ISSN 0968-0446 Bulletin of The Natural History Museum Botany Series THE NATURAL HISTORY MUSEUM VOLUME 23 NUMBER 1 24 JUNE 1993 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History)), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Dr S. Blackmore Editor of Bulletin: Dr R. Huxley Assistant Editor: Mrs M.J. West Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. 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The taxonomy of Piper sections Lepianthes and Radula BRITISH MUSEUM (NATURAL HISTORY) MARGARET CECILIA TEBBS " / J U L 1993 c/o Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 SAB PRESENTED GENERAL LIBRARY CONTENTS Introduction 2 Section Lepianthes Raf 2 Distribution and habitat 2 Morphology 3 Taxonomic treatment 3 Key to the species in section Lepianthes 3 54. Piper umbellatum L 3 55 . P. peltatum L 3 Section Radula Miq 5 Distribution and habitat 5 Morphology 5 Taxonomic treatment 6 Key to the species in section Radula 6 56. Piper marginatum Jacq 7 57. P. cinereumC.DC 10 58. P. multiplinervium C.DC 10 59. P. holdridgeianumW.C. Burger 10 60. P. subflavumC.DC 10 61 . P. via-chicoense Yunck 12 62. P. pseudofuligineum C.DC 12 63. P. bredemeyeri Jacq 13 64. P. dilatatum Rich 13 65. P. thomasiiTebbs 15 66. P. terrabanum C.DC 15 67. P. littorale C.DC 17 68. P. otophorum C.DC 17 69. P. malpasoensis Tebbs 17 70. P. enganyanum Trel. & Yunck 17 71. P. lanceifolium Kunth 19 72. P. aduncum L 19 73. P. mollicomum Kunth 21 74. P. achoteanum Trel 22 75. P. fuligineum Kunth 22 76. P. flavidum C.DC ex Donn.Sm 22 77. P. silvivagum C.DC 24 78. P. pseudoasperifolium C.DC 24 79. P. villiramulum C.DC 24 80. P. hispidum Sw 26 81. P. sancti-felicis Trel 29 82. P. poasanum C.DC 29 83. P. bisasperatum Trel 31 84. P. umbricola C.DC 31 85. P. chrysostachyum C.DC 32 86. P. epigynium C.DC 32 87. P. dotanumTrel 34 88. P. polytrichum C.DC 34 89. P. peracuminatum C.DC 34 90. P. zacatense C.DC 34 91. P. biauritumC.DC 36 M.C. TEBBS 92. P. perhispidum C.DC 36 93 . P. chamissonis (Miq . ) Steud 36 94. P. colonense C.DC 38 95. P. oblanceolatum Trel 38 96. P. hirtellipetiolum C.DC 38 97. P. decurrens C.DC 38 98. P. tenuimucronatumC.DC 40 99. P. carpinteranum C.DC 40 140. P. hostmannianum (Miq.) C.DC 40 101 . P. scalarispicum Trel 42 102. P. curvatipes Trel 42 103. P. jacquemontianum Kunth 42 104. P. biritakTrel 44 105. P. pseudolindenii C.DC 44 106. P. sanctum (Miq.) Schltdl 46 Excluded species 46 References 47 Index 47 SYNOPSIS. The paper presents a taxonomic treatment of Piper sections Lepianthes (formerly the genus Lepianthes) and Radula, and 53 species are recognized, keyed, and described. There are two new species in section Radula, Piper malpasoensis Tebbs and P. thomasii Tebbs. INTRODUCTION This third part of a revision of New World species of Piper deals with sections Lepianthes Raf. and Radula Miq. Section Lepianthes contains two herbaceous or shrubby species with inflorescences clustered together on a contracted, leafless stem, formally in the genus Lepianthes Raf. (Pothomorphe Miq.}. Section Radula contains species with prominent pro- phylls, petioles mostly with ligule-like structures at the base, minute, often ciliate floral bracts, and small, obovoid, trigo- nous or round fruits. Most of the species in this section are shrubs or subshrubs; the remainder are scandent or herba- ceous. DISTRIBUTION AND HABITAT Piper peltatum and P. umbellatum both originated from the New World, but have since become widespread throughout the tropics. Piper peltatum grows in moist places, usually in fairly open positions, such as road- or track sides, edges of pasture, waste ground, and stream sides. It is a lowland plant, generally occurring from sea-level to about 500 m, although it can be found above these heights. Piper umbellatum is a plant of shady clearings in moist forests, cloud forest, and river banks. It is usually found at altitudes over 800 m, and often up to 1600 m. SECTION LEPIANTHES RAF. There are only two species in this section, Piper umbellatum L. and P. peltatum L., separated from the other sections in Piper by their unusual stem development. These species were removed from Piper and placed in the genus Lepianthes by Rafinesque (1838), in Heckeria by Kunth (1839), and in Pothomorphe by Miquel (1844). Examination of the umbel- late inflorescences show that they are not on a common axillary peduncle, but a contracted leafless stem emerging from a sheathing petiole (Burger, 1977). This character is not considered sufficient to maintain Lepianthes as a separate genus, and therefore it has been placed into a section within Piper. Section Lepianthes shares characters with Piper auri- tum Kunth at the end of section Churumayu (Tebbs, 1990), such as large membranous leaves with sheathing petioles, triangular, pale-ciliate bracts, and glabrous, trigonous fruits. Piper marginatum Jacq. and other species in the early parts of section Radula also have large leaves, sheathing petioles, and similar infloresence characters. Section Lepianthes has there- fore been placed between sections Churumayu and Radula. MORPHOLOGY Section Lepianthes contains herbaceous or shrubby plants with large, strongly veined, peltate or cordate leaves with sheathing petioles. The infloresences are clustered together and arise at short intervals from a contracted, leafless stem; at the base of each peduncle are two prophylls, which at first enclose the young inflorescence (Blanc & Andraos, 1983). Inflorescences are slender and erect, with minute, white- ciliate floral bracts and glabrous, trigonous fruits. TAXONOMIC TREATMENT Section LEPIANTHES Raf. Type: Piper umbellatum L. Herbs or shrubs, stems glabrous to sparsely or densely pubescent. Leaves large, membranous, petioles sheathing the stems. Inflorescences clustered together, each one at first enclosed in two prophylls, on a leafless stem emerging from a PIPER IN THE NEW WORLD sheathing petiole. Floral parts densely congested; stamens 2; bracts white-ciliate; fruits obovoid, trigonous, glabrous; stig- mas 3. Open or shady areas, moist secondary forest, cloud forest, road- and stream sides, waste places; 0-2000 m. Key to the species in section Lepianthes la. Shrub or subshrubs; leaves cordate, not peltate 54. P. umbellatum Ib. Herbaceous or subshrubs; leaves ovate to suborbiculate, peltate 55. P. peltatum 54. Piper umbellatum L., Sp. pi. 1: 30 (1753). Type: Habitat in Domingo. Plumier. Fig. 1A, a. Lepianthes umbellata (L.) Raf., Sylva tellur.: 84 (1838). Heckeria umbellata (L.) Kunth in Linnaea 13: 566 (1839). Pothomorphe umbellata (L.) Miq., Comm. phytogr.: 36 (1840). Shrubs or subshrubs, 1-3 m high; stems glabrous to sparsely or densely pubescent, glandular. Leaves 20-40 cm long, 20-40 (-42) cm wide, cordate, membranous, glabrous to minutely puberulent, glandular, apex acute, base deeply lobed. Venation with 3-8 pairs of secondary veins arising from or near the base and 1-2 pairs from the middle part of the midrib, curving to the apex. Petioles sheathing, 10-30 cm long, glabrous to densely puberulent. Prophylls 10-20 cm long, glandular, subtending the peduncles. Inflorescences 4-10 cm long, in clusters of 2-8; peduncles 2-20 mm long, minutely puberulent. Anthers 1-2 mm long. Floral bracts 0.3-0.6 mm wide, triangular, white-ciliate. Fruits 0.5-0.8 mm wide, obovoid, trigonous, glabrous; stigmas 3, sessile. Shady areas in cloud forest, moist secondary forest, hillsides and riverbanks; (400-) 700-2000 m. DISTRIBUTION. Pantropical. Mexico, Guerrero: Paxson et al. 17M833 (BM); Morelos: Pringle 6159 (E); Nayarit: Croat 45328 (MO); Oaxaca: Hernandez & Torres 449 (MO); Temascaltepec: Hinton 4407 (BM); Veracruz: Purpus 2017 (BM). Guatemala, Alta Verapaz: Turckhelm 11258 (E); El Progresso: King 3256 (NY). El Salvador, Achuachapan: Croat 42083 (MO); Santa Ana: Villacorta 179 (LAGU). Honduras, Cortes: Hernandez 5295 (BM); Limpira: Nelson et al. 0265 (MO); Tegucigalpa: von Hagen 1151 (NY). Nicara- gua, Boaco: Stevens 14751 (BM); Esteli: Grijalva & Araquis- tain 614 (BM); Granada: Araquistain & Moreno 612 (BM); Madriz: Moreno 2771 (BM); Matagalpa: Moreno 10209 (MO). Costa Rica, Alajuela: Carvajal 501 (MO); Cartago: Holm & Iltis 118 (NY); Heredia: Croat 35840 (MO); Limon: Burger & Liesner 6893 (NY); Puntarenas: Hepper 34 (BM); San Jose: Burger & Stolze 5382 (NY). Panama, Chiriqui: Croat 66387 (MO); Code: Hammel 3875 (MO); Darien: LeDoux 2640 (NY); Panama: Croat 3373 (NY). Colombia, Dryander 2282 (BM). Venezuela, Miranda: Davidse 4508 (MO). Peru, Huanuco: Mexia 8254 (BM); Loreto: Mexia 6103 (BM). Bolivia, Mapiri: Buchtien 572 (BM). Brazil, Ceara: Gardner 1850 (BM); Minas Gerais: Mexia 4160 (BM); Sao Paulo: Davis et al. 59843 (E). 55. Piper peltatum L., Sp. pi. 1: 30 (1753). Type: Habitat in America calidiore. Fig. lB,b,c. P. pruinosum Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp. 1: 59 (1816). Type: Willd., h.n. 705 (B). P. speciosum Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp. 1: 59 (1816). Type: Willd., h.n. 704 (B). Lepianthes peltata (L.) Raf., Sylva tellur.: 84 (1838). Heckeria peltata (L.) Kunth in Linnaea 13: 565 (1839). H. scutata Kunth in Linnaea 13: 567 (1839). Type: Willd., h.n. 703 (B). H. speciosa (Kunth) Kunth in Linnaea 13: 568 (1839). Pothomorphe peltata (L.) Miq., Comm. phytogr.: 37 (1840). P. scutata (Kunth) Miq., Comm. phytogr.: 37 (1840). P. speciosa (Kunth) Miq., Comm. phytogr.: 37 (1840). P. almirantensis Trel. in Ann. Mo. hot. Gdn 27: 306 (1940). Type: Panama, Bocas del Toro, Cooper 170 (F-holotype). P. baileyorum var. paucispica Trel. in Ann. Mo. hot. Gdn 27: 306 (1940). Type: Panama, Canal Zone. Woodson, Allen & Seibert 1573 (NY!-isotype). P. tecumensis Trel. in Ann. Mo. hot. Gdn 27: 306 (1940). Type: Panama, Canal Zone, Standley 26735 (US- holotype). Herbaceous, sometimes subshrubs, 0.5-1.5 m high; stems glabrous or sparsely puberulent. Leaves 20-40 cm long, 15-30 cm wide, ovate to suborbiculate, membranous, glandu- lar, upper surface minutely puberulent on the veins, apex acute-acuminate, base round to subcordate, peltate. Vena- tion with 10-14 secondary veins arising from or near the petiole attachment, and 1-2 pairs from the middle of the midrib, curving towards the apex. Petioles sheathing, 10-26 cm long, glabrous. Prophylls 10-20 cm long, glandular, subtending the peduncles. Inflorescences 4-10 cm long, in clusters of 2-20; peduncles 3-20 mm long, glabrous. Anthers 0.1-0.2 mm long. Floral bracts 0.3-0.6 mm wide, triangular, white-ciliate. Fruits 0.5-0.8 mm wide, obovoid, trigonous, glabrous; stigmas 3, sessile. Moist places at edges of forest and pastures, road- and track sides, stream sides and waste ground; 0-500 (-800) m. DISTRIBUTION. Pantropical. Mexico, Chiapas: Hoover 159 (MO). Belize, Sibun River: Gentle 1400 (NY). Guatemala, Izabal: Kellerman 7411 (NY); Peten: Molina 15857 (NY). Honduras, Atlantida: Yuncker et al. 8449 (NY); Puerto Sierra: Wilson 128 (NY). Nicaragua, Boaco: Stevens 5917 (BM); Chontales: Stevens 2836 (BM); Granada: Grijalva 1911 (BM); Managua: Stevens 5339 (BM); Matagalpa: Guzman et al. 766 (BM); Rivas: Seymour 1931 (BM); Zelaya: Ortiz 52 (BM). Costa Rica, Cartago: Tonduz 7332 (NY); Heredia: Stevens 13320 (MO); Limon: Davidson & Donahue 8269 (LA); Puntarenas: Kernan 78 (CR); San Jose: Whitmore 60 (NY). Panama, Bocas del Toro: Woodson et al. 1875 (NY); Code: Davidse & Hamilton 23733 (MO); Colon: Nee 6979 (MO); Darien: Antonio 4579 (MO); Panama: Hamilton & D'Arcy 1322 (MO); San Bias: Hamilton & Stockwell 1077 (MO). Colombia, Antioquia: Klevens et al. 17C413 (BM); Barbacoas: Triana 754 (BM); Meta: Philipson et al. 1672 (BM); Santa Marta: Smith 1243 (BM). Venezuela, Carabobo: Alston 5892 (BM). Guyana, Schomburgk 954 (BM). Ecuador, Salanga L: Barclay 629 (BM). Peru, Loreto: Mexia 6102 (BM). Bolivia, Isapuri: Williams 682 (BM). M.C. TEBBS Fig. 1 A: P. umbellatum, habit; a: fruit and bract. B: P. peltatum, habit; b: fruit and bract; c: peduncles and prophylls. PIPER IN THE NEW WORLD Brazil, Amazonas: Mexia 6067a (BM); Para: Moss s.n. (BM). SECTION RADVLA MIQ. The section, as accepted here, follows Miquel's (1844) Artan- the section Radula, which he describes as 'Folia plerumque membranacea, elliptica, ovata, oblonga, raro angusta, inae- quilatera, acuminata, basi inaequalia, supra pilis rigidis ver- ruculisque scabra, raro glabra, subtus pilosa, pilis in pagina sup. sensim deciduis; costata, costis usque ad 1/3-2/3 alt. a basi ortis, adscendentibus, summa ad apicem ducta. Amenta recta vel curvata. Bracteae peltatae, pelta triangulari vel semicirculari ciliata; rarius cucullato-inflexae. Baccae tetra- vel trigono- compressae, subcuneatae.' Piper section Radula contains a number of species which at first glance look very much alike. It is only when characters such as prophylls, bracts, and fruits are examined in detail that differences begin to emerge. The determination of Piper to species is facilitated by using both vegetative and inflores- cence characters. Many species have been reduced to synon- ymy, especially under Piper hispidum Sw. I have endeavoured to see as many type specimens as possible of these species; where types have not been available, plates and photographs of types have been examined, and original descriptions have of course been carefully studied. DISTRIBUTION AND HABITAT Many species in this section are rapid colonisers of cleared land, often forming stands along sides of tracks or in clearings (e.g. P. aduncum L., P. bredemeyeri Jacq., P. hispidum). Piper marginatum Jacq., a widespread species, is commonly found in clearings, often in full sunlight. Piper dnereum C.DC., found from Panama south to Ecuador, is an early coloniser of disturbed ground such as landslides or unstable cliffs, whereas P. littorale C.DC. grows near the sea, often on the strand. Several species are found in wet forest, often at high altitudes (e.g. P. chamissonis (Miq.) Steud. of Mexico and Guatemala, P. holdridgeianum W.C. Burger from Costa Rica and Panama, and P. subflavum C.DC. from Colombia). The climbing P. multiplinervium C.DC. grows in moist forest, reaching and spreading over tree tops. Piper malpasoensis Tebbs is only found in steep moist ravines in southern Mexico; P. chrysostachyum C.DC. grows in seasonally dry evergreen forest, usually on the Pacific side of Nicaragua, Costa Rica, and Panama. Piper dotanum Trel. of Costa Rica grows in the deep shade of forest and P. scalarispicum Trel. is found in Central American cloud forest. MORPHOLOGY Habit The majority of species in this section are shrubs or small trees, 1-6 (-8) m high. Piper multiplinervium and P. sil- vivagum C.DC. are scandent or climbing, ascending by means of adventitious roots at the nodes, while P. hostmanni- anum (Miq.) C.DC. is usually found as a shrub, but may climb if the opportunity arises. Piper thomasii Tebbs is an epiphytic subshrub. Leaves Leaves are cordate, oblong to lanceolate, elliptic or ovate, membranous or coriaceous. The bases are usually unequally attached to the petioles, with one side lower than the other. Occasionally one side is slightly lobed, often concealing the petiole. The upper surfaces of the leaves in this section are typically scabrous, often combined with a sparse to dense indumentum (e.g. Piper aduncum, P. hispidum, P. villiramu- lum C.DC.). Undersides are usually pubescent, especially on the veins. Piper marginatum has glossy leaves, with a distinct margin of pale, minute hairs, whereas P. bredemeyeri has extremely rough, densely brown-pubescent, bullate leaves. Piper dnereum has whitish-pubescent leaves, which are some- times rugose. Venation Several species in this section have palmate venation (actino- dromous or campylodromous), with 3-13 secondary veins arising from the base of the leaf and curving towards the apex (e.g. Piper marginatum, P. sanctum (Miq.) Schltdl.). Piper flavidum C.DC. ex Donn.Sm. has 1-2 pairs of veins arising from the lower part of the leaf, running parallel to the apex, with prominent cross-venation. Piper aduncum and P. lancei- folium Kunth have secondary veins arising from the lower to middle part of the midrib, ascending steeply to the apex. Piper dotanum and P. hirtellipetiolum C.DC. have distinct brochidodromous venation, whereas Piper hispidum and P. poasanum C.DC. have curving or arcuate-ascending second- ary veins arising from the lower to middle part of the midrib. Indumentum and pellucid glands The amount of indumentum is very variable and is often greater in young growth, diminishing as the plant ages. Piper biauritum C.DC. is densely pubescent, often with long multi- cellular hairs, whereas P. hispidum and P. villiramulum are covered with short, pale hairs, especially in young growth. Piper dnereum has white-puberulent stems and leaves. Piper villiramulum has dark, reddish glands visible, especially on the leaf undersides, whereas P. bisasperatum Trel. has copi- ous orange glands on both surfaces. The ligule-like structures present at the base of the petioles are often prominently glandular (e.g. P. chrysostachyum C.DC.) Prophylls The prophylls are generally prominent in this section and usually associated with a ligule-like structure that arises from the base of the petiole. The exceptions are those species with sheathing petioles (P. marginatum, P. dnereum, P. multiplin- ervium, and P. holdridgeianum) where the prophylls are partially or wholly concealed. Piper bredemeyeri has pro- phylls 10-25 mm long, whereas those of P. terrabanum C.DC. are up to 30 mm long. Inflorescences In this section, the inflorescences are erect, arched or curved and are usually pale in colour. The inflorescences of Piper 6 M.C. TEBBS marginatum are erect for about half their length and then 5a. Fruits trigonous from above: arch gracefully. Piper multiplinervium has slender, erect, 6a. Epiphytic subshrubs 65. P. thomasii yellowish spikes with a pleasant fragrance. The inflorescences 6b- Terrestrial herbs, shrubs, or trees: of P. bisasperatum are erect, with pubescent sterile tips up to \ Upper surface of leaves glabrous, often glossy: 4 mm long Those of Piper epigynium C.DC. and P. biauri- 8aQ Leaf bases ,une(lual * au»culate< cove""g the P*i oj»: 9a. Leaves lanceolate, undersides with minute dark turn are coloured pinkish to purple. glands 70 p enganyanum 9b. Leaves elliptic-ovate, undersides eglandular: Floral bracts 10a. Leaves 14-24 x 6-10 cm; inflorescences 6-12 cm long 68. P. otophorum Bracts are triangular or round, usually densely white- or 10b. Leaves 8-12 x 3.5-6 cm; inflorescences 4-6 cm yellow- ciliate with glabrous centres, as in P. poasanum or P. long 69. P. malpasoensis chamissonis. Some species have a pale basal bulge (e.g. Piper 8b. Leaf bases not auriculate, not covering the petioles: chrysostachyum, P. biauritum). Bracts of P. dotanum and P. Ha. Upper surface of fruits glandular: villiramulum are densely puberulent. 12a- Petioles with «gule-Hke structures 3.5 mm long; upper surface of fruits pubescent 61 . P. via-chicoense Fruit 12b. Petioles with ligule-like structures 0.5-1 mm long; upper surfaces of fruits glabrous: Fruits are obovoid, round or trigonous from above, glabrous 13a Leaves membranous; prophylls 20-30 mm (e.g. P. sanctum, P. terrabanum) , densely pale puberulent long 66. P. terrabanum (e.g. P. cinereum, P. flavidum, P. hispidum), or minutely 13b. Leaves subcoriaceous, slightly succulent; pro- granular (e.g. P. achoteanum Trel.). Piper marginatum has phylls 5-10 mm long 67. P. littorale round, dark fruits with a central depression. lib. Upper surface of fruits eglandular 64. P. dilatatum 7b. Upper surface of leaves pubescent: 14a. Stems densely covered with long hairs; inflorescences TAXONOMIC TREATMENT erect: 15a. Leaves narrowly to broadly ovate, upper surface bullate; underside densely covered with yellow- Section RADULA Miq. brown hairs 63. P. bredemeyeri 15b. Leaves elliptic to obovate or rhombic, upper surface Type: Piper radula Kunth not bullate; underside densely covered with pale Herbs, shrubs or small trees, occasionally scandent or climb- yellow hairs , " "• 62' P" Pseudofuligineum , 14b. Stems sparsely to densely covered in short hairs; ing, rarely epiphytic, stems glabrous to densely pubescent. inflorescences arching 71 P lanceifolium Leaves linear, lanceolate, ovate-elliptic, oblong or cordulate, 5b Fmits round or oblong from aboye. glabrous, scabrous or pubescent, often glandular. Venation 16a Upper surface of fruit glabrous, sometimes granular: pinnate or palmate. Petioles mostly with ligule-like structures 17a. Leaves with palmate venation: at the base, occasionally sheathing. Prophylls prominent, 18a. Leaves broadly ovate to elliptic, the bases round to glabrous or pubescent, sometimes glandular, or occasionally shallowly cordate 106. P. sanctum hidden by sheathing petioles. Inflorescences erect or curving, 18b. Leaves oblong-lanceolate to elliptic-lanceolate, the sometimes distinctly arching, rarely pendulous. Floral bracts bases obtuse to sliShtly lobed on one side • triangular, round or semilunar, sparsely to densely ciliate, 105. P. pseudolindenii rarely glabrous. Fruits trigonous or round from above, gla- 17b' 1^eavIesflwlth Pmnate ve"atl°n: 19a. Inflorescences arching, curving, or pendulous: brous to pubescent, sometimes granular or glandular, esty- 20a Upper surface of ,eaves scabrous or dense,y lose; stigmas L-A. puberulent; inflorescences arching or curving, Disturbed areas, often along roads or tracks, secondary *Jti°Ut ^VT* r r- u • 11 u u 21a. Prophylls 5-8 mm long; peduncles 15-30 mm forest, shady wood or field margins, occasionally by the sea; longp B v 73 p> mol,icomum 0-3000 m. jib. Prophylls 20-25 mmlong; peduncles 8-15 mm long 72. P. aduncum Key to the Species in Section Radula 20b- Upper surface of leaves glabrous; inflorescences pendulous, with sterile tips 1-3 mm long la. Petioles partly or completely sheathing the stems: 99. P. carpinteranum 2a. Plants scandent or climbing 58. P. multiplinervium 19b. Inflorescences erect: 2b. Plants shrubs or subshrubs: 22a. Leaves ovate-cordate, more or less sessile, often 3a. Leaves with palmate venation; prophylls hidden by sheath- with blunt apices 23 ing petioles; inflorescences arching or curving: 23a. Inflorescences 5-10 cm long; peduncles 4a. Leaves glossy above, sparsely puberulent, with distinct 6-10 mm long 74. P. achoteanum ciliate margins; fruits glabrous, drying brown or 23b. Inflorescences 3-7 cm long; peduncles black 56. P. marginatum 20-60 mm long 75. P. fuligineum 4b. Leaves dull above, pubescent, without distinct ciliate 22b. Leaves petiolate, elliptic-lanceolate, ovate, or margins; fruits densely white-pubescent oblong to obovate, apices acuminate 24 57. P. cinereum 24a. Leaf surfaces distinctly glandular 25 3b. Leaves pinnately or sub-pinnately veined; prophylls visible, 25a. Upper surfaces of leaves bullate not hidden by sheathing petioles; inflorescences erect , 78. P. pseudoasperifolium 59. P. holdridgeianum 25b. Upper surfaces of leaves smooth 26 26a. Underside of leaves with red or orange Ib. Petioles not partly or completely sheathing the stems: PIPER IN THE NEW WORLD glands ........................ 94. P. colonense 37b. Leaves eglandular ................................... 43 26b. Undersides of leaves lacking red or orange 43a. Floral bracts oblong-triangular; fruits minutely glands .......................................... 27 white-pubescent or granular . 82. P. poasanum 27a. Petioles glabrous, with glabrous ligule- 43b. Floral bracts semi-lunar; fruits densely yellow- like structures . 98. P. tenuimucronatum brown pubescent .... 103. P.jacquemontianum 27b. Petioles villous, with ciliate ligule-like 35b. Upper surface of leaves pubescent, rugose or bul- structure .......... 96. P. hirtellipetiolum late ............................................................. 44 24b . Leaf surfaces eglandular ....................... 28 44a . Leaves with dark red , orange or brown glands 28a. Ligule-like structures on petioles 2-3 mm visible ....................................................... 45 long, glandular ............. 93. P. chamissonis 45a. Petioles glabrous or pubescent, glandular, with 28b. Ligule-like structures on petioles 0.5-1 mm glandular ligule- like structures 4-10 mm long .... long, eglandular ................................. 29 ....................................... 83. P. bisasperatum 29a. Upper surfaces of leaves glossy, gla- 45b. Petioles densely pubescent, with eglandular brous ............................................ 30 ligule-like structures 0.5-1 mm long ............. 46 30a . Leaves with prominent cross-venation ; 46a . Stems densely whitish-pubescent ................. inflorescences with sterile tips ......... 31 .................................... 79. P. villiramulum 31a. Stems glabrous; inflorescences 2-5 46b. Stems densely yellow-brown pubescent ......... (-6) cm long ......... 