I

."7"

.. ,.
A Field Guide
to the Families and Genera
of

Woody Plants of
Northwest South America
(Colombia, Ecuador, Peru)
with supplementary notes on herbaceous taxa

Alwyn H. Gentry

Illustrations by Rodolfo Vasquez

.'

'I
.-

CONTENTS
c

Copyright 1993 by Conservation International

All rights reserved. No part of this book may be reproduced in any form
or by any means without permission in writing from the publisher.

>

FOREWORD ......................................................................................... xi
,

j

ACKNOWLEDGEMENTS .............................................................. xiii
LIST OF FIGURES ............................................................................ xv

Inquiries to the publisher should be directed to the following address:
INTRODUCTION ................................................................................. 1

Department of Conservation Biology
Conservation International
1015 18th St. NW, Suite 1000
Washington, DC 20036 USA

Author's address:

Illustrator'S address:

Alwyn H. Gentry
Senior Curator
Missouri Botanical Garden
P.O. Box 299
St. Louis, MO 63166 USA

Rodolfo Vasquez
Proyecto Flora del Peru
Apartado 280
Iquitos - PERU

KEY OUTLINE ................................................................................... ,. 6
KEY TO FAMILIES
KEY I - Leaves Opposite or Whorled (and Compound)
lA. Leaves bipinnate to biternate (and opposite) ..............................
IB. Leaves simply pinnately compound (and opposite) ....................
IC. Leaves 3-foliolate .......................................................................
ID. Leaves palmately compound (and opposite) ...............................

II
II
13
13

KEY II - Leaves Alternate (and Compound)
lIA. Leaves bipinnate ....................................................................... 14
liB. Leaves simply pinnate (and alternate) ....................................... 14
lIC. Leaves 3-foliolate ...................................................................... 20
liD. Leaves palmately compound (and alternate) ............................. 21
KEY III • Leaves Simple and Opposite (or Whorled)
IlIA. Lianas ........................................... ·...... ·.... ·..... ··.····· .. ·.. ·.. ······· .. · 24
IlIB. Trees and shrubs ....................................................................... 29
KEY IV - Leaves Simple and Alternate
IVA. Trees ........................................................................................ 37
IVB. Lianas with alternate simple leaves ......................................... 60

Library of Congress Cataloging-in-Publication Data No. 92-74686
ISBN 1-881173-01-1

Printed and bound in the United States of America

10 9 8 7 6 5 4 3 2 1

SPECIAL HABIT LISTS AND FIGURES
1. Leafless achlorophyllous parasites and saprophytes ..................... 73
2. Leafless plants with chlorophyllous stems ................................... 73
3. Tendrillate Hanas .......................................................................... 75
4. Anomalous liana stem cross sections ............................................ 77
5. Some common whole-tree branching patterns .............................. 80
6 Ant domatia .................................................................................. 82
7: Stilt roots ...................................................................................... 82
8. Spines .......................................................................................... 84
9. Strongly fenestrated trunks ........................................................... 86

~·"'t<NS

AND GYMNOSPERMS

~ernsd (mostly

Cyalheaceae) .............................................................. 87
yea
aceae.....
..... ........ ....... ............................................................
..
8"';J
E h d
op e raceae ..................................................................................... 92
P n~laceae ......................................................................................... 92
o ocarpaeeae .................................................................................. 92
MONOCOTS

Agavaceae
Araceae
.. .......... ....... .............. ...... ..... ............................................ .. 94
B
I' .......................................................................................... 96
rome
..... .......... ........... ...... ..... ............................................ 105
Burn laeeae
.
Cann~~~::eeae ... ....... .......................................... ........................... 108
Com mer ...................................................................................... 11 f
lOaeeae.......................... ............... ........ .. .............................. . I 11
Cyclanthaceae
Cyperaceae
................. ,.......................................................... 115
Dioscoreace~~·· .. :·· .. ··········.·.··.· .. ·.... ·· .. ·... ' ....................................... 120
Eriocaulaee ............................................................................... 125
..
Oramineae ae ................................. .............................................
. 125

Haemodora.. ·:· .... ·· ...... ·...... ·· .. ·... ·........ ·· ........................................... 128
.
ecae ............................................... ........... .................... 146}
Indaeeae
.
.
..............................................................
Lll!aecae
......................... .. 146
......................................................... ..............................
.
Marantaceae
. 147
Musaeeae , .................................................... .............................. . 147
Orchidace~~"'" ............................ ...... ....... ........... ........................... J50
Palmae
.................................................................................... 152
...............................................................
.
Rapateaceae
...........................
. 180
Smilacaceae ,.: ......................................................... ,....................... 198
..
.. ...................................................... ......................... .. 200
treltlzJaceae
S
Triuridaceae .................................................................................. 200
Ve110Ziace~e ................................................................................... 201
Zingiberaceae,........................
........
.. ......................................................... 20 I
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . . . . . . . . . . . . , •••••• "

. . . . . . . . . . . . . 0>

202

DICOTS

Acanthaccae

Actinidiacea~··········· .. ····· .. ···· .. ·'················· ..................................... 205

Aizoaeeae

................................................................................. 212
AIZateacea~""'"'''''''''''''''''''''''''''''''''''''''''''''''''''''''''''''''''''' .......... 214
··· .. ·.......
.......................................... 215
A maranthaceae
Anacardiaceae ............................................................................... 215
,·······.· ...
............................................. 219
An nonaceae .
..
····
..
····,,·······
················
.. ............................................ 225
A pocynaceae ....
Aquifoliaceae ... :: .............. ,.............................. ,.............................. 238
Araliaceae
................... ,................................
249
~
h

•••••••••••••••••••••••••••

H ................... .

••••••• hu .... H

• • • _._

H

••••••

•••••••••••••• ,

1
1
1
1
1
1
1
vii
Contents
1
Aristolochiaceae ............................................................................. 252
1
Asclepiadaceae ............................................................................... 254
1
Balanophoraceae ............................................................................ 258
1
Basellaceae ..................................................................................... 260
1
Bataceae ......................................................................................... 26 I
1
Begoniaceae .................................................................................... 261
1
Berberidaceae ................................................................................. 263
1
Betulaceae ...................................................................................... 263
Bignoniaceae .................................................................................. 264
1
B'Ixaceae ......................................................................................... 282
1
Bombacaceae .................................................................................. 282
1
Boragmaceae
.
........................................................... ,................ '" '" 292
1
Brunelliaceae .................................................................................. 299
1
Burseraceae .................................................................................... 299
1
Buxaceae ........................................................................................ 304
1
Cactaceae ........................................................................................ 305
Calyceraceae ................................................................................... 312
1
Campanulaceae ............................................................................... 312
1
Capparidaceae ................................................................................ 315
1
Caprifoliaceae ................................................................................ 3 I 9
1
Caricaceae ..................................................................................... , 320
1
Caryocaraceae ................................................................................ 323
1
CaryophyUaceae ............................................................................. 323
1
CeJastraceae .................................................................................... 326
Chenopodiaceae .............................................................................. 328
1
Chloranthaceae .................................................................... ,.......... 330
1
Chrysobalanaceae ........................................................................... 330
1
Clethraceae ..................................................................................... 333
1
Cochlospermaceae .............................. ,........................................... 335
1
Columelliaceae ................................................ ,.............................. 335
1
Combretaceae ................................................................................. 335
Compositae .................................................................................... 339
1
Eupatorieae ............................................................................... 343
1
Vernonieae ................................................................................ 348
1
Astereae .................................................................................... 349
1
In u1eae ...................................................................................... 350
1
Heliantheae ............................................................................... 352
1
Tageteae .... ,............................................................................... 359
1
Liabeae ..................................................................................... 360
Senecioneae .............................................................................. 361
Anthemideae ............................................................................. 363
Mutisieae ...................................................................... .
Lactuceae .......................................... .
Connaraceae ." ....... .

viii

Contents

goriariaceae ................................................................................... 375
omaceae ................................
Crassulacea
..
.. ......................................................"75
'
C 'f
e ................................................................................. 377
rUCI erae ............................... .
Cucurbitaceae
..
. .................................................... . 377

. ............................ .. ............................................. .. 377
C
.
..... ............ ..... ................... ........................................ .. 385
Cunomaceae
'11
......... ........ ........ ................................................. "87
D'yn1 aceae
al ...........
h
.J
la ypet ant aceae ...................... .
387

Dicha etalaceae.
' .............................................. .
DiPtefoca aceae' .......... ·.... ·.... ··· ...... ···· ........ ·.. ·· .... ·.. ··· .. ·.. ··· .. ·.. ·.. ··· 389
Droserace:
........................................................................... 393
....................................................
Ebenace
............................... . 393
Elaeocara:~~~ ................................................................................. 393
.
P
e .................................................... .........................
.
Encacea
. 394
e
........................................................
ErythroxyJac
................................ -3°7
"
eae .............................................. ............................. .. 403
Euphorbiaceae
.................................................. ............ ....... ........... 40"-'
Fagaceae
......................................................... ............................... . 424
Flacourtiaceae
Gentianaceae ................................................................................ 426
.
Geramaceae
. ....................................................... .. ....................... .. 434
.
Gesnenaceae .................................................... ... .......................... .. 437
Goodeniaceae.................................. ............................................... . 438
.
................................................. .............................. .. 445
G ut!lferae
. ........................................................
445
Haloragldaceae ..........
.. ........................... ..
............................... ...................................... . 454
Hamamelidaceae
.
.................................................
Hernandlaceae
......................... .. 456
................................. ,............................................. . 456
Hippocastanaceae
.
............................................ .............................
.
Hlppocrateaceae
. 459
.
.
...............................................
.
Humlnaceae
. . ........................... 459
.
HydrOPhylla~~·~~···· .. ·...... ··············· .. ··············· ..... ,............... ,.......... 466
.
......................................
. 470
I caClnaceae .... .................................................................................
. ·........................... 470
Juglandaceae
.......
.
.............................................
Juhanaceae
' .......................... ..... ...... 47 6
Krameriace~~·· .. · ............................................................................. 476
Labiatae
................................................................................. 478
Lacistematac~·~~· .. ·· ...... ·........ ·...... ·...... ·· .... ·.......... ······ .. · .................. 478
..................................................... ....................... . 484
Lauraceae
. .............................. ,...............................
48
.......................... 4
Le Cythldaceae
.
........... \............ ,............. .
............................................ 497
L egummosae
. :;.: ........ ~ ..................... ,.............. .
................................ 503
C aesalpmloldeae
.......................................
"Imosoldeae
............................. .. 504
M ..
.
...............................
.
............................................ 519
PaplhonOldeae
- .... ·.. ·.............. ·.................. ·.. ·............................. 527
......................................................... ,....... 554
.... ,................ 557
~

Loasaceae .......................................................................... " ........... 558
Loganiaceae ...................................... " ............................................ 560
Loranthaceae .................................................................................. 564
Lythraceae ...................................................................................... 569
Magnoliaceae ........ ,........................................................... ,............ 572
Malesherbiaceae ............................................................................. 574
Malpighiaceae ...................................................................... ,......... 574
Marcgraviaceae ..... ,............................................... ' ........... ,.......... ,. 592
Martyniaceae ..................................... ,.......... ,................................. 595
Melastomataceae ............................................................................ 595
e Iaceae ....................................................................................... 613
Ml'
Menispermaceae ............................................................................. 617
M ontmJaceae
..
.................................................................................. 624
Moraceae ..................................................................................... ,.. 626
yncaceae ., ................................................................................... 637
M'
Mynsttcaceae ................................................................................. 638
Myrsinaceae .............................. ,.................................................... 642
Myrtaceae ....................................................................................... 646
Nolanaceae ,,, ................... ,.............................................................. 657
Nyctaginaceae .................... ,.................................... ,.................... ,. 657
Ochnaceae ........................... ,.......................................................... 662
Olacaceae ....................................................................................... 665
Oleaceae ............................................................................ ,............ 670
Onngraceae ................................................................... ,................. 670
Opiliaceae ...................................................................................... 673
Oxalidaceae .................................................. ,................................. 673
Papaveraceae ............................................... ,.................................. 675
Passifloraceae .................................................................... ,......... ,.. 675
Phytolaccaceae ................................................................................ 679
Pi peraceae ...................................................................................... 683
Plantaginaceae ............................................................................. '" 686
Plumbaginaceae ................................ ,................................ ,............ 688
Polemoniaceae ............................................................................. ,.. 688
Polygalaceae ................................................................................... 689
Portulacaceae .......... ,.......................... " ......... ,................................. 698
Proteaceae ...................................................................................... 702
PyroJaceae .............................. ,................... ,................................... 703
Quilnaceae .................................................... ,.................... ,....... ,.... 704
Rafflesiaceae .................................................................................. 704
Ranunculaceae ................................................................................ 706
Rhamnaceae .................................................................................... 707
Rhizophoraceae .............................................................................. 711
Rosaceae .................... ,.................................................................... 714
Rubiaceae ....................................................................................... 718

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x

FOREWORD

Contents
<

Sabiaceae ....................................................................................... 758
Salicaceae ....................................................................................... 760
Santalaceae ..................................................................................... 760
Sapindaceae .................................................................................... 762
Sapotaceae ...................................................................................... 771
Saxifragaceae ................................................................................. 778
Scrophulariaceae ............................................................................ 782
Simaroubaceae ............................................................................... 783
Solanaceae ...................................................................................... 786
Staphyleaceae ................................................................................. 799
Styracaceae ..................................................................................... 807
Symplocaceae ................................................................................ 807
Theophrastaceae ............................................................................. 812
Th ymelaeaceae ............................................................................... 813
Tiliaceae ......................................................................................... 814
Trigoniaceae ................................................................................... 821
Tropaeolaceae ................................................................................ 823
Turneraceae .................................................................................... 823
Ulmaceae ....................................................................................... 824
Umbelliferae .................................................................................. 826
Urticaceae ...................................................................................... 827
Valerianaceae ................................................................................. 831
Verbenaceae ................................................................................... 833
840
Violaceae ............................................ .
Vitaceae .................................. ....................................................... 845
Vochysiaceae ......... ...... ....... .................. .........................................
.
846
Winteraceae ................................................................................... 848
........... H

............................ .

INDEX: COMMON NAMES ......................................................... 852
INDEX: SCIENTIFIC NAMES ...................................................... 862

Abbreviations used in this book:
C.Am.
incl.

N.Am.
n. temperate
S.Am.
s. temperate
sp. (spp.)
subspp.

Central America
includes
North America
north temperate
South America
south temperate
species
subspecies

Biologists and conservationists who work in tropical countries face
formidable scientific obstacles. Just knowing the biota is a Herculean task.
While a scientist working in a temperate country might easily learn to
identify many of the woody plant species in a single year, a tropical biologist
could easily spend a lifetime in a single small country without mastering the
flora. Tiny Ecuador, for example, has an estimated flora of 13,000 species,
more than all of Europe combined. Neighboring Colombia supports an
estimated 50,000 plants species, virtually a fifth of the entire global flora.
The problem of identifying this floristic abundance is compounded by the
dearth of taxonomic identification materials available to tropical field
biologists. AI Gentry makes a vital contribution to tropical biology and
conservation with this field guide. In tackling northwestern South America,
a region with the most diverse flora on earth, he has created an exhaustive
reference work tailored to meet the needs of field researchers and conservationists alike.
Gentry's near legendary expertise makes him uniquely qualified to
write this ambitious work. For the past quarter of a century, Gentry has
combed the Neotropics collecting and studying plants. His intense devotion
to field work, great stamina, and willingness to botanize under the most
difficult conditions, have enabled him to amass larger collections of plant
specimens from more countries than any other living botanist. Moreover, he
has helped build the plant collections of dozens of national museums and
research institutions around the world. His writings - a body of some 200
scientific papers - are no less impressive a contribution to scientific study.
Gentry has incorporated much of his pragmatic field abilities into this
book and gives us a sorely needed new approach to the identification of
tropical plants. Traditionally, taxonomic determinations have been made
using flower and fruit characteristics. This is a source of considerable
frustration in the field where it is often impossible to find a flowering or
fruiting individual of the most common trees. Instead, Gentry uses sterile
vegetative characters such as leaves, bark, and odor to lead us through the
maze of plant families and genera.
At Conservation International we work at developing the local capacity
to do conservation in the field. The work is urgent. Gentry's encyclopedic
knowledge of the neotropical flora has made him a vital member of
Conservation International's Rapid Assessment Program. He is one of only
a handful of people who can be dropped in the most remote, unexplored
areas of the Neotropics and be able to quickly assess the conservation
significance of the area in terms of species richness and endemism.
It is with great pleasure then, that we publish this innovative, authoritative and, above all, highly useful, volume. Gentry's contribution is the first
in a series of field guides to support conservation biology in the tropics.
AdrianB. Forsvth. Ph.D

