOME. SW region, Chevalier 14598 (P!); Calvario, 970 m, epiphyte, 64 E. FIGUEIREDO Espi'rito Santo5Q59 (LISC!, LISJC!); between LagoaAmelia and Esperanca, 1300 m, epiphyte, Espi'rito Santo5\56 (LISC!, LISJC!); between Sao Carlos and Bom Sucesso, secondary forest, Espi'rito Santo 5 1 90 (LISJC!); Sao Joao dosAngolares, littoral, F.A.E. (Newton) 61 p.p. (BM!), 80 m, F.A.E. (Quintas) 61 p.p. (BM!, COI!, K!-mixed with Barter 1908, LISU!), Newton s.n. p.p. (COI!, P!); Ilheu das Rolas, Greeffs.n. (B?); between Pico and Monte Figo, 6 km S. of Ponta Figo, mountain rainforest, 1200-1800 m, epiphyte, Lejoly 95/6 (LISC!); W. of Pico, 870-1225 m, Monod 1 1906 (BM!); Macambrara, 1200 m, Quintas s.n. (COI!); Ribeira Peixe, between Vila Aida and main house, epiphyte, Rozeira 291 (PO!). PRINCIPE. Pico Papagaio, primary forest, 450 m, epiphyte, Exell 719 (BM!, COI!); Oque Caspar, Newton 25 p.p. (BM!); s.l., Barter 1908 (K!- mixed with F.A.E. 61). DISTRIBUTION. W. tropical Africa to Uganda. Vittaria owariensis Fee, Mem.foug. 3: 2 1 , t. 3, fig. 2 ( 1 852); Alston in Exell, Cat. Vase. PL S. Tome: 87 (1944). SAO TOME. Sao Joao dosAngolares, F.A.E. (Newton)6\ p.p. (BM!), F.A.E. (Quintas) 61 p.p. (BM!, P!), Newton s.n. p.p. (P!), Quintas s.n. (COI!); Vila Concei9§o, 100 m, epiphyte, Matos 7716 (LISC!); Rio Caue, epiphyte, Rozeira 30 (PO!); Rio Manuel Jorge, between Pinheira and Almas, epiphyte, Rozeira 83 (PO!); s.l., Seabra s.n. (COI!). PRINCIPE. Oque Caspar, Newton 25 p.p. (BM!, COI!); Porto Real, Rose 467 (P!); Infante D. Henrique, at the base of Dois Irmaos, epiphyte, Rozeira 480 (PO!); s.l., Welwitsch 13 (BM-not found, K!, LISU!). DISTRIBUTION. W. tropical Africa. DOUBTFUL RECORDS Asplenium hemitomum Hieron. in Engl. in Bot. Jahrb. Syst. 46: 365 (1911); Alston in Exell, Cat. Vase. PL S. Tome: 78 (1944); Benl in Acta Bot. Barcinon. 40: 20 (1991). PRINCIPE ? NOTE. Alston (I.e.) recorded this species based on the collection Barter 1896a (K!) from Principe. Barter 1896 is a mixed collec- tion, containing A. hemitomum and A. eurysorum. Since Barter also collected A. hemitomum in Bioko it is possible that a specimen of this species, from Bioko, was attached to the herbarium sheet with A. eurysorum from Principe. Ceratopteris cornuta (P. Beauv.) Lepr. in Ann. Sci. Nat. (Paris) 19: 103, t. 4A (1830); Alston in Exell, Cat. Vase. PL S. Tome: 92 (1944). PRINCIPE ? Barter (Kuhn, 1868). NOTE. No specimens were seen at BM, K or P. However, at BM there is a specimen collected by Barter during the Niger Expedition, unlocalized, which could have been collected in Principe. Dryopteris manniana (Hook.) C. Chr., Index filic.: 276 (1905); Benl in Acta Bot. Barcinon. 40: 38 (1991). SAO TOME ? NOTE. No specimens were seen. Lastreopsis nigritiana (Baker) Tindale in Contr. New South Wales Natl. Herb. 3: 245 (1963);Tardieu-Blot inAubrev., Fl. Cameroun 3: 280 (1964); Benl in Acta Bot. Barcinon. 40: 51 (1991). Polypodium nigritianum Baker in Hook. & Baker, Syn. fil.: 313 (1867). PRINCIPE ? NOTE. No specimens were seen. Although there are references in the literature to the occurrence of this species in Principe, Tindale (1965) did not record it for Sao Tome and Principe in his monograph of the genus. Lygodium microphyllum (Cav.) R. Br., Prodr.: 162 (1810). Lygodium scandens sensu auct. non (L.) Sw. in /. Bot. (Schrader) 1800(2): 106 (1801); Alston in Exell, Cat. Vase. PL S. Tome: 93 (1944). PRINCIPE ? (Barter, 1860). NOTE. No specimens were seen at BM or K. There are specimens of Lygodium microphyllum collected by Barter in 1859 during the Niger Expedition, some of which are unlocalized and could have been collected on the island. Being a lowland species, L. microphyllum might be extinct due to the extensive cultivation of these areas. Ophioglossum gramineum Willd. in Schrift. Akad. Erfurt: 18, t.l, f.l (1802); Tardieu-Blot in Aubrev., Fl. Cameroun 3: 46 (1964). SAO TOME ? NOTE. No specimens seen. Trichomanes africanum H. Christ in J. Bot. (Morot), ser. 2, 2: 21 (1909); Benl in Acta Bot. Barcinon. 32: 23 (1980). [Vandenboschia africana (H. Christ) G. Kunkel in Nova Hedwigia 6:213(1963)] SAO TOME ? NOTE. No specimens seen. ACKNOWLEDGEMENTS. Thanks are due to Fundagao Calouste Gulbenkian for a research grant. I would also like to thank the directors of BM, K, and P for providing all the facilities during my stay in their herbaria, and also Frederic Badre, Gustav Benl, Josephine Camus, Peter Edwards, Elisa Folhadela, Mary Gibby, Bob Johns, and Alison Paul. REFERENCES Alston, A.H.G. 1944. Pteridophyta. In A. Exell, Catalogue of the vascular plants ofS. Tome (with Principe and Annobon): 57-99. London. 