97. P. decurrens ..................................... 92. P. perhispidum 31b. Stems sparsely to densely yellow- 44b. Leaves eglandular ...................................... 47 pubescent; inflorescences 10-12 cm 47a. Upper surface of leaves covered in long hairs; long ........... 100. P. hostmannianum inflorescences red or purple ..... :91 . P. biauritum 30b. Leaves lacking prominent cross- 47b. Upper surface of leaves minutely pubescent; venation ; inflorescences without sterile infloresences not red or purple .................... 48 tips ........................................... 32 48a. Stems shortly white-pubescent; ligule-like 32a. Leaves lanceolate to elliptic- structures glandular ............. 80. P. hispidum lanceolate; prophylls 7-10 mm long, 48b. Stems with long yellow to brown hairs; ligule- with central line of hairs .................. like structures eglandular ........................ 49 ...................... 101. P. scalarispicum 49a. Floral bracts mostly glabrous, with a few 32b. Leaves elliptic to ovate or rhombic. minute hairs at the base . 88. P. polytrichum prophylls 12-25 mm long, gla- 49b. Floral bracts yellow-ciliate, with glabrous brous ................... 84. P. umbricola centres ............................................. 50 29b. Upper surfaces of leaves moderately to 50a. Prophylls densely puberulent; inflores- densely yellowish-pubescent .............. 33 cences 3-9 cm long ....... 90. P. zacatense 33a. Leaves narrowly elliptic to oblanceolate; 50b. Prophylls hirsute only along midrib; inflo- peduncles 14-25 mm long ................... rescences 10-16 cm long ....................... ......................... 95. P. oblanceolatum .......................... 89. P. peracuminatum 33b. Leaves ovate-lanceolate to elliptic- lanceolate; peduncles 3-5 (-10) mm % K marginatum Jacq.5 /con. pL rar. 2: 2, t. 215 (1786). /?"? •"/ ....... ; ........... 60" P subfl™ Type: West Indies. Jacquin s.n. (G-holotype). 16b. Upper surface of fruit pubescent ............................ 34 Jr 34a. Plants scandent or climbing ........... 77. P. silvivagum rig. ZA,a,D,C. 34b. Plants erect, not scandent or climbing .................. 35 p caudatum Vahl, Eclog. amer. 1: 3 (1796). Type: Rohr 217 35a . Upper surface of leaves glabrous ...................... 36 /£ , hoiotype) 36a. Leaves falcate, linear-lanceolate, with 1-2 pairs of p ^/m™ Kun^ in R BQ & ^^ ^ secondary veins ......................... 76. P. ilavidum co/ioi/:\ T> TI/-//J uu <;no /r> u ^ + ^ 36b. Leaves not falcate, linear-lanceolate, with at least 3 SP- 1: 58 (1816)' TyPe: Wdldeno^ ' Hb 698 (B-holotype). pairs of secondary veins ................................ 37 P- catalpae folium Kunth in Humb. , Bonpl. & Kunth, Nov. 37a. Leaves glandular .................................... 38 gen. sp. 1: 58 (1816). Type: Willdenow Hb. 699 38a. Upper surface of leaves bullate, undersides (B-holotype). lustrous ............................... 104. P. biritak P. alare Ham., Prodr. pi. Ind. occid.: 3 (1825). Type: 38b. Upper surface of leaves smooth, undersides Cayenne. Hamilton s.n. (?-holotype). dull .................................................... 39 Schilleria caudata (Vahl) Kunth in Linnaea 13: 716 (1839). 39a. Ligule-like structures on petioles, 6-18 s cataipaefoiia (Kunth) Kunth in Linnaea 13: 718 (1839). 18 mm ^S ................. 8LP; sanct>-fehcis 5. marginata (Jacq.) Kunth in Linnaea 13: 718 (1839). 39b. Ligule-like structures on petioles, not more , i r> / i r/ „• / Am i -2^ no/im QOO, % ,n Piper patulum Bertol., Fl. guatimal: 407, pi. 36 (1840). See thanSmmlong ................................. 40 40a. Fruits 1-1. 2 mm wide, densely pale yellow- note below. pubescent ................. 102. P. curvatipes Artanthe cataipaefoiia (Kunth) Miq., Syst. piperac.: 382 40b. Fruits 0.4-0.8 mm wide, minutely white (1844). puberulent or glandular .................... 41 A. caudata (Vahl) Miq., Syst. piperac.: 380 (1844). 41a. Leaves with dark glands; prophylls lack- Artanthe marginata (Jacq.) Miq., Syst. piperac.: 381 (1844). ing glands; floral bracts densely puberu- ^ aiar(s (Ham.) Miq., Syst. piperac.: 406 (1844). lent .......................... 87. P. dotanum piper marginatum Var. catalpaefolium (Kunth) C.DC. in 41b. Leaves with pale glands; prophylls glan- DC />rodr 16(1) ' 246 (1869) dular; floral bracts glabrous or minutely p san.'joseanum CDC in Linnaea 37: 351 (1872). Type: Cos^ Rica, Oe^ 893 (C!-holotype) dular 86 p epigynium p- pseudo-margmatum C.DC. in Bull. Herb. Boissier 6: 492 42b. Prophylls 8-15 mm long; fruits eglan- (1898). Type: Ecuador, Sodiro 1/38 (P?-holotype). dular ........... 85 . P. chrysostachyum Fig. 2 A: P. marginatum, habit; a: section of leaf margin; b: prophyll; c: fruit and bract. B: P. cinereum, habit; d: prophyll; e: fruit and bract. C: P. multiplinervium, habit; f: fruit and bract. PIPER IN THE NEW WORLD P. marginatum var. anisatum (Kunth) C.DC. in Urb., Symb. antill. 3: 172 (1902). P. patulum var. cordifolium Trel. in /. Wash. Acad. Sci. 13: 366 (1923). Type: El Salvador, Sonsonate, Standley 22046 (ILL!-holotype). P. uncatum Trel. in/. Wash. Acad. Sci. 13: 367 (1923). Type: El Salvador, Tonacatepeque, Standley 19435 (ILL!- holotype). P. san-joseanum var. minor Trel. in Contr. U.S. natn. Herb. 26: 133 (1929). Type: Costa Rica, Nicoya, Tonduz 13695 (US- holotype, photograph!). P. san-joseanum var. chiriquinum Trel. in Ann. Mo. hot. Gdn 25: 826 (1938). Type: Panama, Chiriquf, Woodson et al. 416 (MO! -holotype). P. san-joseanum var. kobense Trel. in Ann. Mo. hot. Gdn 27: 297 (1940). Type: Panama, Canal Zone, Woodson et al. 1423 (ILL!-holotype). P. san-joseanum var. panamanum Trel. in Ann. Mo. hot. Gdn 27: 297 (1940). Type: Panama, Gorgona Beach, Woodson et al. 1690 (ILL!-holotype). P. san-joseanum var. remediosense Trel. in Ann. Mo. hot. Gdn 27: 297 (1940). Type: Panama, Chiriquf, Woodson etal. 1191 (ILL!-holotype). P. san-joseanum var. tabogense Trel. in ^4nn. Mo. hot. Gdn 27: 297 (1940). Type: Panama, Taboga, Woodson et al, 1531 (ILL!- holotype). P. niceforoi Trel. & Yunck., Piperac. N. South Amer. 1: 79 (1950). Type: Colombia, norte de Santander, Niceforo 49 (ILL!- holotype). P. marginatum var. marginatum forma catalpaefolium (Kunth) Steyerm., Fl. Venezuela 2: 480 (1984). P. marginatum var. niceforoi (Trel. & Yunck.) Steyerm., Fl. Venezuela 2: 482 (1984). Shrubs 2-3 m high, glabrous or slightly pubescent. Leaves 9- 20 (-28) cm long, 5-16 (-24) cm wide, broadly ovate, glossy above, membranous, glabrous or with scattered hairs, under- side often puberulent on veins, margins densely pale-ciliate, apex acuminate, base shallowly to deeply cordate. Venation palmate, with 7-13 nerves arising from the base. Petioles 2-5 cm long, sheathing the stem. Prophylls hidden by sheath- ing petioles, 2-4 mm long, glabrous, papery, minutely glan- dular, apex blunt, sometimes fimbriate. Inflorescences 10-25 cm long, whitish, becoming arched; peduncle 5-12 (-20) mm long, glabrous. Anthers 0.3-0.4 mm long. Floral bracts 0.5-0.8 mm wide, triangular or round, densely ciliate. Fruit 0.5-1 mm wide, obovoid, dark brown, round to oblong from above, glabrous; stigmas 3-4. Hillsides, pasture, disturbed forest, roadsides, and edges of streams; 0-1300 m. DISTRIBUTION. Mexico to Brazil. Mexico, Campeche: Gomez-Pompa 1316 (CAS); Chiapas: Breedlove 28606 (MO); Colima: Ferris 6219 (DS); Oaxaca: Chavelas & Perez 266 (DS); Quintana Roo: Cabrera 609 (CAS); Veracruz: Calzada 894 (CAS); Yucatan: Lundell 1195 (DS). Belize, Cayo: Proctor 29552 (BM). Guatemala, Alta Verapaz: Croat 41520 (MO); Escuintla: Croat 42051 (MO); Peten: Ratcliff42 (CAS); Retalhuleu: Ellis & LeDoux 1128 (DS); Zacapa: Croat 41865 (MO). El Salvador, Ahuachapan: Croat 42067 (MO); La Libertad: Vicente & Villacorta 151 (BM); Santa Ana: Villacorta 180 (BM). Honduras, Comayagua: Soto 41 (BM); Copan: Croat 42518 (MO); Olancho: Carvajal 25 (BM); Santa Barbara: Croat 42759 (MO). Nicaragua, Boaco: Stevens 14708 (BM); Carazo: Moreno 10732 (BM,MO); Chinandega: Fonseca 80 (BM); Granada: Baker 66 (DS); Managua: Stevens 2917 (BM); Masaya: Zelaya 149 (BM); Matagalpa: Stevens et al. 21424 (BM). Costa Rica, Alajuela/ Heredia: Grayum et al. 4134 (BM,MO); Guanacaste: Williams et al. 26596 (DS); Puntarenas: Grayum 4192 (BM,MO). Panama, Canal Zone: Croat 12875 (MO); Chir- iquf: Hamilton et al. 938 (BM,MO); Colon: Sytsma 1611 (BM,MO); Darien: Hammel 1084 (MO); Panama: Hamilton 540 (BM,MO); Perlas Is.: Knapp 3276 (BM,MO); San Bias: Nevers & Herrera 4524 (MO); Veraguas: Antonio 2325 (BM,MO). West Indies, Trinidad: Breedlove 18992 (CAS). Colombia, Boyaca: Lawrance 194 (BM); Los Llanos: Haught 2472 (BM); Santa Marta: Smith 1239 (BM); Sur de Santander: Haught 1421 (BM). Venezuela, Apure: Davidse & Gonzalez 21632 (BM,MO); Aragua: Williams & Alston 173 (BM); Lara: Davidse & Gonzalez 20947 (BM,MO); Miranda: Davidse 4108 (BM,MO); Monogas: Croat 54330 (BM,MO); Portuguesa: Davidse et al. 21466 (BM,MO); Tachira: Dav- idse & Gonzalez 22441 (BM). French Guiana: Sagot 533 (BM). Ecuador, Esmeraldas: Barclay s.n. (BM); Guayas: Camp 3547 (BM,NY). Brazil, Para: Ducke 3494 (BM); Pernambuco: Picket s.n. (DS). Piper marginatum is one of the most widespread species in this section and also one of the easiest to identify. It can be recognized by its glossy, palmately-veined leaves with ciliate margins, sheathing petioles, and white, curving inflores- cences. Piper auritum Kunth also has leaves with ciliate margins and sheathing petioles, but these are oblong-cordate, without palmate venation, and the fruits are trigonous rather than round, as in P. marginatum. Both Piper auritum and P. marginatum have a pleasant, aniseed-like odour when the stems or leaves are crushed. Piper cinereum is closely related to P. marginatum, with cordate leaves, sheathing petioles concealing the prophylls and palmate venation. The leaves, however, are whitish pubescent, the fruits are densely white- puberulent, and the stems, petioles, and peduncles are often streaked with red. In his original description of this species, Bertoloni men- tioned only vegetative characters, failing to describe bracts or fruits which he presumably did not see. He described the leaves as ovate-oblong to cordate-ovate, thin, entire, with a rounded, somewhat unequal base, acuminate apex, 7-9 retic- ulate nerves arising from the base, with dorsal nerves slightly thinner and puberulent. The immature spike was described as being some 5-6 inches (12-15 cm) long, spreading, with flowers small and crowded. The specimen from which his description was made was collected from Esquintla in Guate- mala. As the type has been presumed lost in the Second World War (Duncan, 1983), only Bertoloni's coloured illus- tration has been seen, which lacks the important details of the inflorescence. Standley & Steyermark (1952) noted in the description of P. patulum that 'this is presumably the Piper that has been reported from Guatemala as P. marginatum Jacq.', but did not go into more detail. This account of P. patulum matches the description of P. marginatum. Burger (1971) placed P. patulum Bertol. as a synonym of P. margin- atum Jacq. One of the characters of P. marginatum is the distinct wnite-ciliate margin of the leaves, which is curiously not mentioned by Bertoloni in his original description. All the specimens seen by the present author, originally anno- tated as P. patulum, are actually P. sanctum (Miq.) Schltdl. 10 M.C. TEBBS As the type specimen is missing and the original description and plate are inadequate, it may never be possible to prove the true identity of P. patulum; therefore, for convenience, the name has been retained in the synonymy of P. margina- tum Jacq. 57. Piper cinereum C.DC. in J. Bot., Land. 4: 214 (1866). Type: Colombia, prov. de Choco, Triana s.n. (G-holotype; BM-isotype?). Fig. 2B,d,e. P. candicans Sodiro, Sertula fl. ecuad.: 13 (1905). Type: Ecuador, Esmeraldas, Mille s.n. (?-holotype; ILL!- isotype). Straggling shrubs or subshrubs 1-2 m high, pubescent, with pink or red streaks on stems. Leaves 9-17 cm long, 8-14 cm wide, ovate-cordate, dull green above, paler beneath, whitish pubescent, occasionally rugose, apex acute-acuminate, base deeply cordate. Venation palmate, with 9-13 nerves arising from the base and curving towards the apex, with prominent cross-veins. Petioles 2-7 cm, sheathing, often marked with red. Prophylls hidden beneath sheathing petioles, 3-5 mm long, whitish, papery, pubescent, apex acute. Inflorescences 8-11 cm long, erect or slightly curved; peduncles 12-25 mm long, white pubescent. Anthers 0.1-0. 2 mm long. Floral bracts 0.2-0.4 mm wide, triangular, covered with white hairs. Fruit 0.5 mm wide, obovoid, round to trigonous from above, upper surface white pubescent; stigmas 3-4. Coloniser of disturbed ground, often found on landslide areas or sides of cliffs; 0-700 m. DISTRIBUTION. Panama to Ecuador. Panama, Darien: Whitefoord & Eddy 107 (BM). Colombia, Cauca: Haught 5368 (BM); Choco: Luteyn et al. 10495 (G); El Valle: Killip & Garcia 33360 (BM). 58. Piper multiplinervium C.DC. in /. Bot., Lond. 4: 214 (1866). Type: Colombia, Barbacoas, Triana 27 (G-holotype; BM!- isotype). Fig. 2C,f. P. aragonenseTrel. in Contr. U.S. natn. Herb. 26: 146 (1929). Type: Costa Rica, Aragon near Turrialba, Tonduz 9021 (US-holotype). Scandent or climbing up to 10 m high, glabrous or minutely puberulent. Leaves 6-16 cm long, 5-12 cm wide, glossy, upper leaves ovate-elliptic, lower leaves ovate-cordate, apex acute-acuminate, base obtuse to cordate. Venation with 3-4 secondary veins arising from or near the base, and 1-2 pairs arising from the middle of the midrib, ascending to apex. Petioles 1-3 cm long, sheathing. Prophylls not apparent — if present hidden by the sheathing petioles. Inflorescences 6-16 cm long, erect, yellow-white, lemon-scented; peduncles 1-2 cm long, glabrous or minutely pubescent. Anthers 0.1-0.2 mm long. Floral bracts 0.4-0.6 mm wide, triangular to round, yellow-white ciliate. Fruits 1-1.5 mm wide, obo- void, round from above, glabrous; stigmas 2-3. Moist forest; 0-1000 m. DISTRIBUTION. Nicaragua to Peru. Nicaragua, Zelaya: Miller & Sandino 1206 (BM,MO). Costa Rica, Cartage: Taylor 11429 (NY); Heredia: Burger 8083 (NY); Limon: Whitmore 102 (F); Puntarenas: Greig 174 (F). Panama, Bocas del Toro: McPherson & Allen 9626 (BM); Canal Zone: Gentry 5765 (F); Chiriqui: Werff & Hardeveld 6527 (BM,MO); Code: Croat 67106 (BM,MO); Darien: Mori 7087 (BM,MO); Pan- ama: Garwood 1856A (BM); San Bias: Croat 69237 (BM,MO); Veraguas: Croat 25546 (NY). Colombia, Antio- quia: Shepherd 740 (F); Bogota: Triana 50 (BM); Choco: Gentry & Fallen 17875 (F); Santander: Haught 2177 (BM). Ecuador, Esmeraldas: Madison et al. 5030 (F); Imbabura: Solfs 12525 (F); Los Rfos: Dodson & Gentry 6337 (F). Peru, Huanoco: Schunke 1421 (F); Madre de Dios: Smith & Condon 1070 (F). A vigorous and attractive species, often sprawling over the crowns of trees, and producing many slender, fragrant, yellow- white flowering spikes. The younger leaves are often a different shape from older, lower leaves, a feature noted in other species of Piper, such as P. schiedeanum Steud. Piper multiplinervium has similar leaves, venation, and petioles to the climbing P. nigrum L., an old world species widely cultivated for its fruit, known as the culinary peppercorns. However, P. nigrum has a pendulous inflorescence, dioecious flowers, glabrous bracts and large red to black fruits. 59. Piper holdridgeianum W.C. Burger in Fieldiana Bot. 35: 144 (1971). Type: Costa Rica, Heredia, Burger & Stolze 5776 (F!-holotype; BM!-isotype). Fig. 3A,a,b. P. picardae var. pilinervium Trel. in Feddes Reprium Spec, nov. veg. 23: 309 (1927). Type: Haiti, Ekman 1769 (ILL!- holotype). Shrubs 1-2 m high, stems glabrous or minutely puberulent. Leaves 10-22 cm long, 3.5-13 cm wide, ovate-elliptic to ovate-lanceolate, upper surface glabrous, glossy, dark green, underside glabrous or minutely pubescent, apex acuminate, base round to deeply cordate on older, lower leaves. Vena- tion with 2-4 pairs of secondary veins arising from the lower part of midrib, ascending to apex. Petioles 1- 5 cm long, deeply vaginate, appearing to sheath the stems, glabrous or minutely puberulent. Prophylls 4-10 mm long, blunt, gla- brous, drying dark brown or black. Inflorescences 6-10 cm long, erect; peduncles 10-18 mm long. Anthers 0.2-0.3 mm long. Floral bracts 0.2-0.4 mm wide, triangular or round, white ciliate, with dark, glabrous centres. Fruits 0.7-0.8 mm wide, obovoid, glabrous, fleshy; stigmas 3, sessile, in slight depression. Deep shade in wet forest; 0-1000 m. DISTRIBUTION. Costa Rica, Panama. Costa Rica, Heredia: Raven 20999 (F). Panama, Code: Hammel 3598 (BM,MO); Panama: Croat 14775 (MO); Veraguas: Croat 27617 (MO). Piper holdridgeianum occurs in moist to wet forest, often near streams or rivers. It can be identified by its deeply vaginate petioles, slightly uneven, cordate lower leaves, and upper leaves with little or no basal lobing. This variation of leaf shape on the same plant is quite common in Piper. 60. Piper subflavum C.DC. in J. Bot., Lond. 4: 210 (1866). Type: Colombia, prov. Pasto Ortega, Triana 23 (G-holotype; B!-isotype). Fig. 3B,c,d. PIPER IN THE NEW WORLD Fig. 3 A: P. holdridgeianum, habit; a: prophyll; b: fruit and bract. B: P. subflavum, habit; c: prophyll; d: fruit and bract. C: P. via-chicoense, habit; e: prophyll; f: fruit anf bract. D: P. pseudofuligineum, habit; g: prophyll; h: fruit and bract. 12 Shrubs 2-3 m high, stems densely soft-pubescent with yellow- ish hairs. Leaves 10-17 cm long, 4-8 cm wide, ovate- lanceolate to elliptic-lanceolate, both sides yellowish- pubescent, lower surface densely so especially on veins, apex acuminate, base round to acute, slightly unequal. Venation with 4 pairs of secondary veins arising from the lower to middle part of midrib, ascending sharply to apex. Petioles 3-7 (-10) mm long, densely pubescent, ligule-like structure not apparent, obscured by dense pubescence. Prophylls 10-12 mm long, densely pubescent. Inflorescences 7-9 cm long, erect; peduncles 3-5 (-10) mm long, pubescent. Anthers 0.2-0.3 mm long. Floral bracts 0.7- 0.9 mm wide, triangular with dense fringe of yellowish-white hairs. Fruits 0.9-1.2 mm wide, obovoid, trigonous-round from above, glabrous or slightly granular; stigmas 3. Damp places in clearings; 1000-2300 m. DISTRIBUTION. Colombia. Colombia, El Cauca: Pennell & Killip 8232 (ILL); El Valle: Dry under 64 (ILL); Santander: Killip & Smith 20331 (ILL). Piper subflavum can be recognized by its yellowish-pubescent leaves, densely yellow-ciliate floral bracts and trigonous to round fruits. It is only found at high altitudes in a few areas of Colombia. 61. Piper via-chicoense Yunck. in Ann. Mo. hot. Gdn 37: 67 (1950). Type: Panama, Rio Indio, Chico Trail, Steyer- mark & Allen 17431 (MO!-holotype). Fig. 3C,e,f. Shrubs 1-2 m high, branches arching, stems densely pubes- cent. Leaves 9-16 cm long, 3-5.5 cm wide, elliptic to oblong- lanceolate, upper surface glabrous, underside pubescent, apex acuminate, base unequal, narrowly obtuse. Venation with 3-5 pairs of secondary veins loop-connecting to the apex. Petioles 2- 5 mm long, pubescent, with ligule- like structure 3.5 mm, glandular, densely pubescent. Pro- phylls 10-13 mm long, glandular, densely pubescent. Inflorescences 3-7 cm long; peduncles 5-7 mm long, pubes- cent. Anthers 0.2 mm long. Floral bracts 0.4-0.5 mm wide, densely pale-ciliate, with dark centres. Fruits 0.8-1 mm wide, obovoid, trigonous, upper surface glandular, sparsely pubes- cent; stigmas 3, slender. Premontane rain forest; 200-1000 m. DISTRIBUTION. Panama. Panama, Darien: Duke 8061 (MO); Panama province, Cerro Azul: Dwyer 2085 (MO); Cerro Jefe: Gentry 4870 (MO). 62. Piper pseudofuligineum C.DC. in Linnaea 37: 355 (1872). Type: Costa Rica, Candelaria, Oersted s.n. (C!- holotype). Fig. 3D,g,h P. salinasanum C.DC. in Bull. Soc. r. Bot. Belg. 30(1): 214 (1892). Type: Costa Rica, Salinas, Pittier 2775 (G!- holotype). P. domingense C.DC. in An. Inst. fis.-geogr. C. Rica 9: 161 (1897). Type: Costa Rica, Santo Domingo de Golfo Dulce, Tonduz 10034 (G!-holotype). P. salinasanum var. subscabrifolium C.DC. in An. Inst. fis.- geogr. C. Rica 9: 164 (1897). Type: Costa Rica, Virilla, Tonduz 10126 (G!-holotype). M.C. TEBBS P. dumeticola C.DC. in An. Inst. fis.-geogr. C. Rica 9: 164 (1897). Type: Costa Rica, Boruca, Pittier 4490 (G-holotype; F!-isotype). P. pseudo-dilatatum C.DC. in An. Inst. fis.-geogr. C. Rica 9: 165 (1897). Type: Costa Rica, Punta Mala, Tonduz 6797 (G!- holotype). P. verbenanum C.DC. in An. Inst. fis.-geogr. C. Rica 9: 165 (1897). Type: Costa Rica, Alajuelita, Tonduz 8867, 8870 (G!- syntypes). P. taboganum C.DC. in Smithson. misc. Collns 71(6): 4 (1920). Type: Panama, Taboga Is, Pittier 3529 (US- holotype; G!- isotype). P. bigelovii Trel. in Contr. U.S. natn. Herb. 26: 23 (1927). Type: Panama, Bigelow s.n. (NY!-holotype). P. chagresianum Trel. in Contr. U.S. natn. Herb. 26: 37 (1927). Type: Panama, Chagres, Fendler 268 (US- holotype; MO!-isotype). P. breve C.DC. ex Trel. in Contr. U.S. natn. Herb. 26: 38 (1927). Type: Panama, Canal Zone, Pittier 3787 (US- holotype; NY!- isotype). P. atlantidanum Trel. in /. Wash. Acad. Sci. 19: 329 (1929). Type: Honduras, Atlantida, Standley 56739 (F!-holotype; ILL!- isotype). P. nigricaule Trel. in Contr. U.S. natn. Herb. 26: 158 (1929). Type: Costa Rica, Santo Domingo de Golfo Dulce, Tonduz 9959 (US-holotype; F!-isotype). P. griseo-pubens Trel. in Contr. U.S. natn. Herb. 26: 176 (1929). Type: Costa Rica, Libano, Guanacaste, Standley & Valeria 44879 (US-holotype; ILL! -isotype). P. griseo-pubens var. revocabile Trel. in Contr. U.S. natn. Herb. 26: 176 (1929). Type: Costa Rica, Tilaran, Standley & Valeria 44943 (US-holotype). P. squali-pelliculum Trel. in Contr. U.S. natn. Herb. 26: 178 (1929). Type: Costa Rica: Rio Virilla, San Jose, Tonduz 10126 (US-holotype;F!,NY!-isotypes). P. quadratilimbum Trel. in Publs Field Mus. nat. Hist. (Bot.) 17: 233 (1937). Type: Guatemala, Peten, Lundell 1488 (MICH- holotype; FMsotype). P. deltoideocarpum Trel. in Publs Field Mus. nat. Hist. (Bot.) 17: 344 (1938). Type: Honduras, Comayagua, Yuncker, Dawson & Youse 5654 (ILL!-holotype; F!-holotype). P. clavulispicum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 337 (1937). Type: Costa Rica, El Rodeo, Lankester 1318 (F!-holotype; G!-isotype). P. ponendum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 355 (1937). Type: Costa Rica, Heredia, Brenes 13244 (F!-holotype). P. salutatrix Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 359 (1937). Type: Costa Rica, El General, Skutch 2504 (US-holotype; MO!-isotype). P. vitabile Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 369 (1937). Type: Costa Rica, Buenes Aires, Valeria 894 (F!-holotype). P. canyazasense Trel. in Ann. Mo. hot. Gdn 25: 826 (1938). Type: Panama, between Canazas & Cordillera Central, Allen 185 (ILL!-holotype; MO!-isotype). P. coyolesense Trel. in Publs Field Mus. nat. Hist. (Bot.) 9: 278 (1940). Type: Honduras, Coyoles, Yoro, Yuncker, Koepper & Wagner 8147 (ILL!-holotype; F!,G!- isotypes). P. elasmophyllum Trel. in Publs Field Mus. nat. Hist. (Bot.) 9: 278 (1940). Type: Honduras, Atlantida, Yuncker, Koepper & Wagner 8461 (ILL!-holotype; F!,G!- isotypes). PIPER IN THE NEW WORLD 13 P. subaequilaterum Trel. in Publs Field Mus. not. Hist. (Bot.) 9: 280 (1940). Type: Honduras, Yuncker, Koepper & Wagner 8565 (Gl-isotype). P. amphibium Trel. in Ann. Mo. hot. Gdn 27: 288 (1940). Type: Woodson, Allen & Selbert 1222 (ILL!-holotype; MO!,NY!-isotypes). Shrubs 1-2 (-3) m high, stems densely yellow pubescent, with long multicellular hairs 0.2-1 mm long. Leaves 8-23 cm long, 3-14 cm wide, elliptic to obovate to rhombic, upper surface scabrous, densely long-pubescent, underside densely pale long-pubescent, apex acuminate, base unequally obtuse to rounded. Venation with 4-5 pairs of secondary veins, promi- nent on underside, arising from the lower part of the midrib, arcuate-ascending to apex. Petioles 5-15 (-20) mm long, densely pubescent, with ligule-like structure 0.5-1 mm long. Prophylls 8-16 mm long, acute, puberulent along midrib. Inflorescences 3-11 cm long, erect; peduncles 5-12 mm long, densely pubescent. Anthers 0.2-0.3 mm long. Floral bracts 0.2-0.5 mm wide, triangular, densely pale-ciliate. Fruits 0.7-1 mm wide, obovoid, obpyramidal-trigonous, glabrous; stigmas 3, recurved. Partial shade on hillsides, roadsides, secondary vegetation, and forest edges; 0-1200 m. DISTRIBUTION. Mexico to Venezuela. Mexico, Chiapas: Breedlove 9959 (F); Escuintla: Matuda 16719 (F). Belize, Belize: Dwyer 11488 (MO); Cayo: Gentle 2261 (F); Stann Creek: Spellman 1607 (F). Guatemala, Alta Verapaz: Stand- ley 91745 (F); Escuintla: Standley 64176 (F); Jutiapa: Stand- ley 77560 (F); Peten: Ortiz 1368 (MO); Santa Rosa: Standley 79270 (F). El Salvador, San Vicente: Standley & Padilla 3598 (F). Honduras, Atlantida: Standley 56600 (F); Comayagua: Molina 7036 (F); Olancho: Standley 18175 (F). Nicaragua, Matagalpa: Williams et al. 24010 (F); Zelaya: Seymour 3262 (F). Costa Rica, Guanacaste: Daudenmire 114 (F); Puntare- nas: Burger & Liesner 6534 (F); San Jose: Burger 7618 (F). Panama, Colon: Antonio 1789 (MO); Panama: Greig 204 (BM). Venezuela, Tachira: Berti & Sanchez 982-132 (G). 