1
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ACKNOWLEDGEMENTS
1

1
1
1
Publication of this book has been made possible by the support of the
Pew Scholars Program in Conservation and the Environment I also thank
1
Adrian Forsyth <Uld Conservation International for serving as publisher and
1
distributor for it. Special thanks go to Jane Koplow of Jane Koplow Designs
1
who look a rough manuscript replete with inconsistencies and eased it
1
towards camera· ready copy complete with indices, a process far more demanding in time and editorial skill than either of us envisioned originally.
1
My greatest thanks are reserved for Rodolfo Vasquez, friend and col·
1
league as well as artist, who prepared the illustrations. His unique combi1
nation of skill as both botanist and illustrator has allowed him to capture
1
salient taxonomic characters, often exquisitely. In addition to providing the
illustrations, he is the source of many of the Peruvian vernacular names.
1
Moreover, some of the suggested field recognition characters indicated here
1
are ideas that stem from his observations or that we have worked out together
1
over the course of years of shared field work.
Many colleagues have found time to review the familial treatments of
1
their taxonomic specialties, although not all of them agree with all of my
1
taxonomic decisions. Taxonomists who have reviewed familial treatments
1
include G. Schatz (MO) and P. Mans (U), Annonaceae, T. Croat (MO),
1
Araceae, D. Stevens (MO), Asclepiadaceae, J. Miller (MO), Boraginaceae,
D. Daly (NY), Burseraeeae, D. Hum (K), Caetaceae, R. Faden (US),
1
Commelinaceae, M. Dillon (F), Compositae, E. Forero (NY), Connaraceae,
1
B. Hammel (MO), Cyclanthaceae and Guttiferae, M. Hurt (F), Euphorbia1
ceac, R. Moran (MO), fems, L. Skog (US), Gesneriaceae, G. Davidse (MO),
1
Gramineae, P. Goldblatt (MO), lridaceae, R. Harley (K), Labiatae, H. van
der Wert' (MO), Lauraceae, S. Mori (NY), Lecythidaceae, G. Lewis (K),
1
Leguminosae, 1. Kuijt (LEA), Loranthaceae, W. Anderson (MICH),
1
Malpighiaceae, P. Fryxell (TABS), Malvaceae, S. Renner (AAU) and
1
J. Wurdack (US), Melastomataceae, C. Berg (ARBOHA). Moraceae. J.
1
Pipoly (MO), Myrsinaceae, B. Holst (MO), Myrtaceac, C. Dodson and G.
Carnevali (MO), Orchidaceae, A. Henderson (NY), and W. Hahn (WIS),
1
Palmae, C. Taylor (MO), Rubiaceae, R. Gereau (MO), Rutaceae and
1
Sapindaceae, W. Thomas (NY), Simaroubaceae, M. Nee (NY), Solanaceae,
1
A. Weitzman (US), Theaceae.
1
I also th<Ulk M. Mathias and L. Holdridge for early stimulation of my
interest in identifying tropical plants by vegetative characters; H. Cuadros,
1
C. Dodson, C. Diaz, C. Qrandes, R. Foster, P. Nunez, R. Ort(z, O. Phillips,
1
C. Reynel, A. Repizzo, and J. Zarucchi for contributions of common names
1
or other observations; and graduate students at the University of Turku, San
Marcos University, Washington University, St. Louis, and the University of
Missouri-St. Louis, for helping to test the familial keys.
Finally. thanks are due to Richard C. Schaer!', "~-' •
for generously making aV"nA""~

LIST OF FIGURES
c

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Figure
I

2
3
4
5
6
7
8

9
10
11

12
13
14
15
16
17
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19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40

Parasites and saprophytes
Liana tendrils
Liana cross sections
Tree habits
Domatia
Podocarpaceae and Ephedraceae
Cycadaceae and Gnetaceae
Agavaceae and Amaryllidaceae
Araceae (floating and epiphytic)
Araceae (miscellaneous epiphytic and terrestrial)
Araceae (terrestrial and hemiaquatic)
Bromeliaceae (spiny leaf margins)
Bromeliaceae (nonspiny leaf margins) and Cannaceae
Commelinaceae
Cyclanthaceae (terrestrial)
Cyclanthaceae (hemiepiphytic)
Cyperaceae (A-C)
Cyperaceae (E-U)
Dioscoreaceae
Eriocaulaceae and Gramineae spikelet details
Gramineae (A-Chusquea plus Dissanthelium and Guadua)
Gramineae (Bromus and Chloris-H)
Gramineae (1-0)
Gramineae (P-T)
Haemodoreaceae, Iridaceae, and Liliaceae
Marantaceae
Musaceae
Orchidaceae floral details
Orchidaceae
Palmae (palmate)
Palmae (pinnate taxa with distinctive fruits)
Palmae (spiny - A)
Palmae (spiny - B)
Palmae (Attalea relatives)
Palmae (miscellaneous large pinnate-leaved)
Palmae (stilt roots: Iriarteae)
Palmae (small: Geonoma and relatives)
Palmae (miscellaneous pinnate-leaved)
Rapateaceae, Smilacaceae, Strelitziaceae, and Velloziaceae
Zingiberaceae

Page

74
76
78-79
81
83
90
91
95
99
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103
107
109
113
117
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132
133
145
149
151
154
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181

183
185
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188
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203

xvi

List of Figures

Figure

42
43
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45
46
47
48
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50
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54
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56
57
58
59
60
61
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71
72

73

Acanthaceae (herbs and climbers)
Acanthaceae (herbs - A)
Acanthaceae (herbs - B)
Actinidiaceae, Aizoaceae, Alzateaceae, and Amaranthaceae
(trees and opposite-leaved vines)
Amaranthaceae (herbs and alternate-leaved vine)
Anacardiaceae - A
Anacardiaceae - B
Annonaceae (dehiscent monocarps plus Raimondia)
Annonaceae (syncarpous or appressed monocarps
[see also Raimondia))
Annonaceae (few-carpelled fruits with subsessile monocarps)
Annonaceae (multicarpelled fruits with stipitate monocarps;
broad petals)
Annonaceae (multicarpelled fruits with stipitate monocarps;
narrow or very small petals)
Apocynaceae (trees with opposite leaves and large indehisccnt
fruits)
Apocynaceae (trees with opposite leaves and berries
or dehiscent fruits)
Apocynaceae (trees and shrubs with alternate leaves)
Apocynaceae (Hanas with small flowers or leaves with
midrib glands)
Apocynaceae (lianas with large flowers and nonglandular
leaves)
AquifoHaceae and Araliaceae
Aristolochiaceae and Asclepiadaceae
Asclepiadaceae
Balanophoraceae and Basellaceae
Bataceae, Begoniaceae, Berberidaceae and Betulaceae
Bignoniaceae (trees with indehiscent fruits andlor Simple
leaves)
Bignoniaceae (trees [and a herb) with palmately compound
leaves and dehiscent fruits)
B,ignoniaceae (trees with pinnate leaves, herbaceous vines with
bl~ompound leaves and a simple-leaved hemiepiphyte)
Blgnoniaceae (lianas: A-Ce)
,
Bignoniaceae (lianas: CI-H)
Bignoniaceae (Hanas: L-Man)
Bignoniaceae (lianas: Mar-Pa)
Bignoniaceae (Hanas: Pe-R)
Bignoniaceae (lianas: S-X)
Bixaceae and Bombacac~aP. ('"-,,,\,,,,, 1Dn .. n_'

List of Figures
Page

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237
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251
253
257
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266

Figure

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276
279
280

106
107
108
109
110
111

xvii
Page

Bombacaceae (compound leaves; many stamens)
Bombacaceae (compound leaves; few stamens;
kapok seeds)
Bombacaceae (winged seeds; simple or compound leaves)
Bombacaceae (simple leaves; indehiscent fruits)
Boraginaceae (trees and shrubs)
Boraginaceae (lianas and herbs)
Brunelliaceae and Burseraceae
Burseraceae
Buxaceae and Cactaceae
Cactaceae (erect)
Cactaceae (epiphytic andlor scandent)
Calyceraceae and Campanulaceae
Capparidaceae
Capparidaceae (Tovaria) and Caprifoliaceae
Caricaceae
Caryocaraceae
Caryophyllaceae
Celastraceae
Chenopodiaceae, Chloranthaceae, and Chrysobalanaceae
(shrub: Chrysobalanus)
Chrysobalanaceae (trees)
Clethraceae, Cochlospermaceae, and Columelliaceae
Combretaceae
Compositae (stigmas and head details)
Compositae - A
Compositae - B
Connaraceae
Convolvulaceae (herbs and vines)
Convol vulaceae (Hanas)
,
Coriariaceae, Cornaceae, Crassulaceae, and Cruclferae
Cucurbitaceae (tendrils bifid near apex; sm~ll flower~)
Cucurbitaceae (Hanas and a single-seeded vme; tendnls
branching below middle or si~1Ple),
'
Cucurbitaceae (many-seeded ViOes; tendnls branchmg
below middle or simple)
Cunoniaceae and Cyrillaceae
Dialypetalanthaceae and Dichapetalaceae
Dilleniaceae
Dipterocarpaceae, Droseraceae, and Ebenaceae
Elaeocarpaceae
, '
Ericaceae (shrubs and some distinctive heffileplphytes)
__

_

~..J t... ... ___ :"' .... : ... h"tj3"\

285
287
289
291
294
295
298
301
303
308
309
313
317
318
321
322
324
325
329
331
334
337
341
344
345
368
371
373
376
379
381
382
386
388
391
392
396
398
399

xviii

List of Figures

Figure

113
114
115
116
117
118
119
120
121

122
123
124
125
126
127
128
129
130
131
132

133
134
135
136
137
138

139
140
141
142

Erythroxylaceae
Euphorbiaceae (herbs and succulent shrubs)
Euphorbiaceae (vines and Hanas)
Euphorbiaceae (leaves 3-foliolate or deeply palmately
lobed)
Euphorbiaceae (trees: leaves with peitate scales or stellate
trichomes and lacking petiole glands; latex red or absent)
Euphorbiaceae (trees with 3-veined leaves; mostly with
glands at base of blade)
Euphorbiaceae (Phyllanthoideae: trees with no latex nor
petiole glands)
Euphorbiaceae (trees: pinnate-veined Acalyphoideae; latex
red, watery or absent; mostly with petiole glands)
Euphorbiaceae (trees: pinnate-veined Euphorbioideae;
latex white or watery-white, often poisonous; leaf margins
usually serrate)
Fagaceae and Flacourtiaceae (odd-ball genera: leaves opposite or with reduced secondary veins)
Flacourtiaceae (3-veined leaves)
Flacourtiaceae (branched inflorescences or unusually large
flowers or fruits)
Flacourtiaceae (fasciculate inflorescences; small flowers)
Gentianaceae
Gentianaceae and Geraniaceae
Gesneriaceae (scandent or subwoody)
Gesneriaceae (herbs) and Goodeniaceae
Guttiferae (hemiepiphytic)
G,uttiferae (trees with fleshy capsular fruits; usually with
Stilt roots)
Guttiferae (trees with indehiscent fruit, yellow latex, and
well-developed intersecondary and tertiary veins parallel
to secondaries)
o.uttiferae (shrubs ?r trees with tertiary venation perpendlCular to secondanes; latex scanty or orange)
Haloragidaceae and Hamamelidaceae
Hernandiaceae and Hippocastanaceae
H~ppocrateaceae (dehiscent fruit)
Hlppocrateaceae (indehiscent fruit)
Humiriaceae
Hydrophyllaceae
Icacinaceae (trees: small fruits)
Icacinaceae (lianas)
Icacinaceae (trees: large fruits)

List of Figures
Page

402
406
408
410
412
414

148
149

416
418

421
425
429
431
432
435
436
441
443
447

150
lSI

152
153
154
155
156
157
158
159

450

160

451

161
162

453
455
458
462
464
467
469
472
473

Page

Figure

143
144
145
146
147

163
164
165
166
167
168
169

xix

Julianaceae, Juglandaceae, Krameriaceae
Lacistemataceae and Labiatae
Lauraceae (Cassytha and fruits without cupules)
Lauraceae (opposite leaves)
Lauraceae (4-valved anthers, nonclustered leaves; fruiting
cupule cup-shaped)
Lauraceae (clustered leaves, mostly 2-valved anthers;
fruiting cupule often short)
Lauraceae (2-valved anthers, nonclustered leaves; fruiting
cupule with distinctive rim or raised lenticels or poorly
developed or completely enclosing fruit)
Lecythidaceae (actinomorphic flowers; indehiscent fruits)
Lecythidaceae (fruits either elongate with winged seeds or ,
indehiscent with minute seeds in pulp; flowers zygomorphlc
to subactinomorphic)
Lecythidaceae (fruits broad and woody, without pulp;
flowers zygomorphic)
Leguminosae/Caesalpinioideae (bipi~nate)
,,
Leguminosae/Caesalpinioideae (2-foh,olate or ~Ifld)
LeguminosaelCaesalpinioideae (paripmnate, wmddispersed)
, '
'
Leguminosae/Caesalpinioideae (parlpmnate, non-wmd, ,
'
dispersed [AJ)
Leguminosae/Caesalpinioideae (panpmnate, non-wmddispersed [BJ)
,
Leguminosae/Caesaipinioideae (odd-pmnate) ,
Leguminosae/Mimosoideae (lianas and large mOist forest
trees)
, ,
'th
Leguminosae/Mimosoideae (Inga and blpmnate taxa WI
tiny leaflets)
Leguminosae/Mimosoideae (shrubs and tr~es of d:y areas)
LeguminosaelPapilionoideae (trees; opposite or Simple or
"
3-foliolate leaves)
Leguminosae/Papilionoideae (~ees: wind-dispersed with
one-winged tadpole-shaped frUits)
,
Leguminosae/Papilionoideae (trees: wind-dispersed frolls
with wings surrounding seed)
"
,
IP
Tonoideae
(trees'
fleshy
indehlscent
frUits)
Legummosae apl I
'
,
' )
,
IP
T
noideae
(trees'
fleshy
dehlscent
frUits
Legununosae apllO
'"
[AJ)
, aeIPapilionoideae (trees' dry dehlscent frUits
Legummos
"
f ' [BJ)
,
IP apll
Tonoideae (trees'' dry, dehlscent rUlts
Legummosae
LeguminosaelPapilionoideae (w,oody banas)
__

•• ,

_~...J

___

I •• ~ ....... ,,\

477
481
487
488
491
493

495
498
500
502
506
508
511
514
515
518
521
522
524
530
532
534
537
538
540
542
546
548

Primu1aceae.A Loganiaceae .B Loranthaceae Lythraceae Magnoliaceae. mostly ramiflorous) Meliaceae (fruits capsules with arillate seeds) M. Malesherbiaceae. Oxalidaceae. mostly w' th terminal inflorescences and/or ant dOmatia) 1 Melastomataceae (woody with berry fruits. petiole wiry-pulvinate at apex... 223 Piperaceae 687 1 224 Plantaginaceae. Ficeae) and Myricaceae 1 640 205 Myristicaceae (leaves and fruits) 1 641 206 Myristicaceae (inflorescences and fruits) 1 644 Myrsinaceae 207 1 208 Myrtaceae (Myrcia relatives with paniculate int10rescences 1 and leafy folded cotyledons plus Myrciaria with glomeru649 1 late flowers and thick fleshy cotyledons) (Eugenia and relatives with racemose (or reduced) Myrtaceae 209 1 inflorescence. 4-parted flowers and thick fleshy cotyledons) 651 1 210 Myrtaceae (Psidium and relatives with curved or spiral 1 654 embryo and reduced cotyledons) 1 Myrtaceae (high-altitude genera with small leaves and/or 211 few stanlens. Melastomataceae (woody with berry fruits.. petiole wiry-pulvinate at apex. indehiscent or seeds wmged or leaflets serrate) Menispermaceae (vines or soft Iianas. dorsal connective spurs) Melastomataceae (woody with capsular fruits' ventral connective spurs or inferior ovary) . except Myrciamhes with thick. and Proteaceae 229 701 1 Proteaceae 230 705 Quiinaceae and Ranunculaceae 1 231 708 Rhamnaceae (Hanas) 232 1 709 233 234 235 236 237 Rhamnaceae (trees and shrubs) Rhizophoraceae Rosaceae (trees and shrubs) Rosaceae (herbs and vines) Rubiaceae <:1.. and PolemoOlaceae 691 Polygalaceae (Hanas) 225 1 692 226 Polygalaceae (herbs and trees) 1 695 227 Polygonaceae (trees) 696 1 228 Polygonaceae (herbs or scan dent) 699 1 Portulacaceae..~ . seeds rummate) Monimiaceae and Mo (D ' [pri k1 1 ' raceae orstema [herb] and Poulsenia c es On tWIgS and stipule) Moraceae (Cecropieae) Page 552 555 556 559 562 563 566 571 573 577 579 581 582 585 587 589 590 594 599 600 603 606 609 611 614 615 619 620 623 625 1 1 1 1 1 1 xxi List of Figures 1 1 Page Figure 1 1 630 202 Moraceae (Olmedieae) 1 634 Moraceae (Moreae) 203 635 204 Moraceae (Brosimeae.eJiaceae (fruits large and/or woody. seeds nonruminate) Men~permacea~ (WOody lianas.1""'- -712 ~ . and Papaveraceae 218 1 677 Passifloraceae (exceptPassijtora) 219 1 678 220 Passit10raceae (PassiJlora) 1 681 Phytolaccaceae (herbs and subshrubs) 221 682 1 222 Phytolaccaceae (trees and lianas) 685 1 . Plumbaginaceae. fleshy cotyledons) 1 659 212 Nyctaginaceae (alternate leaves or weedy herbs) 1 660 Nyctaginaceae (opposite-leaved trees and lianas) 213 1 663 214 Ochnaceae 1 666 Olacaceae (calyces fused to fruit or not enlarged) 215 667 1 OIacaceae (calyces enlarged in fruit) and Oleaceae 216 671 1 Onagraceae 217 674 OpiHaceae. seeds nonruminate) Menispermaceae (woody lianas. embryo curved or spiral with reduced cotyle1 655 dons.. Malpighiaceae with 2-3 cocci Malpighiaceae (trees and shrubs with indehiscent fruits) Malpighiaceae (Hanas: fruits with dorsal win as) Malpighiaceae (Hanas: fruit wings from Caly: or both lateral and dorsal) Ml1lpighiaceae (lianas: fruits with lateral Wings) Malvaceae (trees and large shrubs) Millvaceae (herbs and shrubs: capsular fruits or dense bracteate inflorescence) Malvaceae (herbs: cocci [A-P)) Malvaceae (herbs: cocci [S-W)) Marcgraviaceae and Martyniaceae Melastomataceae (Mouriri and woody hemiepiphytes) Melastomataceae (herbs with capsular fruits) Mel~stomataceae (woody with capsular fruits. ~ 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 WI Leguminosae/Papilionoideae (herbs) Lentibulariaceae and LepidobolIyaceae Llnaceae Loasaceae Loganiaceae . petiole flexed at base.- .

