Cecropiaceae: Coussapoa and Pourouma, with an ... - CNCFlora

CECROPIACEAE: COUSSAPOA AND POUROUMA, WITH AN INTRODUCTION TO THE FAMILY C. C. BERG,1 R. W. A. P. AKKERMANS,2 <strong><strong>an</strong>d</strong> E. C. H. v<strong>an</strong> Heusden3 CONTENTS Abstract .................................................................. 2 Introduction to the Family (C. C. BERG) ....... ..................................... ............ 2 History <strong><strong>an</strong>d</strong> Circumscription of the Family ..... ................. ............................ 2 Description of the Family ......... ........................................ .. .............. 3 Key to the Neotropical Genera ................................................ ............. 3 Genus Description of Cecropia .... ......................................................... 3 <strong>Coussapoa</strong> (C. C. BERG AND R. W. A. P. AKKER NS) ............................................ 4 In tro d u ctio n ............................................................................. 4 Taxonom ic H istory ....................................................................... 4 M o rp h o logy ........... ........................ ........................................... 4 A natom y (K . BoNSEN ) .................................................................... 8 Distribution <strong><strong>an</strong>d</strong> Ecology ......................... ......................................... 10 Systematic Position of the Genus ........................................................... 15 Systematic Arr<strong>an</strong>gement of the Species ...................................................... 15 Taxonom y ........ ........................ .............................................. 16 Generic D escription .................................................................. 16 K eys to the Species .................. ........ ......................................... 17 Treatments of the Species <strong><strong>an</strong>d</strong> Subspecies .... .................................... 28 U nnam ed Collections .... ........................ .......... .......................... 108 Names Not Included .................................................................. 109 <strong>Pourouma</strong> (C. C. BERG AND E. C. H. VAN HEUSDEN) .............................................. 110 Introduction ............................................................................. 110 Taxonom ic H istory ....................................................................... 111 M orphology .............................................................................. 111 Phenology ............................................................................... 113 Pollination .................. .... ......................................................... 113 Dispersal ............................................................................... 114 R elations w ith A nts ....................................................................... 114 Distribution <strong><strong>an</strong>d</strong> Ecology ......... ...... ................................ ................... 114 Systematic Position of the Genus .... ..................... .................................. 115 Systematic Arr<strong>an</strong>gement of the Species ...................................................... 115 U ses .................................................................................... 116 Taxonom y ............................................................................... 116 Generic Description .... .. ........................................................ , ... 116 Key to the Species <strong><strong>an</strong>d</strong> Subspecies ...... ................................................ 117 Treatments of the Species <strong><strong>an</strong>d</strong> Subspecies ............................................. 122 Unnamed Collections ............. . ............................................ 193 Names Excluded ................................................................. .... 194 A cknow ledgm ents ............................................................................ 194 Literature Cited .............. .. ...... ................................... ..................... 194 N um erical List of Taxa ......... .............................................................. 196 L ist of E xsiccatae ............................................................................. 197 Index of Vernacular Names . ................................................................... 205 Index of Scientific Names ................................... .............................. .. .. 206 Arboretum <strong><strong>an</strong>d</strong> Bot<strong>an</strong>ical Garden (ARBOHA), University of Bergen, N-5067 Store Milde, Norway. 2 Institute for Systematic Bot<strong>an</strong>y, State University of Utrecht, Heidelbergla<strong>an</strong> 2, 3508 TC Utrecht, The Netherl<strong><strong>an</strong>d</strong>s. 3 Institute for Systematic Bot<strong>an</strong>y, State University of Utrecht, Heidelbergla<strong>an</strong> 2, 3508 TC Utrecht, The Netherl<strong><strong>an</strong>d</strong>s. 4 Institute for Systematic Bot<strong>an</strong>y, State University of Utrecht, Heidelbergla<strong>an</strong> 2, 3508 TC Utrecht, The Netherl<strong><strong>an</strong>d</strong>s. 1