63. Piper bredemeyeri Jacq., Ed. pi. rar. 1: 125 (1815). Type: Venezuela, Caracas, Bredemeyer s.n. (W-holo- type). Fig. 4A,a,b P. radula Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp. 1: 148 (1816). Type: Venezuela, habitat in Provincia Cara- cas, Humboldt & Bonpland s.n. (B-holotype). P. flexuosum Jacq., Ed. pi. rar. 1: 139 (1816). Type: Venezu- ela, Caracas, Bredemeyer s.n. (W-holotype). Steffensia flexuosa (Jacq.) Kunth in Linnaea 13: 644 (1839). S. radula (Kunth) Kunth in Linnaea 13: 650 (1839). Artanthe radula (Kunth) Miq., Syst. piperac.: 426 (1844). A. flexuosa (Jacq.) Miq., Syst. piperac.: 454 (1844). Piper pseudopsisC. DC. in An. Inst.fis.-geogr. C. Rica. 9: 164 (1897). Type: Costa Rica, San Jose, Tonduz 1088, 2847 (G!-syntypes). P. villibracteum C.DC. in Annu. Conserv. Jard. hot. Geneve 21: 233 (1920). Type: Colombia, Santa Marta, Smith 384 (G holotype; BM!-isotype). P. pelliticauleTre\. in Contr. U.S. natn. Herb. 26: 157 (1929). Type: Costa Rica, Alajuela, Standley & Torres 47539 (US- holotype). P. alveolatifolium Trel. in/. Wash. Acad. Sci. 19: 329 (1929). Type: Honduras, Comayagua, Standley 56344 (F-holo- type; ILL!- isotype). P. copacabanense Trel. in Caldasia 1: 88 (1940). Type: Colombia, Copacabana, near Medellin, Daniel 279 (US!- holotype; ILL!- isotype). Shrubs 1-3 m high, stems densely pubescent with yellowish hairs 0.5-1 mm long. Leaves 10-26 cm long, 4-10 cm wide, broadly to narrowly ovate-lanceolate, upper surface bullate, scabrous, hirsute, underside densely brown-pubescent on veins, dark green, apex acuminate, base unequally rounded. Venation with 4-7 pairs of secondary veins arising from the lower part of the midrib, ascending steeply to apex. Petioles 8-18 mm long, densely puberulent, with a ligule-like struc- ture 8 mm long. Prophylls 10-25 mm long, drying dark brown. Inflorescences 6-12 cm long, erect; peduncles 12-24 mm long, densely pubescent. Anthers 0.3-0.4 mm long. Floral bracts 0.8 mm wide, triangular to semi-lunar, densely yellow-ciliate. Fruits 1 mm wide, obovoid, trigonous, glabrous or sparsely to densely pale-puberulent; stigmas 3. Thickets, forest edges, track- and creek-sides; 800-2000 m. DISTRIBUTION. Guatemala to Venezuela. Guatemala, Chiquimula: Molina 25163 (NY); Jalapa: Standley 76781 (F). Honduras, Morazan: Sanchez 83 (BM); Ocotepeque: Molina 22387 (F); Paraiso: Molina 7667 (F). El Salvador, Ahua- chapan: Standley & Padilla: 2698 (F); Santa Ana: Croat 42284 (MO). Nicaragua, Jinotega: Williams et al. 27491 (F); Matagalpa: Williams et al. 23464 (NY). Costa Rica, Alajuela: Smith 1692 (NY); Cartago: Lent 3646 (F); San Jose: Utley 917 (MO). Colombia, Antioquia: Archer 780 (BM); Cauca: Leh- mann 3807 (BM). Venezuela, Trujillo: Alston 6488 (BM). Piper bredemeyeri can be identified by its rough, bullate, ovate-lanceolate leaves, long hairs, triangular to semi-lunar, yellow-ciliate floral bracts, and trigonous fruits. 64. Piper dilatatum Rich, in Act. Soc. Hist. not. Paris 1: 105 (1792). Type: Guadelupe, Bertero s.n. (G-holotype). Fig. 4B,c,d. P. argillicola C.DC. in Annu. Conserv. Jard. hot. Geneve 21: 37 (1920). Type: Colombia, Cordillera Occidentale, Langlass 52 (G!-holotype). P. subsericeum Trel. in Contr. U.S. natn. Herb. 26: 141 (1929). Type: Costa Rica, below Cairo, Limon, Standley & Valeria 48636 (US-holotype). P. echeverrianum Trel. in Contr. U.S. natn. Herb. 26: 172 (1929). Type: Costa Rica, Echeverria, Pittier 2547 (US- holotype). P. obiter-sericeum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 350 (1937). Type: Costa Rica, El Gen- eral, Skutch 2865 (US-holotype; MO!,NY!-isotypes). P. triquetrofructum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 366 (1937). Type: Costa Rica, Rio Pejivalle gorge, Dodge & Thomas 4431 (GH-holotype; MO!- isotype). Spindly shrubs 1-3 m high, stems sparsely to densely puberu- lent. Leaves 11-20 cm long, 4-8 (-10) cm wide, narrowly ovate to elliptic, upper surface smooth or slightly scabrous, underside puberulent, dark green, apex acute-acuminate, base unequally obtuse to rounded. Venation with 3-4 pairs of secondary veins arising from the lower part of the midrib, 14 M.C. TEBBS Fig. 4 A: P. bredemeyeri, habit, a: prophyll; b: fruit and bract. B: P. dilatatum, habit; c: prophyll; d: fruit and bract. C: P. thomasii, habit; e: prophyll; f: fruit and bract. PIPER IN THE NEW WORLD 15 arcuate ascending to apex. Petioles 4-15 mm long, glabrous to puberulent, with ligule-like structure 0.5-2 mm. Prophylls 8-20 mm long, acute, puberulent along midrib. Inflores- cences 6-10 cm long, erect; peduncles 5-10 mm long. Anthers 0.2-0.3 mm long. Floral bracts 0.5-0.7 mm wide, triangular to semi-lunar, pale ciliate. Fruits 0.6-0.8 mm wide, obovoid, trigonous, usually glabrous; stigmas 3, in slight depression. Forest edges, roadsides, river and stream sides; 0-1700 m. DISTRIBUTION. Mexico to Paraguay, W. Indies. Mexico, Cordoba: Bourgeau s.n. (G). Honduras, Atlantida: Yuncker et al. 8461 (MO). El Salvador, Sonsonate: Tucker 1360 (F). Costa Rica, Guanacaste: Williams et al. 26609 (F); Puntare- nas: Lent 3068 (MO). Panama, Chiriqui: Croat 22469 (MO); Colon: Antonio 1940 (MO); Los Santos: Lewis et al. 1549 (MO); Panama: Croat 6997 (MO). Brazil, Bahia: Beta & Hage 219 (K); Mato Grosso: Berg et al. P18507 (K). Para- guay, Sierra de Amambay: Hassler 11039 (K). 65. Piper thomasii Tebbs, sp. nov. Type: Panama, south of El Cope sawmill, Code, 1700 m, Hammel 4157 (MO!- holotype). Fig. 4C,e,f. Fruticulus epiphyticus, caulibus rubris glandulosis pilosis. Folia elliptica coriacea glandulosa petiolis glandulosis, basi pilis flavidis longiusculis induta. Epiphytic subshrubs 1-1.5 m high, stems reddish, glandular, young stems with yellowish-white multicellular hairs. Leaves 5-9 cm long, 2-5 cm wide, elliptic to elliptic-lanceolate, coriaceous, underside glabrous or minutely pubescent, with brown or black glands, apex acuminate, base acute to cuneate. Venation with 2-3 pairs of secondary veins arising from the lower to middle part of the midrib, curving to the apex. Petioles 4-8 (-10) mm long, glandular, usually with long yellowish hairs clustered at the base, obscuring a ligule- like structure 0.5 mm long. Prophylls 5-8 mm long, glandu- lar, glabrous to sparsely pubescent. Inflorescences 2.5-5 cm long, erect; peduncles 6-10 mm long, glabrous to sparsely pubescent, glandular. Anthers 0.1-0.2 mm long. Floral bracts 0.8-1 mm wide, broadly triangular or crescent-shaped, densely yellowish white ciliate. Fruits 1-1.2 mm wide, obo- void, trigonous from above, glabrous, glandular; stigmas 3. Cloud forest, forested slopes, along trails and ridges; 800-1700 m. DISTRIBUTION. Panama. Panama, Bocas del Toro: Croat 69205 (BM,MO); Chiriqui: Hampshire & Whitefoord 161 (BM); Code: Croat 67262 (BM,MO); Panama: McPherson 7438 (BM,MO). The epiphytic habit of this species is unusual in Piper. Piper thomasii can also be identified by its reddish, glandular, long-pubescent stems, glandular coriaceous leaves, and glan- dular petioles with clustered yellowish hairs at the base. It is a species of high forest, endemic to Panama. 66. Piper terrabanum C.DC. in Bull. Soc. r. Bot. Belg. 30 (1): 217 (1892). Type: Costa Rica, Pittier 3604 (G!- holotype). Fig. 5A,a,b. P. dilatatum var. acutifolium C.DC. in Bull. Soc. r. Bot. Belg. 30(1): 217 (1892). Type: Costa Rica, Pittier 3385 (G!- holotype). P. cydophyllum C.DC. in An. Inst. fis.-geogr. C. Rica 9: 167 (1897). Type: Costa Rica, Pittier s.n. (G!-holotype). P. laevifolium C.DC. in An. Inst. fis.-geogr. C. Rica 9: 169 (1897), non Blume (Art. 50c). Type: Costa Rica, Pittier 8586 (G!-holotype). P.falcigerum Trel. in Contr. U.S. natn. Herb. 26: 147 (1929). Type: Costa Rica, El Silencio, near Tilaran, Standley & Valeria 44747 (US-holotype). P. sinuatifolium Trel. in Contr. U.S. natn. Herb. 26: 147 (1929). Type: Costa Rica, Standley & Valeria 45537 (US-holotype; ILL!- isotype). P. sublaevifolium Trel. in Contr. U.S. natn. Herb. 26: 147 (1929). Type: Costa Rica, Boca de Zhorquin, Tonduz 8586 (US- holotype). P. auriculiferum Trel. in Contr. U.S. natn. Herb. 26: 156 (1929). Type: Costa Rica, El Silencio, near Tilaran, Standley & Valerio 44648 (US-holotype). P. celatipetiolum Trel. in Contr. U.S. natn. Herb. 26: 156 (1929). Type: Costa Rica, Capulfn, Alajuela, Standley 40178 (US- holotype). P. disparifolium Trel. in Contr. U.S. natn. Herb. 26: 156 (1929). Type: Costa Rica, Pejivalle, Standley & Valerio 47082 (US- holotype). P. anisophyllum Trel. in Contr. U.S. natn. Herb. 26: 157 (1929). Type: Costa Rica, El General, Pittier 3385 (US!- holotype). P. celatipetiolum var. brenesiiTrel. in Cufod., Arch. hot. Sist. Fitogeogr. Genet. 10: 26 (1934). Type: Costa Rica, Paci- fica, Brenes 80 (ILL! -holotype). P. anisophyllum var. granulatum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot). 18: 331 (1937). Type: Costa Rica, El General, Skutch 2740 (US-holotype; NY!- isotype). P. verruculaepetiolum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 368 (1937). Type: Panama, Chan- guinola Valley, Bocas del Toro, Dunlap 494 (F!- holotype). P. wedelii Yunck. in Ann. Mo. hot. Gdn 37: 56 (1950). Type: Panama, near Chiriqui Lagoon, Wedel 1021 (MO- holotype). Shrubs 1.5-2 (-2.5) m high, glabrous or minutely puberulent. Leaves 15-24 (-30) cm long, 6-10 cm wide, ovate-elliptic to obovate, minutely puberulent on veins below, apex acumi- nate, base unequal, with one side slightly lobed. Venation with 3-6 pairs of secondary veins arcuate-ascending to apex. Petioles 5-12 mm long, glabrous or minutely puberulent, with minute ligule-like structure 0.5 mm long. Prophylls 20-30 mm long, minutely puberulent. Inflorescences erect, 6-12 cm long; peduncles 5-15 mm long, glabrous or minutely puberulent. Anthers 0.2-0.3 mm long. Floral bracts 0.3-0.8 mm wide, triangular, white-ciliate. Fruits 0.5-1 mm wide, obovoid, round- trigonous, glabrous, glandular; stig- mas 3, in slight depression. Shaded sites in forest, edges of roads or streams; 0-1000 m. DISTRIBUTION. Nicaragua to Panama. Nicaragua, Zelaya: Shank & Molina 4866 (F). Costa Rica, Alajuela: Croat 46940 (MO); Cartago: Skutch 4593 (NY); Heredia: Opler 1595 (MO); Puntarenas: Croat 35102 (MO); San Jose: Skutch 2740 (NY). Panama, Bocas del Toro: Kirkbride & Duke 630 16 M.C. TEBBS Fig. 5 A: P. terrabanum, habit; a: prophyll; b: fruit and bract. B: P. littorale, habit; c: prophyll; d: fruit and bract. C: P. otophorum, habit; e: prophyll; f: fruit and bract. PIPER IN THE NEW WORLD 17 (MO); Chiriqui: Croat 21947 (MO); Panama: Croat 27058 (MO); San Bias: Croat 16950 (MO); Veraguas: Mori & Kallunki4138(MO). Piper terrabanum is similar to P. dilatation Rich., but can be distinguished by its larger leaves and prophylls, floral bracts that are triangular, rather than semi-lunar as in P. dilatatum, and glandular fruits. 67. Piper littorale C.DC. in Pittier, Fl. Costaricensis 2: 252 (1898). Type: Costa Rica, bord de la mer, Porto Viejo, Talamanca, Tonduz 8736 (NY!-isosyntype). Fig. 5B,c,d. P. maternale Trel. in Publs Field Must. not. Hist. (Bot.) 18: 349 (1937). Type: Panama, Bocas del Toro, Cooper 210 (F-holotype). Shrubs 1-2 m high, stems sparsely to moderately pubescent, sometimes green or purple-maculate. Leaves 5-15 cm long, 2-6 cm wide, ovate to ovate-elliptic, slightly succulent, subco- riaceous when dry, upper surface glabrous, lustrous, under- side minutely puberulent, apex acuminate, base unequal, obtuse to slightly cordulate. Venation with 3-4 pairs of secondary veins arising from the lower to middle part of the midrib, curving to the apex. Petioles 3-7 mm long, puberu- lent, with ligule-like structure 0.5-1 mm long. Prophylls 5-10 mm long, puberulent along midrib, drying dark brown. Inflorescences 4-8 cm long, erect; peduncles 6-12 mm long, glabrous or minutely pubescent. Anthers 0.2 mm long. Floral bracts 0.5-0.7 mm wide, triangular, densely yellow-ciliate. Fruits 0.7-1 mm wide, trigonous from above, glabrous, glan- dular; stigmas 3. Near seashores, in shady places such as edges of woods or under palms; sea-level. DISTRIBUTION. Nicaragua to Panama. Nicaragua, Rio San Juan: Nelson 5270 (F). Costa Rica, Limon: Burger et al. 10358 (F,MO). Panama, Bocas del Toro: Wedel 2047 (MO); Colon: Croat 10038 (MO). Piper littorale can be found growing on the strand, often in the shade of palm trees. It can be distinguished by its often maculate stems, subcoriaceous, slightly succulent leaves, and glabrous, glandular fruits. 68. Piper otophorum C.DC. in Bull. Soc. r. Bot. Belg. 30(1): 220 (1892). Type: Costa Rica, bois de Siguirres, Pittier 3183 (G!-holotype). Fig. 5C,e,f. P. sperdinum C.DC. in Smithson. misc. Collns 71(6): 1 (1920). Type: Panama, San Bias, Pittier 4348 (US- holotype; G!-isotype). P. milciadesii Trel. & Yunck., Piperac. N. South Amer. 1: 314 (1950). Type: Colombia, El Valle, Killip & Dussan 34732 (US- holotype, photograph!). Shrubs (0.5) 1-2 m high, stems densely yellowish white pubescent. Leaves 14-24 cm long, 6-10 cm wide, asymmetri- cally elliptic to ovate, upper surface smooth or slightly scabrous, glandular, underside puberulent on veins, apex long-acuminate, base unequal, with one lobe distinctly auric- ulate, 1-2 cm across, covering the petiole. Venation with 3-5 pairs of secondary veins arising from the lower part of the midrib, arcuate-ascending to apex. Petioles 3-12 mm long, densely puberulent, with ligule-like structure 0.5 mm long. Prophylls 9-15 (-20) mm long, densely pale-pubescent. Inflo- rescences 6-12 cm long, erect, maroon or brown; peduncles 10-25 mm long, densely puberulent. Anthers 0.2-0.3 mm long. Floral bracts 0.3-0.8 mm wide, triangular, minutely ciliate. Fruits 0.5-0.8 mm wide, obovoid, obpyramidal- trigonous, glabrous, glandular, with a minute depression on upper surface; stigmas 2-3. Shade of moist forest, stream sides; 0-1200 m. DISTRIBUTION. Costa Rica to Colombia. Costa Rica, Limon: Gray urn et al. 8665 (BM,CR); San Jose: Greig 446 (BM). Panama, Bocas del Toro: Antonio 3126 (MO); Colon: Knapp & Sytsma 2394 (BM,MO); Darien: McDonagh et al. 617 (BM); Panama: Gentry & Nee 8648 (MO); San Bias: Nevers & Goncalez 3682 (BM,MO); Veraguas: Antonio 3544 (BM,MO). Colombia, Antioquia: Haught4630 (ILL). Piper otophorum can be recognized by its asymmetric leaves, which have one distinctive auriculate basal lobe, overlapping the petiole. The inflorescences are often brown or maroon. 69. Piper malpasoensis Tebbs, sp. nov. Type: Mexico, Oco- zocoaultla de Espinosa, Chiapas, 550 m, Breedlove & Thome 20762 (MO! -holotype). Fig. 6A,a,b. Frutex 1-2 m altus, caulibus albo-pilosis. Folia elliptico-ovata pilosa, basin auriculata, petiolo albo-piloso. Bacca trigona glabra. Shrubs 1-2 m high, stems white-puberulent. Leaves 8-12 cm long, 3.5-6 cm wide, elliptic-ovate, upper surface glabrous, underside with soft white hairs on veins, apex acute- acuminate, base narrowly unequally lobed, the lobes curved around and concealing the petiole. Venation with 3-5 second- ary veins arising steeply from the lower to middle part of the midrib, curving towards the apex. Petioles 1-4 mm long, white- pubescent, with glabrous ligule-like structure 0.5 mm long. Prophylls 8-10 mm long, narrow, acute, with narrow central strip of white hairs. Inflorescences 4-6 cm long, erect; peduncles 7-10 mm long, white-pubescent. Anthers 0.2-0.4 mm long. Floral bracts 0.3-0.6 mm wide, triangular, with pale centres, shortly white-ciliate. Fruit 0.8-1 mm wide, obovoid, trigonous to round from above, glabrous; stigmas 3. Steep moist ravines in lower montane rain forest; (50-) 100-1200 m. DISTRIBUTION. Southern Mexico. Mexico, Chiapas: Breed- love 34894 (MO); Tabasco: Conrad & Gallegos 2872 (MO, BM). P. malpasoensis can be recognized by its white-puberulent stems and leaves, short erect inflorescences on puberulent peduncles, and glabrous fruits. It is closely related to Piper enganyanum, but that species can be distinguished by its very narrow leaves with dark glands on the undersides, and different venation. The distribution for P. malpasoensis is limited to southern Mexico, whereas P. enganyanum is found in Panama and Colombia. 70. Piper enganyanum Trel. & Yunck. in Piperac. N. South Amer. 1: 292 (1950). Type: Colombia, along Rio Engana, El Valle, Killip 34750 (US-holotype; BM!-isotype). 18 M.C. TEBBS Fig. 6 A: P. malpasoensis, habit; a: prophyll; b: fruit and bract. B: P. enganyanum, habit; c: prophyll; d: fruit and bract. PIPER IN THE NEW WORLD Fig. 6B,c,d. Slender, spindly shrubs 1-2.5 (-3) m high; stems shortly pubescent. Leaves 6-10 cm long, 1-3 cm wide, lanceolate, upper surface glabrous, slightly scabrous, underside with minute dark glands and short hairs on veins, apex narrowly long-acuminate, base unequal, slightly to strongly auriculate, concealing the petioles. Venation with 3-4 pairs of secondary veins mostly arising from along the length of the midrib, loop-connecting to the apex. Petioles 1-3 mm long, pubes- cent, with a minute ligule-like structure 0.5 mm long, mostly obscured by short pale hairs. Prophylls 5-7 mm long, narrow, acute, pubescent along the midrib. Inflorescences 3-5 cm long, erect; peduncles 3-5 mm long, puberulent. Anthers 0.2-0.3 mm long. Floral bracts 0.5-0.7 mm wide, densely white-ciliate with pale, glabrous centres. Fruit 0.8-1 mm wide, obovoid, trigonous from above, glabrous, glandular; stigmas 3, recurved. Rich, moist forest; 200-700 m. DISTRIBUTION. Panama to Colombia. Panama, Chiriqui: Croat 22473 (F,NY). Piper enganyanum is easily recognized by its narrow, lanceo- late leaves with unequal, auriculate bases, concealing the short petioles. 71. Piper lanceifolium Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp. 1: 49 (1816). Type: Peru, Jaen de Bracam- oros, Humboldt & Bonpland s.n. (B-holotype). Fig. 7A,a,b. Artanthe lanceaefolia (Kunth) Miq., Syst. piperac.: 433 (1844). Piper lanceifolium var. acutiusculum C.DC. in DC., Prodr. 16(1): 317 (1869). Type: Ecuador, Jameson 380 (K!- holotype; BM!- isotype). P. friedrichsthalii C.DC. in DC., Prodr. 16(1):327 (1869). Type: Guatemala, Friedrichsthal 1424, (V-holotype; ILL!, K!-isotypes). P. linearifolium C.DC. in Linnaea 37: 355 (1872). Type: Costa Rica, Hacienda Santa Rosa, Oersted s.n. (G-holotype). P. pseudolanceaefolium Trel. in Contr. U.S. natn. Herb. 26: 170 (1929). Type: Costa Rica, Tonduz 12665 (US- holotype; G!- isotype). P. excultum Trel. in Publs Field Mus. nat. Hist. (Bot.) 13: 164 (1936). Type: Peru, San Martin, Llewelyn Williams 7461 (F-holotype; ILL!-isotype). P. lineatum var. leucolepidum Trel. in Publs Field Mus. nat. Hist. (Bot.) 13: 184 (1936). Type: Peru, Killip & Smith 22495 (US-holotype; ILL!-isotype). P. goergeri Trel. in Publs Field Mus. nat. Hist. (Bot.) 18: 344 (1937). Type: Costa Rica, Finca Castillo, Rio Reventa- zon, Dodge & Goerger 9421 (MO-holotype; ILL!- isotype). P. liratinerve Trel. in Ann. Mo. hot. Gdn 24: 186 (1937). Type: Panama, Chiriqui, Seibert 158 (MO!-holotype; ILL, NY!-isotypes). P. alveatum Trel. in Ann. Mo. hot. Gdn 27: 287 (1940). Type: Panama, Bocos del Toro, Woodson et al. 1837 (ILL!- holotype; NY!- isotype). P. arctilimbum Trel. in Ann. Mo. hot. Gdn 27: 288 (1940). 19 Type: Panama, Code, Miller 1813 (US-holotype; ILL!- isotype). Shrub or small tree 1-4 (-6) m high, stems sparsely to densely pubescent. Leaves 10-20 (-22) cm long, 2-8 cm wide, lance- olate to elliptic-lanceolate, greenish brown, upper surface softly pubescent, smooth to the touch, underside pubescent, apex acuminate, base narrow, slightly unequal. Venation with 4-7 pairs of sharply ascending secondary veins, promi- nent on underside. Petioles 4-9 (-12) mm long, with a ligule-like structure 1-2 mm. Prophylls 1^4 cm long, acute, sparsely to densely dull-white pubescent, glandular. Inflores- cences 5-16 (-18) cm long, arching; peduncles 0.5-3 (-5) cm long, pubescent. Floral bracts 0.8-1 mm wide, crescent- shaped, densely white-ciliate. Fruit 0.7-1 mm wide, obovoid, trigonous, glabrous; stigmas 3, linear. Moist forest, damp pastures, roadsides; 0-2800 m. DISTRIBUTION. Costa Rica to Peru. Costa Rica, Alajuela: Lent 1817 (NY); Cartage: Skutch 4620 (MO); Heredia: Godrey 66152a (MO); Limon: Burger & Liesner 6888 (NY); Puntarenas: Semple 81 (MO); San Jose: Skutch 3104 (MO). Panama, Chiriqui: Stern et al. 1063 (MO); Code: Allen 2003 (NY); Panama: Wilbur et al. 11892 (MO). Colombia, El Choco: Killip & Garcia 33579 (BM); Cundinamarca: Killip 33984 (BM); Narino: Ewan 16639 (BM); Valle: Killip 34904 (BM). Ecuador, Napo: Davis 346 (S); Pichincha: Balslev & Boom 2490 (B). Peru, Ayacucho: Killip & Smith 22451 (ILL); Loreto: Klug 3177 (B). P. lanceifolium can be distinguished from P. aduncum, which also has arching inflorescences, by its crescent-shaped bracts, trigonous fruits, and the smooth upper surfaces of its leaves. P. aduncum has extremely rough upper leaf surfaces, triangu- lar bracts, and round fruits. P. friedrichsthalii C.DC. has been reduced to synonymy under P. lanceifolium as it is considered merely to be a narrow-leaved form of this species. The specimens examined for this study show a wide variation in leaf width. The wide-leaved forms sometimes have a prominent stipular structure at the petioles, but this has also been found in plants with narrow leaves. The same type of pubescence is found on the leaves and the prophylls are glandular-pubescent. The inflorescence characters are identical, with crescent-shaped, white-fringed bracts and trigonous, glabrous fruits. 72. Piper aduncum L., Sp. pi. 1: 29 (1753). Type: Jamaica, Houston s.n. (BM!-?lectotype). Fig. 7B,c,d. P. angustifolium Ruiz & Pav., Ft. peruv. 1: 38 (1798). Type Peru, Ruiz Lopez & Pavon s.n. (Madrid-holotype, pho- tograph!). P. elongatum Vahl, Enum. pi. 1: 312 (1805). Type: Peru, Herb. Willd. 659 (B-holotype). P. celtidifolium Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp. 1: 50 (1816). Artanthe adunca (L.) Miq., Comm. phytogr.: 49 (1838). Artanthe elongatum (Vahl) Miq., Syst. piperac.: 434 (1844). A. galleoti Miq., Syst. piperac.: 451 (1844). Type: Mexico, Jalapa, Galleotti s.n. (G-holotype; K!-isotype). Steffensia adunca (L.) Kunth in Linnaea 13: 633 (1839). A. celtidifolia (Kunth) Miq., Syst. piperac.: 452 (1844). Piper pseudo-velutinum \ar.flavescens C.DC. in Bull. Soc. r. M.C. TEBBS Fig. 7 A: P. lanceifolium, habit; a: prophyll; b: fruit and bract. B: P. aduncum, habit; c: prophyll; d: fruit and bract. C: P. mollicomum, habit; e: prophyll; f: fruit and bract. PIPER IN THE NEW WORLD Bot. Belg. 30(1): 203 (1891). Type: Costa Rica, Rio Tileri, near San Jose, Pittier 3175 (G-holotype). P. kuntzei C.DC. in Kuntze, Revis. gen. 3 (2): 274 (1898). Type: Bolivia, Velasco, Kuntze s.n. (S-holotype; G!-isotype). P. fatoanum C.DC. in Smithson. misc. Collns 71(6): 7 (1920). Type: Panama, Colon, Pittier 3935 (US-holotype). P. aduncum var. laevifolium C.DC. in Smithson. misc. Collns 71(6): 8 (1920). Type: Panama, Porto Bello, Colon, Pittier 2438 (US-holotype). P. elongatum var. brachyarthrum Trel. in Contr. U.S. natn. Herb. 26: 37 (1927). Type: Panama, Chiriqui, Maxon 5139 (US- holotype). P. disparispicum Trel. in Contr. U.S. natn. Herb. 26: 170 (1929). Type: Costa Rica, Escasi, San Jose, Standley 32319 (US- holotype). P. aduncifolium Trel. in Contr. U.S. natn. Herb. 26: 171 (1929). Type: Costa Rica, Carrillo, Pittier 1196 (US- holotype). P. anguilliispicum Trel. in Contr. U.S. natn. Herb. 26: 175 (1929). Type: Costa Rica, near San Ramon, Brenes 14197 (US- holotype). P. oblanceolatum \ar.fragilicauleTrel. in Contr. U.S. natn. Herb. 26: 175 (1929). Type: Costa Rica, El Arenal, Guanacaste, Standley & Valerio 45228 (US-holotype). P. submolle Trel. in Contr. U.S. natn. Herb. 26: 178 (1929). Type: Costa Rica, Carmen, Limon, Standley & Valerio 48363 (US- holotype). P.flavescens (C.DC.) Trel. in Contr. U.S. natn. Herb. 26: 184 (1929). Type: Costa Rica, Rio Tileri near San Jose, Tonduz 3175 (US-holotype). P. elongatifolium Trel. in Publs Field Mus. not. Hist. (Bot.) 13: 161 (1936). Type: Peru, Junin, Macbride 5287 (F!- holotype; ILL-isotype). P. elongatum var. pampayacusum Trel. in Publs Field Mus. not. Hist. (Bot.) 13: 162 (1936). Type: Peru, Macbride 5047 (F-holotype; ILL!-isotype). P. lineatum var. hirtipetiolatum Trel. in Publs Field Mus. nat. Hist. (Bot.) 13: 184 (1936). Type: Peru, Macbride 4182 (F-holotype; ILL.'-isotype). P. cuatrecasasiiTrel. in Trab. Mus. Cienc. Nat., Madrid, Ser. Bot. 33: 48 (1936). Type: Colombia, Tolima, Cuatrecasas 2426 (ILL!