nonflattened capsular fruits: calyx lobes not expanded) Rubiaceae (trees with oblong. 3-foliolate. spiny or clustered leaves) Sapotace~e (~etiole base usually not enlarged. and Turneraceae Ulmaceae Urticaceae (herbs) Urticaceae (trees. narrow capsular fruits: calyx lobes not expanded) Rubiaceae (trees with short. ' Tiliaceae (trees: follicular or mammal-dispersed and tndehiscent or capsular) Trigoniaceae. usually flattened capsular fruits. and lianas) Valerianaceae and Verbenaceae (lianas and compoundleaved trees) Verbenaceae (trees: simple leaves) Verbenaceae (shrubs) Verbenaceae (herbs) Violaceae (trees) Violaceae (lianas and herbs) and Vitaceae Vochysiaceae Winteraceae and Zygophyllaceae 796 798 801 802 804 806 809 811 816 817 819 822 825 828 829 832 834 836 839 843 844 847 849 . most1y +/. mostly capsular) .Figure 239 240 241 242 243 244 245 246 247 248 249 250 251 252 253 254 255 256 257 258 259 260 261 262 263 264 265 266 267 268 List of Figures List of Figures xxii Rubiaceae (trees with narrow capsular fruits: epiphytic and hawkmoth-pollinated or terrestrial and with expanded calyx lobes) Rubiaceae (trees with long. dehiscent fruit) Sapotaceae (petiole base usually not enlarged. shrubs. indehiscent fruit) Sapindaceae (trees: simple-pinnate. and small-leaved) Page 725 729 730 732 277 278 735 737 740 743 279 280 281 282 283 284 745 748 751 752 285 286 287 288 289 290 291 756 757 759 761 764 766 768 770 773 774 777 780 781 784 788 790 792 Page Figure 269 270 271 272 273 274 275 276 l xxiii Solanaceae (herbs) Staphyleaceae Sterculiaceae (trees with follicles) Sterculiaceae (trees with indehiscent fruits) Sterculiaceae (shrubs and Hanas) Styracaceae and Symplocaceae Theaceae (small to medium flowers) Theaceae (large solitary flowers).sptnescent . calyx lobes not expanded) Rubiaceae (trees with large indehiscent fleshy fruits and hawkmoth flowers) Rubiaceae (trees with large indehiscent fleshy fruits and small to medium flowers) Rubiaceae (shrubs with 1-2-seeded berries) Rubiaceae (shrubs with several-seeded berries) Rubiaceae (miscellaneous distinctive taxa of shrubs and small trees with indehiscent fleshy fruits) Rubiaceae (herbs) Rutaceae (leaves pinnate or bipinnate) Rutaceae (leaves 3-foliolate to palmately compound) Rutaceae (leaves simple: pachycaul treelets or fleshy indehiscent fruit) Rutaceae (leaves simple or unifoliolate: follicular fruit) Sabiaceae Salicaceae and Santalaceae Sapindaceae (lianas) Sapindaceae (trees: simple. and bipinnate leaves) Sapindaceae (trees: simple-pinnate. Tropaeolaceae. Theophrastaceae and Thymelaeaceae Tiliaceae (shrubs and trees with samaroid fruits) Tiliaceae (trees: nonsamaroid. 2-ranked leaves with ftnely parallel tertialJ' venation) Sapotaceae (petiole base enlarged) Saxifragaceae Scrophulariaceae Simaroubaceae Solanaceae (epiphytic shrubs or hemiepiphytic climbers) Solanaceae (erect trees and shrubs with rotate flowers) Solanaceae (erect trees and shrubs with tubular flowers) Solanaceae (dry-area shrubs'.(or water-)dispersed. wind.

Although these are often somewhat subjective decisions. where such an estimate is available. epiphytic. and Peru. I have been guided by the principal that. I have generally adopted traditional concepts of families and genera. for most families each genus is listed along with a few of its most outstanding characteristics and hints for its differentiation from related taxa. for some wideranging genera only a worldwide estimate of species numbers is available 00 inrlir. For each genus the number of neotropical species is indicated parenthetically. In this connection it is useful for the non taxonomic reader io remember that generic concepts have been more or less at the whim of particular taxonomists. especially if that monographer has been misled in formulating his taxonomic decisions by concern about avoiding paraphyletic groups. Also included oarenthetically.otp. of all the woody plants of Colombia. the generic concept that seems to make the most sense to the taxonomic consumer is likely to be the best one. this means that the families under which specimens are filed in the Missouri Botanical Garden herbarium. the characteristics for recognizing that family are indicated along with an indication of how to recognize its constituent genera. rather than blindly following the most recent monographer. are adopted here. I have made a value judgement in rejecting all of the proposed substitutions of new familial names like Poaceae for Gramineae or Asteraceae for Compositae. even if most of their species are herbaceous. All families that contain woody.O. Ecuador. A few important cultivated or naturalized genera are included parenthetically. or scandent species in northwest South America are included. essentially the old Englerian concepts. with very few exceptions. At the family level. unless compellingly different morphologically. no matter what the current specialist in a particular group may happen to say. When applicable. For each family. I have parenthetically indicated segregate generic names that are here included within a more broadly circumscribed genus. on the other hand. In my opinion it should be the responsibility of the taxonomic consumer to evaluate the evidence and make his/her own decisions as to what generic delimitations most accurately reflect the real world. but only what I judge to be the most convenient technique for a majority of users. At the generic level. without indication of species . Philosophically my taxonomic premise is that the generic limits proposed by a particular monographer constitute a hypothesis. to be accepted or rejected like any other in science. segregate genera should only be recognized when demonstrably polyphyletic. In addition. usually even when only sterile material is available. I have made numerous value judgements as to what genera to recognize. Thus. This does not imply any value judgements as to the status of various groups sometimes recognized as families.INTRODUCTION c > This book aims to make possible the identification to genus.

Mo~e­ over. are 1 always available. kindl" nrnvitl". :0 In ~ener~l. tree vs. Young (eds. Leon and K. herbs are not considered in the introductory keys (although.have included at least synopses of the nonaquatic herbaceous amlhes WhlCh ~~CUf in our area. Vegetative characters. of course... are illustrated. Dillon has been kind enough to provide a tribal key 1 as well as supplementary notes on many of the woody genera of each 1 tribe. Thus. Moreover. Rather I have attempted to talk the user through the relevant character states and possible taxonomic outcome~ much as I would in a field course. that is also indicated. Lima. These include AIls1 mataceae. Most woody genera. M. and a few herbaceous ones.g. scandent. Although the huge family Compositae is mostly herbaceous. fam1lies Gramineae and Compositae. and Xyridaceae. and orientation) that they are of limited practical use. dlose are generally unavoidable in generic delimitation in this 1 family. Kahn. the plants one wants to identify are more often encountered stenle 1 than in fertile condition. myself included.. or epiphytic speclCS. ~o". Butomaceae.~ ~vl. It should also be noted that most of the Peruvian vernacular names are Quechua-derived while most of the Colombian and Ecuadorian names are based on Spanish. Taccaceae. ~or predOm1tl~ntly ~P1Phytic Orchidaceae. 1 placement. Dodson and a previously unpubverSlOn 0 that kev. ~~____ .. h~rbaceous members of a family generally share the features of the famliy s woody representatives that are treated in the keys). in tho~e relatively few cases where distinct names are applied to different speclCs of the same genus." Ho~e. at least to my tropically expcfl1 enced eye..~ this generalized aspect that facilitates full A~. Instead I have listed only those names that I have personally encountered to be used by reliable informants in the respective areas (supplemented by additional similar observations of R. Mayacaceae. Dodson for coastal Ecuador). Although the key is mostly based on technical floral char1 acters. No anempt has been made at a complete catalogue of vernacular names. Elatinaceae. Dr. and these are the only groups included in the key to families. 1U SPfaniSh by C. its taxonomy depends inordinantly on technical floral characters that most 1 nonspecialists. Indeed. mostly in northern Amazonia in Peru. For a treatment of 1 these taxa see B. This book stresses identifications based on leaves. in many ways it is probably eaSIl!r to IdentIty 1 woody tropical plants in sterile condition to family than it is to ident~fy 1 the fertile material to which many systematic botanists tend to restnct 1 themselves.~n in sterile condition. for the very large. Podostemaceae. mostly herbaceous. I suspect thnt ._ 1 1 1 1 1 3 1 Introductioll 1 1 inCluded here.. along with their geographic usage. mostly macroscopically obvious. generic identifications by the 1 nonspecialist will rarely be possible without flowers. Nymphaeaceae. I . In F. apparently have been subjected to much less of the klOd of 1 convergence-inducing selection on taxonomically useful characters than 1 have flowers or fruits. 1 1 Families that are exclusively or essentially aquatic or semiaquatic in our area are omitted entirely from this treatment. an excellent key to genera li~~e~e~n:l~:~lShed.1 I. Lemnaceae. I . On the other hand. with different-sized groups of taxa sharIng a useful set of characters considered together along with their individual defining characters. Najadaceae (and 1 various segregates). and P to Peru. Po1 tamogetonaceae. Ceratophyllaceae. Common names are indicated for many genera. Pontederiaceae. 1992).:. on the other hand. This is true both because of strong convergence by many 1 different families that share a common disperser or pollinator and because 1 the technical characters on which most plant families are defined are so 1 obscure and esoteric (typically involving a determination of ovule number. too.ond listing for many genera that are polymorphic with respect to characters (e. ORSTOM and Inst. ev. ~ .t . pl?S some of the most important herbaceous genera. 1 the leaves. have converged to the extent that they have lost many ~f the Obvious characters for identification. Treatment of the woody and scandent taxa is straight forward. mostly in the coastal lowlands in Colombia and Ecuador.) Plantas Acuaticas del 1 Peru.~ver. Juncaceae. Leon (Calalogo Anatado de las Fanerogamas Acuaticas 1 del Peru. 1 In some other vegetation types like deserts or Mediterranean-climate areas.. It will be noted that the key~ fam~ly (as well as those to genus in the larger families) are not the tradmonal dIchotomous ones of classical taxonomy. useful vegetative chnro"'~~· .. B... 1 Hydrocharitaceae. Callitrichaceae. 1 1 Most neotropical plants are surprisingly easy to identify t~ fami~y. Like the orchids. The illustrations reflect the largely Amazonian material available to the Iquitosbased artist. Vasquez for Amazonian Peru and C. The common plaint of the temperate botanist who encounters his first tropical forest is "The leaves all look just alike. No indigenous ~ames are indicated unless they have been taken up by the mestizo populations of the respective regions. I ha. For both orchids and Compositae. E to Ecuador..ver. I view this "lauraceous" look as a blessing in disguise. these are emphatically nol the uncritical compilations that are found in many monographs or lists of local names. the indicated vernacular names are strongly biased by my field experience. the 1 family is diverse and also includes trees and !ianas. Typhaceae.ve ~~cluded herbaceous genera only when they belong to a fal::11ly With a slglllficant complement of woody. For the 1 Compositae. Frances de Estudios Andinos. I have included a listing only of woo~y taxa. In addition. and at least ~n the 1 rain forest. Most vernacular names are specific only to g~nus. with C referring to Colombia. Iiana) that are used to subdivide various families. find abstruse and difficult to use.

.:~. but have meluded some of the more obvious bark and slash characters here. and habit characters. I am much less adept at such techOlques.tm~?. Thus. generally as a supplementary aid to identification. even a weak one. essentially everyone discovers that they really are able to h~Ck up ra~her SUbtle nuances of plant odors (the maln exception being " ?~to~. section~ . the same. Anyone who has ever observed II good "matero" effortlessly identifying trees with nothing more t~an a machete slash of the bark and a sniff of his nose can begin to apprecIate . wash your hands and look for a nondescript "l~afy" arom I ~rent part of the plant. there are a few families like Buphorbiaceae that are so vegetatively heterogeneous that field identification to family may only be possible with recognition of the constituent genera. coincidentally. the identification is positive.~~. There IS a common epiphyllolls leafy Variants of h as a strong amse odor that is easily confused with some of the produced b w at I here cal. For convenience. Wood anatomists can often identify II piece of wood to genus or family. Frequently a trunk slash must be allowed several minutes for latex to ooze out or to tum to its characteristic color. different twig from a d·~n amse odor. In others. In addition. Also the line between latex and sap is a very fine one and some saps.. In addition to those vegetative characters that m'e most generally useful in family recognition and emphasized in the keys. are here informally called latex.Introduction . unless it is very copious. theu sense of smell). and their relatives).. . an exudate. that can easily be recognized to order and genus but often can only be differentiated to family by using fertile characters or by recognizing the individual genera involved. Also.g~ most families have a few nasty exceptions that have to be le~ned. the red latex of Myristicaceae is in reality sap. that is white or colored may be referred to as latex.. Similarly.t? tn.S for tropical plant identification has complained at first that his/her th:t I~ below ~~erage and una?l~ to dett. nonaromatic leaves h~VlOg serrate or serrulate margins are found almost exclusively in Hlppocrat~aceae. and opposite compound leaves with tendrils are definitively Blgnomaceae. Opposite leaves with punctations strongly suggest Myrtaae . wood. the odor Whenever ~~~ee~~~~~~~ll cells.. Contrary to popular supposition. even at the familial level (e. are not significantly affected by preservatives or other drying techniques. remember that slash and leaf odors are no a ways. with . ~~".. e. liverwort that h or.e. respectively.ct the sometimes rather faint odors bit may be. turning whitish or blackish. that may be extremely useful taxonomically. these color characters. of the leaves of woody Ranalean families). the family is easy to recognize vegetatively but generic differentiation tends to require fertile material. In some families it is very easy to recognize all or most genera by vegetative characters. while the legitimate latex or resin of Burseraceae and Anacardiaceae may be faint and often tends to exude clear. In such cases I have been forced to emphasize the traditional taxonomic differentiations. c: A special word of encouragement on the Sense of smell may prove useful l~ the neophyte. ~~~~?~~~e~. a number of unusual morphological features that may be useful for recognizing particular taxa are noted in the lists and figures that preceed the familial treatment. some characteristics of dried specimens. only on drying..some of these a?ditional characters.g.. and Jianas with simple. a?d If t?e ~eaves ~ave a spic~ odor. while merely suggesting some of the vegetative trends that tend to characterize individual genera. the sweetish odor of most Meliaceae seems restricted to the trunk slash. Although this book is intended primarily as a field manual. of which Myrtaceae and Lauraceae are the most infamous examples. cntlcal fo: recognmng trOPiCal plants vegetatively. opposite compound leaves are enough to suggest B:gnon~aceae. primitive odor and strong bark is diagnostic for mo~t Annonaceae.g. I have also noted. those of the MalvaJes. and these latex traces may be apparent only in the trunk or only in the leaves.. It has .~~ 5 Latex characters can also be tricky. oppOsite.dwate tillS level of slmple characters for recognizing each of the famlhes Included in this book. 'T'''traits include parasitiC and saprophytic JlrOw'" r with chlorophyllous stem!: .mdlvldually. of practice.ates from the crushed leaves of many typical green-bean odor of mos:~~ll~!~r.?gIS:S whose long exposure to formaldehyde has apparently ~:n e . Usually the petiole and midvein are the best places to look for latex. Some species or individuals of some families or genera that are supposed to be characterized by latex have exceedingly minute amounts.g" the characteristic black or olive color of dried Olacaceae.been my experience that any neophyte tropical botanist can learn to re~o:Dlze the ~reat maj?rity of the plants he encounters to family by learnmg to recognlZe a particular combination of only two or three characters. Thus.lthe "primitive" or RanaJean odor (I. It is useful to break or crush both leaves and y t tWigs to check for odor.--" 'u leaf characters. But with a . At the opposite extreme are whole groups of families. the bright red exudate of a number of papilionate legumes. Virtually everyone whom I have instructed in field tech- ~~q~e.o amse. for example. f Finally beware the ad f ' . which Dlay bc taxonomically defInitive. based on fertile material. while an exudate that is clear may be generally considered as sap rather than latex. alt~ou.-. Loranthaceae. I h ve tned . e. many woody families and genera have taxonomically useful bark. There is also a faint nonaromatic plants and ~~s a~. especially the colors of dried leaves. Obviously such characters as latex or odor must be observed or recorded in the field.