1956a. Pteridophyta. In A. Exell, Supplement to the catalogue of the vascular plants ofS. Tome (with Principe and Annobon): 7-9. London. 1956/7. New African ferns. Boletim da Sociedade Broteriana, serie 2, 30: 5-27. 1958. Pteridophyta. In A. Exell &A. Rozeira, Aditamento a Flora das ilhas de Sao Tome e Principe. Conferencia Internacional dos Africanistas Ocidentais 3: 78-81. 1959. Pteridophyta. In A. Exell, Additions to the flora of S. Tome and Principe. Bulletin de I'lnstitut Franfaise d'Afrique Noire, serie A, 21: 440-448. Barter, C. 1 860. Letter from Mr Charles Barter, natural history collector to the Niger Expedition. Journal of the Proceedings of the Linnean Society, Botany. 4: 17-23. Benl, G. 1 978. The Pteridophyta of Fernando Po I. Acta Botanica Barcinonensia 31: 1- 31. 1980. The Pteridophyta of Fernando Po II. Acta Botanica Barcinonensia 32: 1- 34. 1982. The Pteridophyta of Fernando Po III. Acta Botanica Barcinonensia 33: 1- 46. 1 988. The Pteridophyta of Bioko (Fernando Po) IV. Acta Botanica Barcinonensia 38: 1-69. 199 1 . The Pteridophyta of Bioko (Fernando Po) V. Acta Botanica Barcinonensia 40: 1-106. Dubuisson, J.-Y. 1996. Evolutionary relationships within the genus Trichomanes sensu lato (Hymenophyllaceae) based on anatomical and morphological characters and a comparison with rbcL necleotide sequences: preliminary results. In J. Camus et al. (Eds) Pteridology in perspective: 285-287. Kew. PTERIDOPHYTES OF SAO TOME AND PRINCIPE 65 Exell, A. 1944. Catalogue of the vascular plants of S. Tome (with Principe and Annobon). London. Exell, A. & Rozeira, A. 1958. Aditamento a flora das ilhas de Sao Tome e Principe. Conferencia International dos Africanistas Ocidentais 3: 77-91. Figueiredo, E. 1994. Diversity and endemism of angiosperms in the Gulf of Guinea islands. Biodiversity and Conservation 3: 785-793. Kramer, K. 1972. The Lindsaeoid ferns of the Old World-IX Africa and its islands. Bulletin du Jardin Botanique National de Belgique 42: 305-345. Kuhn, M. 1 868. Filices africanae. Lipsiae. Nooteboom, H.P. 1994. Notes on Davalliaceae. II. A revision of the genus Davallia. Blumea39: 151-214. Pichi Sermolli, R.E.G. 1969. Adumbratio Florae Aethiopicae. 16. Marattiaceae. Webbia 23: 329-351. 1985. On the taxonomy and nomenclature of some species of Ctenitis (Aspidiaceae) from Tropical Africa. Webbia 39: 1-28. 1991. On the taxonomy and nomenclature of some species from Tropical Africa of the genus Triplophyllum Holttum (Dryopteridaceae). Webbia 45: 117- 135. Tindale, M.D. 1 965. A monograph of the genus Lastreopsis Ching. Contributions from the New South Wales National Herbarium 3: 249-339. SYSTEMATIC INDEX Accepted names are in roman and synonyms in italic. Acrostichum aureum L. 42, 43 Adiantum mettenii Kuhn 43 Adiantum philippense L. 43 Adiantum raddianum C. Pres 42, 43 Adiantum soboliferum auct. 43 Adiantum vogelii Mett. ex Keyserl. 45 Alsophila camerooniana van camerooniana auct. 51 Alsophila manniana (Hook.) R.M. Tryon 5 1 Alsophila welwitschil (Hook.) R.M. Tryon 51 Anapeltis lycopodioides van owariensis (Desv.) Benl 59 Anisosorus occidentalis (Baker) C. Chr. 52 Anthrophyum immersum (Bory ex Willd.) Mett. ex Hook. 63 Anthrophyum mannianum Hook. 63 Arthropteris monocarpa (Cordem.) C. Chr. 58 Arthropteris obllterata auct. 58 Arthropteris orientalis (J.F. Gmel.) Posth. 58 Arthropteris palisotii (Desv.) Alston 58 Aspidium fraternum Mett. ex Kuhn 47 Aspidium nigritlanum Mett. ex Kuhn 46 Aspidium subquinquefidum P. Beauv. 47 Asplenium aethiopicum (Burm.f.) Bech. 48 Asplenium africanum Desv. 48 Asplenium anisophyllum Kunze 48 Asplenium barteri Hook. 48, 50 Asplenium biafranum Alston & F. Ballard 48 Asplenium cuneatum Lam. 48 Asplenium currorii Hook. 48 Asplenium dregeanum subsp. brachypterum (Kunze ex Houlston & T. Moore) Pic. Serm. 48 Asplenium emarginatum P. Beauv. 48 Asplenium erectum var. usambarense (Hieron.) Schelpe 48 Asplenium eurysorum Hieron. 49, 64 Asplenium exhaustum (H. Christ) Alston 49 Asplenium formosum Willd. 49 Asplenium friesiorum C. Chr. 42, 49 Asplenium geppii Carruth. 