-holotype). P. cumbricola Trel. in Trab. Mus. Cienc. Nat., Madrid, Ser. Bot. 33: 48 (1936). Type: Colombia, La Cumbre, Cua- trecasas 2012, 2427 (ILL-syntypes). P. illudensTrel. in Trab. Mus. Cienc. Nat., Madrid, Ser. Bot. 33: 50 (1936). Type: Colombia, Ibague, Cuatrecasas 2424 (ILL!-holotype). P. intersitum Trel. in Caldasia 1: 86 (1940). Type: Colombia, Antioquia, Daniel 890 (US-holotype). P. aduncum var. brachyarthrum (Trel.) Yunck. in Ann. Mo. hot. Gdn 37: 32 (1950). Shrub or small tree 2-5 (-8) m high, stems sparsely to densely pubescent. Leaves 12-20 cm long, (3-) 5-9 cm wide, oblong- elliptic to lanceolate, yellow-green, upper surface of leaf scabrous, sometimes rugose or bullate, underside pubescent, apex acuminate, base unequally rounded to obtuse, occasion- ally narrowly lobed. Venation with 4-7 pairs of steeply ascending secondary veins. Petioles 2-5 (-8) mm long, pubes- cent, vaginate at base, with a small ligule-like structure 0.5-1 mm long, sometimes reduced to a rim of hairs. Pro- phylls 20-25 mm long, acute, pubescent. Inflorescences 21 5-17 cm long, arching; peduncles 8-15 mm long, sparsely pubescent. Anthers 0.2-0.3 mm long. Floral bracts 0.4-0.7 mm wide, triangular-round, densely yellow-white cil- iate. Fruits 0.8-1 mm wide, obovoid, round from above, glabrous; stigmas 3. Open or disturbed areas, roadsides and edges of woods and streams; 0-1500 m. DISTRIBUTION. Mexico to Brazil, West Indies. Mexico, Chia- pas: Croaf 40344 (MO); Oaxaca: Martinez 883 (B); Veracruz: Rosas 716 (K); Yucatan: Gentle 1533 (K). Belize, Cayo: Croat , 23652 (CAS). Guatemala, Alta Verapaz: Croat 41713 (MO); Izabal: LeDoux et al. 2108 (NY); Peten: Contreras 9904 (K); Santa Rosa: Standley 60467 (F). Honduras, Colon: Bados 105 (BM); Cortes: Croat 42712 (MO); Comayagua: Yuncker et al. 5698 (MO); Morazan: Williams 23286 (CAS); Ocotepe- que: House 031 (BM); Olancho: Corat & Hannon 64179 (MO). El Salvador, San Salvador: Standley 22451 (NY); Sonsonate: Standley 23476 (NY). Nicaragua, Volcan Mom- bacho: Baker 175 (CAS). Costa Rica, Alajuela: Liesneret al. 3467 (MO); Puntarenas: Raven 21726 (DS); San Jose: Brenes 21351 (NY). Panama, Panama: Miller et al. 855 (MO). Colombia, Bogota: Triana 1820 (BM); Choco: Croat 56619 (BM,MO); El Meta: Killip 34357 (BM); Santa Marta: Smith 1229 (BM); Valle: Alston 7967 (BM). Venezuela, Aragua: Alston 5342 (BM); Carabobo: Alston 5738 (BM). Guyana, Mazarumi: Jenman 5290 (BM). Ecuador, Chimborazo: Camp 3244 (BM,NY); Esmeraldas: Jdtiva & Epling 891 (S); Los Rios: Holm-Nielson et al. 2887 (S); Napo-Pastaza: Mexia 7137 (BM). Peru, Chachapoyas: Wurdack 1792 (S); Cuzco: Mexia 8025 (BM); Huanuco: Mexia 8121 (BM); Lima: Mexia 8094 (BM). Bolivia, Nor Yungas: Mexia 7793 (BM). Brazil, Minas Gerais: Mexia 4973 (B); Montenegro: Sehnem 5042 (B); Rio de Janeiro: Bailey 65 (BM). Piper aduncum is a widespread species which can be found from Mexico to Brazil. It is easily recognizable by its arching inflorescences, extremely rough upper surfaces of the leaves, triangular bracts, and round, glabrous fruits. P. lanceifolium also has arching inflorescences, but has smooth upper leaf surfaces, semi-lunar bracts, and trigonous fruits. P.margina- tum, with arching, and P. cinereum with slightly curving inflorescences, have very different leaf venation and fruits. 73. Piper mollicomum Kunth in Linnaea 13: 648 (1839). Type: Brazil, Sellow s.n. (BM!-holotype). Fig. 7C,e,f. Artanthe mollicoma (Kunth) Miq., Syst. piperac.: 438 (1844). Piper pisoense C.DC. in DC., Prodr. 16(1): 278 (1869). Type: Brazil, Riedel 2 (LE-holotype). P. pseudovelutinum C.DC. in DC, Prodr. 16(1): 282 (1869). Type: Brazil, Lhotzky s.n. (G!-holotype). P. olivaceum C.DC. in Linnaea 37: 344 (1872). Type: Brazil, Corcovado, Warming s.n. (C!-holotype). Shrubs 1-2 (-3) m high, stems moderately to densely puberu- lent. Leaves 10-20 cm long, 4-8 (-10) cm wide, lanceolate- elliptic to ovate-elliptic, often rugose, upper surface puberulent, sometimes scabrous, underside dark glandular, densely pubescent, especially on veins, apex acute- acuminate, base unequally obtuse to cordulate. Venation with 4-6 pairs of secondary veins arising from the lower to middle part of the midrib, ascending steeply to the apex. 22 M.C. TEBBS Petioles 5-10 mm long, pale-puberulent. Prophylls 5-8 mm long, pale-puberulent. Inflorescences 7-15 cm long, erect or curved; peduncles (10-) 15-30 mm long, puberulent. Anthers 0.2-0.3 mm long. Floral bracts 0.5-0.8 mm wide, triangular- round, sparsely to densely pale-ciliate. Fruits 0.8-1 mm long, round from above, glabrous, sometimes glandular; stigmas 3. Wooded slopes, banks and track-sides, in partial shade; 0-1500 m. DISTRIBUTION. Colombia, Venezuela, Brazil. Colombia, Bogota: Miguel 1 (ILL); Santander: Killip & Smith 16406 (ILL); Valle: Killip & Hazen 8352 (ILL). Venezuela, Bolivar: Killip 37255 (ILL). Brazil, Ceara: Gardner 1848 (BM); Minas Gerais: Mexia 4617 (BM); Rio de Janeiro: Peckolt 1934 (ILL). Piper mollicomum can at first be mistaken for P. aduncum, with similarly shaped leaves, sometimes curving inflores- cences, and round, glabrous fruits. The inflorescences, how- ever, do not have the distinctive arch of those of P. aduncum. Piper mollicomum can also be distinguished by its generally smaller habit, much more puberulent leaves, without the extremely rough texture of those of P. aduncum, much shorter prophylls, longer peduncles, and often glandular fruits. 74. Piper achoteanum Trel. in /. Wash. Acad. Sci. 19: 328 (1929). Type: Honduras, El Achote near Siguatepeque, Standley 56125 (F!-holotype). Fig. 8A,a,b. P. pictamentum Trel. in Publs Field Mus. nat. Hist. (Bot.) 17: 352 (1938). Type: Honduras, vicinity of Siguatepeque, Yuncker, Dawson & Youse 5902 (ILL!-holotype; MO!- isotype). P. pictamentum var. fluminis Trel. in Publs Field Mus. nat. Hist. (Bot.) 17: 352 (1938). Type: Honduruas, river bank near Siguatepeque. Yuncker, Dawson & Youse 5718 (ILL!-holotype; MO!-isotype). Shrubs 1-2 (-3) m high, stems pinkish especially at nodes, densely pubescent with yellowish white hairs. Leaves 5-12 cm long, 3-6 cm wide, narrowly ovate-cordate, scabrous above, pubescent on underside, yellowish green, apex blunt, base unequally attached to petiole, lower part slightly lobed or cordate. Venation with 3-5 secondary nerves arisng from the lower to middle part of midrib, arising fairly steeply to apex. Petioles 0-3 mm long, pubescent, concealed by lower leaf-base, with ligule-like structure 0.5-1 mm long. Prophylls 6-12 mm long, pale green, pubescent. Inflorescences 5-10 cm long, erect or slightly curved; peduncles 6-10 mm long, pubescent. Anthers 0.2-0.3 mm long. Floral bracts 0.5-0.8 mm wide, centres pale, yellowish white ciliate. Fruits 0.8-1 mm wide, obovoid, round from above, glabrous or minutely granular; stigmas 3, sessile. Rocky hillsides, river banks, subalpine bogs, moist areas in pine forest; 800-1500 m. DISTRIBUTION. Mexico to Honduras. Mexico, Chiapas: Matuda 4474 (NY). Guatemala, Amatitlan: Morales 1165 (F). Honduras, Comayagua: Yuncker et al. 5902 (F); El Paraiso: Standley 25692 (F); Morazan: Rodriguez 499 (F). This species is similar in some ways to P. aduncum, with yellow-green leaves with surfaces that are extremely rough to the touch, pale-ciliate bracts, and round, mostly glabrous fruits. P. achoteanum differs mainly by its smaller, ovate- lanceolate leaves with small basal lobes concealing the short petioles, 6-12 mm long prophyll, and erect or only slightly curving spikes. It is also grows mainly above 800 m, while P. aduncum can be found in lowland areas. 75. Piper fuligineum Kunth in Linnaea 13: 655 (1939). Type: Brazil, Sellow s.n. (B?-holotype). Fig. 8B,c,d. Steffensia fuliginea (Kunth) Kunth in Linnaea 13: 655 (1839). Artanthe amplectans Miq. in Linnaea 20: 155 (1847). Type: Brazil, Minas Gerais, Martins s.n. (M-holotype). Piper palustre C.DC. in DC., Prodr. 16(1): 293 (1869). Type: Brazil, Riedel 125 (K!-isotype). P. amplectans (Miq.) C.DC. in DC., Prodr. 16(1): 293 (1869). Stout herbs or slender shrubs 0.7-1 (-2) m high; stems ridged, pale-pubescent. Leaves 5-15 cm long, 3-7.5 cm wide, broadly to narrowly unequally ovate-cordate, yellowish green, upper surface extremely scabrous, underside crisply puberulent on veins, apex obtuse to moderately acuminate, base unequally cordate. Venation with 3-5 prominent sec- ondary veins mostly arising from the lower part of the midrib, curving steeply to the apex. Petioles 0-4 mm long, with a ligule-like structure 0.5-1 mm long, mostly hidden by hairs. Prophylls 5-8 mm long, pale pubescent, with glabrous mar- gins. Inflorescences 3-7 cm long, erect; peduncles stout, 20-60 mm long, sparsely pubescent. Anthers 0.2-0.3 mm long. Floral bracts 0.7-1 mm wide, densely yellow-ciliate, with yellowish brown, glabrous centres. Fruits 0.8-1 mm wide, round to trigonous from above, glabrous; stigmas 3. Damp ground, edges of streams and creeks, damp places in savanna; 700-1600 m. DISTRIBUTION. Surinam to Paraguay. Surinam, R. Palaime: Wessels Boer 843 (K). Brazil, Goias: Hunt & Ramos 6197 (K); Mato Grosso: Ratter et al. 1535 (K); Minas Gerais: Irwin et al. 26278 (K); Parana: Callejas et al. 1839 (K); Sao Paulo: Mattos 8215 (K). Paraguay, Sierra de Amambay: Hassler 10035 (K). Piper fuligineum is similar to P. achoteanum, with yellowish green leaves, pale yellow-ciliate floral bracts, and round fruits. It can be separated from that species by its much rougher leaf surfaces and shorter inflorescences on peduncles up to 6 cm long. 76. Piper flavidum C.DC. ex Donn.Sm. in Bot. Gaz. 19: 258 (1894). Type: Guatemala, Alta Verapaz, Smith 1744 (G!- holotype). Fig. 8C,e,f,g. Shrubs 0.5-1.5 m high, glabrous, nodes close together. Leaves 7-13 cm long, 0.7-3 cm wide, linear-lanceolate, sometimes falcate, both surfaces glabrous, apex acuminate, base acute to cuneate. Venation with 1-2 pairs of prominent veins arising from lower part of midrib and running parallel to apex, with prominent cross venation. Petioles 2-5 mm long, glabrous, with ligule-like structure 1 mm long. Prophylls 7-10 mm long, glabrous. Inflorescences 2-3 cm long, erect; peduncles 2-6 (-10) mm long. Anthers 0.2 mm long. Floral PIPER IN THE NEW WORLD 23 Fig. 8 A: P. achoteanum, habit; a: prophyll; b: fruit and bract. B: P. fuligineum, habit; c: prophyll; d: fruit and bract. C: P. flavidum, habit; e: narrow leaf; f: prophyll; g: fruit and bract. 24 M.C. TEBBS bracts 0.6-0.8 mm wide, triangular, with densely ciliate mar- gins. Fruits 0.8-1 mm wide, obovoid, round from above, densely pale-puberulent; stigmas 3, linear. Slopes and stream sides in moist forest; 250-1300 m. DISTRIBUTION. Mexico, Guatemala. Mexico, Chiapas: Breedlove 26545 (MO); Tabasco: Rosario & Cowan 2962 (CAS). Guatemala, Alta Verapaz: Tuerckheim 8564 (NY); Izabal: Matuda 3697 (F); Quiche: Skutch 1819 (F). This species is sometimes confused with the narrow-leaved forms of P. lanceifolium. However, the leaves of P. flavidum are often falcate, and have only 1-2 pairs of secondary veins with prominent cross-veining. The round, pale-puberulent fruits and triangular floral bracts further distinguish it from P. lanceifolium, which has trigonous, glabrous fruits and semi- lunar floral bracts. 77. Piper silvivagum C.DC. in An. Inst. fis.-geogr. C. Rica 9: 162 (1897). Type: Costa Rica: forets sur les bords du R. Zhorquin, Tonduz 8595 (US!-holotype; G!-isotype). Fig. 9C,e,f. P. vitibundum Trel. in Contr. U.S. natn. Herb. 26: 38 (1927). Type: Panama, Bocas del Toro, Dunlap 338 (US- holotype, photograph!). P. pseudo-albuginiferum Trel. in Contr. U.S. natn. Herb. 26: 165 (1927). Type: Costa Rica, Cartago, Standley & Valeria 50948 (US- holotype, photograph!). Scandent or climbing to 2 m high, stems slender, glabrous or shortly pale-pubescent on younger growth. Leaves 8-15 cm long, 2.5-6 cm wide, elliptic-lanceolate, upper surface gla- brous to slightly scabrous, underside puberulent on veins, apex acuminate, base unequally narrowly obtuse. Venation with 3-4 secondary veins arising from the lower to middle part of the midrib, ascending steeply to apex. Petioles 3-12 mm long, sparsely to densely pubescent, with a ciliate, ligule-like structure 1-2 mm long. Prophylls 5-15 mm long, minutely puberulent. Inflorescences 6-14 cm long, erect; peduncles 5-15 mm long, glabrous. Anthers 0.1-0.2 mm long. Floral bracts 0.3-0.7 mm wide, triangular to round, white-ciliate, with dark, glabrous centre. Fruits 0.7-1 mm wide, obovoid, minutely pale-puberulent; stigmas 3, sessile. Forest shade; 0-1400 m. DISTRIBUTION. Mexico, Costa Rica to Peru. Mexico, More- los: Croat & Hannon 65771 (MO). Costa Rica, Cartago: Lent 861 (NY); Heredia: Opler 1642 (MO); Limon: Burger 8489 (NY). Panama, Chiriqui: Hammel 2240 (BM). Ecuador, Esmeraldas: Mexia 7792 (B,BM). Bolivia, Nor Yungas: Mexia 7792 (BM). Peru, Cuzco: Mexia 8063 (BM,K); Mariscal Caceres: Schunke 3725 (F,NY); Loreto: Mexia 6382 (B). Piper silvivagum is one of a small group of species in Piper with a scandent or climbing habit. In section Radula there is only one other species with this habit, P. multiplinervium, which can be distinguished from P. silvivagum by its extremely vigorous growth, glabrous, ovate to cordate leaves with sheathing petioles, and yellow, scented inflorescences. P. silvivagum mostly occurs from Costa Rica southwards and only one specimen from Mexico has been positively identified as this species. 78. Piper pseudoasperifolium C.DC. in DC., Prodr. 16 (1): 318 (1869). Type: Mexico, Oaxaca, Boissier s.n. (G!- holotype). Fig. 9B,c,d. P. cartagoanum C.DC. in Linnaea 37: 350 (1872). Type: Costa Rica, Cartago, Oersted s.n. (S-holotype; C!-isotype). P. vestitifolium C.DC. in Bot. Gaz. 70: 183 (1920). Type: Guatemala, near Cajabon, Alta Verapaz, Cook & Griggs 651 (F-holotype; G!-isotype). P. indignum Trel. in /. Wash. Acad. Sci. 19: 333 (1929). Type: Honduras, Siguatepeque, Standley 55990 (F!- holotype) . P. micoense Trel. in /. Wash. Acad. Sci. 19: 334 (1929). Type: Guatemala, Sierra del Mico, Izabal, Kellerman 6715 (F!-holotype). P. sibunense Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 12: 408 (1936). Type: Belize, Gracie Rock, Gentle 1562 (MICH-holotype; MO!-isotype). Shrubs 1-3 m high, stems densely white- or yellow- pubescent. Leaves 8-22 cm long, 3-8 cm wide, ovate- lanceolate to elliptic-lanceolate, bullate, glandular, upper surface scabrous, shortly hispid, underside densely pubescent on veins, apex narrowly acuminate, base unequally rounded to obtuse. Venation with 4-5 pairs of secondary veins arising from the lower two-thirds of the midrib, curving to the apex. Petioles 4-10 mm long, densely pubescent, with hispid ligule- like structure 1-3 mm long. Prophylls 12-15 mm long, pubes- cent along midrib. Inflorescences 7-9 cm long, erect; peduncles 3-8 mm long, pubescent. Anthers 0.2-0.3 mm long. Floral bracts 0.7-0.9 mm wide, densely yellow- to white-ciliate. Fruits 0.8-1 mm wide, round from above, gla- brous; stigmas 3. Moist forest, thickets, pine woods; 0-1500 m. DISTRIBUTION. Mexico to Costa Rica. Guatemala, Alta Ver- apaz: Standley 69370 (F); Huehuetenango: Standley 82813 (F); Peten: Steyermark 46195 (F). 79. Piper villiramulum C.DC. in Smithson. misc. Collns 71: 11 (1920). Type: Panama, Loma de la Gloria near Fato, Colon, Pittier4083 (US-holotype; NY!-isotype). Fig. 9A,a,b. P. talamancanum Trel. in Contr. U.S. natn. Herb. 26: 173 (1929). Type: Costa Rica, Shirores, Talamanca. Tonduz 9274 (US-holotype; F-isotype). P. laevius (C. DC). Trel. in Contr. U.S. natn. Herb. 26: 174 (1929). P. comatum Trel. in Contr. U.S. natn. Herb. 26: 175 (1929). Type: Costa Rica, Rio Corazal, Tonduz 9932 (US- holotype). P. granulatum Trel. in Contr. U.S. natn. Herb. 26: 175 (1929). Type: Costa Rica, Buenas Aires, Pittier 3593 (US-holotype). P. leucophlebium Trel. in Contr. U.S. natn. Herb. 26: 176 (1929). Type: Costa Rica, Tilaran, Standley & Valeria 44277 (US-holotype; ILL!-isotype!). P. capacibracteum Trel. in Contr. U.S. natn. Herb. 26: 183 (1929). Type: Costa Rica, between Aserri and Tarbaca, Standley 41397 (US-holotype; F!,ILL-isotypes). P. cayoense Trel. ex Standl. in Publs Field Mus. nat. Hist. PIPER IN THE NEW WORLD 25 Fig. 9 A: P. villiramulum, habit; a: prophyll; b: fruit and bract. B: P. pseudoasperifolium, habit; c: prophyll; d: fruit and bract. C: P. silvivagum, habit; e: prophyll; f: fruit and bract. 26 (Bot.) 12: 407 (1936). Type: Belize, Cayo, Bartlett 13070 (MICH-holotype; KMsotype). P. tikalense Trel. in Publs Field Mus. nat. Hist. (Bot.) 17: 234 (1937). Type: Guatemala, Peten, Bartlett 12595 (ILL!- holotype). P. yalochanum Trel. in Publs Field Mus. nat. Hist. (Bot.) 17: 234 (1937). Type: Guatemala, Peten, Bartlett 12849 (ILL!-holotype). P. bocasense Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 333 (1937). Type: Panama, Changuinola Val- ley, Dunlap 234 (F!-holotype). P. yapeanum Trel. in Ann. Mo. hot. Gdn 25: 827 (1938). Type: Panama, Darien, Allen 351 (ILL!-holotype; MO!- isotype). P. gamboanum var. yapense Trel. in Ann. Mo. hot. Gdn 27: 291 (1940). Type: Panama, Yape, Darien, Allen 854 (ILL!-holotype). Shrubs or small trees (1-) 2-3 m high, stems usually densely whitish-pubescent. Leaves 7-24 cm long, 2-9 cm wide, ovate to elliptic-ovate or elliptic-lanceolate, upper surface scabrous, often rugose or bullate, sparsely pubescent, underside densely pubescent, with minute dark red or brown glands visible, apex acuminate, base unequally obtuse or rounded. Venation with 3-5 pairs of secondary veins arising from the lower part of the midrib and ascending fairly steeply to apex. Petioles 3-10 mm long, densely pubescent, with a minute, ligule-like structure up to 0.5 mm long, hidden by dense hairs. Prophylls 8-20 mm long, acute, pubescent with gla- brous margins. Inflorescences 6-11 cm long, erect; peduncles 4-8 mm long, sparsely to densely puberulent. Anthers 0.1-0.2 mm long. Floral bracts 0.3-0.6 mm wide, triangular, densely white-ciliate. Fruits 0.6-0.8 mm wide, obovoid, pubescent, with prominent glands on sides; stigmas 3, sessile. Shade at edge of forest and thickets, road or track sides; 0-1000 m. DISTRIBUTION. Mexico to Ecuador. Mexico, Chiapas: Croat 47174 (BM,MO); Nayarit: Croat 45346 (BM,MO); Veracruz: Croat & Hannon 63089 (BM,MO). Belize, Belize: Dwyer 12684 (MO). Guatemala, Alta Verapaz: Standley 70240 (F); Baja Verapaz: Lundell & Contreras 19653 (F); Izabal: Stevens et al. 25488 (BM); Peten: Lundell & Contreras 20535 (F); Quiche: Proctor 25209 (MO); Zacapa: Harmon & Fuentes 1862 (F). Honduras, Colon: Guerra 128 (BM); Islas de la Bahia: Molina 20737 (F); Olancho: Croat & Hannon 64143 (BM,MO). Nicaragua, Boaco: Stevens 5918 (BM,MO); Jinotega: Araquistain & Castro 1856 (F); Zelaya: Marshall & Neill 6473 (MO). Costa Rica, Alajuela: Molina et al. 17414 (F); Cartago: Rodriguez 426 (MU); Limon: Burger & Liesner 6873 (F); Puntarenas: Burger & Liesner 6575 (F); San Jose: Whitmore 54 (F). Panama, Bocas del Toro: Correa et al. 4004 (MO); Colon: Croat 67321 (BM); Panama: Atencio 24 (F); San Bias: Nevers & Herrera 4523 (MO). Colombia, Santa Marta: Smith 1228 (S). Ecuador, Esmeraldas: Fagerlind & Wibom s.n. (S). Piper villiramulum is often confused with P. hispidum. It can be distinguished by its rugose or bullate leaves, densely pubescent stems, petioles, and prophylls, and fruits with prominent glands. The petioles of P. hispidum have more prominent ligule-like structures than those of P. villiramu- lum, which are difficult to see amongst the dense pubescence. M.C. TEBBS 80. Piper hispidum Sw., Prodr. Veg. Ind. Occ.: 15 (1788). Type: Jamaica, Swartz s.n. (M!-holotype; BM!- photograph). Fig. 10A,a,b. P. scabrum Sw., Ft. Ind. occid. 1: 59 (1797). Type: Jamaica, Wiles s.n. (B?-holotype; BM!-photograph). P. hispidum Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp. 1:50(1816). Steffensia hirsuta (Sw.) Kunth in Linnaea 13: 640 (1839). 5. scabra (Sw.) Kunth in Linnaea 13: 640 (1839). Artanthe olfersiana Miq., Syst. piperac.: 445 (1844). Type: Brazil, Campos Vittoria, Bahia, Sellow s.n. (B-holotype). A. scabra (Sw.) Miq., Syst. piperac.: 446 (1844). Piper hirsutum var. tonduzii C.DC. in Bull. Soc. r. Bot. Belg.3Q(l): 203 (1892). Type: Costa Rica, Asseri, Pittier 1270 (G!-holotype). P. alluvicola C.DC. in Annu. Conserv. Jard. hot. Geneve 21: 313 (1920). Type: Mexico, Michoacan & Guerrero, Langlasse 664 (G!- holotype). P. erectamentum C.DC. in Smithson. misc. Collns 71(6): 10 (1920). Type: Panama, Los Siguas Camp, Pittier 3191 (US-holotype; G!-isotype). P. scabrilimbum C.DC. in Candollea 1: 121 (1923). Type: Panama, Pittier 5584 (not located). P. bayamonanum Trel. in Feddes Reprium Spec. nov. veg. 23: 9 (1926). Type: Cuba, Bayamon, Eggers 4689 (B-holotype). P. sabanillanum Trel. in Feddes Reprium Spec. nov. veg. 23: 9 (1926). Type: Cuba, Sabanilla, Ekman 10659 (EHH- holotype). P. sumideranum Trel. in Feddes Reprium Spec. nov. veg. 23: 9 (1926). Type: Cuba, Sumidero, Ekman 18211 (EHH- holotype). P. maestranum Trel. in Feddes Reprium Spec. nov. veg. 23: 10 (1926). Type: Oriente, Ekman 14347 (EHH- holotype). P. williamsii Trel. in Contr. U.S. natn. Herb. 26: 32 (1927). Type: Panama, Marraganti, Williams 986 (US-holotype; NY!- isotype). P. trachydermum Trel. in Contr. U.S. natn. Herb. 26: 33 (1927). Type: Panama, Sutton Hayes 791 (NY!- holotype). P. killipiTrel. in Contr. U.S. natn. Herb. 26: 33 (1927). Type: Panama, El Boquete, Killip 3549 (US-holotype, photo- graph!). P. killipivar. calderanumTrel. in Contr. U.S. natn. Herb. 26: 33 (1927). Type: Panama, Rio Caldera, Chiriqui, Killip 3544 (US-holotype, photograph!). P. cataractarum Trel. in Mem. N.Y. hot. Gdn 7: 224 (1927). Type: Bolivia, Bopi river, Rusby 672 (NY-holotype; ILL!-isotype). P. gonagricum Trel. in Contr. U.S. natn. Herb. 26: 171 (1929). Type: Costa Rica, Heredia, Standley & Valeria 49082 (US-holotype; ILL!-isotype). P. pejivallense Trel. in Contr. U.S. natn. Herb. 26: 171 (1929). Type: Costa Rica, Pejivalle, Standley & Valeria 46727 (US-holotype, photograph!). P. genuflexum Trel. in Contr. U.S. natn. Herb. 26: 172 (1929). Type: Costa Rica, Santa Rosa del Copey, Tonduz 11687 (US-holotype; NY!-isotype). P. pergeniculatum Trel. in Contr. U.S. natn. Herb. 26: 172 (1929). Type: Costa Rica, Canas Gordas, Pittier 11033 PIPER IN THE NEW WORLD 27 Fig. 10 A: P. hispidum, habit; a: prophyll; b: fruit and bract. B: P. sancti-felicis , habit; c: prophyll; d: bract and fruit. 28 M.C. TEBBS (US-holotype, photograph!). P. pavasense Trel. in Contr. U.S. natn. Herb. 26: 173 (1929). Type: Costa Rica, Las Pavas, San Jose, Pittier 3188 (US-holotype, photograph!). P. scalpens Trel. in Contr. U.S. natn. Herb. 26: 176 (1929). Type: Costa Rica, San Jose, Tonduz 10154 (US-isotype, photograph!). P. caudatifolium Trel. in Contr. U.S. natn. Herb. 26: 177 (1929). Type: Costa Rica, Las Pavas, Pittier 3191 (US- holotype). P. Inhorrescens Trel. in Contr. U.S. natn. Herb. 26: 177 (1929). Type: Costa Rica, Las Pavas, Standley 36083 (US-holotype). P. torresanum Trel. in Contr. U.S. natn. Herb. 26: 177 (1929). Type: Costa Rica, Rio Torres, San Francisco de Guadalupe, Tonduz & Pittier 8971 (US-holotype). P. trichophlebium Trel. in Contr. U.S. natn. Herb. 26: 177 (1929). Type: Costa Rica, Asserf, Tonduz 1270 (G!- holotype). P. valetudinari Trel. in Contr. U.S. natn. Herb. 26: 178 (1929). Type: Costa Rica, San Jose, Tonduz 7235 (NY!- isotype). P. carminis Trel. in Contr. U.S. natn. Herb. 26: 179 (1929). Type: Costa Rica, Carmen, Limon, Standley & Valeria 48383 (US- holotype). P. coronatibracteum Trel. in Contr. U.S. natn. Herb. 26: 179 (1929). Type; Costa Rica, Hamburg Finca, Limon, Standley & Valeria 48844 (US-holotype). P. baculiferum Trel. in Contr. U.S. natn. Herb. 26: 180 (1929). Type: Costa Rica, El Arenal, Standley & Valeria 45052 (US-holotype). P. punctiunculatum Trel. in Contr. U.S. natn. Herb. 26: 180 (1929). Type: Costa Rica, Tilaran, Standley & Valeria 45712 (US-holotype, photograph!). P. abuginiferum Trel. in Contr. U.S. natn. Herb. 26: 181 (1929). Type: Costa Rica, El Muneco, Cartago, Standley & Torres 51102 (US-holotype). P. injucundum var. praecalvinervium Trel. in Contr. U.S. natn. Herb. 26: 181 (1929). Type: Costa Rica, El Muneco, Cartago, Standley & Torres 51101 (US- holotype, photograph!). P. injucundum var. praepubinervium Trel. in Contr. U.S. natn. Herb. 26: 181 (1929). Type: Costa Rica, El Muneco, Cartago, Standley & Valeria 51067 (US- holotype, photograph!). P. lanatibracteum Trel. in Contr. U.S. natn. Herb. 26: 182 (1929). Type: Costa Rica, El Muneco, Cartago, Standley 33419 (US-holotype, photograph!). P. lanosibracteum Trel. in Contr. U.S. natn. Herb. 26: 182 (1929). Type: Costa Rica, Rio Reventado, Standley & Valeria 49457 (US-holotype, photograph!). P. phanerolepidum Trel. in Contr. U.S. natn. Herb. 26: 182 (1929). Type: Costa Rica, El Muneco, Cartago, Standley 33570 (US-holotype, photograph!). P. pullibracteatum Trel. in Contr. U.S. natn. Herb. 26: 182 (1929). Type, Costa Rica, La Estrella, Cartago, Standley 39236 (US-holotype, photograph!). P. curridabatanum Trel. in Contr. U.S. natn. Herb. 26: 183 (1929). Type: Costa Rica, between San Pedro Monies de Oca & Curridabat, San Jose, Standley 32787 (US- isotype). P. fusco-bracteatum Trel. in Contr. U.S. natn. Herb. 26: 183 (1929). Type: Costa Rica, Escasii, San Jose, Standley 32631 (US-holotype). P. rivi-vetusti Trel. in Ann. Mo. hot. Gdn. 24: 186 (1937). Type: Panama, Chiriqui, Seibert 192 (MO!