2 Flora Neotropica ABSTRACT Berg, C. C. (Arboretum <strong><strong>an</strong>d</strong> Bot<strong>an</strong>ical Garden [ARBOHA], University of Bergen, N-5067 Store Milde, Norway), R. W. A. P. Akkerm<strong>an</strong>s <strong><strong>an</strong>d</strong> E. C. H. v<strong>an</strong> Heusden (Institute for Systematic Bot<strong>an</strong>y, State University of Utrecht, Heidelbergla<strong>an</strong> 2, 3508 TC Utrecht, The Netherl<strong><strong>an</strong>d</strong>s). <strong>Cecropiaceae</strong>: <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>. Flora Neotropica 51: 1-000. 1990. The present monograph comprises the revisions of two of the three Neotropical genera of the <strong>Cecropiaceae</strong>: <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>. For <strong>Coussapoa</strong> 46 species are recognized at present; 15 of them have been described as new during the present study. All or most species are hemi-epiphytes. They have entire leaves <strong><strong>an</strong>d</strong> are dioecious. The small simple flowers are borne in uni- to pluricapitate inflorescences. The staminate flowers have one stamen or two or three connate stamens. The fruits are small. The genus r<strong>an</strong>ges from southern Mexico to south-eastern Brazil <strong><strong>an</strong>d</strong> is absent in the West Indies. Most species are components of lowl<strong><strong>an</strong>d</strong> rain forest habitats. Few species are wide-spread, most species have relatively small or even very small (known) r<strong>an</strong>ges of distribution. The species are morphologically clearcut <strong><strong>an</strong>d</strong> most of them rather uniform. For several reasons, e.g., lack of staminate material, the species are (provisionally) arr<strong>an</strong>ged in alphabetical sequences. Little is known about the biology of the genus. For <strong>Pourouma</strong> 25 species are presently recognized. Four taxa have been described as new during the preparation of this revision. All species are terrestrial, usually small to medium-sized trees. They have entire to palmately lobed to parted leaves <strong><strong>an</strong>d</strong> are dioecious. The pistillate flowers are pedicellate <strong><strong>an</strong>d</strong> are arr<strong>an</strong>ged in repeatedly br<strong>an</strong>ched to subumbellate inflorescences. The small staminate flowers are pedicellate or sessile <strong><strong>an</strong>d</strong> occur + loosely glomerate or in globose heads. The stamens are free in all species but one. The fruits are relatively large. The genus r<strong>an</strong>ges from Central America to southeastern Brazil <strong><strong>an</strong>d</strong> is absent in the West Indies. Most of them occur in + disturbed <strong><strong>an</strong>d</strong>/or + open places in lowl<strong><strong>an</strong>d</strong> rain forest. Besides a number of clear-cut <strong><strong>an</strong>d</strong> slightly variable species the genus comprises a number of (very) variable <strong><strong>an</strong>d</strong> + complex species: P. bicolor, P. cucura, P. gui<strong>an</strong>ensis, P. melinonii, P. mollis, <strong><strong>an</strong>d</strong> P. tomentosa. The former three <strong><strong>an</strong>d</strong> the latter three species are often difficult to separate from each other (<strong><strong>an</strong>d</strong> some other species). These six taxa <strong><strong>an</strong>d</strong> some others show a considerable plasticity, especially <strong>with</strong> regard to the leaf shape, leaf dimension, <strong><strong>an</strong>d</strong> composition of the indument, <strong><strong>an</strong>d</strong> most prominently so in the Upper Amazon Basin. Little is known about the biology of the genus. P. cecropiifolia is in cultivation as a fruit tree. INTRODUCTION TO THE FAMILY The present monograph is comprised of re- visions of two Neotropical genera of the Ce- cropiaceae, <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>. To place these revisions in a proper setting, there is pro- vided first a history <strong><strong>an</strong>d</strong> circumscription of the family, along <strong>with</strong> a brief discussion of the char- acters that separate the family from the related Moraceae <strong><strong>an</strong>d</strong> Urticaceae, a description of the family <strong><strong>an</strong>d</strong> the type genus, Cecropia, <strong><strong>an</strong>d</strong> a key to the Neotropical genera. History <strong><strong>an</strong>d</strong> Circumscription of the Family The family was proposed by Berg (1978a) <strong><strong>an</strong>d</strong> accepted by Cronquist (1981). It comprises six genera: three (Cecropia, <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>) that are Neotropical, two (Mus<strong>an</strong>ga <strong><strong>an</strong>d</strong> Myri<strong>an</strong>thus) Afric<strong>an</strong>, <strong><strong>an</strong>d</strong> <strong>an</strong> Asi<strong>an</strong>-Australasi<strong>an</strong> genus (Poikilospermum). The majority of the members of the family constituted subfamily Conocephaloideae in Engler's Moraceae (Engler, 1889), but Poikilospermum <strong><strong>an</strong>d</strong> some of the species presently included in it were included in the Urticaceae. Chew Wee-lek (1963) proposed r<strong>an</strong>king the four microspermous genera (Cecropia, <strong>Coussapoa</strong>, Poikilospermum <strong><strong>an</strong>d</strong> Mus<strong>an</strong>ga) under the Urticaceae <strong><strong>an</strong>d</strong> leave the two macrospermous genera (Myri<strong>an</strong>thus <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>) in the Moraceae. Corer (1962), however, regarded all six genera as members of the Urticaceae. The creation of a new family appeared to be <strong>an</strong> appropriate resolution to <strong>an</strong> unsatisfactory situation. Several features suggest that the <strong>Cecropiaceae</strong>