48 Asplenium hemitomum Hieron. 64 Asplenium hypomelas Kuhn 49 Asplenium inaequilaterale Bory ex Willd. 49 Asplenium lividum Mett. ex Kuhn 49 Asplenium longicauda Hook. 49 Asplenium megalura var. molleri (Hieron.) Tardieu 49 Asplenium molleri Hieron. 49 Asplenium nigritianum Hook. 49, 50 Asplenium paucijugum F. Ballard 50 Asplenium protensum auct. 49 Asplenium quintasil Gand. 49 Asplenium sandersonii Hook. 50 Asplenium unilaterale Lam. 50 Asplenium variabile Hook. 50 Asplenium variabile var. paucijugum (F. Ballard) Alston 50 Athyrium newtonii (Baker) Diels 50 Belvisia spicata (L.f.) Mirb. ex Copel. 59 Blechnum attenuatum (Sw.) Mett. 51 Blotiella currorii (Hook.) R.M. Tryon 51 Blotiella glabra (Bory) R.M. Tryon 52 Blotiella mannii (Baker) Pic. Serm. 52 Bolbitis acrostichoides (Afzel. ex Sw.) Ching 56 Bolbitis auriculata (Lam.) Ching 56 Bolbitis fluviatilis (Hook.) Ching 56 Callipteris prolifera (Lam.) Bory 50 Ceratopteris cornuta (P. Beauv.) Lepr. 64 Christella dentata (Forssk.) Brownsey & Jermy 62 Christella hispidula (Decne.) Holttum 62 Coniogramme africana Hieron. 54 Ctenitis attenuata Pic. Serm. 47 Ctenitis buchholzii (Kuhn) Alston 47 Ctenitis cirrhosa (Schum.) Ching 42, 45 Ctenitis fraterna (Mett.) Tardieu 47 Ctenitis lanuginosa (Willd. ex Kaulf.) Copel. 45 Ctenitis nigritiana (Mett. ex Kuhn) Alston 46 Cyathea camerooniana var. currorii Holttum 5 1 Cyathea manniana Hook. 51 Cyathea welwitschii Hook. 5 1 Cyclophorus spissus var. continentalis Hieron. ex Engl. 60 Cyclosorus afer (H. Christ) Ching 63 Cyclosorus elatus (Mett. ex Kuhn) Alston 63 Cyclosorus oppositifolius (Hook.) Tardieu 63 Cyclosorus striatus (Schum.) Ching 62 Davallia chaerophylloides (Poir.) Steud. 51 Davallia denticulata var. denticulata (Burm.f.) Mett. ex Kuhn 51 Davallia repens (L.f.) Kuhn 51 Dicranopteris linearis (Burm.f.) Underw. 53 Didymochlaena truncatula (Sw.) J. Sm. 45 Diplazium arborescens (Bory) Sw. 50 Diplazium proliferum (Lam.) Thouars 50 Drynaria laurentii (H. Christ ex De Wild. & T. Durand) Hieron. 59 Drynaria volkensii Hieron. 59 Dryopteris crinobulbon (Hook.) C. Chr. 45 Dryopteris dentata (Forssk.) C. Chr. 62 Dryopteris fragilis (Baker) C. Chr. 62 Dryopteris henriquesii (Baker) C. Chr. 63 Dryopteris lanuginosa (Willd. ex Kaulf.) C. Chr. 45 Dryopteris manniana (Hook.) C. Chr. 64 Dryopteris nigritiana (Mett. ex Kuhn) Kuntze 46 Dryopteris oligantha auct. 45 Dryopteris oppositifolia auct. 63 Dryopteris oppositifolia (Hook.) C. Chr. 63 Dryopteris pentheri (Krasser) C. Chr. 45 Dryopteris protensa (Afzel. ex Sw.) C. Chr. 47 Dryopteris quadrangularis (Fee) Alston 62 Dryopteris securidiformis (Hook.) C. Chr. 47 Dryopteris securidiformis var. nana Bonap. 47 Dryopteris striata (Schum.) C. Chr. 62 Dryopteris variabilis var. barteri (Hook.) Alston 47 Dryopteris venulosa Kuntze 63 Elaphoglossum acrostichoides (Hook. & Grev.) Schelpe 52 Elaphoglossum aubertii (Desv.) T. Moore 53 Elaphoglossum chevalieri H. Christ 53 Elaphoglossum clarenceanum auct. 53 Elaphoglossum isabelense Brause 53 Elaphoglossum petiolatum subsp. salicifolium (Willd. ex Kaulf.) Schelpe 53 Elaphoglossum salicifolium (Willd. ex Kaulf.) Alston 53 Gleichenia linearis (Burm.f.) C.B. Clarke 53 Gonocormus mannii (Hook, ex Hook. & Baker) G. Kunkel 55 Grammitis ebenina (Maxon) Tardieu 53 Grammitis molleri (Baker) Schelpe 42, 53 Grammitis nigrocincta Alston 42, 53 Grammitis sp. 53 Grammitis tomensis Schelpe 42, 54 Histiopteris incisa (Thunb.) J. Sm. 52 Humata repens (L.f.) Diels 51 Huperzia mildbraedii (Herter) Pic. Serm. 57 Huperzia ophioglossoides (Lam.) Rothm. 42, 57 Huperzia phlegmaria (L.) Rothm. 57 Huperzia verticillata (L.f.) Trevis. 57 Huperzia warneckei (Herter ex Nessel) Pic. Serm. 57 Hymenolepis spicata (L.f.) C. Presl 59 Hymenophyllum capillare Desv. 54 Hymenophyllum ciliatum var. boryanum (Willd.) Mett. ex Kuhn 54 Hymenophyllum hirsutum (L.) Sw. 54 Hymenophyllum kuhnii C. Chr. 54 Hymenophyllum mildbraedii (Brause ex Brause & Hieron.) Alston 55 Hymenophyllum polyanthos var. kuhnii (C. Chr.) Schelpe 54 Hymenophyllum polyanthos var. mildbraedii (Brause ex Brause & Hieron.) Schelpe 42, 55 Hymenophyllum splendidum Bosch 55 Hypolepis sparsisora (Schrad.) Kuhn 52 Lastreopsis aff. currori (Mett. ex Kuhn) Tindale 46 Lastreopsis nigritiana (Baker) Tindale 64 Lomariopsis guineensis (Underw.) Alston 56 Lomariopsis hederacea Alston 56 Lomariopsis