-holotype). P. aquacalientisTrel. ex Standl. in Publs Field M us. nat. Hist. (Bot.) 18: 330 (1937). Type: Costa Rica, SW. of Agua Caliente, Stork 1317 (F-holotype). P. articulosum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 332 (1937). Type: Costa Rica, San Ramon to La Paz, Brenes 6060 (F!-holotype). P. humoense Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 346 (1937). Type: Costa Rica, Pejivalle Farm, Cartago, Dodge & Thomas 4371 (GH-holotype; MO!- isotype). P. subasperatum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 362 (1937). Type: Costa Rica, San Jose, Skutch 2157 (US-holotype; NY!-isotype). P. meritum Trel. in Publs Field Mus. nat. Hist. (Bot.) 9: 279 (1940). Type: Honduras, Atlantida, Yuncker, Koepper & Wagner 8363 (ILL!-holotype; MO!-isotype). P. barbirostre Trel. in Ann. Mo. bat. Gdn 27: 288 (1940). Type: Panama, Chiriqui, Allen 1365 (ILL!-holotype; MO!-isotype). P. casitense Trel. in Ann. Mo. hot. Gdn 27: 289 (1940). Type: Panama, Casita Alta, Chiriqui, Woodson, Allen & Seibert 978 (ILL!-holotype; NY!-isotype). P. cerro-puntoense Trel. in Ann. Mo. hot. Gdn 27: 289 (1940). Type: Panama, Chiriqui, White 52 (ILL!- holotype; MO!-isotype). P. konkintoense Trel. in Ann. Mo. bat. Gdn 27: 293 (1940). Type: Panama, Bocas del Toro, Woodson, Allen & Seibert 1894 (ILL!-holotype; NY!-isotype). P. margaretae Trel. in Ann. Mo. hot. Gdn 27: 293 (1940). Type: Panama, Chiriqui, White 158 (ILL!-holotype; MO!-isotype). P. pervicax Trel. in Ann. Mo. hot. Gdn 27: 296 (1940). Type: Panama, near Volcan de Chiriqui, Woodson, Allen & Seibert 843 (ILL!-holotype). P. argentamentum Trel. & Yunck. in Piper ac. N. South Amer. 1: 256 (1950). Type: Colombia, Istmina, Choco, Killip 35452 (US-holotype; BM!-isotype). P. rivialbi Trel. & Yunck. in Piperac. N. South Amer. 1: 265 (1950). Type: Colombia, Tio Digua Valley, El Valle, Killip 34723 (US-holotype; BM!-isotype). P. phaeophyllum Trel. in Fieldiana Bot. 24: 316 (1952). Type: Guatemala, above Tamahu, Alta Verapaz, Standley 70911 (Fl-holotype). Shrubs to 4 m high, stems sparsely to densely white- pubescent. Leaves 7-18 (-21) cm long, 3-10 cm wide, ovate to elliptic-oblong, yellowish green, scabrous above, some- times with short whitish hairs, underside sparsely to densely pubescent, especially on veins, apex acuminate, base unequally obtuse or rounded. Venation with 3-5 pairs of secondary veins curving to apex. Petioles 4-18 mm long, sparsely to densely pubescent, with a glandular, sometimes ciliate ligule-like structure 1-3 mm long. Prophylls 6-20 mm long, acute, sparsely to densely pubescent. Inflorescences 6-12 cm long, slender, erect or slightly curving; peduncles 4-20 mm long. Anthers 0.2 mm long. Floral bracts 0.3-0.6 mm wide, triangular to round, densely white-ciliate. Fruits 0.8-1 mm wide, obovoid, oblong to round from above, pubescent; stigmas 3, sessile. Disturbed areas, moist thickets, edges of woods and track- sides; 0-2000 m. PIPER IN THE NEW WORLD DISTRIBUTION. Throughout tropical America. Mexico, Chia- pas: Breedlove 6194 (F); Nayarit: Miller & Tellez 3239 (MO). Belize, Toledo: Gentle 7291 (F). Guatemala, Baja Verapaz: Contreras 11269 (K); Huehuetenango: Standley 81395 (F); Izabal: Lundell & Contreras 19346 (K); Peten: Contreras 9835 (K); Retalhuleu: Standley 66691 (F). Honduras, Comay- agua: Molina 7111 (F); Copan: Molina 11749 (F); Morazan: Molina 6125 (F); Paraiso: 11315 (F). Nicaragua, Managua: Gamier 1669 (F). Costa Rica, Alajuela: Molina et al. 17123 (F); Cartago: Utley 801 (CR); Guanacaste: Chacon 2170 (MO); Heredia: Croat 68396 (MO); Limon: Burger 8444 (F); Puntarenas: Liesner 3068 (MO); San Jose: Greig 442 (BM). Panama, Code: Croat 67464 (BM); Darien: Garwood et al. 2056a; Panama: Greig 214 (BM). Colombia, Choco: Kittip 35034 (BM); El Meta: Killip 34463 (BM); El Valle: Killip 35564 (BM). Venezuela, Bolivar: Croat 53979 (MO), Cara- bobo: Alston 5753 (BM); Caracas: Linden 121 (BM); Lara: Davidse & Gonzalez 21058 (BM); Orinoco basin: Rusby 58 (BM). Ecuador, Esmeraldas: Mexia 8415 (B); Leon: Mexia 6726 (BM); Los Rfos: Mexia 6631 (BM); Napo-Pastaza: Mexia 7157 (BM). Peru: Mathews 1706 (BM). Bolivia, Yun- gas: Buchtien 753 (BM). Brazil, Matogrosso: Moore 123 (BM); Para: Huber 1348 (BM). Probably the most confused and confusing of the taxa in this section and often difficult to identify, especially without mature inflorescences. The vegetative plasticity of P. hispi- dum has given rise to the description of numerous taxa. Many of these have now been reduced to synonymy, mainly by Trelease & Yuncker (1950). The main characters of this species are scabrous, ovate-elliptic leaves (often pale or yellowish green in colour especially when dry), petioles with distinct, often ciliate ligule-like structures, acute prophylls, slender, erect or slightly curving inflorescences, and small, obovoid, pubescent, often green fruits. Piper villiramulum and P. sancti-felicis are probably the most closely related species to P. hispidum, but can be separated by the densely pubescent stems, rugose, glandular leaves and fruits of P. villiramulum, and the long prophylls and ligule-like structures drying dark brown of P. sancti-felicis. 81. Piper sancti-felicis Trel. in Contr. U.S. natn. Herb. 26: 35 (1927). Type: Panama, Chiriqui, Pittier 5124 (US- holotype, photograph!). Fig. 10B,c,d. P. spicilongum Trel. in Contr. U.S. natn. Herb. 26: 177 (1929). Type: Costa Rica, Santo Domingo de Golfo Dulce, Tonduz 9962 (US-holotype; F!-isotype). P. rectamentum Trel. in Contr. U.S. natn. Herb. 26: 180 (1929). Type: Costa Rica, Guanacaste, Standley & Vale- ria 45237 (US-holotype, photograph!). P. fraguanum Trel. in /. Wash. Acad. Sci. 19: 332 (1929). Type: Honduras, La Fragua, Standley 55730 (F!- holotype). P. tentatum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 365 (1937). Type: Costa Rica, Alajuela, Brenes 15032 (F-holotype; NY!-isotype). P. pseudo-viridicaule var. nievecitanum Trel. in Ann. Mo. hot. Gdn 27: 296 (1940). Type: Panama, Bocas del Toro, Woodson, Allen & Seibert 1829 (ILL!-holotype; NY!- isotype). P. variitrichum Yunck. in Ann. Mo. hot. Gdn 37: 37 (1950). 29 Type: Panama, Darien, Terry & Terry 1428 (MO!- holotype). Shrubs 1.5-4 m high, stems densely pale-puberulent. Leaves 12-20 cm long, 5-10 cm wide, widely elliptic to obovate, upper surface scabrous, underside with hairs on veins, and minute dark glands, apex acute-acuminate, base unequally rounded to obtuse. Venation with 4-5 pairs of secondary veins arising steeply from the upper to middle part of the midrib, curving to apex. Petioles 6-15 mm long, pubescent, with ligule-like structure 6-18 mm long, glabrous or with scattered hairs, drying dark brown. Prophylls 12-22 mm long, drying dark brown, prominently veined, midrib pubes- cent. Inflorescences 8-15 cm long, erect; peduncles 5-10 mm long, pubescent. Anthers 0.1-0.2 mm long. Floral bracts 0.2-0.5 mm wide, triangular, pale-ciliate, with dark centres. Fruits 0.5-0.8 mm wide, obovoid, sides with prominent glands, oblong or square from above, with white or yellowish hairs; stigmas 3, minute. Forest, edges of streams, damp thickets, and roadsides; 0-1000 m. DISTRIBUTION. Mexico to Venezuela, West Indies. Mexico, Chiapas: Purpus 7301 (BM); Veracruz: Williams 8821 (F). Belize, Toledo: Schipp 504 (BM). Guatemala, Izabal: Stand- ley 72887 (F); Peten: Ortiz 1129 (F); Quetzaltenango: Stand- ley 84609 (F); Suchitepequez: Standley 66888 (F). Honduras, Atlantida: Mitchell 11 (F); Comayagua: Molina 8141 (BM); Olancho: Standley 18439 (F). Nicaragua: Bluefields: Proctor et al. 27293 (F); Chontales: Narvaez 3358 (BM); Nueva Segovia: Atwood et al. 6831a (F); Zelaya: Shank & Molina 4934 (F). Costa Rica, Alajuela: Burger & Stolze 4962 (F); Cartago: Croat 36526 (MO); Heredia: Greig 612 (BM); Limon: Robles 2143 (CR); Puntarenas: Raven 21482B; San Jose: Greig 445 (BM). Panama, Bocas del Toro: Cooper 165 (F); Chiriqui: Nee 10665 (MO); Code: Gracia 49 (F); Darien: Garwood 2166 A (BM); Herrera: Sytsma & Darcy 3261 (MO); Panama: Saiz 33 (F); Veraguas: Mori et al. 7531 (MO). This species can be distinguished from P. hispidum by its large, leaf-like prophylls, and ligule-like structures almost equalling the length of the prophylls, both drying to dark brown. The leaves of Piper sancti-felicis are dark green, drying to brown, while those of P. hispidum are a pale yellowish green, even when dry. 82. Piper poasanum C.DC. in Bull. Soc. r. Bot. Belg. 30(1): 206 (1892). Type: Costa Rica, vallee du Rio Poas, Pittier 2386 (G!-holotype). Fig. HA,a,b. P. palmasanum C.DC. in Smithson. misc. Collns 71(6): 3 (1920). Type: Panama, Cuesta de las Palmas, Chiriqui, Pittier 3225 (US- holotype, photograph!) P. pexum Trel. in Contr. U.S. natn. Herb. 26: 140 (1929). Type: Costa Rica, Rio Blanco, San Jose, Standley 41887 (US-holotype, photograph!). P. silvanorum Trel. in Contr. U.S. natn. Herb. 26: 169 (1929). Type: Costa Rica, Las Nubes, San Jose, Standley 38500 (US-holotype, photograph!). P. crisp atimargine Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 339 (1937). Type: Costa Rica, Palmira del Naranjo, Brenes 3499 (F-holotype). M.C. TEBBS Fig. 11 A: P. poasanum, habit; a: prophyll; b: fruit and bract. B: P. bisasperatum, habit; c: prophyll; d: fruit and bract. C: P. umbricola, habit; e: prophyll; f: fruit and bract. PIPER IN THE NEW WORLD 31 Shrubs or small trees 1-3 (-4) m high, stems sparsely to densely yellow-pubescent. Leaves 8-18 cm long, 4-9 cm wide, elliptic to ovate, coriaceous, upper surface glabrous, underside with hairs on veins, apex acuminate, base slightly unequal, rounded to subcordate. Venation with 3-4 pairs of deeply impressed secondary veins mostly arising from the lower part of the midrib, arcuate ascending to apex. Petioles 10-20 mm long, with ligule-like structure 10-15 mm long, usually caducous. Prophylls 15-20 (-30) mm long, pale, pubescent along midrib. Inflorescences 3-12 cm long, erect, with short sterile tip; peduncles 6-18 (-22) mm long, glabrous or pubescent. Anthers 0.3-0.4 mm long. Floral bracts (0.5-) 1-1.2 mm wide, deeply triangular, pale whitish or yellowish ciliate with a dark, glabrous centre. Fruits 1-1.5 mm wide, obovoid, round from above, glabrous or minutely white- pubescent or granular; stigmas 3. Partial shade in moist or wet forest; 1500-2600 m. DISTRIBUTION. Costa Rica, Panama. Costa Rica, Alajuela: Lent 1661 (BM); San Jose: Burger & Stolze 5203 (F). Panama, Chiriqui: Davidson 368 (F). Piper poasanum is a plant of high altitudes. It can be recognized by its rather coarse leaves with deeply impressed venation, and ciliate floral bracts with dark glabrous centres. When crushed, the leaves have a spicy aromatic odour. 83. Piper bisasperatum Trel. in Contr. U.S. natn. Herb. 26: 173 (1929). Type: Costa Rica, Cerro de la Carpintera, Cartage, Standley 35723 (US-holotype). Fig. llB,c,d. P. blepharilepidum Trel. in Contr. U.S. natn. Herb. 26: 160 (1929). Type: Costa Rica, Tilaran, Standley & Valeria 46217 (US-holotype). P. coactoris Trel. in Contr. U.S. natn. Herb. 26: 161 (1929). Type: Costa Rica, Alajuela, Standley & Torres 47935 (US-holotype). P. pubens Trel. in Contr. U.S. natn. Herb. 26: 163 (1929). Type: Costa Rica, La Estrella, Cartago, Standley 39329 (US-holotype, photograph!). P. emollitum Trel. in Contr. U.S. natn. Herb. 26: 181 (1929). Type: Costa Rica, El Silencio, Guanacaste, Standley & Valeria 44650 (US-holotype). P. ventoleranum Trel. in Contr. U.S. natn. Herb. 26: 184 (1929). Type: Costa Rica, Volcan Poas, Standley 34712 (US-holotype). P. austinii Trel. in Publs Field Mus. nat. Hist. (Bot.) 18: 1546 (1938). Type; Costa Rica, Tapesco de Zarcero, Smith H435 (F!-holotype). P. austinii var. aequilaterum Trel. in Publs Field Mus. nat. Hist. Bot. 18: 1546 (1938). Type: Costa Rica, Zaote de San Carlos, Smith H860 (F!-holotype). Shrubs, rarely small trees, 1.5-5 (-5) m high, stems glandu- lar, glabrous or pubescent, with whitish multicellular hairs up to 1.5 mm long. Leaves 10-24 cm long, 4-11 cm wide, elliptic to ovate-lanceolate, upper surface scabrous, glabrous to sparsely pubescent, sometimes bullate, underside sparsely to densely pubescent, especially on veins, both surfaces with orange glands visible, apex narrowly acuminate, apiculate, base unequally obtuse or slightly rounded. Venation with 3-5 pairs of secondary veins mostly arising from the lower to middle part of the midrib, ascending fairly steeply to apex. Petioles 5-12 (-16) mm long, glandular, glabrous or pubes- cent, with ligule-like structure 4-10 mm long, with prominent orange glands and sparse hairs on margin. Prophylls 15-30 mm long, acute, pale, glabrous or sparsely pubescent, with prominent orange glands. Inflorescences 6-12 cm long, erect, with pubescent tip 2-4 mm long; peduncles 5-12 (-20) mm long, glandular, glabrous or pubescent. Anthers 0.1-0.2 mm long. Floral bracts 0.4-0.6 mm wide, triangular, white-ciliate with dark glabrous centre. Fruits 1-1.2 mm wide, obovoid, oblong from above, pale puberulent; stigmas 3, in slight depression, sessile. Shady, moist places in open forest, forested slopes near streams, shaded road and track sides; 800-2000 m. DISTRIBUTION. Mexico to Venezuela. Mexico, Chiapas: Breedlove 35413 (F); Tabasco: Cowan & Nino 3381 (MO); Veracruz: Burnett et al. 92 (BM,MO). Costa Rica, Cartago: Burger & Gentry 9200 (F); Guanacaste: Burger & Pohl 7824 (F); Heredia: Antonio 755 (F,BM); Puntarenas: Burger & Gentry 8549 (F,MO); San Jose: Burger & Stolze 5356 (F). Panama, Chiriqui: Croat 26834 (MO). Colombia, Amazonas: Plowman & Martin 130 (S). Venezuela, Tachira: Berti & Pena 552-779 (G). Piper bisasperatum can be distinguished by its scabrous leaves with long, sharp apices, large prophylls and ligule-like struc- tures at the petioles, all with prominent orange glands, and its occurence at altitudes above 800 m. It is closely related to P. chrysostachyum which has similarly shaped leaves, prophylls and ligule-like structures. However, P. chrysostachyum has pale coloured bracts with whitish gibbous bases, whereas those of P. bisasperatum are densely pale-ciliate, with dark, glabrous centres. Piper chrysostachyum also tends to grow in dryer areas and at lower altitudes. 84. Piper umbricola C.DC. in Bull. Soc. r. Bot. Belg. 30(1): 215 (1891). Type: Costa Rica, Rodeo de Pacaca, Pittier 3238 (G!-holotype). Fig. HC,e,f. P. brachistopodium C.DC. in Bot. Gaz. 70: 182 (1920). Type: Costa Rica, Tucurrique, Tonduz 13143 (US). P. nodosum C.DC. in Bot. Gaz. 70: 185 (1920). Type: Costa Rica, Canas Gordas, Pittier 11072 (F!-isotype). P. disparipes Trel. in Contr. U.S. natn. Herb. 26: 162 (1929). Type: Costa Rica, El Silencio, Tilaran, Standley & Vale- ria 44766 (US-holotype). P. imparipes Trel. in Contr. U.S. natn. Herb. 26: 163 (1929). Type: Costa Rica, La Verbena, Standley & Valeria 32244 (US-holotype). P. papulatum Trel. in Contr. U.S. natn. Herb. 26: 163 (1929). Type: Costa Rica, Capulin, Alajuela, Standley 40181 (US-holotype). P. injucundum Trel. in Contr. U.S. natn. Herb. 26: 181 (1929). Type: Costa Rica, El Muneco, Standley & Valeria 51351 (US-holotype). P. captum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 335 (1937). Type: Costa Rica, El General, Skutch 2158 (US-holotype; K!,MO!-isotypes). P. pustulicaule Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 357 (1937). Type: Costa Rica, El General, Skutch 2938 (US-holotype; MO!-isotype). Shrubs 1-3 m high, stems glabrous or minutely puberulent. 32 M.C. TEBBS Leaves 11-24 cm long, 4-9 cm wide, elliptic to ovate or rhombic, upper surface dark, glossy, glabrous, underside minutely pubescent on veins, apex acuminate, base unequally obtuse or rounded. Venation with 3-4 pairs of secondary veins arising steeply towards apex. Petioles 4-15 mm long, glabrous or minutely puberulent, with ciliate ligule-like struc- ture 1-2 mm long. Prophylls 12-25 mm long, acute, glabrous, drying dark brown or black. Inflorescences 5-11 cm long, erect; peduncles 2-8 (-12) mm long, glabrous. Anthers 0.1-0.3 mm long. Floral bracts 0.4-0.6 mm wide, triangular, minutely ciliate, with glabrous centres. Fruits 0.6-0.9 mm wide, obovoid, round to oblong from above, glabrous; stig- mas 3, in central depression. Moist forest; 500-1500 m. DISTRIBUTION. Mexico to Costa Rica. Mexico, Chiapas: Spellman et al. 181 (BM); Nayarit: Croat 45325 (BM,MO); Oaxaca: Croat & Hannon 65585 (MO); Veracruz: Gentry et al. 32551 (BM,MO). Honduras, Comayagua: Molina 25468 (F); Cortez: Yuncker 4874 (F). Nicaragua, Granada: Stevens 10849 (F). Costa Rica, Guanacaste: Standley & Valeria 44228 (F); Puntarenas: Grayum & Sleeper 5898 (BM,MO). 85. Piper chrysostachyum C.DC. in Bull. Soc. r. Bot. Belg. 30(1): 207 (1892). Type: Costa Rica, near San Mateo, Pittier 4013 (G!-holotype). Fig. 12A,a,b. P. stenocladum C.DC. in An. Inst. fis-geogr. C. Rica 9: 162 (1897). Type: Costa Rica, forests of Boruca, Tonduz 6747, pro parte (G!-holotype). P. davidianum C.DC. in Smithson. misc. Collns 71(6): 9 (1920). Type: Panama, Dos Bocas, Colon, Pittier 4210 (US-holotype). P. callibracteum C.DC. in Smithson. misc. Collns 71(6): 13 (1920). Type: Panama, Chiriqui, Pittier 2940 (US!- holotype). P. chamissonis var. rubellibracteum C.DC. in Smithson. misc. Collns 71(6): 13 (1920). Type: Panama, El Boquete, Chiriquf, Pittier 2899 (US-holotype). P. nitidifolium C.DC. in Smithson. misc. Collns 71(6): 14 (1920). Type: Panama, El Boquete, Chiriqui, Maxon 4943 (US-holotype; G!-isotype). P. diquisanum C.DC. in Bot. Gaz. 70: 185 (1920). Type: Costa Rica, Diquis, Pittier 10567 (US-holotype; G!-isotype). P. surubresanum Trel. in Contr. U.S. natn. Herb. 26: 148 (1929). Type: Costa Rica, Rio Surubres, San Mateo, Biolley 17353 (US- holotype). P. vicinum Trel. in Contr. U.S. natn. Herb. 26: 157 (1929). Type: Costa Rica, Tonduz 6635 (US-holotype). P. alajuelanum Trel. in Contr. U.S. natn. Herb. 26: 158 (1929). Type: Costa Rica, Nuestro Amo, Alajuela, Jimenez 988 (US-holotype). P. verruculigerum Trel. in Contr. U.S. natn. Herb. 26: 165 (1929). Type: Costa Rica, Quebrada Serena, Guana- caste, Standley & Valeria 46239 (US-holotype; ILL!- isotype). P. hanckeli Trel. ex Standl. in Publs Field Mus. not. Hist. (Bot.) 18: 345 (1937). Type: Costa Rica, Guanacaste, Dodge, Hanckel & Thomas 6384 (GH-holotype; MO!- isotype). P. luridispicum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 348 (1937). Type: Costa Rica, El Rodeo, Lankester 1322 (F-holotype). P. papulaecaule Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 352 (1937). Type: Costa Rica, Guanacaste, Dodge & Thomas 6286 (GH-holotype; MO!-isotype). P. rubripes Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 358 (1937). Type: Costa Rica, Guanacaste, Dodge, Hanckel & Thomas 6382 (GH-holotype; MO!- isotype). P. tacaresense Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 364 (1937). Type: Costa Rica, Tacares, Valeria 302 (F!-holotype). Shrubs 1-3 m high, stems glabrous or minutely puberulent. Leaves 10-22 cm long, 4-9 cm wide, elliptic to ovate- lanceolate, upper surface glabrous, slightly scabrous, under- side glabrous or minutely pubescent, both sides glandular, apex acute-acuminate, base unequally obtuse to rounded. Venation with 3-5 pairs of secondary veins arising from the lower to middle part of the midrib, curving to apex. Petioles 5-15 mm long, with a ligule-like structure 1-2 mm long, glandular, sparsely ciliate. Prophylls 8-15 (-18) mm long, acute, glabrous or minutely pubescent, glandular, drying to dark brown or black. Inflorescences 5-12 cm long, erect, yellow or white, sometimes with short sterile tips; peduncles 6-14 (-18) mm long, glabrous. Anthers 0.1-0.2 mm long. Floral bracts 0.2-0.4 mm wide, triangular, glabrous or minutely ciliate, with a yellowish or white bulge at the base. Fruits 0.5-0.8 mm wide, obovoid, oblong or round from above, minutely yellow-puberulent; stigmas 3, sessile, in slight depression. Seasonally dry evergreen forest mostly on Pacific side; 0-1300 (-1500) m. DISTRIBUTION. Nicaragua to Panama. Nicaragua, Granada: Williams & Molina 20003 (F); Jinotega: Hawkes et al. 2175 (F); Rivas: Neill & Vincelli 3212 (F); Zelaya: Pipoly 4684 (MO). Costa Rica, Alajuela: Haber 1487 (MO); Guanacaste: Valeria 1145 (F); Puntarenas: Burger et al. 10634 (F); San Jose: Burger & Liesner 7153 (F). Panama, Chiriqui: Croat 13560 (MO); Code: Gentry 7420 (F, MO); Veraguas: Mori & Kallunki 6141 (MO). The inflorescences with minute floral bracts bulging basally and the yellow-puberulent fruits distinguish Piper chrysos- tachyum from other species. 86. Piper epigynium C.DC. in Linnaea 37: 346 (1872). Type: Costa Rica, Turrialba, Oersted 858 (C!-holotype). Fig. 12B,c,d. P. villosisquamulum Trel. in Contr. U.S. natn. Herb. 26: 158 (1929). Type: Costa Rica, La Hondura, Standley 37799 (US-holotype). P. villistipulum Trel. in Contr. U.S. natn. Herb. 26: 162 (1929). Type: Costa Rica, El Murieco, S. of Navarra, Cartago, Standley 33438 (US-holotype). P. subdivaricatum Trel. in Contr. U.S. natn. Herb. 26: 163 (1929). Type: Costa Rica, La Hondura, San Jose, Stand- ley 36489 (US-holotype). Shrubs or small trees, 1.5-6 m high, stems glabrous. Leaves 12-26 cm long, 4-9 cm wide, elliptic to narrowly ovate, glandular, upper surface glabrous, sometimes slightly sca- brous, dark green, underside with pale hairs on the veins, PIPER IN THE NEW WORLD 33 Fig. 12 A: P. chrysostachyum, habit; a: prophyll; b: fruit and bract. B: P. epigynium, habit; c: prophyll; d: fruit and bract. C: P. dotanum, habit; e: prophyll; f: fruit and bract. 34 M.C. TEBBS apex acuminate, base unequally obtuse or rounded. Venation with 3-4 pairs of secondary veins arising mostly from the lower to middle part of the midrib. Petioles 4-14 (-20) mm long, with glandular ligule-like structure 1-3 mm long. Pro- phylls 14-30 (-40) mm long, glandular, sparsely puberulent along midrib. Inflorescences 6-15 cm long, erect, sometimes pinkish to purple in early stages; peduncles 8-16 mm long, glabrous. Anthers 0.1-0.2 mm long. Floral bracts 0.2-0.3 mm wide, triangular, with pale bulge at base, gla- brous or with minute hairs. Fruits 0.4-0.6 mm wide, obovoid, minutely puberulent, glandular; stigmas 3, sessile. Cloud forest, ravines, open areas, stream and trail sides in moist forest; 600-2000 m. DISTRIBUTION. Guatemala to Panama. Guatemala, San Mar- coas: Williams et al. 26970 (F). Nicaragua, Granada: Croat 39087 (MO). Costa Rica, Alajuela: Liesner 4761 (MO); Heredia: Nee 14007 (F); Puntarenas: Dinerstein 3; San Jose: Burger & Gentry 9082 (F). Panama, Chiriqui: van der Werff 6751 (MO); Veraguas: Croat & Folsom 34270 (MO). Piper epigynium can be recognized by its often showy brick- red inflorescences and large, glabrous, glandular prophylls. It is only found at altitudes above 600 m. 87. Piper dotanum Trel. in Contr. U.S. natn. Herb. 26: 165 (1929). Type: Costa Rica, Santa Maria de Dota, San Jose, Standley 41739 (US-holotype; ILL!-isotype). Fig. 12C,e,f. Shrubs 1-4 m high, stems glabrous, slender, spreading. Leaves 5-13 cm long, 1.5^4- cm wide, elliptic-lanceolate to ovate-lanceolate, glabrous, upper surface glossy, slightly sca- brous, underside with prominent dark glands, apex acumi- nate, base unequally rounded. Venation with 3 pairs of secondary veins arising from the lower to middle part of the midrib, loop-connecting to apex. Petioles 2-7 mm long, gla- brous, with a sparsely ciliate, ligule-like structure 1-2 mm long. Prophylls 6-14 mm long, glabrous, drying to dark brown or black. Inflorescences 4-6 cm long, cream-coloured, erect, with short sterile tips; peduncles 4-12 mm long, gla- brous. Anthers 0.1-0.2 mm long. Floral bracts 0.2-0.4 mm wide, triangular, densely puberulent. Fruits 0.5-0.7 mm wide, obovoid, oblong to round from above, glabrous to minutely puberulent; stigmas 3, sessile. Deep shade in forest; 500-1800 m. DISTRIBUTION. Costa Rica. Costa Rica, Alajuela: Brenes 20417 (NY); Guanacaste: Garwood et al. 797 (BM); Puntar- enas: Haber 4042 (MO). Piper dotanum is a slender, rather spindly plant growing in forest shade in the drier regions of Costa Rica. It can be identified by its small, glossy, glabrous leaves, glabrous prophylls, prominent ligule-like structure, and slender, cream-coloured inflorescences. 88. Piper polytrichum C.DC. ex A. Schroed. in Candollea 3: 138 (1926). Type: Costa Rica, Canas Gordas, Pittier 11070(G!