Quiinaceae. AraJiaceae. Allophylus. Bignoniaceae. Acanthaceae. 3-foliate (page 13) B~gnoniaceae. Cochlospermaceae IIDb. Compositae.l~~~~~~:t pf:~~~~)'(CTerminal "bud" on. Brunelliaceae. Com positae. SIMPLE AND OPPOSITE lIlA. Rutaceae. Trees Palmae. Guttiferae. Spines Le . Rutaceae. Loranthaceae. aborted rachis tip (Sapindaceae. Amaranthaceae. Zygophyllaceae. Gesneriaceae. Palmae. Trigoniaceae. Rutaceae ID. Bipinnate to biternate (page 11) Bignoniaceae. Quiinaceae. Vochysiaceae. Sapindaceae. Palmately compound and alternate (page 21) IIDa. Verbenaceae. Palmae Cycadaceae IIB I. Hevea. Hlppocastanaceae. 7 Lepidobotryaccac. PUllctatiolls Myrtaceae. Sapindaceae. Valerianaceae lilAc. Compositae. Compositae. Rhizophoraccae IIIBb. Cunoniaceae. Leguminosae IIBf. Leguminosae m. Nyctaginaccae. 3(-7)-veilled leaves with parallel cross veills Melastomataceae . (Euphorbiaceae). Rutaceae. Palmately compound (page 13) Bignoniaceae. expanded petiole base (Polylepis). Staphyleaceae (Huertea). IlBi g~mo~lae. Juglandaceae. Simply pinnate (page 11) Bign?ni~ceae. Sapindaceae. Burseraceae IlBd. Sapindaceae. Leguminosae IIBc. Miscellaneous Hippocrateaceae. Verbenaceae II. Stipules (or stipule scars) Rubiaceae. Lythraceac. Simaroubaceae nco 3-foliolate and alternate (page 20) Ilea. Lianas (page 24) IIlAa. Odor of esselltial oils or turpelltille Rutaceae. Eveu-pilmate Le' . Rubiaceae. Rutaceae. Sweetish odor ill trullk Meliaceae IlBe. COMPOUND AND OPPOSITE IA. Staphyleaceae. Anacardiaceae. Leguminosae) IlBk. Valerianaceae IC.Key Outline KEY OUTLINE c I. rachis 1 onnaraceae. Cycadaceae IlBb. Melastomataceae. Anacardiaceae (Tapirira). Vitaceae. Araceae. Sabiaceae (OphiocOIyon). Dioscoreaceae. Guttiferae.h~. Trees and shrubs (page 29) IIIBa. Passifloraceae. Simaroubaceae). (Compositae) IliAd. Trees Anthodiscus. Araliaceae. Ranunculaceae. 3(-7)-veilled. Chloranthaceae. Saxifragaceae nIB. Ranunculaceae. Anacardiaceae. Euplassa). PUI/ctations Rutaceae. Ranunculaceae. Malpighiaceae. MeJastomataceae. Latex ' ~~~~~daceae. Isce aueous useful features ~~~:~e~:~. Leguminosae. Coriariaceae. AscJepiadaceae. (Menispermaceae) no. Meliaceae lIB. Connaraceae. (Rhizophoraceae) IIlBd. Mehaceae. Combretaceae. Sapindaceae. Rutaceae. Sapindaceae. Dialypetalanthuceae. apical tendril (Polemoniaceae. Guttiferae lllBc. Meliaceae. Verbenaceae. Legummosae. Rutaceae IICb. guminosae. Vines Leguminosae. Proteaceae. Latex Apocynaceae. Caricaceae. Parallel vellatioll Palmae. Julianaceae. Cucurbitaceae. Milky latex Apocynaceae. Rallk odor Juglandaceae. Vitaceae. Simply Pinnate (page 14) IlBa. Vines Cucurbitaceae. Sterculiaceae. Gnetaceae. Caryocaraceae. (Dioscoreaceae). ALTERNATE AND COMPOUND ITA. Cycadaceae J'. Burseraceae. Bombacaceae.leaves Loganiaceae. T-shaped tric/tomes Malpighiaceae IlIAa. Bipinnate (page 14) Leguminosae. NOlldescript Meliaceae (Trichilia). Convolvulaceae. Bitter taste Simaroubaceae IlBg. Capnfollaceae. LegumiIIBi. Erythrina. Cyclanthaceae III. Moraceae.

Clethraceae. Euphorbiaceae IIlBh. Campanulaceac. Leaves parallel-veilled or lacking secolldary veins Podocarpaceae. Chloranthaceae. Rhamnaceae. (Urticales) " IVAf· Unequal petioles Araliaceae. Ebenaceae. Verbenaceae. Latex Sapotaceae. Myristicaceae. Euphorbiaceae. Clethraceae. (Urticaceae) IVAp. (Burseraceae). Lecythidaceae. Acanthaccae. Thymelaeaceae (Leguminosae) (MalVales). Capparidaceae. Phytolaccaceae. (Hernandiaceae). Violaceae IVAi. Buxaceae. (Mal vales). Cyrillaceae. VlOlaceae . Thickened or flexed petiole apex Dipterocarpaceae. Cactaceae. Rosaceae. Guttiferae. (Araliaceae). Flacourtiaceae. Leguminosae. (Moraceae). (Menispermaceae) ~egomaceae. few May tenus). (Flacourtiaceae). Theophrastaceae. Boraginaceae. (Winteraceae) IVAc. Flacourtiaceae. (Myrsinaceae). Celastraceae (GYlwlOsporia. Icacinaceae. (Violaceae) IVAI. Hippocrateaceae. Chloranthaceae. Composi tae. Papaveraceae. Euphorbiaceae. Gesneriaceae. Anacardiaceac. Ochnaceae. Rhamnaceae. Flacourtiaceae. Elaeocarpaceae. AqUifohaceae (few /lex). (Sabiaceae). (Euphorbiaceae). Gramineae. Serrate margillS Hippocrateaceae. Olacaceae. Cornaceae. (Guttiferae). Humiriaceae. Euphorbiaceae. Compositae. (Annonaceae) [VAb. Celastraceae. Phytolac~aceae. Malvaceae. Stipules IV. Styracaceae IVAm. Linaceae. Euphorbiaceae. Annonaceae. Lepidote or stellate trichomes (Annonaceae). Lacistemataceae. Euphorbiaceae. Euphorbiaceae. Alzateaceae. Hamamehdaceae. Cunoniaceae. Solanaceae. (Sterculiaceae) Celastraceae. Loganiaceae. Amaranthaceae (Pleuropetalum). Polygalaceae. Myricaceae. Onagraceae. Bombacaceae. Cochlospermaceae. (Sapotaceae).8 Key Outline Key Outline 9 11IBe. (Leguminosae). Malpighiaceae. Myrsinaceae. (Chrysobalanaceae). (Leguminosae). Trees (page 37) IVAa. Combretaceae. Caprifoliaceae. Fagaceae. Dilleniaceac. Moraceae. (Myristicaceae). (Buxaceae). (Palmae) IVAII. Lacistemataceae. Violaceae. Rosaceae. Palmately veined (or 3-veil/ed) (I) Malvaleall Pulvillus Tiliaceae. Verbenaceae. Rhamnaceae. Boraginaceae. Lacistemataceae. Theaceae IVAk. Flacourtiaceae. Labiatae. Petiole glal/ds Chrysobalanaceae. Melastomataceae. Brunelliaceae. Odor of esselltial oils Piperaceae. (Rhamnaceae) IVAI!. Urticaceae. Erythroxylaceae. (Euphorbiaceae). (Compositae). Icacinaceae. SIMPLE AND ALTERNATE IVA. Amphitecna). Parallel tertiary vellatioll Opiliaceae. Icacinaceae. (Chrysobalanaceae). Hummaceae. Flacourtiaceae. Columelliaceae. Caprifoliaceae. (Passifloraceae). Serrate margills Actinidiaceae. Combretaceae. Nyctaginaceae). Magnoliaceae. (Rutaceae). Euphorbiaceae. Compositae 11IBf Glands 011 twig at petiole base Vochysiaceae 11IBg. Aquifoliaccae. Rhamnaceae. Araliaceae. Apocynaccae. Symplocaceae. Icacinaceae. (Myricaceae). (monocots). Columelliaceae. Rhizophoraceae. Strong bark (Annonaceae). IVAe. Chrysobalanaccae. Fagaceae. Oleaceac. Rhizophoraceac. Lauraceae. Sabiaceac. Celastraceae. (Solanaceae). Olacaceae. Betulaceae. Lythraceae. Winteraceae. Solanaceae. Myristicaceae. Verbenaceae. (Magnoliaceae). Capparidaceae. (Anacardiaceae). Bixaceae (2) Without swollel/ pulvillus Ulmac:ae. (Icacinaceae) [VAd. Sterculiaceac. Rhamnaceae. PUllctations Flacourtiaceae. NOlie of above . Myrtaceae. (Leguminosae). Loganiaceae. Lauraceae. Saxifragaceae. Flacourtiaceae. (Meliaceae) IVAj. (Elacocarpaceae). Compositae. . Lecythidaceae. . Blgnoniaceae (Crescentia. Acanthaceae. Odor of essential oils Monimiaceae. (Oiacaceac). (Polygonaceae). (Caricaceae). Elaeocarpaceae. (Flacourtiaceae). Monimiaceae. Hernandiaccac. Olacaceae. Olacaceae IVAo. Spilles or spine-tipped leaves Berberidaceae. Theaceae. Euphorbiaceae. (Dilleniaceae). Elaeocarpaceae. (Theophrastaceae). (Solanaceae). Capparidaceae. Elaeocarpaceae IVAg. Ochnaceae. (Rosaceae). (Moraceae). Dichapetalaceae. Thcophrastaceae. Boraginaceae. Theaceae. Polygonaceae. Goodeniaceac. Dichape~a!aceae. Miscellal/eous Nyctaginaceae. COllical termil/al stipule Moraceae.

10 Key Outline KEY TO FAMILIES c IVB. (Phytolaccaceae). Leaves bipinnate to biternate (and opposite) Bignoniaceae . plants Leguminosae . (Malvaceae). Bipinnate only in herbs: leaflets serrate. (Urticaceae). Rhamnaceae. conspicuously jointed. Leaves simple and alternate Hints for compound leaves: If in doubt. (Dilleniaceae). Compositae. Telldrils absellt (I) Parallel veills Araceae. Compositae. Pleol1otoma. Vitaceae. Violaceae. Digomphia [I sp. Dioscoreaceae. Zygophyllaceae. Cyclanthaceae. Ulmaceae. Memora. Plumbaginaceae. Lianas (page 60) IVBa. Cactaceae. Compositae aromatic. Marcgraviaceae. In deciduous species. Leaves compound and alternate III.A few PW'kia species with glandular m'ea on top of petiole and characteristic legume pulvinus and pulvinuli. Lauraceae (5) Petiolar or lamilla base glallds Euphorbiaceae (6) Palmately 3(-5)-veilled Sterculiaceae. (Onagraceae) > The families may be divided into four main vegetative groups: I. Convolvulaceae. (Annonaceae). (Caricaceae). In fallen leaflets. (Olacaceae). Campanulaceae. Cucurbitaceae. Juglandaceae. Tecoma. Loasaceae. Lcguminosac IVBb. Leaves simple and opposite IV. Leaves simply pinnately compound (and opposite) Useful differentiating characters include: the rachis (winged in Cunoniaceae.Clematis: lianas with deeply multicostate stems and sensitive petiole. Bignoniaceae (few genera. Only one important family (Bignoniaceae) plus a few other small families and miscellaneous genera or species are characterized by opposite compound leaves. Phytolaccaceae. Amaranthaceae. and Juglandaceae. Rhamnaceae. and presence and type of stipules. . angled and more or less grooved above in Bignoniaceac. (Basellaceae). KEY I LEAVES OPPOSITE OR WHORLED (AND COMPOUND) . (Boraginaceae) (3) Deeply lobed alld/or peltate Tropaeolaceae. (Caricaceae). especially in Staphyleaceae). Tmdrils Passifloraceae. Solanaceae. lB. (Basellaceae). few Arrabidaea species. Euphorbiaceae.)) -. Dichapetalaceae. Sapindaceae). (Menispennaceae). (Leguminosae). Very weak interpetiolar line (V -shaped in Digomphia). (other monocots) (2) Serrate leaves DilIcniaceae. Polygalaceae. Urticaceae.Jacaranda. Boraginaceae. IA. thiCk twigs tend to indicate compound leaves. BruneIliaceae. (Olacaceae) (8) Spilles (Ulmaceae). Ericaceae. Euphorbiaceae. Icacinaceae. Compositae. Celastraceae. entire and subsessile in Digomphia. leaflets peliolulate and ~harnlv serrate in Tecoma. Leaves compound and opposite II.the easiest category. (Polygonaceac). Gramineae. look for the axillary bud to determine whether the leaf is really compound or not. Menispermaceae. Ranunculaceae . asymmetric leaf bases often suggest origin [rom a compound leaf. (Solanaceae). type of marginal serrations and pubescence of the leanets. Smilacaceae. (Aristolochiaceae) (4) Primitive odor Aristolochiaceae. Basellaceae. Moraceae. Euphorbiaceae (7) Latex Convolvulaceae. Solanaceae (9) NOlle of the above Polygonaceae. Ericaceae. (Olacaceae). presence or absence of interpetiolar line or ridge (absent only in three families that are usually alternateleaved: Rutaceae. Polygalaceae. Hernandiaceae. (Euphorbiaceae).

Hida/goa has unde- Zygophyllaceae - Caryocaraceae (only Cmyocar: lowland) . the sessile leaflets with coarsely serrate to remotely dentate margins. A few Cl e1l1atls .Few species.Trees. sessile on strongly angled rachiS more or less grooved above.<0 'I ..Onl~ Amyris: with conspicuously punctate leanets. Taraiea).. all montane. thick-based.Vines or lianas.A fe with 0 and . Hippocastanaceae (only Billia: montane) . Rutaceae (Metrodorea) . similar to bignons but very different deeply costate-ribbed stems and evenly costate-striate twigs. BruneIliaceae . The flISt three families listed below are mostly vines when 3-foliolate (also herbs in Compositae and trees with lepidote scales in Tabebuia of Bignoniaceae). Rutacea~ . Q~iinaceae - Very dry areas.p· . Matayba apetaia with margins entire. .~n"n. distinctive small leafy caducous stipules. branchlets with a conspicuous rubiac-like terminal stipule.1. Verbenaceae (Vitex) .. Ranunculaceae . may have occasional variants (or species) which are 3-foliolate. with opposite Simply pinnate leaves. and with closely finely serrate or serrulate leaflet margins.. usually closely serrate and with prominulously reticulate venation below and numerous secondary veins making obtuse angle with midvein.Leaflet bases rounded to cuneate. veloped terminal leaflets forming kind of tenuous grappling hooks. in so~e s~ecies be~oming stipulelike. interpetiolar ridge usually with small subulate stipulelike projections. w a ellana specICs. e. . are vines h~~~~~~. rachis usually winged. ID. petlOlulate.~~~l. .Only Sambucus: montane. (sometimes very small) to broadly oval. green-bean odor.. Leaves palmately compound (and opposite) Bignoniaceae (most neotropical trees of the family) . the rest are trees. leaflets often serrate or serrulate). oblong. round-tipped. Only Touroulia: leaflets often incompletely separated with bases usually decurrent on rachis. almost always more or less densely pubescent leaves.Montane except in Choc6. at least in part. lower leaflets smaller.species . Bignoniaceae (most lianas of the family are basically 3-foliolate) Unique in the terminal leaflet of some leaves replaced by a tendril (or tendril scar) Ranunculaceae (only a few Clematis) .Trees. twigs and petiole hispid or the tWigs wllh conspICuous round white lenticels.Montane. Caprifoliaceae .f'p. not tapering into petiolule (except Godmania with characteristic rank vegetative odor [cf.Trees. especially Bidens)Characteristic composite odor. very characteristic in hollowed petiole base with Iigulelike dorsal projection. petlolule short. typical rank walnutlike odor. completely entire leaflets. glabrous. always petiolulate leanets with stipel-like glands between upper leaflets.Key to Families 12 Key I Staphyleaceae - Only TUl1Jinia with glabrous leaves on conspicuously jointed rachis. Cunoniaceae . lacking tendrils but petioles or petiolules sometimes twining. typical legume pulvinus and pulvinuli. Sa~indac~ae .g.Very few genera (PlatymisciulIl. many species are characterized by distinctive gland pair at apex of petiole (unique in opposite-leaved taxa). horse urine]). dr~~ng~lac~ish. Compositae (Hidalgoa [vine] and several herbs. with secondary veins rather close together and ending in spinose teeth. leaflets entire. 3-foliolate taxa. Leaves 3-foliolate All groups listed as simply pinnate above and some of those listed as palmately compound below. 13 IC. "O. Vaierianaceae ."g!n~~~:"l:~. il I Verbenaceae (Vitex) •• _1:1 M Leaflet bases taper into indistinct petiolule c1iffers from almost all area Tabebuia ~n". are our only hanas . acute to acununate. pubescence usually of stellate trichomes or lepidote scales.Vegetatively very similar to Caryocar but lacks petiolar glands and elongate terminal stipules. ~ I . Leguminosae .. leaf bases usually gradually tapering into indistinct petioles is unique among opposite. finely and rather unevenly serrate. Juglandaceae - Only Alfaroa: montane.:~: ~~S~~~~iShed?y. serrate margins.