hederacea auct. 56 Lomariopsis nigrescens Holttum 56 Lomariopsis warneckei (Hieron.) Alston 42, 56 Lonchitis currorii (Hook.) Mett. ex Kuhn 51 Lonchitis glabra Bory 52 Lonchitis gracilis Alston 52 Lonchitis mannii (Baker) Alston 52 Lonchitis occidentalis Baker 52 Loxogramme lanceolata (Sw.) C. Presl 56 Lycopodiella cernua (L.) Pic. Serm. 57 Lycopodium cemuum L. 57 Lycopodium clavatum L. 42, 57 Lycopodium mildbraedii Herter 57 Lycopodium phlegmaria L. 57 Lycopodium verticillatum L.f. 57 Lycopodium warneckei (Herter ex Nessel) Alston 57 Lygodium microphyllum (Cav.) R. Br. 64 Lygodium scandens auct. 64 Macrothelypteris henriquesii (Baker) Pic. Serm. 63 Marattia fraxinea Sm. 57, 58 Marattia robusta Alston 57, 58 Metathelypteris fragilis subsp. guineensis Benl 62 Microgonium erosum (Willd.) C. Presl 55 66 E. FIGUEIREDO Microgramma owariensis (Desv.) Alston 59 Microlepia speluncae (L.) T. Moore 42, 52 Microsorum punctatum (L.) Copel. 59 Microtrichomanes digitatum (Sw.) Copel. 55 Nephmdium subquinquefidum var. elongatum Hook. 47 Nephrolepis biserrata (Sw.) Schott 58 Nephrolepis pumicicola F. Ballard 58 Nephrolepis undulata (Afzel. ex Sw.) J. Sm. 58 Odontosoria chinensis var. divaricata H. Christ 52 Oleandra annetii auct. 58 Oleandra annetii Tardieu 59 Oleandra distenta Kunze 58 Ophioglossum costatum R. Br. 59 Ophioglossum gramineum Willd. 64 Ophioglossum reticulatum L. 59 Pellaea doniana Hook. 62 Phymatosorus scolopendria (Burm.f.) Pic. Serm. 60 Pityrogramma calomelanos var. calomelanos (L.) Link 42, 54 Platycerium stemaria (P. Beauv.) Desv. 60 Pleopeltis excavata (Bory ex Willd.) Sledge 60 Pleopeltis macrocarpa (Bory ex Willd.) Kaulf. 60 Pneumatopteris afra (C. Chr.) Holttum 63 Pneumatopteris oppositifolia (Hook.) Holttum 63 Pneumatopteris venulosa (Kuntze) Holttum 63 Polypodium cultratum var. elasticum (Bory ex Willd.) Baker 54 Polypodium ebeninum Maxon 53 Pol\podium henriquesii Baker 63 Polypodium irioides Poir. 59 Polypodium lycopodioides auct. 59 Polypodium molleri Baker 53 Polypodium nigritianum Baker 64 Polypodium oosorum Baker 54 Polypodium phymatodes L. 60 Polypodium preussii Hieron. 60 Polypodium villosissimum Hook. 54 Pseudophegopteris henriquesii (Baker) Holttum 63 Psilotum nudum (L.) P. Beauv. 60 Pteridium aquilinum (L.) Kuhn 52 Pteris atrovirens Willd. 60, 61 Pteris atmvirens auct. 61 Pteris biaurita auct. 61 Pteris burtonii Baker 60, 61 Pteris catoptera auct. 61 Pteris communata auct. 6 1 Pteris dentata subsp. flabellata (Thunb.) Runemark 61 Pteris linearis Poir. 61 Pteris paucipinnata Alston 61 Pteris pteridioides (Hook.) F. Ballard 61 Pteris quadriaurita auct. 61 Pteris similis Kuhn 61 Pteris togoensis Hieron. 61 Pteris tripartita Sw. 54, 61 Pteris vittata L. 42, 61 Pyrrosia lanceolata (L.) Farw. 60 Selaginella cathedrifolia Spring 61 Selaginella mannii Baker 61 Selaginella molleri Hieron. 62 Selaginella molliceps Spring 62 Selaginella monodii Alston 42, 62 Selaginella myosurus (Sw.) Alston 62 Selaginella squarrosa Baker 42, 62 Selaginella thomensis Alston 62 Selenodesmium rigidum (Sw.) Copel. 56 Sphaerostephanos elatus subsp. thomensis Holttum 42, 63 Sphenomeris chinensis var. divaricata (H. Christ) Kramer 52 Sphenomeris chusana var. divaricata (H. Christ) Tardieu 52 Stenoloma chinense var. divaricatum (H. Christ) Alston 52 Tectaria angelicifolia (Schum.) Copel. 46 Tectaria buchholzii (Kuhn) Copel. 47 Tectaria camerooniana (Hook.) Alston 46 Tectaria fernandensis (Baker) C. Chr. 47 Thelypteris henriquesii (Baker) Tardieu 63 Thelypteris mauritiana subsp. thomensis (Holttum) Viane 63 Trichomanes africanum H. Christ 64 Trichomanes borbonicum Bosch 55 Trichomanes chamaedrys Taton 55 Trichomanes crispiforme Alston 55 Trichomanes cupressoides Desv. 56 Trichomanes digitatum Sw. 55 Trichomanes erosum Willd. 55 Trichomanes giganteum auct. 55 Trichomanes mannii Hook, ex Hook. & Baker 55 Trichomanes melanotrichum Schltdl. 55 Trichomanes pyxidiferum var. melanotrichum (Schltdl.) Schelpe 55 Trichomanes radicans Sw. 55 Trichomanes rigidum Sw. 56 Triplophyllum attenuatum (Pic. Serm.) Pic. Serm. 47 Triplophyllum buchholzii (Kuhn) Holttum 47 Triplophyllum fraternum var. elongatum (Hook.) Holttum 47 Triplophyllum heudelotii Pic. Serm. 47 Triplophyllum jenseniae (C. Chr.) Holttum 42, 47 Triplophyllum principis Holttum 47 Triplophyllum protensum (Afzel. ex Sw.) Holttum 47 Triplophyllum securidiforme (Hook.) Holttum 47 Triplophyllum securidiforme var. nanum (Bonap.) Holttum 47 Triplophyllum subquinquefidum (P. Beauv.) Pic. Serm. 47 Vandenboschia africana (H. Christ) G. Kunkel 64 Vandenboschia borbonica (Bosch) G. Kunkel 55 Vandenboschia pyxidifera (L.) Copel. 