-holotype). Fig. 13A,a,b. Shrubs 1-3 m high, stems covered with thin yellowish multi- cellular hairs up to 2 mm long. Leaves 6-17 cm long, 2-7 cm wide, narrowly elliptic to ovate or ovate-lanceolate, upper surface dark green, slightly scabrous with scattered whitish hairs, underside densely pubescent on the veins, apex acumi- nate, base unequally obtuse or rounded. Venation with 3-5 pairs of secondary veins arising from the lower to middle part of the midrib, ascending fairly steeply to apex. Petioles 4-7 mm long, densely pubescent, ligule-like structure 0.5 mm long, hidden by hairs. Prophylls 10-20 mm long, acute, hirsute along the midrib, drying dark brown. Inflorescences 5-9 cm long, erect; peduncles 5-7 mm long, densely pubes- cent. Anthers 0.2-0.3 mm long. Floral bracts 0.2-0.5 mm wide, triangular to round, mostly glabrous with a few minute pale hairs at the base. Fruits 0.6-1 mm wide, obovoid, oblong from above, glabrous or sparsely pubescent; stigmas 3. Shade of moist forest; 600-1200 m. DISTRIBUTION. Costa Rica to Panama. Costa Rica, Cartago: Skutch 4657 (NY); Puntarenas: Davidson 7158 (NY); San Jose: Burger & Liesner 7113A (F). Panama, Colon: Dressier 3854 (F); Darien: Gentry Mori 13905 (F); Panama: Maas & Dressier 712 (F). Piper polytrichum can be confused with P. biauritum because of similar indumentum and leaf-shape. However, P. biauri- tum has red or purple inflorescences and pale, glandular prophylls, whereas P. polytrichum has green or whitish inflorescences and smaller prophylls drying to a dark brown. 89. Piper peracuminatum C.DC. in Smithson. misc. Collns 71(6): 9 (1920). Type: Panama, Dos Bocas, Rio Fato valley, Pittier 42W (US-holotype; G!-isotype). Fig. 13B,c,d. P. fusco-granulatum Trel. in Contr. U.S. natn. Herb. 26: 180 (1929). Type: Costa Rica, Hacienda de Zent, United Fruit Co. 269 (US-holotype). Shrubs or small trees 2-3 m high, stems hirsute with yellowish hairs 0.2-1.5 mm long. Leaves 15-27 cm long, 8-13 cm wide, elliptic-obovate to ovate, upper surface glossy, slightly sca- brous, sparsely and minutely puberulent, underside pubes- cent on veins, apex acuminate, base unequally obtuse or slightly lobed on one side. Venation with 4-5 pairs of secondary veins arising from the lower to middle part of the midrib, ascending fairly steeply to apex. Petioles 7-16 mm long, pubescent, ligule-like structure minute or absent. Pro- phylls 10-14 mm long, acute, hirsute along midrib, drying dark brown. Inflorescences 10-16 cm long, erect; peduncles 10-22 mm long, pubescent. Anthers 0.3-0.4 mm long. Floral bracts 0.4-0.7 mm wide, yellow-ciliate with glabrous centres. Fruits 0.5-0.8 mm wide, obovoid, oblong or round from above, minutely puberulent; stigmas 3, recurved. Lowland moist forest; 0-500 m. DISTRIBUTION. Costa Rica to Panama. Panama, Code: Lewis et al. 5526 (MO); Panama: Kennedy & Dressier 2966 (MO). 90. Piper zacatense C.DC. in An. Inst. fis-geogr. C. Rica 9: 161 (1897). Type: Costa Rica, Boca Zacate, Pittier 6828 (W-holotype; G!-isotype). Fig. 13C,e,f. P. zacatense var. percaudatum C.DC. in An. Inst. fis-geogr. PIPER IN THE NEW WORLD Fig. 13 A: P. polytrichum, habit; a: prophyll; b: fruit and bract. B: P. peracuminatum, habit; c: prophyll; d: fruit and bract. C: P. zacatense, habit; e: prophyll; f: fruit and bract. 36 C. Rica 9: 161 (1897). Type: Costa Rica, Golfo Dulce, Pittier 9914 (G!-holotype). Shrubs or slender trees; stems densely pubescent with yellow- brown hairs 0.5-2 mm long. Leaves 16-28 cm long, 7-12 cm wide, elliptic to rhombic, upper surface dark, smooth or slightly scabrous, minutely puberulent, underside pubescent on veins, apex long-acuminate, base unequally obtuse or rounded. Venation with 4-5 pairs of secondary veins arising from the lower to middle part of the midrib, curving steeply to apex. Petioles 6-12 mm long, densely pubescent, with a minute ligule-like structure 0.5 mm long. Prophylls 12-15 mm long, acute, densely puberulent. Inflorescences 3-9 cm long, erect; peduncles 8-14 mm long, minutely pubes- cent. Anthers 0.2-0.3 mm long. Floral bracts 0.3-0.6 mm wide, yellow-ciliate with glabrous centres. Fruits 0.7-1 mm wide, obovoid, oblong from above, sparsely puberulent; stigmas 3, in central depression. Lowland moist forest; 0-500 m. DISTRIBUTION. Costa Rica. P. zacatense is very rare, and not well represented in her- baria. Only type material has been seen for this study. 91. Piper biauritum C.DC. in An. Inst. fis-geogr. C. Rica 9: 161 (1897). Type: Costa Rica, forest of Xirores, Tala- manca, Tonduz 9270 (G!-holotype). Fig. 14A,a,b. P. tortuosipilum Trel. in Contr. U.S. natn. Herb. 26: 148 (1929). Type: Costa Rica, El Muneco, Cartago, Standley & Valeria 51026 (US-holotype). P. insolens Trel. in Contr. U.S. natn. Herb. 26: 156 (1929). Type: Costa Rica, Santa Maria de Dota, San Jose, Standley & Valeria 43306 (US-holotype, photograph!; ILL-isotype). Shrubs 1-2 m high, stems pubescent with long, pale, multicel- lular hairs. Leaves 11-25 cm long, 5-12 cm wide, elliptic to ovate, both surfaces sparsely to densely covered with long white hairs, especially on veins beneath, apex acute- acuminate, base unequally obtuse to cordulate. Venation with 4-5 pairs of secondary veins arising from the lower to middle part of the midrib. Petioles 4-10 (-20) mm long, pubescent, ligule-like structure minute or absent. Prophylls 15-20 (-25) mm long, acute, pale, glandular, with long pale hairs. Inflorescences 7-12 cm long, reddish or purple, erect; peduncles 8-22 mm long, glabrous or pubescent. Anthers 0.1-0.2 mm long. Floral bracts 0.4 mm wide, triangular, with pale bulge and pale hairs at the base, or glabrous. Fruits 0.8-1 mm wide, obovoid, round or oblong from above, minutely puberulent; stigmas 3, sessile. Moist slopes; 0-1600 m. DISTRIBUTION. Costa Rica to Panama. Costa Rica, Alajuela: Burger et al. 107724 (F); Cartago: Greig 380 (BM); Limon: Burger & Antonio 10999 (CAS); Puntarenas: Skutch 5268; San Jose: Davidse et al. 23179 (MO). Panama, Bocas del Toro: Croat & Grayum 60136 (MO); Chiriqui: Churchill 5381 (BM, MO); Darien: Hammel et al. 16173 (BM,MO); Vera- guas: Nee 9903 (MO). 92. Piper perhispidum C.DC. in Bot. Gaz. 70: 183 (1920). M.C. TEBBS Type: Costa Rica, near San Juan de San Ramon, Tonduz 17771 (G!-holotype). Fig. 14C,e,f. P. pileatum Trel. in Contr. U.S. natn. Herb. 26: 184 (1929). Type: Costa Rica, forests of Copey, Tonduz 11895 (NY- holotype; F!,MO!-isotypes). P. pileatum var. obliquum Trel. in Contr. U.S. natn. Herb. 26: 184 (1929). Type: Costa Rica, El Copey, Tonduz 11675 (US-holotype, photograph!). P. rugosifolium Trel. in Contr. U.S. natn. Herb. 26: 185 (1929). Type: Costa Rica, Finca Las Concavas, Cartago, Standley 41533 (US-holotype, photograph!). Shrubs 1-3 m high, stems covered with yellow-brown multi- cellular hairs up to 2.5 mm long. Leaves 10-24 cm long, 2.5-9 cm wide, elliptic-ovate to lanceolate, bullate, scabrous above, both surfaces covered with long hairs, apex acumi- nate, base unequally obtuse or rounded. Venation with 3-5 pairs of secondary veins arising from the lower to middle part of the midrib. Petioles 3-10 (-20) mm long, densely pubes- cent, with pubescent ligule-like structure 1 mm long. Pro- phylls 10-20 mm long, acute, puberulent with glabrous margins. Inflorescences 6-11 cm long, erect; peduncles 8-14 mm long, densely villous. Anthers 0.2-0.3 mm long. Floral bracts 0.4-0.6 mm wide, triangular, densely pale- pubescent. Fruits 0.5-1 mm wide, obovoid, minutely puberu- lent, sides glandular; stigmas 3, sessile, in slight depression. Moist forest; 1000-2000 m. DISTRIBUTION. Costa Rica. 93. Piper chamissonis (Miq.) Steud., Nomencl. hot. ed.2, 2: 340 (1841). Fig. 14D,g,h. Artanthe chamissonis Miq., Syst. piperac.: 457 (1844). Type: Mexico, Hacienda de la Laguna, Veracruz, Schiede s.n. (B-holotype; G!-drawing). Shrubs 1-2.5 m high, stems glabrous or sparsely pubescent. Leaves 13-21 cm long, 5-9 cm wide, elliptic to obovate, upper surface dark green, glabrous, lower surface pale with yellowish white hairs on veins, apex narrowly acuminate, base slightly unequal, cuneate. Venation with 3-5 secondary veins arising from lower to middle part of midrib, loop- connecting to apex. Petioles 8-15 mm long, glabrous to sparsely pubescent, with pale-glandular, glabrous ligule-like structure 2-3 mm long. Prophylls 10-20 mm long, narrow, glabrous or minutely puberulent, drying dark. Inflorescences 6-10 cm long, erect; peduncles 8-12 mm long, glabrous or sparsely pubescent. Anthers 0.4-0.5 mm long. Floral bracts 0.8-1 mm wide, triangular, densely pale-ciliate, with dark centre. Fruits 1 mm wide, obovoid, oblong-trigonous from above, glabrous; stigmas 3, sessile. Moist forest, damp thickets, and wooded ravines; 900-2000 (-3000) m. DISTRIBUTION. Mexico to Guatemala. Guatemala, Quezalt- enango: Standley 68161 (F); San Marcos: Standley 68946 (F). Piper chamissonis is only found at high altitudes in moist, wooded or thicketed areas. It can be recognized by its elliptic to obovate leaves with yellowish white hairs on the veins of the undersides, prominent prophylls, pale-ciliate bracts, and PIPER IN THE NEW WORLD 37 Fig. 14 A: P. biauritum, habit; a: prophyll; b: fruit and bract. B: P. colonense, habit; c: prophyll; d: fruit & bract. C: P. perhispidum C. DC., habit; e: prophyll; f: fruit and bract. D: P. chamissonis, habit; g: prophyll; h: fruit and bract. 38 glabrous fruits. Piper chrysostachyum has similar leaves but very different floral bracts. 94. Piper colonense C.DC. in Smithson. misc. Collns 71(6): 11 (1920). Type: Panama, Rio Fato valley, Colon, Pittier 4221 (US!-holotype; BM!-isotype). Fig. 14B,c,d. P. culebranum C.DC. ex A. Schroed. in Candollea 3: 136 (1926). Type: Panama, Rio Culebra, Pittier 4154 (G!- holotype). P. varablancanum Trel. in Publs Field Mus. nat. Hist. (Bot.) 18: 1547 (1938). Type: Costa Rica, Vara Blanca de Sarapiqui, Skutch 3205 (US-holotype; MO!,NY!- isotypes). Shrubs or spindly tree 1.5-6 (-7) m high, stems sparsely to densely pubescent. Leaves 12-25 cm long, 4-10 cm wide, oblong-elliptic to oblanceolate, upper surface glaucous, gla- brous or with sparse hairs, underside usually with conspicu- ous red or orange glands, and long hairs on veins, apex acute-acuminate, tapering abruptly, base slightly unequal, round to obtuse. Venation with 3-4 pairs of secondary veins ascending steeply to apex. Petioles 5-10 mm long, pubescent, with ligule-like structure to 1 mm long. Prophylls 6-20 mm long, sparsely pubescent, glandular, drying dark brown or black. Inflorescences 6-12 cm long, erect; peduncles 5-12 mm long, glabrous or sparsely pubescent. Anthers 0.2-0.3 mm long. Floral bracts 0.7-1 mm wide, triangular, yellowish-ciliate, with glabrous centre. Fruits 1-1.2 mm wide, obovoid, round or oblong from above, fleshy, glabrous; stigmas 3, sessile. Disturbed or cleared areas of forest, along trails and road- sides; 0-1600 m. DISTRIBUTION. Nicaragua to Panama. Nicaragua, Zelaya: Bunting & Licht 1252 (F). Costa Rica, Alajuela: Utley 2235 (F); Cartago: Holm & Iltis 76 (F); Heredia: Burger & Stolze 5758 (F); Limon: Burger & Liesner 7006 (F); San Jose: Skutch 4931 (NY). Panama, Chiriqui: Croat 22343 (MO); Colon: Hammel 4492 (BM,MO); Panama: Mori 7744 (BM,MO). 95. Piper oblanceolatum Trel. in Contr. U.S. natn. Herb. 26: 175 1929). Type: Costa Rica, Los Ayotes, Tilaran, Stand- ley & Valeria 45558 (US-holotype, photograph!). Fig. 15A,a,b. Shrubs or small trees 2-8 (-10) m high, stems glabrous to sparsely pubescent, with yellowish hairs 0.5-1 mm long. Leaves 10-18 cm long, 3-^6 cm wide, narrowly elliptic to oblanceolate, upper surface dark green, slightly scabrous, glabrous or minutely pubescent, underside with whitish hairs on veins, apex long-acuminate, base unequally obtuse or cuneate. Venation with 3-5 pairs of secondary veins arising from the lower part of the midrib, arcuate ascending to apex. Petioles 2-8 mm long, glabrous or sparsely pubescent, with a ciliate ligule-like structure 0.5 mm long. Prophylls 10-22 mm long, acute, puberulent along midrib, glandular, drying dark brown or black. Inflorescences 5-10 cm long, erect; pedun- cles 14-25 mm long, glabrous to sparsely pubescent. Anthers 0.2-0.3 mm long. Floral bracts 0.4-0.6 mm wide, narrowly triangular, upper part glabrous, glandular, lower sparsely ciliate. Fruits 0.5-0.8 mm wide, obovoid, oblong from above; M.C. TEBBS stigmas 3, in slight depression. Shade of moist forest; 600-2000 m. DISTRIBUTION. Nicaragua to Costa Rica. Nicaragua, Jinotega: Standley 10697 (F). Costa Rica, Alajuela: Lent 1673 (MO); Monteverde: Feinsinger s.n. 'Y' (F). 96. Piper hirtellipetiolum C.DC. in Smithson. misc. Collns 71 (6): 3 (1920). Type: Panama, vicinity of David, Chiriqui, Pittier 2932 (US-holotype; G!,NY!-isotypes). Fig. 15B,c,d. P. tapianum Trel. in Contr. U.S. natn. Herb. 26: 30 (1927). Type: Panama, near Tapia River, Maxon & Harvey 6709 (US-holotype; ILL!-isotype). Shrubs 1-2 m high, stems glabrous or sparsely covered with white multicellular hairs. Leaves 8-14 cm long, 2.5-5 cm wide, lanceolate, glabrous, glandular, upper surface glossy, underside dull, apex acuminate, base acute to obtuse. Vena- tion with 2-4 pairs of secondary veins arising from lower to middle part of midrib, loop-connecting to apex. Petioles 3-5 mm long, villous, with a sparsely ciliate, ligule-like structure 0.5-2 mm long. Prophylls 6-12 mm long, acute, glabrous. Inflorescences 3-6 cm long, erect; peduncles 3-10 mm long, puberulent. Anthers 0.2-0.3 mm long. Floral bracts 0.8-1 mm wide, triangular, densely yellow-ciliate. Fruits 1 mm wide, obovoid, round from above, glabrous; stigmas 3, sessile. Moist remnant forest, edges of streams, roadsides; 0-1000 (-2000) m. DISTRIBUTION. Panama. Panama, Canal Zone: Miller 1790 (MO); Chiriqui: Him & Gordon 270 (MO); Code: Miller 1814 (MO); Herrera: Stern et al. 33618 (MO); Panama: Duke 12565 (MO); Perlas Is.: Johnston 754 (MO); Taboga Is.: Pi«zer3618(NY). 97. Piper decurrens C.DC. in /. Bot., Lond. 4: 215 (1866). Type: Costa Rica, monte Candelaria, Hoffmann 853 (B-holotype, photograph!) Fig. 15C,e,f. P. leptoneuron C.DC. in Bot. Gaz. 70: 184 (1920). Type: Costa Rica, Santa Clara, Pittier 10675 (G!-holotype). P. gracilipedunculum Trel. in Contr. U.S. natn. Herb. 26: 148 (1929). Type: Costa Rica, Fraijanes, Alajuela, Standley & Torres 47519 (US-holotype). Shrubs 1-3 (-5) m high, stems glabrous. Leaves 6-12 (-16) cm long, 2.5-7 cm wide, elliptic to obovate, glossy, glabrous, apex acute-acuminate, base cuneate. Venation with 2-3 pairs of secondary veins arising from lower to middle part of midrib, with prominent cross-veins. Petioles 3-12 mm long, glabrous, with ligule-like structure 1 mm long. Pro- phylls 6-20 mm long, glabrous. Inflorescences 2-5 (-6) cm long, with sterile tip 1-2 mm long; peduncles 6-25 mm long. Floral bracts 0.5-1 mm wide, triangular, white-ciliate, with pale centres. Anthers 0.5 mm long. Fruits 1-2 mm wide, obovoid, round from above, glabrous; stigmas 2-3. Moist, shady places in cloud forest and lower rain forest; 700-2000 m. DISTRIBUTION. Mexico, Costa Rica. Mexico, Chiapas: PIPER IN THE NEW WORLD 39 Fig. 15 A: P. oblanceolatum, habit; a: prophyll; b: fruit and bract. B: P. hirtellipetiolum, habit; c: prophyll; d: fruit and bract. C: P. decurrens, habit; e: prophyll; f: fruit and bract. 40 M.C. TEBBS Breedlove & Raven 13022 (CAS). Costa Rica, Alajuela: Lent 2371 (F); Guanacaste: Burger & Gentry 9108 (F, BM); Heredia & San Jose: Antonio 111 (MO); Puntarenas: Burger & Gentry 8581 (MO). Piper decurrens can be recognized by its small, glabrous, glossy leaves and short inflorescences with sterile tips. It is very similar in appearance to P. tenuimucronatum, but can be distinguished by its much shorter ligule-like structure on the petiole, pale-centred bracts, and larger fruits. Only one specimen has been seen from Mexico, growing at 1700 m. It displays all the characteristics of P. decurrens, although the leaves are larger than usual. 98. Piper tenuimucronatum C.DC. in Smithson. misc. Collns 71(6): 12 (1920). Type: Panama, Los Siguas Camp, Chiriqui, Maxon 5421 (US-holotype; G!-isotype). Fig. 16A,a,b. P. infraleucumlrel. in Contr. U.S. natn. Herb. 26: 33 (1927). Type: Panama, Sutton Hayes 793 (NY!-holotype). P. tractifolium Trel. in Contr. U.S. natn. Herb. 26: 166 (1929). Type: Costa Rica, Heredia, Standley & Valeria 52052 (US-holotype). P. perfugii Trel. in Ann. Mo. hot. Gdn 27: 295 (1940). Type: Panama, Chiriqui, Woodson, Allen & Seibert 928 (ILL!- holotype; MO!-isotype). Shrubs 1-3 m high, stems glabrous. Leaves 6-12 cm long, 2-5 cm wide, elliptic to narrowly ovate, glandular, glabrous, upper surface dark green, lustrous, underside whitish green, apex acuminate, sometimes apiculate, base unequally obtuse or acute. Venation with 2-13 pairs of secondary veins mostly arising from the lower half of the midrib. Petioles 5-11 mm long, glabrous, with a glabrous, ligule-like structure 2-10 mm long. Prophylls 8-20 (-25) mm long, pale, glabrous. Inflores- cences 2.5-8 cm long, erect, with short sterile tips; peduncles 10-20 mm long, glabrous. Anthers 0.4-0.5 mm long. Floral bracts 0.6-0.8 mm wide, triangular, pale-ciliate, with a dark, glabrous centre. Fruits 1 mm wide, obovoid, round to oblong from above, glabrous; stigmas 3, sessile. Moist forest, wooded slopes; 1000-2000 m. DISTRIBUTION. Guatemala to Panama. Guatemala, Alta Verapaz: Williams et al. 40468 (F). Costa Rica, Alajuela: Smith 434 (MO); Cartago: Burger 7600 (F); Guanacaste: Davidse et al. 23355 (MO); San Jose: Grayum & Sleeper 6113 (BM,MO). Panama, Bocas del Toro: McPherson 9591 (MO); Chiriqui: Dwyer et al. 523 (K); Panama: Tyson 4035 (MO). Piper tenuimucronatum is confined to moist forest above altitudes of 1000 m. It has much larger ligule-like structures on the petioles than those of P. decurrens and darker floral bracts; otherwise there is very little difference between them. However, unless more specimens are collected which show intermediates, they are best maintained as separate species. 99. Piper carpinteranum C.DC. in An. Inst. fis.-geogr. C. Rica 9: 165 (1897). Type: Costa Rica, La Carpintera, Pittier434$ (G!-holotype). Fig. 16B,c,d. P. ejuncidum Trel. in Contr. U.S. natn. Herb. 26: 164 (1929). Type: Costa Rica, Heredia, Standley & Valeria 52220 (US-holotype). P. rotundibaccum Trel. in Contr. U.S. natn. Herb. 26: 164 (1929). Type: Costa Rica, San Jose, Standley 42902 (US-holotype, photograph!). P. rotundibaccum var. fraijanesanum Trel. in Contr. U.S. natn. Herb. 26: 164 (1929). Type: Costa Rica, Alajuela, Standley & Torres 47669 (US-holotype, photograph!). P. zonulatispicum Trel. in Contr. U.S. natn. Herb. 26: 164 (1929). Type: Costa Rica, Cartago, Standley & Valeria 51116 (US-holotype, photograph!). Shrubs 1-2 (-3) m high, stems sparsely to densely puberu- lent. Leaves 8-15 cm long, 3-7 cm wide, narrowly ovate to elliptic, glandular, upper surface glabrous, underside paler, puberulent on veins, apex long-acuminate, base unequal, lower side slightly lobed, overlapping petiole. Venation with 3-4 pairs of secondary veins arising from the lower part of the midrib, curving towards apex. Petioles 2-8 mm long, densely puberulent, with a ligule-like structure 1-5 mm long. Pro- phylls 10-20 mm long, puberulent along the midrib. Inflores- cences 2-5 (-8) cm long, erect, becoming pendulous, with sterile tip 1-3 mm long; peduncles 1-3 cm long, minutely puberulent. Anthers 0.3-0.4 mm long. Floral bracts 0.6-1 mm wide, triangular to semi-lunar, white- to yellow- ciliate with glabrous centres. Fruits 0.8-1.2 mm wide, obo- void, round to trigonous from above, glabrous; stigmas 3, recurved. Shade in moist forest, edges of woods; 1000-2500 m. DISTRIBUTION. Costa Rica to Panama. Costa Rica, Alajuela: Smith 1040 (F); Cartago: Standley 39812 (F); Heredia: Lent 223 (F); San Jose: Burger & Liesner 6205 A (F). Panama, Chiriqui: Kirkbride 122 (MO). Piper carpinteranum C.DC. can be separated from P. tenuimucronatum by its pubescent stems, leaves with unequal, slightly lobed bases, puberulent petioles with shorter ligule-like structures, and pubescent prophylls. 100. Piper hostmannianum (Miq.) C.DC. in DC., Prodr. 16(1): 287 (1869). Fig. 16D,g,h. Artanthe hostmannianum Miq. in /. Bot., Lond. 4: 465 (1845). Type: Surinam, Hostmann 116 (K!-holotype). Shrubs, occasionally scandent, 1-3 (-5) m high, stems sparsely to densely yellow-pubescent. Leaves 13-24 cm long, 6-12 cm wide, elliptic-lanceolate to ovate, upper surface glossy, glabrous, underside pale-pubescent on veins, pale- glandular, apex acuminate, base unequally obtuse or sub- cordate. Venation with 4-6 secondary veins arising from lower two-thirds of midrib, curving to the apex. Petioles 5-10 mm long, yellow-pubescent, with ciliate ligule-like structures 2-5 mm long. Prophylls 6-12 mm long, densely yellow-pubescent along midrib. Inflorescences 10-12 cm long, erect, sometimes with sterile tips 1-2 mm long; pedun- cles 8-15 mm long, glabrous to sparsely pubescent. Anthers 0.2-0.3 mm long. Floral bracts 0.5-0.8 mm wide, triangular or round, pale-ciliate, with darker glabrous centres. Fruits 0.8-1.2 mm wide, obovoid, oblong or round from above, glabrous or slightly granular; stigmas 3, sessile. Upland forest, disturbed areas along roads and railways; 0-1000 m. PIPER IN THE NEW WORLD 41 Fig. 16 A: P. tenuimucronatum, habit; a: prophyll; b: fruit and bract. B: P. carpinteranum, habit; c: prophyll; d: fruit and bract. C: P. scalarispicum, habit; e: prophyll; f: fruit and bract. D: P. hostmannianum, habit; g: prophyll; h: fruit and bract. 42 M.C. TEBBS DISTRIBUTION. Venezuela to Brazil. Venezuela, Atabapo: Liesner 17872 (MO); Atures: Steyermark et al. 122312 (NY); Bolivar: Liesner & Gonzalez 5850 (NY). Guyana, Kaieteur Plateau: Cowan & Soderstrom 2250 A (NY); Mazaruni R.: De la Cruz 2279 (NY); Pomeroon: De la Cruz 1827 (MO); Tumatumari: Gleason 307 (NY). Surinam, Lely mts.: Linde- man et al. 675 (NY). French Guiana, Cayenne: Lescure 107 (NY); Pompidou-Papaichton: Sastre 4051 (NY); Saul: Mori & Boom 14906 (MO). Peru, Loreto: Lleras et al. P17112 (MO). Bolivia, Beni: Boom 4074 (MO); Pando: Prance et al. 5870 (MO). Brazil, Amazonas: Nasdmento 551 (NY); Man- aus: Lowe 4233 (E); Mato Grosso: Berg et al. P18395 (US); Para: Cid & Ramos 1156 (NY); Rondonia: Prance et al. 5955 (MO). Piper hostmannianum is very similar vegetatively to P. jac- quemontianum, with large, glossy leaves and yellow pubes- cence. These species can be distinguished by their different inflorescences. Those of P. jacquemontianum are shorter, with semi-lunar, densely yellow-ciliate floral bracts and larger round fruits covered with a distinctive, felty brown indumen- tum. Although usually found growing as a shrub, P. hostman- nianum often has spreading stems that are capable of climbing upwards, presumably by means of adventitious roots at the nodes. 101. Piper scalarispicum Trel. in Publs Field Mus. nat. Hist. (Bot.) 17: 353 (1938). Type: Honduras, El Achote, SW. of Siguatepeque, Yuncker, Dawson & Youse 6156 (ILL!-holotype; F!,MO!-isotypes). Fig. 16C,e,f. P. nubigaudens Trel. in Publs Field Mus. nat. Hist. (Bot.) 17: 351 (1938). Type: Honduras, range above El Achote, Yuncker, Dawson & Youse 5991 (ILL!-holotype; F!-isotype). P. segoviarum Standl. & L.O.Williams in Ceiba 1: 142 (1950). Type: Nicaragua, Jinotega, Standley 9912 (F!- holotype). P. brujoense Trel. & Standl. in Fieldiana Bot. 24: 290 (1952). Type: Guatemala, Chiquimula, Steyermark 31049 (F!- holotype). Weak shrubs 1-2 (-4) m high, with pubescent stems. Leaves 5-12 (-15) cm long, 1.5-4 cm wide, elliptic-lanceolate to lanceolate, upper surface dark green, glabrous, undersurface with hairs on veins, apex acuminate, base slightly unequal. Venation with 2-4 pairs of secondary veins arising from lower to middle part of midrib, loop-connecting to apex. Petioles 3-6 mm long, pubescent, with ligule-like structure 0.5 mm long. Prophylls 7-10 mm long, drying dark, pubescent along midrib. Inflorescences 3-8 cm long, erect; peduncles 7-10 mm long, glabrous or sparsely pubescent. Anthers 0.3 mm long. Floral bracts 0.8-1 mm wide, narrowly triangu- lar to semi-lunar, yellowish white ciliate, with longer hairs on lower margins. Fruits 0.8-1 mm wide, obovoid, oblong-round from above, glabrous; stigmas 3. Cloud forest; 1300-2500 m. DISTRIBUTION. Guatemala to Nicaragua. Guatemala, Baja Verapaz: Lundell & Contreras 18965 (MO); Zacapa: Steyer- mark 29884 (F). Honduras, Cortes: Molina 8211 (F); Inti- buca: Molina 6408 (F); La Paz: Molina 14078 (F); Morazan: Molina 8525 (F); Ocotepeque: Molina 31403a; Paraiso: Molina 26148 (F). El Salvador, Santa Ana: Molina 12642 (F). Nicaragua, Jinotega: Standley 10672 (F); Matagalpa: Williams et al. 23527 (F). 102. Piper curvatipes Trel. in Publs Field Mus. nat. Hist. (Bot.) 17: 231 (1937). Type: Guatemala, Peten, Lundell 3122 (ILL!-holotype; F!