Usually serrate leaflets with characteristic walnut odor. young growth often conspicuously whitish. swollen petiolules (whole length of petiolule uniformly cylindric) and petiole base.spiny with spines shorter and thicker-based than in pinnateleaved spiny palms. Bipinnate leaves and spines on trunk.Canned-meat odor is lypical of Roupala. branches.These two families usually have a fairly strong turpentine-like or mangolike odor but are very difficult to tell apart vegetatively. ~Meliace~e) (Melia) .Usually more weakly turpentine-odored or with a somewhat sweetish mangolike odor (only with a very faintly mangolike odor in the common Tapirira guianensis which might be taken for a Trichilia with too many leaflets). or rachises are unique to mimosoid legumes and some species of Caesalpinia. trees or herbs. check bOlh families. Matayba. look for spines on trunk or stems (unique in simply pinnate taxa) and typically punctate leaflets.Cedrela typically has a somewhat garliclike odor but always entire leaflets. Leguminosae .A tree cultivated and also escaped. usually with one side flat (even concave). Araliaceae (Sciadodendron) . charcteristic large flat-topped inflorescence.Key to Families 14 KEY II LEAVES ALTERNATE (AND COMPOUND) . Odor of esse1ltial oils or turpelltille Rutaceae .) .Small second-growth tree of dry areas. ---------------------------IlBb. Meliaceae .Trees. Leaflets with parallel vellatioll ~~Jnae. Anacardiaceae and Burseraceae . Proteaceae . Rallk odor Juglandaceae (montane) . at least along margin below. Sapindaceae . ITA. Talisia.-.Several families in this group are very difficult to separate on the basis of sterile characters. the other margin serrate.) from some Meliaceae (Trichilia) and a few nonaromatic Anacardiaceae (Tapirira). characterIzed by thm serrate leaflets. Rutaceae (Dictyo!oflUl) . sometimes with a watery latex which dries black (wounded trunks often stained with black). especially some Sapindaceae (Cupallia. mostly with compound juvenile leaves. but often with bitter taste and the strongly alternate leaflets becoming much smaller toward leaf base. often +/. like nonspiny bipinnate legumes these taxa are characterized by the typical cylindrical legume pulvinus and pulvinulus and (in mimosoids) by development of an often elaborate gland on the dorsal side of the lower petiole or sometimes between the lower rachises. IO?kS much m?re like Araliaceae than Meliaceae but twigs much thmner than Scwdodendron and lacking characteristic ligulelike base.Vines with basically ternate leaves. Connaraceae (lianas and occasionally treelets) lack the odor but are otherwise indistinguishable vegetatively from Leguminosae). fiB. (Vitaceae) (a few Cissus spp. Leaves bipinnate (sometimes ternately so) (and alternate) Leguminosae .Vines and lianas (usually basically ternate and usually with milky latex and sometimes characteristic compound stem anatomy) or trees (Dilodendron with many small serrate leaflets and the typical aborted rachis apex of the family). leaflets with marginal punctations. thick branches and very large leaves with a ligulelike dorsal projection on petiole base.Zanthoxylum.Vines. may be consi- . etc. Anacardiaceae . Leaves simply pinnate (and alternate) llBa. Some Picramnia species (Simaroubaceae) have similarly pulvinate petioles and petiolules. Easy to recognize by distinctively round.Similar only to palms from which the leaflets differ in being more coriaceous and having the parallel leaflet veins all equal. the trunk slash has a characteristic odor resembling low-quality canned beef. leaf-opposed tendrils and enlarged often reddish nodes are characteristic. the leaflets are extremely asymmetrical. ~e only woody monocots with pinnately compound Key II 15 Cycadaceae . ---------------------------I1Bc.Most legumes have a characteristic more or less rank green-bean-like odor.

---------. (Apparently a few people don't taste tbe . even in bark. or leaves. ---------------------------IIBf. Palmae . almost always with milky latex either in the twigs or as few widely scattered droplets from the bark slash. most pinnate-leaved rutacs are species of Zanthoxylum and most of these have spines on the trunks..• nn no. also often with sninp. Eve1l-pinnate leaves Leguminosae .. the punctatlOns often rather linear. bltte~ pnnclpal of Simaroubaceae.Subwatery latex in Toxicodendron (montane. Sweetish odor . h ~n . thick spines on yplCal. (black resinous latex in many Old World species).16 Key to Families Key II dered a derived version of burseracs differentiated by the technical character of one anatropous ovule per locule or a single ovule in ovary. usually only with paired s lpU ar spmes and usually with red latex.. n• . 17 (Sapindaceae) . excluding mGngolike odors) Meliaceae . . and several other c~esalpi?ioid genera. Machaerium.Red latex in a number of papili..(Look both asainst strong fisht and out of it.) Rutaceae .sae - Machaerium. (Burseraceae) . all with typical ~"T"ln"O onrl nnJvinnJj and often with LYDlCallegume odor. almost always in exceedingly inconspicuous scattered droplets in stem slash...A few genera of legume have punctate leaves. Spines - -------------- (Rare ill Simply pi1l1late taxa) ~~g~mino. Anacardiaceae .Latex sometimes present in twigs. Leguminosae . Very bitter taste Simaroubaceae . Latex (rare ill compoulld-leaved taxa) Sapindaceae .. Swartzia. !::c:a~ - Zanthoxylum.. at least in part.(In trunk only. Distinctive in the few thick branches with terminally clustered leaves and serrate leaflets.Essentially an anacard reduced to wind-pollination. alllianas.The majority of Zamia species have short spines on the petiole and/or rachis. .. and two per locule. characterized by milky latex..-------IIBg. Burseraceae . have branchderived spines or short spines on angles of stem..Most Meliaceae arc characterized by a faint but distinctly sweetish odor from the trunk slash (but Cecb'eta has a very different rank garlic like Odor). in our area occurring only in very dry parts of the western Andean slopes of Peru. Burseraceae . has branch spines._.. these typically continuing to exude and forming cloudy white drippings below trunk wounds. technically separated from Anacardiaceae hy ovules pendulous. highly allergenic) and few other genera. Cycadaceae .------------------lIBe.~. Pterocarpus. L' .. ' -----. Dussia. ---------------------------lIBIz.Several pinnate-leaved palm genera have spiny trunks and/or leaves.Most species of Rutaceae are punctate at least along leaflet margin. e wig ~s c ewed.lnga (unique in being even-pinnate and with gl~llIds between all leaflets).Bursera orinocensis. mostly lianas. .. Another helpful hint: If you suspect Simaroubaceae get someone else to taste for you or be sure you have water handy!). always trees in Neotropics..A very few Paullinia species.Usually strongly turpentine-odored. egum~nosae .lIBi.. the latex drying whitish around trunk wounds. punctatlOns are often restricted to the sinuses oj marginal teeth or se11'ulations.Most simaroubacs are characterized by a bitter taste wh th t .. somelimes very tamtly m caesalpinioid Dialium). ~ullctatiolls . branchlets.. epitropous.Latex present in most lianas (characterized also by bifurcating inflorescence-derived tendril and frequently compound wood) but never in trees. distinctive in its strongly pungent aroma. Cassia (often with glands between basal patr or pairs of leaflets). check your taste buds on a known specimen. ---------------------------lIBd.onate gen~r~ (e.lnTBn n_~ . tends to dry blackish and sometimes visible around old wounds. Julianaceae . --------.

especially Guarea with typical apical "bud".umes and connaracs which are extremely hard to disting~ISh vegetatIVe.'" . The leaflets are enti:'e.. and thickened flexed petIolules and often suhwoody petiole bases.Huertea has membra~aceous asymmetric leaflets with distinctive serrulate-glandular margms.Some herbaceous and vine Solanum species have winged rachises.Several unrelated genera have winged rachis: Inga. __ . often with incompletely divided leaflets. Legumes can be eIther trees or hanas. odor. mostly with compound wood and/or bicompound leaves. is atypical in the family in odd-pinnate leaves. Nondescript .Cobaea is a cloud-forest vine with tenuous leaflets and a much-branched terminal leaf-tendril. Sabiaceae . Often there is at least a faint trace of a mangohke . Miscellaneous useful characters for genera or common species with pinnately compound a/temate leaves (1) Winged rachis -Individual species of many genera and families: Leguminosae .. Simaroubaceae . the small sessile oblong leaflets. and there is usually a sweetish odor from the trunk slash..) Meliaceae . confuse wilh T. Pal mae and Cycadaceae .Quassia amara is characterized by mostly 5-foliolate leaves and biller taste.Mutisia has the leaf rachis ending in a tendril. Solanaceae .~t Rnecies dries with a characteristic reddIsh tmt.Most meliacs except Trichilia are even-pinnate. oddpinnate). have winged rachises. . coriaceous... : . _ 1 ~ _ •• (4) Naked rachis apex - (5) Petiole base expanded to form sheath surrounding twig- Polylepis (Rosaceae).illal "bud" of ullfolding leaflet pair at tip of rachis (see IllustratIOn) ..18 Key to Families Key II 19 Meliaceae . latex. ... m our area connaracs are nearly always Iianas (rarely treelets).One tree with even-pinnate leaves with distinctively subalternate leaflets (Sapindus saponaria) and many Iianas (Paullinia and Seljania). arc also characteristic. Sapindaceae .ly (an~ to some extent of Picranmia).. one speclCs distinctive in glandular area on petiole (cf. Compositae . Swartzia (few species. unfortunately the commonest Meliaceae genus. (~) Term. etc..Guarea(Meliaceae). usually conspicuously pubescent below.Most Sapindaceae are basically even-pinnate but with alternate leaflets and a very characteristic aborted rachis apex at base of what often appears to be a terminal leaflet. no spines.Herbaceous Vicia (and some species of bipinnate Entada) has the leaf rachis ending in tendril. Stapbyleaceae .Even-pinnate but very distinctive in their parallel-veined leaflets (see above).Odorless anacards (e. Dipteryx (only subwinged). most obvious in juveniles. restricted to the highest altitude Andean forests has a very characteristic growth form with leaves borne on short shoots and having an enlarged petiole base that surrounds the twig. punctations. usually in conjunction with strongly serrate leaflets. ---------------------------/lBj.ck obvious latex are very nondescript and espeCially easy to.(Odd-pinnately compound. generally lack a noticeable vegetative odor and never have red latex.A few Burse/'a species.OphiocOlyon has smooth. ---------------------------llBk. a/temate. Burseraceae .· h'I' odor.IIC I 10. distinctive in their strongly aromatic odor. Leguminosae . Tapirira) that also la. ('''mm''nP. Tree Sapindaceae (especially Cupania and Matayba) with pinnate leaves almost always have a very characteristic aborted rachis apex extended as a small projection at base of what appears to be a terminal leaflet. Lepidobotryaceae have a unifoliolate leaf with legumelike pulvinulus. The . (3) Ulliformly cylilldrical pulvilluli alld pulvillus _ Typical of ne~ly all leg. even-pinnate with glands between all leaflet pairs. Meliaceae . olive-grayishdrying leaflets. (6) Tendril from apex of rachis Polemoniaceae .g. very irregular leaflet n~mbers (some leaves often 1(-2)-foIiolate). Sapindaceae . mimosoid legumes). Most legumes have the typical legume green-bean odor ann m~n" ('###BOT_TEXT###quot;.Trichilia..Guarea pter'orachis is even-pinnate with many thick leaflets and broad coria ceo us rachis wings. Anacardiaceae . Eup/assa (Proteaceae) has a similar rachis apex.

few Fevillea species) are recognizable by the divided spirally coiling tendrils that make a right angle with base of petiole. andlor cucurbit odor.20 Key to Families Key II Simaroubaceae . Leaves 3-foliolate - Usually blunt apex and crenatc leaflet margins. (Dioscoreaceae) (Dioscorea) .Most 3-foliolate climbers are papilionate legumes characte~ized by typical legume odor. all are characterized by the pellucid punctations and most have a more or less citruslike vegetative odor.Only a few species of Dalechampia. ITD. Connaraceae leaflets mostly have finely prominulous venation and a characteristic chartaceous texture that is subtley different from those of legume climbers.Very like legumes in the cylindrical pulvinuli and pulvmus but without a green-bean odor and alwavs lacking stioels Menispermaceae . all differ from cucurbits in having the tendril arising opposite the petiole base and are also usually distinctive in swollen nodes.Nonbitter simaroubs are often characterized by legumelike cylindrical pulvinuli. with the typical legume cylindrical pulvinus and pulvinuli. margin always entire. Leaves palmately componnd (and alternate) IIDa. and by tendency for remotely toothed margins andlor scabrous surface andlor large glands near apex of petiole (Fevillea). are 3-foliolate. a 3-~oli?l~te I. Lophostigma. Erythrilla (Leguminosae) - Usually with spiny trunks and branchlets. Hevea (Euphorbiaceae) . Sapindaceae . (Euphorbiaceae) .At least one menisperm ~as.Several rutac genera have 3-foliolate leaves.Urvillea. all more or less herbaceous.The great majority oj 3-Joliolate climbers are legumes (the leaflets ulliformly entire. Thinouia. a second 3-foliolate euphorb genus. Cayaponia. nc. they are characterized by the bifurcating inflorescence-derived Sapindaceac tendril and usually irregularly coarsely toothed margin. Leguminosae . in seasonally inundated forest. IICa.Easily recognized by the latex. A few other vine genera (all but Psiguria usually with pinnate or bipinnate or simple and palmately lobed leaves) have somc individual species with alternate 3-foliolatc leaves: Allthodiscus (Caryocaraceae) - Cucurbitaceae . . are 3·foliolate vines. very rarely with velY broad entire lobes) and Sapindaceae (the leaflets nearly always somewhat serrate or dentate). progressively smaller toward base of the rachis. and red latex. Ii' " I I. at least in some species.3-foliolate cucurbits (Gul'Unia. the anomalous stem cross section with concentric flngs IS dlstmcl1ve. Picramnia can be distinguished from legumes by the typical alternate leaflets.Four fairly common tree genera plus miscellaneous genera and species oj Rutaceae have consistently 3-Joliolate alfemate leaves.Several Cissus species have 3-foliolate leaves. Piranhea is common in central Amazonia Vitaceae . a few species have simple leaves. charac· terized by the rather thickish and usually curved and angled base of the long petiole. ---------------------------IICb. . Rutaceae . Allophy/us (Sapindaceae) - Usually acute or acuminate apex and toothed (or entire) margins. Psiguria. Villes .A few species of Dioscorea. Most 3-foliolate Connaraceae have alternate basal leaflets (rare in legumes). Trees . Con?araceae .ear. The commonest of these have a characteristic 4-angled subwinged branchlet. and a few species of Seljania and Paullinia have uniformly 3-foliolate leaves. Alternate consistently 3-foliolate leaves are not very common although they may occur as variants in basically pinnately compound-leaved individuals (or species or genera). uniformly cylindrical pulvinuli and pulvmus and often with (unique) stipels at base of lateral pulvinuli. 21 and red latex. Trees Palmae _ Fan palms are our only arborescent plants with palmately compound leaves with parallel-veined segments.

whether with or without spines.One or two rare Dioscorea species have palmately compound leaves. Araliaceae .. Cochlospermaceae - One Cochlospermum has very bombaclike palmately compound leaves. Several genera have spines on the trunk (at least when young) (a unique combination except for JacO/'aria).22 Key II Key to Families Rutaceae . Cyclanthera) have palmately compound leaves.Most species of Merremia have palmately compound leaves. and usually in finely and closely parallel secondary venation. Bombacaceae . characterized by distinctive petiolar glands. One spineless genus has the leaflets continuous with the digitately parted petiole apex (unique in Malvales).g. One species of Paullinia has palmate~y 5-foliolate leaves. The tendril arises at 90° angle from petiole base. is the only palmately compoundleaved tree with milky latex in our area. leaves very dlstmcl1ve m thelf parallel venation. lacking petiolar glands except in Siolmatra. without tendrils and with distinctively angulate petiole with thickened curved base.Syngonium and a few atypical species of Philodendron and Anthurium are hemiepiphytic climbers with palmately compound leaves. Sapindaceae - Araceae . tendrils are absent in Convolvulaceae. adventitious attachment roots. IlDb. The tendril is axillary.Most area araliacs have palmately compound leaves. usually large. characterized by compound stem anatomy and milky latex. one Bolivian) also have Bom bacaceae-like compound leaves.A few species of various genera (e. conspicuously hairy and borne on pachycaul treelet with maroon cauliflorous flowers and cacao-like fruit. ------- 23 . Convolvulaceae . Cucurbitaceae . characterized by the punctations and more or less pungent or citruslike odor.Two area Passiflora species have palmately compound leaves. have. Cyclanthaceae Sterculiaceae - Herrania always has palmately compound leaves. Caricaceae . the latter with dark brown latex in the young twigs and both with stilt roots and the conspicuous conical Moraceae stipule.Some species of Angostura and Raputiu (and cxtralimital species of Esenbeckia and Casimiroa) have palmately compound leaves. usually irregularly divided and with distinctive rank cucurbit odor. Vines . deeply bifid.One genus (Jacaratia). only a few of species become large trees (characterized by tan leaf undersurface). Passifloraceae .Most bombacs have palmately compound leaves. several species have spiny trunks and resemble Bombacaceae except for the latex and lack of a pulvinus. the petiole apex expanded as in Pseudobombax but without the Malvalean pulvinus and leaflets thinner than in Pseudobombax. very different from the above taxa in succulent stems. Nearly all climbing Cycl~nt. Dioscoreaceae .(Most vines with palmately compound leaves are ten- drillate and most are atypical members o/predominantly simple-leaved taxa). rather than truly compound.hac~ae. bombacs are often unusually large emergents with distinctively swollen thick trunkS. characterized by the rank or medicinal odor and the thickly triangular ligule projecting up from the more or less clasping petiole base. always with a Malvalean pulvinus at petiole apex. Moraceae - One common Cecropia and one common Pourouma have the leaves completely split into separate leaflets. two extralimital neotropical species of Sterculia (one Mexican.