55 Vandenboschia radicans (Sw.) Copel. 55 Vittaria guineensis Desv. 63 Vittaria owariensis Fee 64 Xiphopteris cultrata (Willd.) Schelpe 54 Xiphopteris oosora (Baker) Alston 42, 54 Xiphopteris villosissima (Hook.) Alston 54 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 2 No. 1 New Himalayan species of Pedicularis with special reference to those from the eastern Himalaya. P.C. Tsoong. 1955. Pp. 1- 34. Facsimile edition. £4.35 No. 2 Mosses of Dominica, British West Indies. E.B. Bartram. 1955. Pp. 37-49. Mosses of the Ecuadorian Andes collected by P.R. Bell. E.B. Bartram. 1955. Pp. 51-64. Facsimile edition. £4.00 No. 3 Novitates Himalaicae-1. F. Ludlow & W.T. Stearn. 1956. Pp. 65-81, 8 plates, 1 1 figs. Facsimile edition. £5.25 No. 4 Saxifraga of the Himalaya 1. Section Kabschia. H. Smith. 1958. Pp. 83-129, 14 figs. £9.65 No. 5 The Polypodiaceae and Grammitidaceae of Ceylon. W.A. Sledge. 1960. Pp. 131-158, 4 figs. Facsimile edition. £3.90 No. 6 Allium and Milula in the central and eastern Himalaya. W.T. Stearn. 1960. Pp. 159-191, 4 plates, 10 figs. Facsimile edition. £4.35 No. 7 The identity of Isopyrum aquilegioides. L.G. de Beer & W.T. Stearn. 1960. Pp. 193-202, 3 figs. Facsimile edition. £3.15 No. 8 On the geographical relationships of the angiosperm flora of New Guinea. R. Good. 1960. Pp. 203-226, 1 fig. Facsimile edition. £3.60 No. 9 Saxifraga of the Himalaya II. Some new species. H. Smith. 1960. Pp. 227-260. 9 plates, 17 figs. Facsimile edition. £5.65 No. 10 New species of Taraxacum from the Himalayan region. J.L. van Soest. 1961. Pp. 261-273, 8 plates. Facsimile edition. £6.15 No. 1 1 The athyriod ferns of Ceylon. W.A. Sledge. 1962. Pp. 275-323. 3 plates. Facsimile edition. £5.25 No. 12 The genus Epilobium in the Himalayan region. PH. Raven. 1962. Pp. 325-382, 7 plates, 13 figs. Facsimile edition. £5.90 Volume 3 No. 1 A revision of the genera Buchenavia and Ramatuella. A.W. Exell & C.A. Stace. 1963. Pp. 1-46, 5 figs. Facsimile edition. £5.15 No. 2 The diatom genus Capartogramma and the identity of Schizostauron. R. Ross. 1963. Pp. 47-92, 2 plates, 38 figs. Facsimile edition. £5.15 No. 3 Angiosperms of the Cambridge Annobon Island Expedition. A.W. Exell. 1963. Pp. 93-1 18, 10 plates. Facsimile edition. £3.75 No. 4 A revision of the genus Petrorhagia. P.W. Ball & V.H. Hey wood. 1964. Pp. 1 19-172, 3 plates, 22 figs. Facsimile edition. £4.90 No. 5 Marine algae of Gough Island. Y.M. Chamberlain. 1965. Pp. 173-232, 4 plates, 80 figs. £12.50 No. 6 The Ceylon species ofAsplenium. W.A. Sledge. 1965. Pp. 233- 277, 1 plate, 3 figs. Facsimile edition. £5.15 Volume 4 No. 1 Cuticular studies as an aid to plant taxonomy. C.A. Stace. 1965. Pp. 1-78, 5 plates, 10 figs. Facsimile edition. £7.20 No. 2 The genus Elaphoglossum in the Indian peninsula and Ceylon. W.A. Sledge. 1967. Pp. 79-96. Facsimile edition. £3.25 No. 3 Fungi of recent Nepal expeditions. F.L. Balfour-Browne. 1968. Pp. 97-141, 4 figs. Facsimile edition. £3.75 No. 4 No. 5 No. 6 No. 7 A synopsis of Jamaican Myrsinaceae. W.T. Stearn. 1969. Pp. 143-178, 8 plates, 25 figs. £4.55 The Jamaican species of Columnea and Alloplectus (Gesneriaceae). W.T.Stearn. 1969. Pp. 179-236, 8 plates, 29 figs. £6.40 New or little known Himalayan species of Swertia and Veratrilla (Gentianaceae). H. Smith. 1970. Pp. 237-258, 16 plates, 7 figs. £8.25 A survey of the tropical genera Oplonia and Psilanthele (Acanthaceae). W.T. Stearn. 1971. Pp. 259-323, 10 plates, 18 figs. £12.40 Angiosperms of the islands of the Gulf of Guinea (Fernando Po, Principe, S. Tome, and Annobon). A.W. Exell. 1973. Pp. 325-411. £12.50 Volume 5 No. 1 The dryopteroid ferns of Ceylon. W.A. Sledge. 1973. Pp. 1-43, 4 figs. £6.90 No. 2 New Himalayan and Tibetan species of Corydalis (Papaveraceae). F. Ludlow & W.T. Stearn. 1975. Pp. 45-69, 15 plates, 14 figs. £8.40 No. 3 The marine algae of Trinidad, West Indies. W.D. Richardson. 1975. Pp. 71-143, 12 plates, 2 figs. £13.45 No. 4 A revision of the Macaronesian genus Argyranthemum Webb ex Schultz Bip. (Compositae-Anthemideae).C.J. Humphries. 