-isotype). Fig. 17A,a,b. Shrubs 1-2.5 m high, stems slender, pale-glandular, slightly granular. Leaves 8-12 cm long, 2-4.5 cm wide, oblong- lanceolate, pale-glandular, glabrous, upper surface lustrous, underside with prominent veins, apex acute-acuminate, base unequally obtuse. Venation with 2-3 pairs of prominent secondary veins arising from the lower part of the midrib, ascending steeply to the apex. Petioles 3-8 mm long, gla- brous, with ligule-like structures 1-2 mm long. Prophylls 5-12 mm long, acute, glabrous. Inflorescences 2-4 cm long, erect or slightly curved; peduncles 4-9 mm long, glabrous. Anthers 0.2-0.3 mm long. Floral bracts 0.6-1 mm wide, triangular, densely yellow-ciliate, with dark centres. Fruits 1-1.2 mm wide, obovoid, round from above, densely pale- yellow pubescent; stigmas 3, sessile. Moist forest; 0-200 m. DISTRIBUTION. Mexico to Guatemala. Mexico, Chiapas: Shi- lom Ton 2604 (CAS). Belize, Cayo: Gentle 2606 (F). Guate- mala, Peten: Molina 15586 (F). Piper curvatipes is similar to P. jacquemontianum, with densely pubescent fruits and ciliate bracts on a stout inflores- cence. It can be distinguished by its much smaller, glabrous, glandular leaves, not as glossy as those of P. jacquemon- tianum, and its glabrous prophylls. 103. Piper jacquemontianum Kunth in Linnaea 13: 631 (1839). Type: Insula S. Domingo, Jacquemont s.n. (B-holotype). Fig. 17B,c,d. P. panamense C.DC. in J. Bot., Lond. 4: 216 (1866). Type: Panama, Chagres, Fendler 270 (K!-holotype; BM!,F!- isotypes). P. citrifolium sensu C.DC. in DC., Prodr. 16(1): 270 (1869). Type: Porto Rico, Blaumers.n. (G!-holotype). P. pilibaccum C.DC. in Bot. Gaz. 70: 179 (1920). Type: Costa Rica, Matamba, Nicoya Peninsula, Cook & Doyle 702 (US-holotype, photograph!). P. uvitanum C.DC. in Bot. Gaz. 70: 182 (1920). Type: Costa Rica, Limon, Pittier 12690 (G!-holotype). P. subcitrifolium C.DC. in Bot. Gaz. 70: 186 (1920). Type: Guatemala, Santa Rosa, Smith 3827 (US-holotype; G!-isotype). P. barbulatum C.DC. ex A. Schroed. in Candollea 3: 135 (1926). Type: Costa Rica, Nicoya, Tonduz 13697 (G!- holotype). P. orosianum Trel. in Contr. U.S. natn. Herb. 26: 143 (1929). Type: Costa Rica, Orosi, Cartago, Standley 39764 (US- holotype, photograph!). P. tabanicidum Trel. in Contr. U.S. natn. Herb. 26: 162 (1929). Type: Costa Rica, Tilaran, Standley & Valeria 45676 (US-holotype). P. aeruginosibaccum Trel. in J. Wash. Acad. Sci. 19: 336 (1929). Type: Honduras, near La Ceiba, Atlantida, PIPER IN THE NEW WORLD 43 Fig. 17 A: P. curvipes, habit; a: prophyll; b: fruit and bract. B: P. jacquemontianum, habit; c: prophyll; d: fruit and bract. C: P. biritak, habit; e: prophyll; f: fruit and bract. 44 M.C. TEBBS Standley 56735 (F-holotype; ILLl-isotype). P. dedititium Trel. in /. Wash. Acad. Sci. 19: 331 (1929). Type: Honduras, Quebrada Seca, Yoro, Standley 53937 (F-holotype; ILLl-isotype). P. onerosum Trel. in /. Wash. Acad. Sci. 19: 335 (1929). Type: Honduras, near Tela, Atlantida, Standley 53696 (F-holotype; ILL!-isotype). P. vexans Trel. in J. Wash. Acad. Sci. 19: 336 (1929). Type: Honduras, near Tela, Atlantida, Standley 54742 (F-holotype; ILL!-isotype). P. dimorphophyllum Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 12: 407 (1936). Type: Belize, Gentle 1387 (MICH-holotype; ILLl-isotype). P. gentlei Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 12: 407 (1936). Type: Belize, Corazal, Gentle 1077 (MICH-holotype; NY!-isotype). P. plumbeicolorTrel. in Publs Field Mus. nat. Hist. (Bot.) 17: 233 (1937). Type: Guatemala, Peten, Lundell 2554 (ILLI-holotype). P. catalinianum Trel. in Publs Field Mus. nat. Hist. (Bot.) 18: 335 (1937). Type: Costa Rica, Catalina, Guanacaste, Stork 2779 (ILLI-holotype). P. siquirresenseTre\. in Publs Field Mus. nat. Hist. (Bot.) 18: 361 (1937). Type: Costa Rica, Siquirres, Stork 2251 (ILLI-holotype). P. evulsipilosum Trel. in Publs Field Mus. nat. Hist. (Bot.) 9: 279 (1940). Type: Honduras, La Ceiba, Atlantida, Yuncker, Koepper & Wagner 8015 (ILLI-holotype; MOI-isotype). P. tumidipedunculum Trel. in Publs Field Mus. nat. Hist. (Bot.) 9: 281 (1940). Type: Honduras, Yoro, Yuncker, Koepper & Wagner 8151 (ILLI-holotype; MOI-isotype). P. gleasonii Yunck. in Bull. Torrey hot. Club 75: 287 (1948). Type: Guyana, Gleason 857 (NY-holotype). P. jactatum Trel. in Fieldiana Bot. 24: 305 (1952). Type: Guatemala, Alta Verapaz. Standley 90614 (Fl- holotype). Shrubs 2-3 (-4) m, young stems brown-pubescent. Leaves (3-) 10-22 cm long, 2.5-11 cm wide, narrowly lanceolate to broadly ovate-elliptic, upper surface glossy and mostly gla- brous, underside shortly pubescent on veins, apex acuminate, base unequally rounded, sometimes slightly lobed. Venation with 3-5 secondary veins mostly arising from the lower part of the midrib, curving to apex. Petioles 6-16 mm long, vaginate at base, occasionally with a minute ligule-like structure. Prophylls 10-20 mm long, acute, pubescent. Inflorescences 4-8 cm long, 2-4 mm wide in fruit; peduncles 3-12 mm long. Anthers 0.2-0.4 mm long. Floral bracts 0.7-1 mm wide, semi-lunar or triangular, densely yellowish-ciliate. Fruits 1-1.5 mm wide, obovoid, round from above, with dense covering of yellowish brown hairs; stigmas 3, linear, in slight depression at top of fruit. Cleared or disturbed areas, stream sides and partial shade in forest or dense scrub; 0-1500 m. DISTRIBUTION. Mexico to Panama. Mexico, Chiapas: Croat 40307 (MO); Laughlin 2566 (DS); Escuintla: Matuda 16720 (F); Quintana Roo: Tellez 1951 (BM); Tabasco: Cowan 29925 (DS); Veracruz: Rosas 1371 (F). Belize, Belize: Proctor 29546 (BM); Cayo: Sutton et al. 314 (BM); Honey Camp: Lundell 556 (DS). Guatemala, Alta Verapaz: Tuerckheim 1904 (NY); Escuintla: Standley 89278 (F); Izabal: Contreras 9973 (K); Peten: Contreras 8999 (K); Santa Rosa Standley 78378 (F). Honduras, Colon: Saunders 518 (MO); Cortes: Blackmore & Chorley 4003 (BM); Intibuca: Molina 13942 (F); Olancho: Molina: 8318 (F): Ruatan Island: Gaumer 47 (K). Nicaragua, Boaco: Stevens 9261 (MO); Chinandega: Baker 26 (DS); Zelaya: Stevens 18714 (BM, MO). Costa Rica, Cartago: Liesner 14389 (MO); Guanacaste: Tonduz 13697 (BM). Panama, Colon: Croat 36890 (BM, MO). Piper jacquemontianum is easily recognizable by its lustrous leaves and short, stout inflorescences with densely yellow brown pubescent fruits. It is related to P. curvatipes of Belize and Guatemala which has a similar inflorescence structure, but short, lanceolate, glandular leaves. 104. Piper biritak Trel. in Fieldiana Bot. 24: 289 (1952). Type: Guatemala, near Coban, Alta Verapaz, Standley 69345 (F!-holotype). Fig. 17C,e,f. Shrubs 1-2 m high, stems spreading, densely pubescent. Leaves 12-16 (-21) cm long, 2-5 cm wide, lanceolate, glan- dular, bullate, upper surface glossy, glabrous, undersurface lustrous, pubescent on veins, apex long-acuminate, base unequally rounded. Venation with 3-4 prominent secondary veins arising from the lower to middle part of the midrib, loop-connecting to apex, prominently reticulate. Petioles 3-10 mm long, pubescent, with pubescent ligule-like struc- ture 1-3 mm long. Prophylls 7-15 mm long, obtuse, minutely pubescent. Inflorescences 5-6 cm long, whitish or green, erect; peduncles 4-6 mm long, pubescent. Anthers 0.3-0.4 mm long. Floral bracts 0.6-0.8 mm wide, triangular, pale-ciliate with darker centres. Fruits 0.8-1 mm wide, obo- void, round from above, granular; stigmas 3. Rocky hillsides and disturbed forest, on limestone; 900-1500 m. DISTRIBUTION. Guatemala. Guatemala, Alta Verapaz: Croat 41387 (MO), Standley 91690 (F). Piper biritak is confined to the upland areas of Alta Verapaz. In the original description, the author states that the spikes are unknown. There is a collection from the same area (Croat 41387) which vegetatively matches the type and which has reasonably mature inflorescences. The inflorescence descrip- tion was taken from this specimen. The lanceolate leaves of this species are distinctive, lustrous on both the upper and under surfaces, and markedly bullate. 105. Piper pseudolindenii C.DC. in Linnaea 37: 336 (1872). Type: Oersted 888 (C.'-holotype). Fig. 18A,a,b. P. pseudo-lindenii var. magnifolium C.DC. in Linnaea 37: 336 (1872). Type: Costa Rica, Naranjo, Oersted s.n. (S-holotype). P. virillanum C.DC. in An. Inst. fis-geogr. C. Rica. 9: 158 (1897). Type: Costa Rica, bord du R. Virilla, Tonduz 9831, 10127 (G-syntypes). P. udimontanum C.DC. inAnnu. Conserv. Jard. hot. Geneve 21: 306 (1920). Type: Mexico, above Cuernavaca, Prin- gle 6828 (G-holotype; cml-isotype). P. pertractatum Trel. in Contr. U.S. natn. Herb. 26: 130 (1929). Type: Costa Rica, Nuestro Amo, Alajuela, Jimenez s.n. (US-holotype, photograph!). PIPER IN THE NEW WORLD 45 Fig. 18 A: P. pseudolindenii, habit; a: large leaf; b: prophyll; c: fruit and bract. B: P. sanctum, habit; d: prophyll; e: fruit and bract. 46 M.C. TEBBS P. magnifolium (C.DC.) Trel. in Contr. U.S. natn. Herb. 26: 131 (1929). Type: Costa Rica, Naranjo, Oersted s.n. (US-holotype). P. colaphitolerans Trel. in Publs Field Mas. nat. Hist. (Bot.) 17: 231 (1937). Type: Guatemala, Lake Peten, Lundell 3827 (ILL!-holotype). P. humorigaudens Trel. in Ann. Mo. hot. Gdn 27: 293 (1940). Type: Panama, north rim of El Valle de Anton, Code, Allen 1833 (ILL!-holotype). P. oblique-ovatum Trel. in Publs Field Mus. nat. Hist. (Bot.) 22: 10 (1940). Type: Guatemala, Concepcion, Chimalt- enango, Johnston 557 (F!-holotype). P. stillans Trel. & Standl. in Fieldiana Bot. 24: 325 (1952). Type: Guatemala, Quiche, Skutch 1822 (F!-holotype). Shrubs 1-4 m high, stems glabrous or minutely puberulent. Leaves 6-17 (-22) cm long, 2-7 (-9) cm wide, oblong- lanceolate to elliptic, slightly asymmetric, apex acuminate, base obtuse to slightly lobed and overlapping the petiole. Venation palmate, with 3-5 secondary veins arising from the base and ascending steeply to apex. Petioles 1-10 (-15) mm long, glabrous or minutely pubescent, with minute ligule-like structure. Prophylls 3-6 mm long, apex acute, glabrous. Inflorescences 4-10 cm long, minutely pubescent; peduncles 6-12 (-15) mm long, minutely pubescent. Anthers 0.2-0.3 mm long. Floral bracts 0.4-0.6 mm wide, triangular or round with ciliate margins. Fruits 1-2 mm wide, obovoid, round or rhomboid from above, glabrous; stigmas 3-4, ses- sile. Montane, deciduous and evergreen forest, damp ravines, edges of forest tracks and river banks; 0-2200 m. DISTRIBUTION. Mexico to Panama. Mexico, Chiapas: Croat 47402 (BM,MO); Jalisco: Breedlove & Almeda 45635 (CAS); Michoacan: Arsene 8445 (BM); Morelos: 6828 (BM); Oax- aca: Calderon 525 (BM); Veracruz: Calderon 1734 (CAS). Belize, Cayo: Lundell 6483 (DS). Guatemala, El Quiche: Proctor 25254 (MO); Izabal: Croat 41842 (MO); Peten: Contreras 5841 (DS). Honduras, Atlantida: Izaguirre 1 (MO); Comayagua: Molina 23334 (BM); El Paraiso: Molina 11916 (BM); Intibuca: Bonilla 100 (BM); Olancho: Nelson & Romero 4642 (BM); Santa Barbara: Cruz G-088 (MO). El Salvador, Santa Ana: Croat 42400 (MO); Sonsonate: Croat 42211 (MO). Nicaragua, Boaco: Stevens 9260 (BM,MO). Costa Rica, Cartago: Grayum & Sleeper 3274 (BM,MO); Puntarenas: Davidse 24468 (BM,MO); San Jose: Tonduz 7145 (BM). Panama, Chiriqui: Proctor 31822 (MO); Code: Alston 8721 (BM); Los Santos: Croat 34531 (BM,MO); Panama: Croat s.n. (BM,MO). Piper pseudo-lindenii can be distinguished by its slightly asymmetric, palmately veined leaves and the rather lumpy appearance of its infructescence. It is a fairly common species of disturbed forest, often growing at high altitudes. The original description of Piper oblique-ovatum Trel. was based on a sterile specimen collected by Johnston in Guate- mala. This plant has a shrubby, glabrous habit, with lanceolate-ovate leaves and the base slightly lobed, overlap- ping the petioles. In the Latin diagnosis of P. oblique- ovatum, the leaves are described as follows: 'folia lanceolato- ovata vel saepius late ovata-acuta, basi gibboso-cordata, lobo majore petiolum occulante.' This is very similar to that of P. pseudolindenii C.DC.: 'foliis breviter petiolatis lanceolato- oblongis apice acuminatis, acumine acuto, basi inaequalis gibbosis, latere majore rotundato minore acute.' The type material shows that both species have the same sort of leaf venation and inflorescence structure. Examination of a wide range of specimens has shown that plants with larger basal lobes tend to occur at lower altitudes. 106. Piper sanctum (Miq.) Schltdl. in DC., Prodr. 16(1): 330 (1869). Type: Mexico, Atlacomulco, Schiede 105 (?B- holotype). Fig. 18B,c,d. Artanthe sancta Miq., Syst. piperac.: 339 (1844). Piper papantlense C. DC. in DC., Prodr. 16(1): 338 (1869). Type: In sylvis Papantlae, Fischer 74 (?B-holotype). P. diandrum C.DC. in Linnaea 37: 364 (1872). Type: Mexico, Pital, Liebmann 55 (C!-holotype). P. venulosum Trel. in Contr. U.S. natn. Herb. 26: 132 (1929). Type: Costa Rica, San Ramon, Brenes 14192 (US- holotype, photograph!). P. dissimulans Trel. in Contr. U.S. natn. Herb. 26: 133 (1929). Type: Costa Rica, Tucurrique, Tonduz 12773 (US-holotype, photograph!). P. heterophlebium Trel. ex Standl. in Publs Field Mus. nat. Hist. (Bot.) 18: 345 (1937). Type: Costa Rica, San Jose, Skutch 2293 (US-holotype). Shrubs or small tree 1.5-4 (-5) m high, stems glabrous. Leaves 10-20 cm long, 7-16 cm wide, broadly ovate to ellip- tic, pale green, glabrous, apex acute-acuminate, base round to shallowly cordate on older, lower leaves. Venation pal- mate, with 3-7 prominent secondary veins arising from the leaf-base and ascending fairly steeply to apex. Prophylls 5-15 (-20) mm long, slender, apex acute, glabrous. Petioles 10-20 mm long. Inflorescences 7-22 cm long, erect, becom- ing pendulous in fruit; peduncles 12-25 mm long. Anthers 0.1-0. 2 mm. Floral bracts 0.3-0.5 mm wide, triangular to round, margins ciliate. Fruits 1 mm wide, obovoid, round from above, glabrous; stigmas 3-4. Wet forest, river banks, disturbed ground; 0-2500 m. DISTRIBUTION. Mexico to Costa Rica. Mexico, Chiapas: Breedlove 34522 (DS); Hidalgo: Croat & Hannon 66004 (BM,MO); Morelos: Croat & Hannon 65774 (BM,MO); Veracruz: Calderon 2232 (CAS). Belize, Toledo: Proctor 36075 (BM). Guatemala, Baja Verapaz: Tuerckheim 111704 (BM). Honduras, Colon: Ramos 193 (BM); Morazan: Molina 826 (BM). Nicaragua, Chontales: Grijalva & Rios 3451 (BM); Granada: Sandino 2941 (BM); Managua: Stevens 5409 (BM); Rivas: Sandino 4289 (BM); Zelaya: Stevens 6931 (BM). El Salvador, Ahuachapan: Berendsohn & Villacorta 1112 (BM); La Libertad: Berendsohn et al. 1044 (BM). Costa Rica, Guanacaste: Garwood et al. 682 (BM); San Jose: Tonduz 12773 (BM). Excluded species Piper gaudichaudianum Kunth in Linnaea 13: 638 (1839)-(as Steffensia gaudichaudiana) . The original description indicates that vegetatively this taxon is very like P. aduncum L. Specimens at the BM, named by Yuncker (1972) as P. gaudichaudianum, are actually P. aduncum L., with scabrous upper leaf surfaces, arching inflorescences, and round gla- brous fruits. As the holotype (Gaudicaud & Luschnath s.n.) was not available for this study, and judging from the original PIPER IN THE NEW WORLD 47 description, presumably has immature inflorescences, P. gau- dichaudianum has been excluded from this study. ACKNOWLEDGEMENTS. My grateful thanks are due to Dr A.J. Born- stein, Southeast Missouri State University, for advice and helpful comments. I would also like to thank the following herbaria for the loan of their specimens — B, C, CAS, E, F, G, ILL, K, MO, MU, S. REFERENCES Blanc, P. & Andraos, K. 1983. Remarques sur la dynamique de croissance dans le genre Piper L. (Piperaceae) et les genres affines. Adansonia 3: 259-282. Burger, W.C. 1971. Piperaceae. Flora Costaricensis. Fieldiana Bot. 35: 5-277 '. — 1977. The Piperaceae and the Monocots. Alternate hypotheses for the origin of monocotyledonous flowers. Bot. Rev. 43: 345-393. Duncan, T. 1983. Antonio Bertoloni's herbarium and the Florula Guatimalen- sis types. Taxon 32: 299. Kunth, K. 1839. Familie der Piperaceen. Linnaea 13: 561-726. Miquel, F.A. 1844. Systema Piperacearum. Rotterdam. Rafinesque, C.R. 1838. Sylva Telluriana. Philadelphia. Standley, P.C. & Steyermark, J.A. 1952. Flora of Guatemala. Fieldiana Bot. 24: 312-313. Tebbs, M.C. 1990. Revision of Piper (Piperaceae) in the New World. 2. The taxonomy of Piper section Churumayu. Bull. Br. Mus. nat. Hist. (Bot.) 20: 193-236. Trelease, W. & Yuncker, T.G. 1950. Piperaceae of Northern South America 1: 1-434. Urbana. Yuncker, T.G. 1972. The Piperaceae of Brazil. Hoehnea 2: 19-366. INDEX Accepted names are in roman and synonyms in italic, new names are in bold. Artanthe adunca (L.) Miq., 19 A. alaris (Ham.) Miq., 7 A. amplectans Miq., 22 A. catalpaefolia (Kunth) Miq., 7 A. caudata (Vahl) Miq., 7 A. celtidifolia (Kunth) Miq., 19 A. chamissonis Miq., 36 A. elongatum (Vahl) Miq., 19 A. flexuosa (Jacq.) Miq., 13 A. galleoti Miq., 19 A. hostmannianum Miq., 40 A. lanceaefolia (Kunth) Miq., 19 A. marginata (Jacq.) Miq., 7 A. mollicoma (Kunth) Miq., 21 A. olfersiana Miq., 26 A. radula (Kunth) Miq., 13 A. sancta Miq., 46 A. scabra (Sw.) Miq., 26 Heckeria peltata (L.) Kunth, 3 H. scutata Kunth, 3 H. speciosa (Kunth) Kunth, 3 H. umbellata (L.) Kunth, 3 section Lepianthes Raf . , 3 Lepianthes peltata (L.) Raf., 3 L. umbellata (L.) Raf., 3 Piper abuginiferum Trel., 28 P. achoteanum Trel., 22 P. aduncifolium Trel., 21 P. aduncum L., 19 P. aduncum var. brachyarthrum (Trel.) Yunck., 21 P. aduncum var. laevifolium C.DC., 21 P. aeruginosibaccum Trel., 42 P. alajuelanum Trel., 32 P. alare Ham., 7 P. alluvicolaC.DC.,26 P. alveatumTrel., 19 P. alveolatifolium Trel., 13 P. amphibium'Tre}., 13 P. amplectans (Miq.) C.DC., 22 P. anguilliispicum Trel., 21 P. angustifolium Ruiz & Pav., 19 P. anisatum Kunth, 7 P. anisophy Hum Trel., 15 P. anisophyllum var. granulatum Trel. ex Standl., 15 P. aquacalientis Trel. ex Standl., 28 P. aragonenseTrel., 10 P. arctilimbum Trel., 19 P. argentamentum Trel & Yunck., 28 P.argillicolaC.DC.,13 P. articulosum Trel. ex Standl., 28 P. atlantidanum Trel., 12 P. auriculiferumTrel., 15 P. austiniiTrel., 31 P. austinii var. aequilaterum Trel., 31 P. baculiferum Trel., 28 P. bailey orum var. paucispica Trel., 3 P. barbirostrelrel., 28 P. barbulatum C.DC. ex A. Schroed., 42 P. bayamonanum Trel., 26 P. biauritumC.DC.,36 P. bigelovii Trel., 12 P. biritakTrel., 44 P. bisasperatum Trel., 31 P. blepharilepidum Trel., 31 P. bocasense Trel. ex Standl., 26 P. brachistopodium C.DC., 31 P. bredemeyeri Jacq., 13 P. breve C.DC. ex Trel., 12 P. bmjoense Trel. & Standl., 42 P. callibracteum C.DC., 32 P. candicans Sodiro, 10 P. canyazasenseTrel., 12 P. capacibracteum Trel., 24 P. captum Trel. ex Standl., 31 P. carminisTrel., 28 P. carpi liter ami in C.DC., 40 P. cartagoanum C.DC., 24 P. casitenseTrel., 28 P. catalinianum Trel., 44 P. catalpaefolium Kunth, 7 P. cataractarum Trel., 26 P. caudatifolium Trel., 28 P. caudatum Vahl, 7 P. cayoense Trel. ex Standl., 24 P. celatipetiolum Trel., 15 P. celatipetiolum var. brenesii Trel., 15 P. celtidifolium Kunth, 19 P. cerro-puntoense Trel., 28 P. chagresianum Trel., 12 P. chamissonis (Miq.) Steud., 36 P. chamissonis var. rubellibracteum C.DC., 32 P. chrysostachyunt C.DC., 32 P. cinereumC.DC., 10 P. citrifolium sensu C.DC., 42 P. clavulispicum Trel. ex Standl., 12 P. coactoris Trel., 31 P. colaphitolerans Trel., 46 P. colonenseC.DC.,38 P. comatum Trel., 24 P. copacabanenseTrel., 13 P. coronatibracteum Trel., 28 P. coyolesenseTrsl., 12 P. crispatimargine Trel. ex Standl., 29 P. cuatrecasasiiTKl., 21 P. culebranum C.DC. ex A. Schroed., 38 P. cumbricola Trel., 21 P. curridabatanum Trel., 28 P. curvatipes Trel., 42 P. cyclophyllum C.DC., 15 P. davidianum C.DC., 32 48 M.C. TEBBS P. decurrensC.DC.,38 P. dedititium Trel., 44 P. deltoideocarpumTrel., 12 P. diandrumC.DC.,46 P. dilatatumRich., 13 P. dilatatum var. acutifolium C.DC., 15 P. dimorphophyllum Trel. ex Standl., 44 P. diquisanum C.DC., 32 P. disparifolium Trel., 15 P. disparipes Trel., 31 P. disparispicum Trel., 21 P. dissimulans Trel., 46 P. domingense C.DC., 12 P. dotaniim Trel., 34 P. dumeticolaC.DC., 12 P. echeverrianum Trel., 13 P. ejuncidum Trel., 40 P. elasmophyllum Trel., 12 P. elongatifolium Trel., 21 P. elongatum Vahl, 19 P. elongatum var. brachyarthrum Trel., 21 P. elongatum var. pampayacusum Trel., 21 P. emollitum Trel., 31 P. enganyanum Trel. & Yunck., 17 P. epigynium C.DC., 32 P. erectamentum C.DC., 26 P. evulsipilosum Trel., 44 P. excultum Trel., 19 P. falcigerum Trel., 15 P.fatoanumC.DC.,21 P. flavescens (C.DC.) Trel., 21 P. flavidum C.DC. ex Donn.Sm., 22 P. flexuosum Jacq., 13 P. fraguanum Trel., 29 P.friedrichsthaliiC.DC., 19 P. fuligineum K. until, 22 P. fusco-bracteatum Trel., 28 P. fusco-granulatum Trel., 34 P. gamboanum var. yapense Trel., 26 P. gaudichaudianum Kunth, 46 P. gentlei Trel. ex Standl., 44 P. genuflexum Trel., 26 P. gleasonii Yunck., 44 P. goergeriTre\., 19 P. gonagricum Trel., 26 P. gracilipedunculum Trel., 38 P. granulatum Trel., 24 P. griseo-pubens Trel., 12 P. griseo-pubens var. revocabileTre}., 12 P. hanckeli Trel. ex Standl., 32 P. heterophlebium Trel. ex Standl., 46 P. hirsutum var. tonduzii C.DC., 26 P. hirtellipetiolum C.DC., 38 P. hispidum Kunth, 26 P. hispidum Sw., 26 P. holdridgeianum W.C. Burger, 10 P. hostmannianum (Miq.) C.DC., 40 P. humoense Trel. ex Standl., 28 P. humorigaudens Trel., 46 P. illudens1ie\.,2\ P. imparipes Trel., 31 P. indignum Trel., 24 P. infraleucum Trel., 40 P. inhorrescens Trel., 28 P. injucundum Trel., 31 P. injucundum var. praecalvinervium Trel., 28 P. injucundum var. praepubinervium Trel., 28 P. insolens Trel., 36 P. intersittumTrel., 21 P. jacquemontianum Kunth, 42 P. jactatum Trel., 44 P. kiUipiTnl.,26 P. killipi var. calderanum Trel., 26 P. konkintoense Trel., 28 P. kuntzeiC.DC.,21 P. laevifolium C.DC., 15 P. laevius(C. DC). Trel., 24 P. lanatibracteum Trel., 28 P. lanceifolium Kunth, 19 P. lanceifolium var. acutiusculum C.DC., 19 P. lanosibracteum Trel., 28 P. leptoneuron C.DC., 38 P. leucophlebium Trel., 24 P. linearifolium C.DC., 19 P. lineatum var. hirtipetiolatum Trel., 21 P. lineatum var. leucolepidum Trel., 19 P. HratinerveTiel., 19 P. HttoraleC.DC., 17 P. luridispicum Trel. ex Standl., 32 P. maestranum Trel., 26 P. magnifolium (C.DC.) Trel., 46 P. malpasoensis Tebbs, 17 P. margin at u m Jacq., 7 P. marginatum var. catalpaefolium (Kunth) C.DC., 7 P. margaretae Trel., 28 P. marginatum var. anisatum (Kunth) C.DC., 9 P. marginatum var. marginatum forma catalpaefolium (Kunth) Steyerm., 9 P. marginatum var. niceforoi (Trel. & Yunck.) Steyerm., 9 P. maternaleTrel., 17 P. meritum Trel., 28 P. micoense Trel., 24 P. milciadesii'Tre\. & Yunck., 17 P. mollicomum Kunth, 21 P. multiplinervium C.DC., 10 P. niceforoi Trel. & Yunck., 9 P. nigricauleTrel., 12 P. nitidifolium C.DC., 32 P. nodosum C.DC., 31 P. nubigaudens Trel., 42 P. obiter-sericeum Trel. ex Standl., 13 P. oblanceolatum Trel., 38 P. oblanceolatum var. fragilicaule Trel., 21 P. oblique-ovatum Trel., 46 P. olivaceum C.DC., 21 P. onerosum Trel., 44 P. otophorumC.DC., 17 P. orosianum Trel., 42 P. palmasanum C.DC., 29 P. palustreC.DC.,22 P. panamense C.DC., 42 P. papantlense C. DC, 46 P. papulaecaule Trel. ex Standl., 32 P. population Trel., 31 P. patulum Bertol., 7 P. patulum var. cordifolium Trel., 9 P. pavasense Trel., 28 P. pejivallense Trel., 26 P. pelliticaule Trel., 13 P. peltatum L., 3 P. peracuminatum C.DC., 34 P. perfugii Trel. , 40 P. pergeniculatum Trel., 26 P. per hispidum C.DC., 36 P. pert ractatu in Trel., 44 P. pervicax Trel., 28 P. pexum Trel., 29 P. phaeophyllum Trel., 28 P. phanerolepidum Trel., 28 P. picardae var. pilinervium Trel., 10 P. pictamentum Trel., 22 P. pictamentum var. fluminis Trel., 22 P. pileatum Trel., 36 P. pileatum var. obliquum Trel., 36 P. pilibaccum C.DC., 42 P. pisoenseC.DC.,2l P. plumbeicolor Trel., 44 P. poasanum C.DC., 29 P. polytrichum C.DC. ex A. Schroed., 34 P. ponendum Trel. ex Standl., 12 P. pruinosum Kunth, 3 P. pseudo-albuginiferum Trel., 24 P. pseudoasperifolium C.DC., 24 P. pseudo-dilatatum C.DC., 12 P. pseudofuligineum C.DC., 12 P. pseudolanceaefolium Trel., 19 P. pseudolindenii C.DC., 44 P. pseudo-lindenii var. magnifolium C.DC., 44 P. pseudo-marginatum C.DC., 7 PIPER IN THE NEW WORLD 49 P. pseudopsis C.DC., 13 P. pseudovelutinum C.DC., 21 P. pseudo-velutinum var. flavescens C.DC., 19 P. pseudo-viridicaule var. nievecitanum Trel., 29 P. pubens Trel., 31 P. pullibracteatum Trel., 28 P. punctiunculatum Trel., 28 P. pustulicaule Trel. ex Standl., 31 P. quadratilimbumTrel., 12 P. radula Kunth, 13 P. rectamentum Trel., 29 P. rivi-vetustilK\., 28 P. rivJfl/W Trel & Yunck., 28 P. rotundibaccum Trel., 40 P. rotundibaccum var. fraijanesanum Trel., 40 P. rubripes Trel. ex Standl., 32 P. rugosifolium Trel., 36 P. sabanillanum Trel., 26 P. salinasanum C.DC., 12 P. salinasanum var. subscabrifolium C.DC., 12 P. salutatrix Trel. ex Standl., 12 P. sancti-felicisTrel.,29 P. sanctum (Miq.) Schltdl., 46 P. san-joseanum C.DC., 7 P. san-joseanum var. chiriquinum Trel., 9 P. san-joseanum var. kobense Trel., 9 P. san-joseanum