. malpighs. swollen jOinted nodes (acanths. Several genera have pair of stipulelike appendages (glands) on petiole or pair of thicker glands near petiole apex. T-shaped trichomes Malpighiaceae _ T-shaped trichomes (= malpighiaceous hairs) give rise to a macroscopically sericeous look. amaranths. hippocrats. Pisonia with leaves mostly on short-side branches. not always obvious to naked eye. Look for: serrate or serrulate margins (Hippocrateaceae). some Gesneriaceae. Gnetum. asperous surface (Petrea [Verbenaceae]). Coriariaceae _ The only neotropical species is Coriaria ruscifolia. Melastomataceae. especially on petioles (and buds) and are definitive among plants with opposite leaves. Coriaria. Most malp~ghs have tannish petioles (from the T-shaped trichomes) a~d browOlsh or tannish twigs. Compositae.. Three-veined leaves . Compositae.---------------IIlAb. Sometimes with watery .Petrea. ---------------------------IlIAa. even on the petioles. Chomelia. Only Hippocrateaceae (plus a very few comps and a few montane forest oddballs of miscellaneous families) are ever serrate or ~er~ulate. thick and cOiled 10 a smgle plane. easy to recognize by the opposite 3-veined leav~s. and combretacs.Strychnos.. Combretum.. unlike combretacs and hippocrats. . Apocynaceae lianas always have milky latex.. :'i I .. they also usually have green petioles and twigs.oltat1Og ~~rtex. The four main liana families with opposite simple leaves are apocs. 3-veined from base (Strychnos. these lackmg 10 other 3-veined taxa. The entire leaves ~e . Rubiaceae (Uncaria.. Prionostemma. the latter with [slow] resin from cut stem). Valerianaceae Serrate or serrulate margins . sometimes with branch spines on stem.d along thin opposite branchlets giving the effect of a mull1foholate p10nately compound leaf. Combretaceae mostly have consp~cuous ngldly parallel tertiary connecting veins adjacent secOnd~f1es. a good 1Odlcator for most entire-leaved hippocrats. Combretum. Lianas A majority of lianas have simple opposite leaves. IlIA. at least in the leaves and young stems and center of stem. Petiolar enations or "glands" - Malpighiaceae. . a feature rare in malpighs and nonexistent in hippocrats (which may ?ave parallel tertiary venation but then almost parallel to the secondanes). several Compositae. palmately veined leaves. a very distinctive and common weedy Andean species with small.lack sericeous indumentum and may have an exf. Hydrangea. petiolar glands andlor sericeous petiole (and/or other parts) with T-shaped trichomes (Malpighiaceae).. spines (several rubiacs mostly with paired spines from leaf-axils). Valerianaceae.dve~n and perpendicularly connected by finer cross velOs. I 24 Key III Key to the Families 25 KEY III Some unusual features of opposite-leaved lianas: Spines . the former usually woody.allel to the ml. or above base (Valerianaceae and Compositae [usually aromatic]). Grows at higher altitudes than StJychnos and the leaf arrangement is very different. when present these are a sure familial indicator. the young branchlets of Combretaceae are generally hOllow-centered and even the large stems tend to have one or more small mucilage-secreting channels in extreme center. Melastomataceae). the latter mainly herbaceous). if in doubt use a hand lens. submilky latex approaching that of some ascleps 10 texture.arrange. Some malpigh genera have petiolar glands which may be stipulelike enations on the petiOl~ (Hiraea) or secretory glands at the petiole apex (especially Sttgmaphyllon).. LEAVES SIMPLE AND OPPOSITE (OR WHORLED) Randia). Malpighs with nonglandular petioles might be confused with Combretaceae but the petioles and twigs of the latte~ ~e usually brown.. Prionostemma (Hippocrateaceae. Melastomataceae ..Pisonia.Easy 10 recognize by the leaves with ?ne ~o four pairs of longitudinal veins arcuately subpar. Stems sometimes fragment into cables ..Hippocrateaceae. the others never do (though a kind of thin watery latex may be present). some Compositae). 3-veilled leaves Loganiaceae _ The only neotropicalliana genus is Strychnos. Coriaria. sessile.very characteristic tendrils are usually present. Asperous leaves . milky latex (Apocynaceae and Asclepiadaceae. rather s?ars~ but .

more cross sec tIOn .g. Valerianaceae (Valerial/a) . and smooth-surfaced with immersed fine venation. Leaves usually with characteristic glands on midrib above. . The stems are usually very flexible and smoothbarked except for conspicuously raised lenticcls. Stems of some genera strongly anomalous. Arrabidaea. and they tend to be hemiepiphytic. w~l1ch may be ~ SCrOPh). The two families are very difficult to distinguish vegetatively and are sometimes united into the same family.1formly simple leaves. a phenomenon which also occurs in a few malpighs. at least at base of midrib. often with conspicuous Simple trlchomes. Pubescence (If present) of simple trichomes. but usually also with some tendnllate 2-foltolate leaves) have simple-leaved species or varian. Cornbretaceae . Acanthaceae .Tribe Liabeae climbers have milky latex but usually triangular or serrate leaves and/or winged petioles. or less membranaceous leaves.ch are 3-veined from base.The best vegetative character may be the usually hollow or secretory (discolored when dry) stem center of young branches and the tendency to have even older stems with I or 3 mucilage-oozing canals in extreme center. all have rather strongly coriaceous leaves with Clusia-like venation. -----.Leaves usually serrate or at least +/. Commonest liana genus is Uncaria with paired curved aXillary splOes. most other scandent comps with opposite leaves tend to be clambering. A few species have rather watery latex. most scan dent species are more or less toothed but a few are quite entire.Abundant milky latex is unique to this and the following family among lianas or vines with opposite leaves. glabrous. IliAd.boPP?site and ~ few species actually have alternate leaves (these species With ste~ splOes). the leaves tend to be somewhat su. Miscellal/eous Hippocrateaceae .ornpositae . The hemteplp es g .Only a few opposite-leaved genera (e. Thicke~ed.Nearly all climbin<>I:> Gesneriaceae are subwoody Gesneflacea . Liabeae climbers lack odor but have milky leaves. hytrowing appressed to a U'ee trunk."". Wulffia) are actually lianas. the margins are usually entire but may be (usually more or less irregularly) toothed. Asclepiadaceae .:10. an interpetiolar line or ridge is never present..Guttiferae climbers rarely have conspicuous latex.! . e . ----------------------------IlIAc.A few compound-I~aved gene~'a (e."" . more or less herbaceous vines.1 1 Ii 26 Key to the Families Key III 27 Cornpositae . leaves are usua IIy mem bran ac eous or succulent and are often either serrate or . C.---------------------- Rubiaceae . some Mikania.ts whi.Essentially a herbaceous version of Apocynaceae and often indistinguishable vegetatively.. this especially pronounced in entire-leaved genera.serrulate' the only lowland liana family (except a few subwoody Gesneriacea~ and aromatic sUb-3-veined Comno~it"p. . has . One genus has reddish or pinkish watery latex. some Mikania species have the leaves 3-veined from base but the leaves then cordate.) with nnnAOlto on __ "'" .One liana genus (Mendol1cia) and several genera which include clambering vines. typically with a few irregular reddish concentric circles interconnected by spokes. distinctive in having the petiolar bases united to [ann a kind of nodal sheath.Vine genera have the typical interpetiolar rubiac stipules but these are not always obvious in many I~ana tax~.g.. strikingly conaceous and With a chatact~I~Sl1C appressed-conical "pseudostipular" axillary bud. Also characterized by typical bent tendril-like lateral branches which twist around support (but sometimes occur in other families). also in usually drying black. on. The combination of opposite leaves 3-veined above the base and aromatic odor (also frequently more or less asperous) is unique to comps. When the tertiary venation is more or less parallel it tends to also be parallel to the secondary veins unlike Com bretaceae. and adventitIOus roots along stem. Guttiferae . Bignoniaceae . and the surface often distinctively asperous.e hemiepiphytic genus (Schlegelca. more or less ~wollen fragile branchlets are tYPIcal of all acanth cltmbers. Milky latex Apocynaceae . Fibrous bark and leaves typically with rigidly parallel often somewhat raised ter~iru: venation and brown petioles (sometimes with more or less exfoliatlOg cortex) are typical. Leaves of nonserrate genera are coriaceous and often dry with a characteristic grayish-olive color and are typically coriaceous. I d' d Mendoncia has a soft easily broken stem with an amazmg y Issecte · an d often with corky bark'' also.The leaves of Climbing species are 3-veined above the base just as in Compositae. no des an d rather . In general woody lianas are apocynacs while herbaceous climbers are mostly asclepiads.

and resin oozing from cut stem with concentri~ rings In cross section are diagn l' h uninitiated will re . 29 lIIB. Gnetaceae (Olletllm) . Nyctaginaceae .Vochysia usually has whorled leaves and/or rather thick-based stipules (other genera have stipule apex broken off to leave characteristic gland. Both often have squarish stems or twigs. :ss s~ri~eous. interpetiolar lines are also usually present. Trees and shrubs (with leaves simple and opposite [or whorled]) [lIBa. prominent subulate axIllary bud scales usually present. These should be in plane at right angle to two uppermost leaves and leave an interpetiolar line when they fall. Chloranthaceae . Y ra~gea IS a thick woody liana leaved species sim~l. Malpighiaceae .. the ba. no es to form conspicuous ochrea-like joint- Rubiaceae . these almost always visible at least on the petioles and young twigs. Ippocrats. early-caducous stipule usually sericeous. Stipules (or stipule scars!) Verbenaceae -. but the margin entire. Iresl11e has hollow twigs. . os ~c c aracters. and Aeglphda. Dialypetalanthaceae .Two opposite-leaved genera become lianas in our area. .. Rhizophoraceae . large triangular pair of interpetiolar stipules. characterized by acanthlike swollen nodes (sometimes ~ runh~n ~bove.Two genera 0 f more or less woody oppositeIeaved climbers In our area I · ' usuall ' lesme an d Pfaffia (other genera are Ch ~ pr~strate or scrambling herbs.The tree genera have intrapetiolar stipules in the axil between the petiole and twig (looking like ligular dorsal projection from petiole base). Trigoniaceae . these always separate (usually fused in Rubiaceae) and often rather long and subfoliaceous.) . .k IS usually smoothish and light colored. Quiinaceae . the fluted-based trunk with thick reddish fibrous bark also distinctive. nodes coria~eous leaves.Swo IIen. more or less Jomted .Rhizophora. " Amaranthaceae ' . differs from Rubiaceae in the usually serrate or serrulate leaf margin (deeply incised in juveniles of few species).r ~~I~ :tth entlIe or se~'fate leaves.Rather s· '1 ally whitish I Iml ar to Malplghlaceae because of the usuwith a dense :a~r~f or. the leaves arc often membrana~eous and/or noticeably simple-pubescent in Aegiphila. discolored twO ' .. with Sterigmapetalum having mostly whorled leaves with tannish-puberulous petioles and caducolls narrowly triangular-pubescent stipules between them and Cassipourea in the leaves usually obscurely and remotely denticulate or serrate (unlike Rubiaceae) and the small. the other (Colignonia) woody only i~ cloud forests and characterized by the tendency to verticillate leaves (and the very Characteristic Hydrangea-like inflorescence bracts)' both have concentric rings of anomalous growth.Interpetiolar stipules are present at least 99% of the time. the other two opposite-leaved genera are less striking. (Note: there is no way the Salada and relatedCoh~n1Ze thiS as a gymnosperm. restricted to coastal mangroves. these differing from the few rubiacs with similar stipules by being fused (usually bifid in Rubiaccae except Capil'ona with conspicuously smooth red bark). the plants are easily distinguished by the strong Ranalean odor and the serrate leaves. npubescent specIeS tend to have hollow or in having an ~~~~~~~:~I~~~i. :wo genera of climbers. The only banas other than Petrea with asperous opposite leaves arc Prionostemma aspera (hippocrat) and some comps (3-veined above base).In our area H d . Vochysiaceae . see also alternate-leaved h am~ssoa .The other main family with interpetiolar stipules. it looks much like .The single species similar to Quiinaceae in the completely separate. (sometimes hemie i h' . see below). Petrea with nspcrous leave~. one (Pisonia) characterized by spines and rather clustered not clearly opposite leaves.28 Key III Key to the Families strongly anisophyllous (sometimes strikingly red-tipped or reddish below or with red apical "eyespots").~. entirebases connected across d u laceae when stenle but with petiolar sheath. node when dried) and by the evenly striate-ribbed ranc ets. if stipules are not readily apparent check terminal bud to see if it is enclosed by caducous stipules. but never T-shaped on!fl ~~we -like trlchom~s and/or simple trichomes. narrowly triangular. puberulous leaf undersides.mbretaceae but differ from that family Saxifragaceae . The main familial vegetative charactetistic is the presence of malpighiaceous or T-shaped hairs.The more or less swollen node has a stipulelike sheath. is utterly distinctive as the only mangrove with stilt roots.

differing from Myrtaceae in secondary and tertiary venation of the thick-coriaceous leaves completely invisible.Very distinctive in typical terminal bud and colored latex.Rhizophora. Lacks the typical guttifer terminal bud (i. the petioles of terminal leaf pair not hOllow-based with terminal bud growing [rom within cavity except in a few species with very profuse latex). ---------------------------lIIBd. leaves. . ' and the small C~riac ~ c. Strychnos (few shrubby species). but unifoliolate.. . I. The latex may not be very obvious. . Chloranthaceae .. have oppoSIte leaves. f b th families are unlike the Ranales.IIlBc.Leaves thinner than in most myrtac~ and .. Melastomataceae . 'i .. A few other famIlies h~ve 3-veined bases but lack the typical cross veins (Delostoma [Blgnoniaceae.The very characteristic leaf venation ~~es this one of the easiest families to identify. . Rutaceae - 31 .ooks almost exactly like Myrtaceae and may have punctatlOns: It dIffers from myrtacs in the somewhat jointcd nodes. typical terminal bud not developed only when latex strongly colored. the latter usually by oran"e a mOIst orangish area just inside bark where latex should be. White. ---------..Opposite-leaved Compositae aref nearly T ) nd have veined above the base (unlike any of the above amlles a more pungent odors. peeling bark. try breaking a leaf and twig as well as the trunk slash. Verbenaceae an d LabJ'atae . R I' leaves an d are usu ally aro hat minty) and less pungent than m the ana es. Terminal bud characteristically from between the hollowed-out leaf bases.lose-to?ether secondary and tertiary venation ending in a submargm.These two families IS clearer an sweeter matic but the odor .. 1 I' . mostly opposite. unknown in Myrlaceac) or fro. aro(oft~n somebwrS of these families usually have tetragonal branchlets mattc mem e . I' llIBe.--------- 'i.. Lauraceae . aractenzed by mostly upland habitat latex (or . some myrtacs that are not obviously punctate (also beware Mourll'l [see below]).al vem.(Only two Ranalean families are characterized by aromatic opposite leaves. Strongly aromatic specIes 0 0 mostly herbs or subshrubs. Latex white and free-flowing (red in some species with alternate leaves). Odor of essential oils .. keyed out above on account of the terminal stipule. eous encOld leaflets..m.Mouriri completely lacks the ascending veins of other mel~stom~s.. most notably peculiar 3-veined Cmyodaphnopsis. 1 i II i 'I I' . ---------------------------- Myrtaceae .raceae..) Monimiaceae . stilt roots are rather common.. Andean]).(Some genera alternate-leaved and many are climbers).A very few atypical Lau.. always 3C omposJ'tae . haved opposite . Guttiferae .Shrubby H ypellcul1l and some Vismia species hwc punctate leaves the forme h .e.. usually slowerflowing than in apocs..' Guttiferae . can have leaf punctations.. Latex Apocynaceae .Hedyosmum (our only genus) has very characteristic swollen nodes with stipulelike sheath.-----.Siparuna. ..with more ascending secondary veins and absence of collecting vem.30 Key to the Families Key III IlIBb.Usually further characterized by more or less parallel and c. Many have smooth. differs from Guttiferae in lacking latex and the small glandular p ct (' a un a IOns. ------------. 3(-7)-veilled leaves with parallel crosS veills more or less perpendicular to main veins Melastomataceae . latex white only when the leaf bases form conspicuous chamber. some have aro~altc leaves. cream. The only possible confusion comes from a very few gultlfers that bave punctations (but also latex.I'. usually puberulous or ~ith lepidote scales' the second genus (Mollinedia) often lacks obvIOUS odor but is cha'racterized by very distinctive leaf with few very separated marginal teeth. ch~actenzed by sheathing guttifer-Iike petiole bases in which the apIcal bud is protected. Raven' has " ..Gaiadendron is an exclusively montane freestanding tree with conspicuous punctations. or orange. ne upland Amyris specics has small. la Opposlle SImple leaves WIth punctatlOns.. . Also beware of Mouriri which lacks the typical venation and looks almost exactly like Myrtaceae.. PUI/ctatiolls Loranthaceae . Lythraceae (AdenQria and Pehria) . I . latex commonly yellow. I (Rhizophoraceae) .