1976. Pp. 145-240, 2 plates, 26 figs. £14.20 No. 5 Frank Ludlow (1885-1972) and the Ludlow-Sherriff expedi- tions to Bhutan and south-eastern Tibet of 1933-1950. W.T. Stearn. 1976. Pp. 243-268, 1 fig. Reliquiae botanicae himalaicae. F. Ludlow. 1976. Pp. 269-289, 7 plates, 8 figs. Facsimile edition. £11.10 No. 6 Studies in the genus Hypericum L. (Guttiferae). 1 . Infrageneric classification. N.K.B. Robson. 1977. Pp. 291-355, 9 figs. £14.20 No. 7 Sphagnales of tropical Asia. A.Eddy. 1977. Pp. 357^45, 4 plates, 17 maps, 25 figs. £17.80 Volume 6 No. 1 The handwriting of Joseph Banks, his scientific staff and amanuenses. J.B. Marshall. 1978. Pp. 1-85, 62 figs. £18.30 No. 2 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. II. Phaeophyta. J.H. Price, D.M. John & G.W. Lawson. 1978. Pp. 87-182, 1 fig. £24.40 No. 3 The lichenicolous Hyphomycetes. D.L. Hawksworth. 1979. Pp. 183-300, 47 figs. £24.40 No. 4 The species of Chisocheton (Meliaceae). D.J. Mabberley. 1979. Pp. 301-386, 3 plates, 10 figs. £24.40 Volume 7 No. 1 The distribution of Padina pavonica (L.) Lamour. (Phaeophyta: Dictyotales) on British and adjacent European shores. J.H. Price, I. Tittley & W.D. Richardson. 1979. Pp. 1-67, 3 plates, 2 figs. £17.40 No. 2 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. 111. Rhodophyta (Bangiophyceae). D.M. John, J.H. Price, C.A. Maggs, G.W. Lawson. 1979. Pp. 69-82, 1 fig. £5.40 No. 3 A revision of the genus Anacyclus L. (Compositae: Anthemi- deae). C.J. Humphries. 1979. Pp. 83-142, 27 figs. £15.60 Volume 8 No. 1 The Thelypteridaceae of Ceylon. W.A. Sledge. Pp. 1-54, 5 figs. 1981. £15.15 No. 2 Studies in the genus Hypericum L. (Guttiferae) 2. Characters of the genus. N.K.B. Robson. 1981. Pp. 55-226, 73 figs. £33.55 No. 3 A revision of the lichen family Thelotremataceae in Sri Lanka. M.E. Hale, Jr. 1981. Pp. 227-332, 20 figs. £24.80 No. 4 Vascular plant collections from the Tristan da Cunha group of islands. E.W. Groves. Pp. 333^20, 33 figs. £21.40 Volume 9 No. 1 The lichenicolous Coelomycetes. D.L. Hawksworth. 1981. Pp. 1-98, 36 figs. £22.70 No. 2 The genus Callithamnion (Rhodophyta: Ceramiaceae) in the British Isles. PS. Dixon & J.H. Price. 1981. Pp. 99-141,5 figs. £12.50 No. 3 Parmelia subgenus Amphigymnia (lichens) in East Africa. H. Krog & T.F.V. Swinscow. 1981. Pp. 143-231, 31 figs. £21.05 No. 4 The genus Selaginella in tropical South America. A.H.G. Alston, A.C. Jermy & J.M. Rankin. 1981. Pp. 233-330, 18 figs. £23.05 Volume 10 No. 1 Taxonomic studies in the Labiatae tribe Pogostemoneae. J.R. Press. 1982. Pp. 1-89, 33 figs. £21.85 No. 2 The typification of Hudson's algae: a taxonomic and nomenclatural reappraisal. L.M. Irvine & PS. Dixon. 1982. Pp. 91-105. £5.40 No. 3 Seaweeds of the Faroes. Various authors. 1982. Pp. 107-225, 13 figs. £27.15 No. 4 The lichen genus Steinera. A.M. Henssen & P.W. James. 1982. Pp. 227-256, 24 figs. £9.70 Volume 11 No. 1 The algae of Lightfoot's Flora scotica. PS. Dixon. 1983. Pp. 1-15, 2 figs. £5.55 No. 2 A taxonomic study of the lichen genus Micarea in Europe. B.J. Coppins. 1983. Pp. 17-214, 57 figs, 28 maps. £37.75 No. 3 The hepatics of Sierra Leone and Ghana. E.W. Jones & A.J. Harrington. 1983. Pp. 215-289, 8 figs. £18.15 No. 4 Studies in the Corallinaceae with special reference to Fosliella and Pneophyllum in the British Isles. Y.M. Chamberlain. 1983. Pp. 291^63, 89 figs. £33.45 Volume 12 No. 1 A revision of the Morinaceae (Magnoliophyta-Dipsacales). M.J. Cannon & J.F.M. Cannon. 1984. Pp. 1-35, 9 figs. £11.40 No. 2 An introduction to fern genera of the Indian subcontinent. C.R. Fraser- Jenkins. 1984. Pp. 37-76, 1 fig. £12.50 No. 3 A revision of African Sphagnales. A. Eddy. 1985. Pp. 77-162, 47 figs. £23.05 No. 4 Studies in the genus Hypericum L. (Guttiferae) 3. Sections 1. Campylosporus to 6a. Umbraculoides. N.K.B. Robson. 1985. Pp. 163-325, 24 figs, 34 maps. £39.20 Volume 13 No. 1 The lichen genus Usnea subgenus Neuropogon. F.J. Walker. 1985. Pp. 1-130, 39 figs. £31.85 No. 2 Cytotaxonomic studies of the ferns of Trinidad. A.C. Jermy & T.G. Walker. 1985. Pp. 131-276, 69 figs. £31.50 No. 3 Some genera of the Biddulphiaceae (diatoms) with interlocking linking spines. R. Ross & PA. Sims. 1985. Pp. 277-381, 33 plates. £28.75 Volume 14 No. 1 Cytological observations on Indian subcontinent and Chinese Dryopteris and Polystichum (Pteridophyta: Dryopteridaceae). M. Gibby. 