var. m/norTrel., 9 P. san-joseanum var. panamanum Trel., 9 P. san-joseanum var. remediosense Trel., 9 P. san-joseanum var. tabogense Trel., 9 P. scabrilimbum C.DC., 26 P. scabrum Sw., 26 P. scalarispicum Trel., 42 P. scalpens Trel., 28 P. segoviarum Standl. & L.O.Williams, 42 P. sibunense Trel. ex Standl., 24 P. silvanorum Trel., 29 P. silvivagumC.DC.,24 P. sinuatifolium Trel., 15 P. siquirresense Trel., 44 P. spedosum Kunth, 3 P. sperdinum C.DC., 17 P. spicilongum Trel., 29 P. squali-pelliculum Trel., 12 P. stenocladum C.DC., 32 P. sritfa/w Trel. & Standl., 46 P. subaequilaterum Trel., 13 P. subasperatum Trel. ex Standl., 28 P. subcitrifotium C.DC., 42 P. subdivaricatum Trel., 32 P. subflavumC.DC., 10 P. sublaevifolium Trel., 15 P.submoUeTKl.,21 P. subsericeum Trel., 13 P. sumideranum Trel., 26 P. surubresanum Trel., 32 P. tabanicidum Trel., 42 P. taboganumC.DC., 12 P. tacare?ert.se Trel. ex Standl., 32 P. talamancanum Trel., 24 P. tapianum Trel., 38 P. tentatum Trel. ex Standl., 29 P. tenuimucronatum C.DC., 40 P. terrabanumC.DC., 15 P. thomasii Tebbs, 15 P. tikalenseTrel., 26 P. torresanum Trel., 28 P. tortuosipilum Trel., 36 P. trachydermum Trel., 26 P. tractifolium Trel., 40 P. trichophlebium Trel., 28 P. triquetrofructum Trel. ex Standl., 13 P. tumidipedunculum Trel., 44 P. udimontanum C.DC., 44 P. umbellatum L., 3 P. umbricolaC.DC.,31 P. uncatum Trel., 9 P. wv/Yanw/rtC.DC.,42 P. valetudinari Trel., 28 P. varablancanum Trel., 38 P. variitrichum Yunck., 29 P. ventoleranum Trel., 31 P. venulosum Trel., 46 P. verbenanum C.DC., 12 P. verruculaepetiolum Trel. ex Standl., 15 P. verruculigerum Trel., 32 P. vestitifolium C.DC., 24 P. vejca/w Trel., 44 P. via-chicoense Yunck., 12 P. vicinum Trel., 32 P. villibracteum C.DC., 13 P. villiramulum C.DC., 24 P. villistipulum Trel., 32 P. villosisquamulumTre\.,32 P. virillanumC.DC.,44 P. vitabile Trel. ex Standl., 12 P. vitibundum Trel., 24 P. wedelii Yunck., 15 P. williamsiiTre\.,26 P. yalochanum Trel., 26 P. yapeanum Trel., 26 P. zacatenseC.DC.,34 P. zacatense var. percaudatum C.DC., 34 P. zonulatispicum Trel., 40 Pothomorphe almirantensis Trel., 3 P. baileyorum var. paucispica Trel., 3 P. pe/tata(L.)Miq.,3 P. scutata (Kunth) Miq., 3 P. speciosa (Kunth) Miq., 3 P. tecumensis Trel., 3 P. umbellata (L.) Miq., 3 section Radula Miq., 6 Schilleria catalpaefolia (Kunth) Kunth, 7 5. caudata (Vahl) Kunth, 7 5. marginata (Jacq.) Kunth, 7 Steffensia adunca (L.) Kunth, 19 5. flexuosa (Jacq.) Kunth, 13 5. fuliginea (Kunth) Kunth, 22 5. hirsute (Sw.) Kunth, 26 5. radula (Kunth) Kunth, 13 S. scabra (Sw.) Kunth, 26 Bull. nat. Hist. Mus. Land. (Bot.) 23(1): 51-54 Issued 24 June 1993 Mounting techniques for the preservation and analysis of diatoms STEPHEN J. RUSSELL Department of Botany, The Natural History Museum, London SW7 5BD INTRODUCTION In order that diatoms can be safely stored and preserved, they are mounted and displayed on microscope slides. These slides are curated to provide a scientific, taxonomic, and historical reference collection. The method presented here will provide new workers with a comprehensive schedule for the prepara- tion of consistent, good quality slides. EQUIPMENT The only specialized equipment required is a low power, wide field dissection microscope and a small revolving table for ringing coverslips. Ideally the microscope should be fitted with forearm rests which allow total freedom of wrist and hand movement. All other pieces of equipment can be cheaply acquired or handmade. Above all the operator should be comfortable and all the equipment arranged logically within easy reach. The working area should be well lit, clean, and free from draughts and distraction; on occasion difficult mounts require great con- centration and patience. METHOD The preparation, mounting, and finishing steps are laid out below. Slide preparation 1. Take a new microscope slide and with a moist tissue dipped in scouring powder gently abrade one surface. (This keys the glass surface ready for mounting.) Thoroughly clean the slide in warm soapy water to remove the scouring powder and any grease, fingerprints, etc. Rinse the slide in distilled water and store under ethanol spirit. 2. Polish a clean coverslip (No. 0. 13 mm diameter) with a lens tissue or fibre-free cloth. 3. Place a small drop of distilled water on the centre of a revolving table and place the coverslip onto the water. A sufficiently small drop of water will spread under the cover- slip and cause it to adhere to the revolving surface. Carefully centralize the coverslip on the table. 4. Trim a fine paint brush to a point and using a quality Indian ink, scribe a circle of the required size. To do this, gently spin the coverslip and touch the glass surface at one point with the tip of the paint brush lightly loaded with ink. Practice will enable rings of the desired size and thickness to be made. Put the prepared coverslip to one side to dry. If required, batches of slides and coverslips can be prepared prior to a mounting session. Mounting 5. Take a prepared slide, leave to air dry and wipe with a lens tissue. Using a marked template, spot the centre of the unsecured slide surface with a fibre tipped pen. 6. Using a fine paint brush, apply as thin a smear of mountant glue as possible to the abraded surface. Gentle sweeping of the glue over the spotted area with the brush will remove any dust, airborne or static contamination. 7. Using a pig's eyelash or finely drawn glass fibre, pick up the diatom(s) and with gentle manipulation mount in the desired orientation over the ink spot. The viscosity of the glue will allow several mounted diatoms to be gently pushed together in a discrete area for comparative viewing, even under eventual high power microscopic examination. 8. Gently waft the slide over a low spirit lamp flame. The glue evaporates and hardens to a transparent film invisible under microscopic analysis. This will secure the diatoms as orientated during the rest of the slide preparation. 9. Take a prepared coverslip and with the ringed surface uppermost place a drop of mountant in the centre. Invert the coverslip and gently position the ringed area above the ink spot on the slide. Alternatively the mountant may be applied to the slide surface. Allow the mountant to spread under the coverslip as it settles under its own weight, guiding it into position if necessary. Do not press the coverslip down under any circumstances — damage to the diatom is guaranteed! Finishing steps 10. Where time is not a limiting factor, the slide may now be placed in a hot 60°C oven and left to cure. For exceptionally fragile specimens or precious material this is probably the only safe way of making a slide. Gentle hardening of the mountant removes the need for further processing until the specimen is safely preserved. It takes about a week to cure a slide completely. 11. In many cases enclosed frustules trap air during mount- ing, which causes considerable aberration of detail under microscopic examination. To remove trapped air bubbles and considerably speed up the slide making process, the moun- tant can be rapidly hardened by cooking the slide. After step 52 S.J. RUSSELL 9, gently warm the slide by wafting over a low spirit flame. This gentle warming reduces the viscosity of the mountant and the spirit diluent bubbles out of solution, rapidly curing the mountant. At the same time the trapped air is normally forced out of the specimen, replaced by the less viscous mountant. Any small air pockets which remain frequently shrink and disappear as the slide cools. Persistent air pockets may require several warming and cooling cycles. Gently tapping the slide on a hard surface may dislodge 'stuck' bubbles, but beware, this has the additional risk of dislodging the glued specimen(s), or in extreme cases, causing actual physical damage. 12. Once the slide has cooled to room temperature and the mountant is fully cured, any excess mountant which has bubbled out around the edges of the coverslip may be removed. This is done firstly, by gentle chipping and scraping with a pointed scalpel, and then by wiping away any final traces with a toluene soaked wipe. 13. Polish the slide carefully, removing any remaining ink spot under the slide. Do not press hard on the coverslip surface as the diatoms may break under the pressure. 14. The final step in the process is to examine critically the mounted specimen(s) under a high magnification. If the slide is acceptable then it should be labelled with as much informa- tion as possible. The Natural History Museum has purpose printed labels which are gummed to the slide and where available, the following information is written in ink using a very fine mapping pen: the date of mounting, the mounter's name or initials, the type of mountant used, a specimen identification (if known), the locality or collection site, a core number (if applicable), date of collection, and collector's name. The annotated slide labels are stuck to the slide and orientated such that they can be read in the slide trays without needing to be removed. MOUNT ANTS In the past a succession of mountants have found favour with slide makers. Their use today is restricted on the grounds of health and safety. The properties of a good mountant are that it should be safe, indefinitely stable, and totally transparent when cured. The mountant should be readily available, and have a viscosity which may be altered to a required consis- tency. The mountant which fulfills these criteria and is used consistently in the diatom section at The Natural History Museum is Naphrax (N.B.S., 3 Betts Avenue, Martlesham Heath, Ipswich). Naphrax is supplied as a ready-to-use liquid diluted in toluene. Naphrax may be too fluid for some mounts as supplied, but a small sample left in a warm oven will increase in viscosity. Conversely, after a while the mountant may become too thick; dilution with toluene will remedy the problem. MOUNTING GLUE The glue used (step 6) should be totally transparent when dried, and of a viscosity that allows diatoms to be manipu- lated during the positioning part of the slide making proce- dure. Once in position the glue should be such that it may be dried quickly, ensuring the secure positioning and orientation of the specimens during the rest of the slide preparation. The glue used in the diatom section is prepared in the following way. A saturated solution of gum tragacanth in sterile dis- tilled water is prepared and left to stand for at least a week in an airtight container. The clear supernatant layer is removed, and glycerol 10% w/v added. To prevent microbial growth a few crystals of phenol may be added or an anti-fouling agent. It is only neccessary to prepare a small amount of glue at a time; 10 mis can last as long as a year with constant use. ADDITIONAL NOTES More specialized techniques for the mounting of diatoms require a certain amount of manual dexterity, especially where large numbers of specimens are mounted on a single slide. Coverslips or slides etched with grids remove the need for ringing as an aid to specimen location. Mounting selected diatoms on the coverslip, instead of on the slide, is sometimes preferred, as this allows precise positioning within the ringed area. The diatoms are presented fractionally nearer the microscope lens and in some instances this can afford greater image resolution under high power examination. The potential drawback to this is that if the glue is not applied thinly enough then puddling occurs at the point of diatom adherence and considerable aberration of the image results. Many specimens are very fragile and/or have thin delicate silica processes and when mounted on thin edges will smash under the weight of a settling coverslip. To help prevent this, small pieces of broken coverslip or sponge spicules may be glued around the coverslip edge to support the weight during slide making. Sometimes the mountant will shrink during the curing process; also, where insufficient mountant has been used air pockets form. This in itself may not interfere with micro- scopic examination, but if the whole undersurface of the coverslip is not evenly supported the coverslip will flex, with an increased tendency for it to crack, especially during slide cleaning. In the event of such a problem occurring, it is possible to save the slide by placing a small drop of mountant close to the coverslip near the air pocket and gently warming the slide. The mountant will be drawn under the coverslip and should displace any air pockets. If a coverslip is broken and the specimen needs to be saved it is possible to repair a slide. Place the whole slide in a container and cover it in neat mountant diluent. The mountant will become soft and even- tually melt, dissolving sufficiently to allow the coverslip to be gently lifted free. The specimen may remain glued to the slide, in which case remount with a fresh coverslip. In some cases, the glue will dissolve and the diatom should be recovered, washed in diluent and further washed in a descending alcohol series dilution into water, then dried and the mounting procedure repeated. When the diatom frustule is composed of very fine pores, the viscosity of the mountant, even during warming, will not allow it to penetrate the diatom. Thinning the mountant should overcome the problem, but extra mountant to account for the additional spirit will need to be applied. Alternatively add a drop of spirit directly to the specimen on the slide, which will enable the mountant to penetrate the specimen more easily. DIATOM MOUNTING TECHNIQUES 53 Fig.l. A Coscinodiscus fossil diatom slide, photographed using differential interference contrast photomicrography under high magnification (x!600). This demonstrates the high resolution of surface detail possible using the mounting technique described. 54 S.J. RUSSELL At all times when preparing slides be aware of the potential volatility of the mountant and diluent, especially when using naked flames. Potential fire risk may be avoided by having small volumes in tightly sealed bottles at the work station and bulk supplies suitably stored elsewhere. When curing slides take care not to boil the mountant excessively, as this will uncontrollably bounce the coverslip and smash the specimen. Extreme cases of boiling, or where dilute mountant is used can cause the evacuated vapour to flash ignite. Although this is not in itself dangerous (the amount of mountant present is too small), a surprised technician may drop the slide. Avoid placing hot slides on cold surfaces after curing as they will crack. Patient waiting until the slide cools is best, but bridging the slide across two wooden blocks will free the hands. STREWN SLIDES An alternative to individually mounted specimens, particu- larly for routine sample analysis (after sample cleaning, for example) or population monitoring, is that of strewn mount- ing. The basic slide making technique is similar to that for selected slides, but differs in the following way. A cleaned diatom sample in distilled water is shaken gently and a small amount is pipetted through a large bore onto the centre of a glass slide or cover-slip (No.O. 19 mm diameter) and left to dry, either in a warm 37°C oven or air dried. Drying a strew too quickly (i.e. using a flame) sets up small convection currents and the specimens, especially the smallest, aggregate into a crust at the air/water interface making analysis difficult. Preparation of the strews by drying down overnight before mounting will enable good evenly distributed strews to be made. This method, if followed carefully, will enable the new- comer to produce diatom (or similar organisms) slides of good quality. Like all skills the initial results may seem poor but with practice ambitious mounts will be achieved. 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 1 No. 1 Contributions to our knowledge of Old World Araliaceae. W.R. Philipson. 1951. Pp. 3-20. An undescribed species of Mastlchodendron (Sapotaceae) from Barbados and Antigua. H.E. Box & W.R. Philipson. 1951. Pp. 21-24, 1 plate. £2.80 No. 2 Out of print. No. 3 Notes on Podostemaceae for the revision of the Flora of West Tropical Africa. G. Taylor. 1953. Pp. 51-79, 14 figs. £2.15 No. 4 Notulae criticae and floram hispaniae pertinentes, 1. V.H. Heywood. 1954. Pp. 81-122. £2.95 Nos 5-8 Out of print. Volume 2 No. 1 Out of print. No. 2 No. 3 Out of print. Novitates Himalaicae-1. F. Ludlow & W.T. Steam. 1956. Pp. 65-81, 8 plates, 11 figs. £2.15 Saxifraga of the Himalaya 1. Section Kabschia. H. Smith. 1958. Pp. 83-129, 14 figs. £4.50 Out of print. Out of print. A revision of the genus Petrorhagia. P.W. Ball & V.H. Heywood. 1964. Pp. 119-172, 3 plates, 22 figs. £1.95 Marine algae of Gough Island. Y.M. Chamberlain. 1965. Pp. 173-232, 4 plates, 80 figs. £6.00 Out of print. Out of print. A synopsis of Jamaican Myrsinaceae. W.T. Steam. 1969. Pp. 143-178, 8 plates, 25 figs. £3.65 The Jamaican species of Columnea and Alloplectus (Gesneriaceae). W.T.Stearn. 1969. Pp. 179-236, 8 plates, 29 figs. £5.15 New or little known Himalayan species of Swertia and Veratrilla (Gentianaceae). H. Smith. 1970. Pp. 237-258, 16 plates, 7 figs. £3.75 A survey of the tropical genera Oplonia and Psilanthele (Acanthaceae). W.T. Steam. 1971. Pp. 259-323, 10 plates, 18 figs. £5.95 Angiosperms ofthe islands of the Gulf of Guinea (Fernando Po, Principe, S. Tome, and Annobon). A.W. Exell. 1973. Pp. 325-411. £6.00 Volume 5 No. 1 The dryopteroid ferns of Ceylon. W.A. Sledge. 1973. Pp. 1-43, 4 figs. £3.00 No. 2 New Himalayan and Tibetan species of Corydalis (Papaveraceae). F.Ludlow & W.T. Stearn. 1975. Pp. 45-69, 15 plates, 14 figs. £3.85 No. 4 Nos 5-12 Volume 3 Nos 1-3 No. 4 No. 5 No. 6 Volume 4 Nos 1-3 No. 4 No. 5 No. 6 No. 7 No. 3 The marine algae of Trinidad, West Indies. W.D.Richardson. 1975. Pp. 71-143, 12 plates, 2 figs. £6.80 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. £7.25 No. 5 Frank Ludlow (1885-1972) and the Ludlow-Sherriff expeditions 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. £5.25 No. 6 Studies in the genus Hypericum L. (Guttiferae). 1. Infrageneric classification. N.K.B. Robson. 1977. Pp. 291-355, 9 figs. £7.25 No. 7 Sphagnales of tropical Asia. A.Eddy. 1977. Pp. 357-445, 4 plates, 17 maps, 25 figs. £9.25 Volume 6 No. 1 The handwriting of Joseph Banks, his scientific staff and amanuenses. J.B. Marshall. 1978. Pp. 1-85, 62 figs. £9.50 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. £15.00 No. 3 The lichenicolous Hyphomycetes. D.L. Hawksworth. 1979. Pp. 183-300, 47 figs. £15.00 No. 4 The species of Chisocheton (Meliaceae). D.J. Mabberley. 1979. Pp. 301-386, 3 plates, 10 figs. £15.00 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. £9.00 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. £2.20 No. 3 A revision of the genus Anacyclus L. (Compositae: Anthemideae). C.J. Humphries. 1979. Pp. 83-142, 27 figs. £8.00 Volume 8 No. 1 The Thelypteridaceae of Ceylon. W.A. Sledge. Pp. 1-54, 5 figs. 1981. £7.75 No. 2 Studies in the genus Hypericum L. (Guttiferae) 2. Characters of the genus. N.K.B. Robson. 1981. Pp. 55-226, 73 figs. £23.50 No. 3 A revision of the lichen family Thelotremataceae in Sri Lanka. M.E. Hale, Jr. 1981. Pp. 227-332, 20 figs. £14.50 No. 4 Vascular plant collections from the Tristan da Cunha group of islands. E.W. Groves. Pp. 333-420, 33 figs. £12.50 Volume 9 No. 1 The lichenicolous Coelomycetes. D.L. Hawksworth. 1981. Pp. 1-98, 36 figs. £13.25 No. 2 The genus Callithamnion (Rhodophyta: Ceramiaceae) in the British Isles. P.S. Dixon & J.H. Price. 1981. Pp. 99-141, 5 figs. £6.00 No. 3 Parmelia subgenus Amphigymnia (lichens) in East Africa. H. Krog & T.F.V. Swinscow. 1981. Pp. 143-231, 31 figs. £12.25 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. £13.50 Volume 10 No. 1 Taxonomic studies in the Labiatae tribe Pogostemoneae. J.R. Press. 1982. Pp. 1-89, 33 figs. £12.75 No. 2 The typification of Hudson's algae: a taxonomic and nomenclatural reappraisal. L.M. Irvine & P.S. Dixon. 1982. Pp. 91-105. £2.25 No. 3 Seaweeds of the Faroes. Various authors. 1982. Pp. 107-225, 13 figs. £16.75 No. 4 The lichen genus Steinera. A.M. Henssen & P.W. James. 1982. Pp. 227-256, 24 figs. £4.50 Volume 11 No. 1 The algae of Lightfoot's Flora scotlca. P.S. Dixon. 1983. Pp. 1-15, 2 figs. £2.30 No. 2 A taxonomic study of the lichen genus Micarea in Europe. B.J. Coppins. 1983. Pp. 17-214, 57 figs, 28 maps. £27.75 No. 3 The hepatics of Sierra Leone and Ghana. E.W. Jones & A.J. Harrington. 1983. Pp. 215-289, 8 figs. £10.50 No. 4 Studies in the Corallinaceae with special reference to Fosliella and Pneophyllum in the British Isles. Y.M. Chamberlain. 1983. Pp. 291-463, 89 figs. £24.50 Volume 12 No. 1 A revision of the Morinaceae (Magnoliophyta- Dipsacales). M.J. Cannon & J.F.M. Cannon. 1984. Pp. 1-35, 9 figs. £5.40 No. 2 An introduction to fern genera of the Indian subcontinent. C.R. Fraser-Jenkins. 1984. Pp. 37-76, 1 fig. £6.00 No. 3 A revision of African Sphagnales. A. Eddy. 1985. Pp. 77-162, 47 figs. £13.50 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. £28.80 Volume 13 No. 1 The lichen genus Usnea subgenus Neuropogon. F.J. Walker. 1985. Pp. 1-130, 39 figs. £19.80 No. 2 Cytotaxonomic studies of the ferns of Trinidad. A.C. Jermy & T.G. Walker. 1985. Pp. 131-276, 69 figs. £22.00 No. 3 Some genera of the Biddulphiaceae (diatoms) with interlocking linking spines. R. Ross & P. A. Sims. 1985. Pp. 277-381 , 33 plates. £20.00 Volume 14 No. 1 Cytological observations on Indian subcontinent and Chinese Dryopteris and Polystichum (Pteridophyta: Dryopteridaceae). M. Gibby. 1985. Pp. 1-42, 78 figs. £6.00 No. 2 A redisposition of the species referred to the ascomycete genus Microthelia. D.L. Hawksworth. 1985. Pp. 43-181, 73 figs. £24.00 No. 3 A classification of the genus Cryopteris (Pteridophyta: Dryopteridacaee). C.R. Fraser-Jenkins. 1986. Pp. 183-218, 4 figs. £5.60 No. 4 Evolutionary cladistics of marattialean ferns. C.R. Hill & J.M. Camus. 1986. Pp. 219-300, 27 figs. £14.50 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. £23.50 No. 2 Cytology of the fern flora of Madeira. I. Manton, J.D. Lovis, G. Vida & M. Gibby. 1986. Pp. 123-161, 12 plates. £7.50 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, 1 17 maps. £38.00 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. £20.00 No. 2 The lichen genus Ramalina in Australia. G.N. Stevens. 1987. Pp. 107-233, 15 plates, 31 figs. £22.50 No. 3 An annotated list of vascular plants collected in the valleys south of Mt Everest. G. Miehe. 1987. Pp. 225-268, 4 figs. £8.50 No. 4 Further genera of the Biddulphiaceae (diatoms) with interlocking linking spines. R. Ross & A.Sims. 1987. Pp. 269-311, Opiates. £8.00 Volume 17 No. 1 Studies in Pseudocyphellaria (lichens) 1. The New Zealand species. D.J. Galloway. 1988. Pp. 1-267, 124 figs. £51.00 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. £28.00 No. 2 No. 3 No. 4 No. 5 Erik Acharius and his influence on English lichenology. D.J. Galloway. 1988. Pp. 149-194, 18 figs. £8.80 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Phodophyta (Florideae) 2. Genera G.J.H. Price, D.M. John & G.W. Lawson. 1988. Pp. 195-273, 1 fig. £14.80 Some Cretaceous and Palaeogene Trinacria (diatom) species. P.A. Sims & R. Ross. 1988. Pp. 275-322, 13 plates. £9.10 A monograph of Dryopteris (Pteridophyta: Dryopteridaceae) in the Indian subcontinent. C.R. Fraser-Jenkins. 1989. Pp. 323-477, 79 figs. £30.00 Corydalis (Papaveraceae: Fumarioideae) in Nepal. M. Liden. 1989. Pp. 479-538, 26 figs. £11.20 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. 5-61, 36 figs. A revision of the Solarium nitidum group (section Holo- phylla 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. 103-112, 8 figs. The application of names of some Indian species of Ocimum and Geniosporum (Labiatae). J.R. Press & V.V. Sivarajan. 1989. Pp. 113-116, 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. 117-158, 41 figs. £48.00 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. £45.00 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. £25.00 Volume 21 No. 1 Historical and taxonomic studies in the genus Titanoderma (Rhodophyta, Corallinales) in the British Isles. Y.M. Chamberlain. 1991. Pp. 1-80, 247 figs. 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. 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. £37.00 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 Solatium section Geminata (Solanaceae) from Cerro del Torra 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. £37.50 CONTENTS 1 Revision of Piper (Piperaceae) in the New World 3. The taxonomy of Piper sections Lepianthes and Radula M.C. Tebbs 51 Mounting techniques for the preservation and analysis of diatoms S.J. Russell bulletin of The Natural History Museui BOTANY SERIES Vol. 23, No. 1, June 1993