IIIBg. . at least on occasion: Viburnum (Caprifoliaceae) -Leaves with few strongly ascending veins.32 . with conspicuously floccose indument on leaf undersides and twigs.. distinctive in the small grayish-sericeous leaves with opposite petioles strongly connected by line or flap of tissue. Several other mostly exclusively IIwntanefamilies also have taxa with opposite serrate leaves. whItish-margined. woody neotropical lowla s IS ~ ~are one. I .lpule-enclosed apical bud immediq e. stl'ong interpetiolar line.Mostl lian ~ h as but a few are trees and Cheilocliby tertiary venation more anI ave serrate leaves. stlpules.th~ typIcally short petioles. margins vary from quite ee ~~ex and strictly pinnate venation.Mollinedia has the teeth usually very widely separated (typically only one or two per side) and rather sharp. commonest understor tr y as OpposIte leaves and is one of the the nodes noticeabl/joi~~e~enera of ~any forests. easily recognized by the strongly 3-veined leaves with glands in the lower vein axils. (Aphelandra [Acanthaceaej) .. I ste late-tomentose below d ac enzed by leaves white or tan shape.. Characterized by tendency to have a small .I Key to the Families IIIBf Glallds 011 Key III twig at base of petiole Vochysiaceae . Siparuna also is frequently toothed but easy to recognize by the Ranalean odor. (Brullellia [Brunelliaceaej) - The occasional simple-leaved species.g. also very distinctive in the flexed.. Serrate (serrulate) mar . Clethra). nate and opposite leaves s charactenzed by a mixture of alteralmost never uniformly ~even . (Alc1lOmea pearcei ([Euphorbiaceaej) - A montane cloud-forest taxon with mostly opposite leaves. above) Afiew'add't' °l~hanthaceae which are easily recognized . The dentate' species with n rre to rather shallowly and coarsely suband so~e of the pub more serrate leaves tend to be more pubescent ... is small tree. Verbenaceae . at least the leaf below and twigs.Sloanea i . dodromous strikingly f f (sec~ndary vems few and brochiI ar rom margm' . arac cnSllc glands from thc fallen stipules _ _ es c aractenze most Vochysiaceae (except Vochysia). but the leaves are but non pulvinate petiol PPOsile. like compound-leaved congeners. sparsely and bluntly serrulate or more or less bluntly few-toothed toward apex. and the ately subtended by obJi uacute ~t.Very ch t" or stipule bas h . (Eleaocarpaceae) . Monimiaceae . Ilona erbace fi '1' ( have some viny (see abo) b ous ami les e. serru ate (resembling si '1 I ' margm mostly remotely blunter).. thus. when fertile distinguished by narrow terminal raceme or spike (cf.. are closely serrate and with prominulously reticulate venation below and numerous secondm'y veins making obtuse angle with midvein. nium can be both a tree Violaceae . puberulous at least below. escent species have conspICuous . ---------.Abatia (always pubescent.Buddleja (m I weedy species B a l ' ost y Andean upland except for one ' ... Desjontainia has spa~ Usually narrow or somewhat rhombic G mose margi lik h Omara have +/_ serrulate mar'" ns e Oily. Hedyosmum (Chloranthaceae) . Gesneriaceae) leaves.The comblllatlOn of opposite s an serrate margin' .. caducous triang~l ar y f7w-to~thed Mollinedia but teeth olar line. 33 Flacourtiaceae . andfound only in eight Melastomataceae and Chi nd fal1l/11es beSIdes the Quiinaceae. looking almost exactly like typical alternate-leaved species of the genus.Typically with more or less tetragonal stem and aromatic odor. Interpetiolar lines lacking. ve or su woody members with opposite serrate Hippocrateaceae . simple leave d gills . Columellia (ColumelJiaceae) - Mostly entire-leaved (with apical spine or apicule) but sometimes with a few thickened subterminal teeth. nerzcana) char t .A few montane species have conspicuously spinose leaf margins. persistent leafy Rhizophoraceae .Cassipourea .. it is characterized or ess parallel and perpendicular to midvein. Peltanthera and "lnS. usually with floccose trichomes very like Callicarpa) rather sharply serrate.----.As noted above unmistakable in the sheathing node and strong Ranalean odor. (cf.on the same branch. interpetiolar ridge. Most woody verbenacs are entire but usually serrateleaved CallicO/pa. u ar termmal slIpule pair leaving interpetiLoganiaceae .Rinorea usuall h . . (Wei1l1l1allllia [Cunoniaceaej) - A few species are simple-leaved but otherwise remarkably similar to the pinnate species in the rather coarsely toothed margins and distinctive small leafy caducous stipules.

usually rather membranaceous and somewhat aromatic. It differs from myrtacs.Only two real tree genera (Bravaisia and T~'ichan­ 1 thera. .In some Myrtaceae the punctations are not very evident. h ous teet towar . has the leaves 1 Capn. interpetiolar lines or ridges lacking except in Lajoet/sia (with close-together secondary veins prominulous above and below. serrate margins) (Lacking Nyctaginaceae . Loganiaceae . oblanceolate leaves. essential oils.Vegetatively heterogeneous. Alzateaceae (close to Lythraceae) . stnilght. L - • O -- 1 .!:o~~~: 1 entire or merelv serrulate. The best sterile character is the reddish-brown pubescent terminal bud. Cornaceae . the secondary veins immersed or slightly prominuJous below. Except for Lhe spiny-margined species.. Most acanlhs characterIzed by con~pl­ cuously jointed nodes.Usually with tetragonally angled young twigs andlor longitudinally exfOliating.Usually with tendency to tetragonal branchlets ~nd 1 raised petiole attachments. • Viburnum exclusively montane.fohaceae .. 1 1 Hippocrateaceae . the.o:i~~::~~om most acanths in lacking cystooften pubescent leaves "nIJ 1 liths and interpetiolar lines. at 1 least of forest taxa.. eav . holly) upland Aphelan I'a 1 species.0 midvein).genera mclud. Miscellaneous opposite simple-leaved trees latex. ven~t1on :nore or 1 less perpendicular \. one genus glabrous with spinose holIyUke ll1argms.Rhamnidillm and some Colubri?a have opposite 1 leaves the former characterized by the close. Yd'" the few strongly ascending 1 d apex) charactenze vy . even when in Dower or fruit. to a well-developed submarginal collecting vein. 1 1 Only Besleria become small soft-. 1 . often reddish twig bark in older branchlets. leaves thick-coriaceous.:::o~:~~~~g~~eti~~e:' y 1 generiads mostl hav~ n. exclusively mn n ' to Viburnum. veins of the family and the pale leaf under~urface.ge very 1 thick-coriaceous entire leaves with Immersed flOC venation and 1 drying a characteristic dull olive. is II large Amazonian lowland tree with opposite leaves having a legumelike odor and cylindtical legumelike petioles. several genera have serrate leaves.Comus.II/Bh.e 1 shrubby or small tree species. except for lacking stipules. ' uall with at least a few inconspicusometimes subentlre (though us . always with conspicuous interpeuolar hne. oval Witil rounded apex and base and very short petiole. (POI!galaceae) . 1 Most acanths have an obvious interpetio!ar line. 11 than in acanths)' 1 our area (these usually with entire leaves sma er W d' tm 1 although several serrate-Ieav. Leaves.wooded trees in · Gesnenaceae . in the jointed nodes. several of Lhe t~~a 1 lacking this line are spinose-margined (cr. The combination of somewhat succulent. 1 1 1 1 1 1 Key III 35 1 1 Verbenaceae . but never truly serrate. 1 Acanthaceae . both with very weak wood) although several . Lythraceae . most notably. each adjacent pair separated by a well-developed intersecondat'Y)' Physocalymma is very distinctive in entire-margined strongly scabrous leaves. secondary veins and tendency to . Myrtaceae . latter 1 by conspicuous glands at base of lamina (typically In basal auncles).Mouriri looks much more like Myrtaccac in vegetative condition than like typical 3-7-veined melastomes.Two genera are sometimes tr~es. another a pachycaul treelet with large coriaceous entire. often different sized and/or subopposite blackish-drying leaves and rufescent terminal bud immediately indicates Nyctaginaceae.. Melastomataceae . a few genera are herbs the commonest of which ha+le'! ' s a fmlOa whorl of 4 leaves subtending the inflorescence.ed ~~!~. They are usually characterized (as are the punctate-leaved taxa) by the straight often rather close-together secondary and intersecondary veins that end almost perpendiculat. these always pubescent. be +/_ puberulous at least on vews. parallel sec1 ondar.. leaf base typically attenuate onto petlo~c 1 and in many species (most Citherexylon) with an elongate gland In 1 the laminar attenuation on either side of petiole apex.Rather nondescript and might be confused with Psyehotria or similar rubiacs. which vegetatively lJas nothmg whatsoever to suggest this family. ' (look like ShOft black lines) often present on upper leaf surface (also 1 in Urticaceae). one 1 (Cheiloclinium) usually with finely crenate.Polygala sclel'Oxylon. sometimes with stellate trich~mes (BlIddleja). the ot~er (SalaelG: ~lth lat. 1 Rhamnaceae .Easily distinguished vegetatively by very strongly tetragonal thickish twigs and jointed stems with strong interpetiolar ridges. ou~ ac~~th~. swollen when fresh and contracted when dned. . fW"" .-serrate ~argms (also distinguished by conspicuously parallel tertiary.

II " 36 strongly ascending secondary veins and petiole bases decurrent onto the somewhat angled tannish puberulous twig. Trees (Guttiferae) .This "grab bag" category constitutes by far the largest and generally the most nondescript group. has petioles usually somewhat thickened at base (cf. also check the twig bark itselD. serrate margins (uncommon in tropical forest species). .Chionanthus. . strong bark (pull a leaf off a twig to see if a strip of bark comes with it.Latex (in Neotropics) always white and milky.I I.' II . unless they are fertile and technical characters are used. Sapindaceae petiolules). how these are arranged. IVAa. Malpighia and many species of Bunchosia have neither obvious stipules nor interpetiolar lines and are often characterized instead by pair of ocellar glands near base of lamina below. be sure to check both trunk and leaves since sometimes obvious latex is apparent only in one 01' the other. I. a few Vismia species (which lack the typical hollowed petiole base) may not always show the orange latex but there is always a hint of orange color under the bark where the latex should be.. Malpighiaceae . margins always entire and latex of slash usually emerging in discrete droplets. Colume\liaceae .Sloanea is characterized by a mixture of alternate and opposite leaves. 3-veined base (frequently suggests Mal vales). has one species on limestone outcrops in northern Columbia. IVA. Margins sometimes bluntly irregularly toothed (see also above). In vines . whether petioles are thickened at base or apex or of unequal lengths. has a conspicuous interpetiolar line (sometimes accentuated into an actual flap of tissue) like Viburnum but is very distinctive in the small grayishsericeous always apiculate or spine-tipped leaves.) (Elaeocarpaceae) . (Buxaceae) .' Key IV Key to the Families 37 KEY IV LEAVES SIMPLE AND AL TERNA TE . breaking the midveins or petioles of several leaves as well as young twigs. Leaves typically with base of petiole enlarged (petiole more or less pop-bottle shaped) and with numerous parallel secondary veins. odor of essential oils (Ranalean odor). in this group there will be many plants which end up as family indets.OccaSionally lacks apparent latex (Tovomitopsis.. sub-3veined leaves and the petiole bases attenuating into strong ridges on the thus irregularly 6-angled twig.Columellia. even on same branch.(Look carefully. Like the species with stipules. conical terminal stipules (usually = Moraceae). latex (a few Convolvulaceae lianas have latex). the leaf blade either pubescent or else rather narrow and oblong. and what color the latex is. Never with conical terminal stipule or glands on petiole. they arc also vegetatively characterized by the typical T-shaped trichomcs at least on petioles and young branchlets. exclusively montane. glands at tip of petiole (usually Euphorbiaceae or Flacourtiaceae). also very distinctive in the flexed but nonpulvinate petiole apex and strictly pinnate venation.' I I ! . ' In trees -Look for (in approximate order of importance): latcx. I [. twigs lacking interpetiolar lines. glands on petiole (especially common in Passifloraceae). some Chrysochlamys). often with scattered round raised white lenticels. spines. Latex (and alternate simple leaves) . but then with the typical hollowed-out Guttiferae petiole bases that form a protective chamber for the developing bud. almost never uniformly opposite. . petiole length and flexion. with very characteristic coriaceous. Oleaceae .Buxus.A few genera of shrubs and small trees lack obvious intrapetiolar stipules (as do the lianas). In flower unmistakable in only 2 anthers and very narrow petals. Sapotaceae .Look for (in approximate order of importance): tendrils (only ten families have true tendrils and the type of tendril is usually specific to a given family). sometimes not very apparent but almost always visible in either trunk slash or leaves (if not both). punctations (and the undersurface texture which accompanies punclations in Myrsinaceae). note whether the trunk slash has discrete latex droplets. Some genera lack the typical ~etiole but these mo~t­ Iy have finely parallel tertiary and secondary vems (the extremes with leaves similar to Clusia). In preceding groups any sterile woody plant should be identifiable to family.

l~ck of R~nalean odor separates from Myristicaceae. Pausandra and some Croton have bright red latex. other genera with conspicuous trunk latex. ed families have young leaves ro e.. a feature never found in Sapotaceae. Conical termillal stipuie (+/. Resinous black-drying latex is sometimes present 10 simple-leaved species.38 . Look for a slightly longish. ceae have conical terminal stipulcs Polygonaceae . somewhat similar to the Moraceae stipllle. . Ompha/ea (liana in our area) has a cloudy latex that turns rather purplish.. a good character for some of these species (series Nitida) is the conspicuously fenestrated trunk. Conical terminal stipules and the scar from these stipules usually obvious (and definitive for Moraceae). Euphorbiaceae .Exclusively high-altitude Bocconia is unique among alternate-leaved taxa in its orange latex. Trap/us g I b s 'dered present by stretching the ImagmatlOn. II similar to the conical ternunal branch apex and are super ICla Y stioule of Moraceae. lack of petioiar glands.. long petioles with flexed apices and often of different lengths. Anacardiaceae .nd almost universal in Moraceae. this not always visible 10 mdlVldual trees.. ) A number of genera of Theaceae (Th ceae and Myrsmaceae II d t ea . also Characterized by the large irregularly pinnatifidly lobed leaves. Papaveraceae .. Note: Although latex is considered characteristic of Euphorbiaceae. a and Myrslnaceae and relat fi . Olacaceae ..D Illllys.. ~yristicaceae . white and milky (usually) to somewhat watery (Minquartia)..' . this most obvious around old trun. . and in a few varying to tannish yellow (some Nauc/eopsis) or dark brown (Pourowna). inconspicuous terminal stipule. The margins are always entire (at least in our area). '' Key to the Families Moraceae .Most species have compound leaves. and a pair of glands near petiole apex are good indicators of Euphorbiaceae and are unique to this family among species with alternate simple leaves and latex.---------. Serrate (or serrulate) leaf margins. Alternate-leaved apocs usually have white and milky free-flowing latex but this may be bright red in some species of A~pidosperma and orangish or pinkish in others (one Aspidosperma has both white and bright red latex in the same tWigs!). also can on y e con I . where recognizable as Moraceae only by the typical leaf venation..Latex of over half of species a unique tan shade (exactly the color of "cafe con leche"). Aspidosperma often has little or no latex in the trunk.. Many species of Aspidosperma are extremely difficult to distinguish from Sapotaceae vegetatively. has an Winteraceae . (Chrysobalanaceae) . Leaf venation very characteristic with the brochidodromous lower secondary veins closer together andlor joining mid vein at different angle from others... the only potentlal confuSlOn.At least one species of Annonaceae ?1~y h~ve a faint trace of red latex (Unonopsis f!oribunda).:ith a distinct trace of r~ddish latex. distinctively curved (putatively U-shaped) and somewhat apically thickened petiole. Very easy to distmgUlsh from other families with occasional species or Jl:enera with Key IV 39 red latex by the typical myristicaceous branching. the stipule is not always ObVIO~S Q!:d lI1. . M - The combination of milky latex and conical terminal Sti~~~. The exception is Tl'ophis where neaher latex nor stipule may be discernible. v. but many other species with milky while latex (usually only watery in Trophis). Campanulaceae ..[VAb.definitive for Mor~ceae) -:A ain look carefully. Apocynaceae . at least ~n Neotropics.. It can be distingUished from Myristicaceae by its strong bark.k wounds and not usually evident in fresh slashes of the leaves or tWigs.. Mostly herbs but a few higher-altitude shrubby trees (actually overgrown herbs). . (Annonaceae) . espeCially when drying [except Osteophloeum which is ~ersl~tently straw-colored but still dries reddish]).~n~chrea rather than falling cleanly but these rupture to form as in Moraceae. and presence of Ranalean odor.Usually with red latex (only in trunk. b t is aromatic and that falls to leave a conspIcuous nodal flng.Very ~ar~ly. lacking latex and strongly aromatic.Latex present only in few species and usually present only in leaves and petioles. this sometlmes rather watery at first but almost always SOon becoming obviousl~ red. many species have no latex at all.Relatively few apocs have alternate leaves.~~~at falls to leave a distinct scar) is definitive [or a..some POly. • 1: I has a Moraceae-like terminal slipule Magnobaceae a awn~ . often caustic and harmful to eyes.: " . 'I ". leaves of most species of this family have a characteristic grayish or tanniSh-green color when dry. u nonlactiferous. Look for Sl1pules on the young twigs or their scars.White to cream milky latex typically present.. h' h fi 'ally note that otherfamities may have young leaves w IC are super ICI .Always with white and milky latex.