1985. Pp. 1^2, 78 figs. £12.50 No. 2 A redisposition of the species referred to the ascomycete genus Microthelia. D.L. Hawksworth. 1985. Pp. 43-181, 73 figs. £34.25 No. 3 A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae). C.R. Fraser- Jenkins. 1986. Pp. 183-218, 4 figs. £11.75 No. 4 Evolutionary cladistics of marattialean ferns. C.R. Hill & J.M. Camus. 1986. Pp. 219-300, 27 figs. £24.65 Volume 15 No. 1 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 1. Genera A-F. J.H. Price, D.M. John & G.W. Lawson. 1986. Pp. 1-122, 1 fig. £33.60 No. 2 Cytology of the fern flora of Madeira. I. Manton, J.D. Lovis, G. Vida & M. Gibby. 1986. Pp. 123-161, 12 plates. £14.70 No. 3 A revision of the lichen genus Xanthoparmelia in Australasia. J.A. Elix, J. Johnston & P.M. Armstrong. 1986. Pp. 163-362, 42 figs, 117 maps. £42.90 Volume 16 No. 1 Studies in the genus Hypericum L. (Guttiferae) 7. Section 29. Brathys (part 1). N.K.B. Robson. 1987. Pp. 1-106, 14 figs, 25 maps. £28.75 No. 2 The lichen genus Ramalina in Australia. G.N. Stevens. 1987. Pp. 107-233, 15 plates, 31 figs. £32.20 No. 3 An annotated list of vascular plants collected in the valleys south of Mt Everest. G. Miehe. 1987. Pp. 225-268, 4 figs. £16.50 No. 4 Further genera of the Biddulphiaceae (diatoms) with interlock- ing linking spines. R. Ross & A.Sims. 1987. Pp. 269-31 1, 13 plates. £15.60 Volume 17 No. 1 Studies in Pseudocyphellaria (lichens) 1 . The New Zealand species. D.J. Galloway. 1988. Pp. 1-267, 124 figs. £55.65 Volume 18 No. 1 An illustrated catalogue of the type specimens in the Greville diatom herbarium. D.M. Williams. 1988. Pp. 1-148, 74 plates. £38.00 No. 2 Erik Acharius and his influence on English lichenology. D.J. Galloway. 1988. Pp. 149-194, 18 figs. £17.05 No. 3 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 2. Genera G. J.H. Price, D.M. John & G.W. Lawson. 1988. Pp. 195-273, 1 fig. £25.10 No. 4 Some Cretaceous and Palaeogene Trinacria (diatom) species. PA. Sims & R. Ross. 1988. Pp. 275-322, 13 plates. £17.55 No. 5 A monograph of Dryopteris (Pteridophyta: Dryopteridaceae) in the Indian subcontinent. C.R. Fraser- Jenkins. 1989. Pp. 323- 477, 79 figs. £34.10 No. 6 Corydalis (Papaveraceae: Fumarioideae) in Nepal. M. Liden. 1 989. Pp. 479-538, 26 figs. £19.35 Volume 19 A new species of Maytenus (Celastraceae) in Ethiopia. Sebsebe Demissew. 1989. Pp. 1-3, 1 fig. Central American Araliaceae - a precursory study for the Flora Mesoamericana. M.J. Cannon & J.F.M.Cannon. 1989. Pp. 5- 6 1,36 figs. A revision of the Solarium nitidum group (section Holophylla pro parte): Solanaceae. S. Knapp. 1989. Pp. 63-102, 21 figs. Six new species of Solatium sect. Geminata from South America. S. Knapp. 1989. Pp. 103-1 12, 8 figs. The application of names of some Indian species of Ocimum and Geniosporum (Labiatae). J.R. Press & V.V. Sivarajan. 1989. Pp. 11 3- 11 6, 4 figs. Revision of Piper (Piperaceae) in the New World 1 . Review of characters and taxonomy of Piper section Macrostachys. M.C. Tebbs. 1989. Pp. 1 17-158, 41 figs. Facsimile edition. £52.45 Volume 20 No. 1 Studies in the genus Hypericum L. (Guttiferae) 8. Sections 29. Brathys (part 2) and 30. Trigynobrathys. N.K.B. Robson. 1990. Pp. 1-151, 22 figs, 46 maps. Facsimile edition. £49.25 No. 2 The marine algal flora of Namibia: its distributions and affinities. G.W. Lawson, R.H. Simons and W.E. Isaac. 1990. Pp. 153-168, 1 fig, 7 plates. The infrageneric classification of Gentiana (Gentianaceae). T- N. Ho and S.-W. Liu. 1990. Pp. 169-192, 13 figs. Revision of Piper (Piperaceae) in the New World. 2. The taxonomy of Piper section Churumayu. M.C. Tebbs. 1990. Pp. 193-236, 49 figs. £34.10 Volume 21 No. 1 Historical and taxonomic studies in the genus Titanoderma (Rhodophyta, Corallinales) in the British Isles. Y.M. Chamber- lain. 1991. Pp. 1-80, 247 figs. £42.35 No. 2 Early collections of the Holy Thorn (Crataegus monogyna cv. Biflora). A.R. Vickery. 1991. Pp. 81-83, 1 fig. A taxonomic study of the species referred to the ascomycete genus Leptorhaphis. B. Aguirre-Hudson. 1991 . Pp. 85-192, 76 figs. The typification and identification of Calymperes crassilimbatum Renauld & Cardot (Musci: Calymperaceae). L.T. Ellis. 1991. Pp. 193-194, 1 fig. £42.35 Volume 22 No. 1 An account of southern Australian species of Lithophyllum (Corallinaceae, Rhodophyta). Wm. J. Woelkerling and S.J. Campbell. 1992. Pp. 1-107, 63 figs. £41.25 No. 2 Palynological evidence for the generic delimitation of Sechium (Cucurbitaceae) and its allies. J.L. Alvarado, R. Lira-Saade & J. Caballero. 1992. Pp. 109-121. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 3. Genera H-K. J.H. Price, D.M. John & G.W. Lawson. 1992. pp. 123-146. Two new species of Solarium section Geminata (Solanaceae) from Cerro del Torr in western Colombia. S. Knapp. 1992. Pp. 147-152. Fissidens ceylonensis Dozy & Molkenb. (Musci: Fissidentaceae) and some allied taxa from southern India. L.T. Ellis. 1992. Pp. 153-156, 2 figs. New species of Piper (Piperaceae) from Central America. M. Tebbs. 1992. Pp. 157-158. Studies on the Cretan flora 1 . Floristic notes. N.J. Turland. 1992. Pp. 159-164. Studies on the Cretan flora 2. The Dianthus juniperinus complex (Caryophyllaceae). N.J. Turland. 1992. Pp. 165-169. £41.25 Volume 23 No. 1 Revision of Piper (Piperaceae) in the New World 3. The taxonomy of Piper sections Lepianthes and Radula. M.C. Tebbs. 1993. Pp. 1-50, 18 figs. Mounting techniques for the preservation and analysis of diatoms. S.J. Russell. 1993. Pp. 51-54. 1 fig. £43.25 No. 2 New taxa of Gentiana (Gentianaceae) from Western China and the Himalayan region. T.-N. Ho and S.-W. Liu. 1993. Pp. 55- 60. 2 figs. New combinations, names and taxonomic notes on Gentianella (Gentianaceae) from South America and New Zealand. T.-N. Ho and S.-W. Liu. 1993. Pp. 61-66. Studies in Hypericum: validation of new names. N.K.B. Robson. 1993. Pp. 67-70. Generic monograph of the Asteraceae-Anthemideae. K. Bremer and C.J. Humphries. 1993. Pp. 71-177. 12 figs. £43.25 Volume 24 No. 1 Pre-Linnaean references for the Macaronesian flora found in Leonard Plukenet's works and collections. J. Francisco-Ortega, A. Santos-Guerra and C.E. Jarvis. Pp. 1-34. Studies on the lichen genus Sticta (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. 9 1-100. £43.25 No. 2 Observations on the benthic marine algal flora of South Georgia: a floristic and ecological analysis. D.M. John, P.J.A. Pugh and I. Tittley. Pp. 101-1 14. Studies in Pseudocyphellaria (Lichens) IV. Palaeotropical species (excluding Australia). D.J. Galloway. Pp. 1 15-160. Morphology and ecology of seedlings, fruits and seeds of Panama: Bixaceae and Cochlospermaceae. N.C. Garwood. Pp. 161-172. A study of Bixa (Bixaceae), with particular reference to the leaf undersurface indumentum as a diagnostic character. R.E. Dempsey and N.C. Garwood. Pp. 173-180. £43.40 Volume 25 No. 1 A revision of Rutilaria Greville (Bacillariophyta). R. Ross. Pp. 1-94. William Roxburgh's St Helena plants. Q.C.B. Cronk. Pp. 95-98. £43.40 No. 2 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 5. Genera P. G.W. Lawson, W.J. Woelkerling, J.H. Price, W.F. Prud'homme Van Reine and D.M. John. Pp. 99-122. A new species of Odontorrhynchos (Orchidaceae, Spiranthinae) from Boliva. D.L. Szlachetko. Pp. 123-125. Linnaeus's interpretation of Prospero Alpino's De plantis exoticis, with special emphasis on the flora of Crete. N.J. Turland. Pp. 127-159. Book review. M.G. Gilbert. P. 1 6 1 . £43.40 Volume 26 No. 1 A morphological study of Chaetoceros species (Bacillariophyta) from the plankton of the Pacific ocean of Mexico. D.U. Hernandez-Becerril. 1996. Pp. 1-73. £43.40 No. 2 Studies in the genus Hypericum L. (Guttiferae) 6. Sections 20. Myriandra to 28. Elodes. N.K.B. Robson. 1996. Pp. 75-217. £43.40 Volume 27 No. 1 Notes on the diatom species Tetracyclus castellum (Ehrenb.) Grunow with a description of Tetracyclus pseudocastellum nov. sp. D.M. Williams. Pp. 1-5. A new species of Calymperes (Musci: Calymperaceae) from Peninsular Malaysia. L.T. Ellis. Pp. 7-9. A phylogenetic conspectus of the tribe Hyoscyameae (Solanaceae). A.L. Hoare and S. Knapp. Pp. 1 1-29. A revision of Solanum section Ptemidea: Solanaceae. S. Knapp and T. Helgason. Pp. 3 1-73. 1997. £43.40 No. 2 Systematics of Pogostemon (Labiatae) G.R. Bhatti and M. Ingrouille. Pp. 77-147. 1997 £43.40 CONTENTS 1 Morphology and ecology of seedlings, fruits and seeds of Panama: Vochysiaceae Nancy C. Garwood 17 A revision of the genus Mandragora (Solanaceae) S. Ungricht, S. Knapp and J.R. Press 41 The pteridophytes of Sao Tome and Principe (Gulf of Guinea) Estrela Figueiredo tural History Musei BOTANY SERIES Vol.28, No. 1, June 1998