note complete rings around twig at nodes from the distinctive caducous stipule that completely covers terminal bud. leaf base often strikingly asymmetric. are intermediate but stellate (or lepidote) trichomes are rare (mostly Duguetia) and sericeous pubescence is common only in Xylopia. typically with at least the hi~t of an inv~lu­ tion of the margin and sometimes with a distinctly Involute auncle on each side. Typically. when present is usually sericeous with appressed simple trichomes or softly rufcscenl. the only significant South American genus. Typically Lauraceae have short elliptic leaves with glossy shiny surfaces. Lauraceae leaves typically have shinier surfaces than do the other two families.. The vegetatIVe odor IS ranker than in most Lauraceae in which 3-ve~ned taxa also differ in shorter petioles. At least five other taxa with simple alternate leaves have. typically with prominent stilt roots. all the gre~n-twlgged annonacs have strong bark but only some of the brow. in others the reverse may be true. ~f '! .Swollen nodes with shoot proceeding from leaf axil. Very many Lauraceae are distinguished by the way the leaf blade gradually merges with the petiole apex. when trees (usually small). Hernandiaceae . as usual. Annonaceae are intermediate and overlap with both vegetatively. in Myristicaceae.g. with little or no leaf odor. the bark slash is aromatic. As a group these are easy to recognize by their "primitive" odor and many of these families are vel)' common and important in neotropical forests. Although there is little room for confusion between Myristicaceae and Lauraceae. Myristicaceae.Most Rana/ean plants have alternate simple leaves and most have more or less conspicuous rank or tlllpentiny odors. Most Annonaceae have strong bark ("cargadero" = useful for lying cargo) a featur~ not found in Myristicaceae or Lauraceae. Annonaceae . and relatively few.long often somewhat different-length petioles. often sU'ongly ascending and not strictly parallel secondary veins. Lauraceae never have such a phyllotaxy and their leaves are often irregularly spaced along the branches with a definite clustering towards the branchlet apex. In basal lateral vein pair curving upward rather than being straight or curving outward. and relatively glossy shiny surfaces. Magnoliaceae . Odor of essential oils (Rana/ean odor) . The beginner almost always complains of either not having an adequately developed sense of smell or of being unable to discriminate nuances of different vegetative odors. and usually rufescent (to whitIsh on the leaf undersurface).n-twlgged ones.Only Drimys (montane) which has the leaves strongly whitish below and lacking noticeable secondary veins. that of Annonaceae tends to be slightly rank. and many close-together parallel secondary veins. Key IV 41 I This leaves three very large and very important Ranalean families Annonaceae. frequently very conspicuous. odors that might be confused with the Ranalean group: Most slmpleleaved species of Anacardiaceae (c. especially. Myristicaceae trichomes are either stellate or 2-branched (. Frequently a twig split longitudinally will give off a more easily detectable odor than the leaves themselves.Mostly montane. SPICY. Both Myristicaceae and Annonaceae (but not Lauraceae) are characterized by myristicaceous branching with the lateral branches at right angles to the trunk and the evenly spaced leaves 2-ranked (except Tetrameranthus) along these or along their lateral branches. Differ from similar Arah~ceae in lacking conspicuously smaller short-petioled leaves and. odor tends to be peppery (not surprisingly since pepper comes from this family). 4]). relatively long petioles.commonly have either green or brown twigs. Also distinctive among ~anales in .which are easy to tell apart when fertile but can be confusing when sterile. relatively long petioles. Also. In some Lauraceae. and Lauraceae . Lauraceae tWigs are typically green while those of Myristicaceae are brownish. Don't despair .T-shaped). an important warning: There is a common epiphyllous leafy liverwort with a rather licoricelike smell. Distinctive spicate inflorescence. Families with Ralla/ean odors -All with completely entire margins except for a very few somewhat lobed-leaved (but never serrate) species. The odor of Lauraceae is usually either clear.Three-veined (occasionally peltate or subpeltate and rarely somewhat 3-10bed in part) leaves are unique in aromatic ~analean taxa except for a very few atypical Lauraceae. Piperaceae . Myristicaceae have relatively long oblong leaves with dull surfaces. the lateral branches tend to be clustered and appear to have an almost whorled arrangement ("myristicaceolls branching" [Fig. Winteraceae . the petiole is conspicuously grooved above in Talauma. Annonaceae. Usually shrubs. Learn that anise odor well and eliminate it from consideration.40 Key to the Families IVAc. that of Myristicaceae is usually more pungently turpentiny and typically not very strong.with a little bit of practice you really can (usually) pick out the Ranalean families by the combination of simple alternate leaves and their odor. and almost sweetish or foetid and unpleasant. Anacardium) have a morc . short petioles. and the pubescence.

Flowers distinctive by numerous stamens with filaments fused around style into staminal column and/or an epicalyx. Mostly differing from Slcrculs and tiliacs in combination of more broadly ovate leaves with serrate or lobed margins and from most other Malvales in less developed pulvinus. When sterile. pulvmus.t usually included in Malvales but has a similar. Petioles with apical pulvinar thickenillg (or with leafy stipules) (= Malvales) -Perhaps the main palmately veined group of plants. Although recognition to order on vegetative characters is :asy. I I . are best characterized by fused filaments. respectively. a feature shared in Malvales only with Malvaceae. The bark slashes of tree Malvalesall tend to have a mucilaginous secretion which can be felt when fresh or seen as globules after afew hours. the Hamamelidae and other three-veined families do not. the latter in their mostly herbaceous habit. Bombacaceae. Simple-leaved Leguminosae (Bocoa.The three genera with 3-v. Tiliaceae. Leaves palmately 3(-9)-veined at base (and alternate and simple) . The only definitive difference from Malvaceae is the absence of spinulose pollen. except Guazuma which has leaves more jaggedly serrate than in any Tiliaceae. Sterculiaceae) or Hamamelidae (especially Ulmaceae. .The majority of taxa with palmate basal ve ins (he re refe rred to as "3-veined") belong to one of two quite unrelated main groups: Malvales (Malvaceae. Lecointea. a feature never found in Bombacaceae. Shrub sterculs (Le.The relatively few simple-leaved genera. entire-leaved tiliacs (except a few genera rare in our area) have the lower leaf surface canescent. though shorter. appressed-stellate indumentum. Bombacaceae . the fomer distinctive in their !oliaceous stipules. Key IV 43 Tiliaceae .42 Key to the Families strongly turpentiny odor. and black-drying color. a character combination not found in simple-leaved bombacs and only in a few Theobroma species in Sterculiaceae (from which entire-leaved Apeiba species can be differentiated by longer more slender petioles). The Malvalean woody taxa have petioles with a distinctive swollen apical pulvinus. Malvaceae are mostly herbs and subshrubs. by stellate (or lepidote) trichomes and the velY distinctive petiole apex which is more or less swollen and pulvinar. as do the few simple-leaved species of Burseraceae. with the woody species in our area serrate -ma rgme. ---------------------------IVAd. as an order also characterized by strong barkjibers. Araliaceae have aromatic leaves and some are reminiscent of some Lauraceae. Tiliaceae. the former differing from Malvaceae and most Bombacaceae in having 2-celled anthers. Sterculiaceae . Elaeocarpaceae. reddish. It is also characterized by scattered. Bombacaceae are all trees and the simple-leaved ones we entire (velY weakly sublobed in Ochroma). exclusively large trees. Urticaceae). Sterculiaceae.eined leav~s are ~is­ tinctive in the order in persistent foliaceous stIpules and III lacklllg the typical Malvalean pulvinus. they differ prominently in their varying petiole lengths. Only EiaeoCal1JaCeae and most Malvaceae lack the typical pulvinus. although the stamen tube often differs from Mulvaceae in being fused only at basco Malvaceae _ Essentially the herbaceous counterpart of Bombacaceae with which they share the distinctive feature of fused filaments. Bixa IS nO. The latter also have prominently flexed petiole apices indicating their compound-leaved affinities. IVAd(I). most Malvalean shrubs) have serrate leaves (see Tiliaceae above for distinguishing characters). Bixaceae _ Bixa closer to Flacourtiaceae than Malvales. some Swartzia) have the distinctive green-bean odor typical of their family as well as a petiole that assumes the round pulvinate-cylindrical form typical of the pulvinuli of compound-leaved legumes. separation of the individual Malvalean families without flowers l~ frequently problematic. peltate scales below (but lacks the typical Malvalean stellate trichomes). d T'I' lzaceae ( mostly serrate) and Sterculiaceae (trees mostly entire except Guazuma) include both large trees and small weedy shrubs. which differ in being mostly herbs and shrubs. but shorter. Alternate-leaved weedy Compositae are mostly not strongly aromatic. Flowers characterized by multiple stamens arranged in single whorl and with free filaments. differ from stercul shrubs by more crenate marginal serrations and a tendency to 3-lobed leaves. Shrub genera CO/'chorus and Triumfetta.. The flowers can have fused or distinct filaments. has a distinct apical pulvi~us similar to that of the Malvales. and with some Sterculiaceae. Dendrobangia (Icacinaceae) has a more medicinal odor than typical of Ranalean families and is also characterized by a grooved petiole.Most serrate Malvalean trees are Tiliaceae (see also sterculiaceous Guazuma). The most definitive difference is spinulose pollen. and slmpl~-leaved Bombacaceae are reliably differentiated only by jirst knowm~ the ?enera.Tree genera differ from mosttiliacs in being entireleaved (or palmately lobed or compound). Elaeocarpaceae .

Neosprucea and Lunania have the leaves more strongly 3-veined to near apex than do most other taxa with alternate3-veined leaves. but the commonest erect species has entire leaves recognizable by the noticeably asymmetric base that characterizes most Ulmaceae. entire leaf margins. ~ernandiaceae . in tree taxa always serrate. Strong bark .ExtraJimital Curupira has 3-veined leaves. !runk with swollen nodes. brachycalyx).wrinkled usually tannish-drying twig bark and main lateral vein pair straight or curving slightly outward rather than upward. B. ---------------------------IVAe. but in our area only when opposite. differing from elaeocarps with similar stipules by the COmblnallOn of longer slender petioles and serrate margin. a combination unique among area trees (although also In some extralimital Oreopanax). Begoniaceae . the common Prockia has distinctive semicircular foliaceou~ stipules. stro~g. Euphorbiaceae . the common Trema has asperous leaves with fine close-together teeth. Buxaceae . Flacourtiaceae . Leaves nearly always With. (Menispermaceae) . the main lateral vein pair curving upward unlike similar araliacs. leaves.Palmately lobed with serra~e margins. Some milky latex usually present.. drying olive. see above) or at base of lamina (usually in axils of basal vein pair below) or have stellate or peltate trichomes (Croton) or are deeply palmately lobed.Conspicuously 3-veined euphorbs mostly have glands at apex of petiole (sometimes also with latex. Urticaceae . shallowly Jaggedly serrate.. cinnamomum lacks spines but has oblong leaves 3-veined all the way to apex.Most taxa with pinnate venation. ~o. Three-veined species of Dendropanax differ from nonpeltate Hernandia in +/. and this is a very useful character for several families. Petioles lacking apical pulvinus Ulmaceae . (Caricaceae) - Key IV 45 subpeltate (many Hemandia).g. can be identified b~ the. Faint but characteristic "huasca" odor. (Olacaceae) . Lecythidaceae _ Differs from other strong-barked families in bark of trunk peeling off in layers rather than as single unit. coriaceous with smooth surface. sub-3veined from above base. mostly simple. twig bark.Zizyphus has conspicuously 3-veined leaves. and vertical flber hnes In a very shallow bark slash. Cochlospermaceae (Cochlospermum) . Araliaceae .(Pull off a leaf and see if a long strip of bark comes off with it.st like Araliaceae but usually either with two main lateraJ veInS ariSing slightly below base of lamina (Gyrocarpus) or the base (Leguminosae [Bau/Zillia]) .44 Key to the Families IVAd(2).g. if odor not apparent.A few species of Bauhinia have the two leaflets completely fused (e. o:ten greenish.Styloceras. Thymelaeaceae) otherwise nondescript. Celtis is often spiny and has leaves with coarse rather irregular teeth. usuaJly rank-smelling (but the odor not clearly Ranale~n).Begonia parviflora is our only erect woody Begonia. but Trerna and Celtis have 3-veined alternate leaves. some of them (e. Hamam~lidaceae . the petioles always of equal lengths. serrate or serrulate margins and distinctive secondary \and usually Intersecondary) veins that turn up and fade out at margIns. Rhizophoraceae .Leaves long-petioled and entire (in part 3-lobed In Gyrocarpus). (Annonaceae) _ Keyed out above under plants with primitive odors.Characterized by leaves with rank odor and of dramatically different sizes and with petioles of different length.Close to Ulmaceae but leaves usually with cystoliths in upper surface andlor with stinging hairs. I . leaves large.Two of the commonest 3-veined flacourt genera h~ve ~ v~ry characteristic pair of glands at petiole apex (euphorbs With slmllar glands differ in having latex or leaves larger and more broadly ovate). the species mostly in dry areas where also characterized by spines. very asymmetric. moistforest Z. Rhamnaceae . Some Colubrina have 3-veined leaves.A few Abuta species are trees with 3-veined leaves and longish petioles with wiry flexed apex. if 3-veined from exact base.Our only genus has distinctively oblong-ovate lea~es wlth somewhat asymmetric base and a short dorsally grooved pellole. Oreopanax usually either epiphytic or with leaves palmately lobed and conspicuously tannish-pubescent below.Anisophyllea of Amazonian sandy soil areas has distinctive oblong 3-5-plinerved leaves.) All neotropical species with strong bark fibers have altemate.

Characteristic green outer layer in bark slash. (Leguminosae [Bauhiniaj) . unlike Clethra or Curatella.ac h~ve alterna..~. unlike Sabiaceae.Distinctive in the densely tannish-stellate tomentum of the leaf undersurface.~e serrate leaves. (Mal vales and Urticales) . Petiole glands Chrysobalanaceae . they can usually be recognized by the red gritty-textured mner bark and/or small inconspicuous stipules on young twigs. IVAh. petiole base not enlarged.Maytelll~.Some Chrysobalanaceae species have a pair of lateral glands at extreme apex of petiole or at extreme base of leaf ?lade below. Clethraceae . and Verbesina (typically d~eply pmnatel. very coriaceous. (S~erculiaceae [Sterculia]) . Gavarrettia). "'e"saria no us conaceou "" narrow. ----- ------------------------ IVAg.Numerous straight parallel secondary veins. Aquifoliaceae . Celastraceae . stellate-pubescent.Usually conspicuously coriaceous with faint blackish tracing of tertiary venation and/or blackish dots below on light green undersurface.The Malvalean and Urticalean families. . Didymocistus. keyed out above on account of 3-veined leaves.Some Colubrina species have pair of large glands at extreme base of lamina. Euphorbiaceae . but lowland species often entire.All taxa with pair of glands near petiole apex have latex and/or are conspicuously 3-veined (see above). the entire trunk bark peeling off when pulled (as opposed to peeling in layers in Lecythidaceae). usually rather remotely serrate or serrulate. . sometimes only toward apex. nonserrate taxa with unequal petioles also have the flexed petiole apex (Nealchornea. . Dodecastigma. also charactenzed by leaves clustered at tips of ascending short-shoot branchlets or branch tips. Serrate (or serrulate) margins Actinidiaceae .Very distinctive in the thick homogeneous bark that strips as a unit from entire twig. s leaves . typicalJy in basal auricles. gray leaves).The combination of serrate leaf margins with conspicuously different-length petioles having t1exed apices is definitive for Euphorbiaceae. 47 (Euphorbiaceae) . and thus. Capparidaceae .The flacourt genera with glands at apex of petiole have conspicuously 3-veined leaves (see above). Cmyodendron. leaves more oblong and/or petioles more wiry than in other taxa with unequal petioles.Although most tree Boraginaceae have en~ire leaves. (Flacourtiaceae) . Dilleniaceae (Curatella) Leaves asperous. Sagotia.Doubly toothed leaf margin (with teeth over secondary vein endings slightly larger). the only family with thick strong nonlaycred homogeneous bark. restricted to open savannas.Rather few arborescent Composit. trichomes simple. ---------------------------IVA/. Boraginaceae .Most tree combretacs (except most Terminalia) ha~e a dlstmcl!ve pair of glands on upper petiole surface. Sene/eldera. the main lowland genus. usually has conaceous olive-drymg leaves. serrate. in effect at petiole apex. (Rhamnaceae) . Unequal petioles AraIiaceae - Leaves with rank vegetative aroma. Betulaceae . Rhamnaceae) and a few arborescent Cordia and Toume/ortw species have serrate or serrulate margins.'I 46 Key to the Families Key IV 1 I Thymelaeaceae . Compositae .Twig usually irregularly angled from decurrent petiole base and often zigzag and/or gre~nish whe~ fresh . Saccellium (with ascending close-together secondary ve1l1~.Although Sterculia petioles are consPIcuously unequal. surface frequently rough-pubescent.Erect Bauhinia brachycalyx with completely fused leaflets has surprisingly strong Lecythidaceae-like bark. Garcia.~ lobed) being the most frequently e?countered. the genus is keyed out above on account of the Malvalean pulvinus. Combr~ta~ea~ . LIke most othct comps these can be recognized by thelf rather pungent aroma.Petioles unequal only when leaves terminally clustered. Baccharis (usually shrubby and chmactenzed by rest(with shallowly remotely . Pogonophora. cr. are also characterized by strong bark fibers.

. Sabiaceae .. . like other members of the family they are characterized by the typical. High Andes only. Vantanea) have crenate . and C"Itlonea and usually with conspiCUOUS axil!' d . strongly ascending secondary veins. (Myrsinaceae) . I III.Unique leaves strongly yellow gland-dotted. fairly straight. an perpendicular to midvein. exclusively montan~. twigs strongly ridged from decurrent petiole base.M b lion parallel d e?l ranaceous to chartaceous. .Rhamnus has leaves with pinnate venation and finely crenate-serrulate on margins. but more coriaceous and the petiole base thickened and often woody. Similar to Saul'Ouia. ~argms and festooned-brochidodromous venation Young leaves at Soot apex rolled into narrow cone. 'I. the leaves are less festooned-brochidodromous than Symplocos. typically with spic. and several' shrubs (Aeldoton. rigidly parallel.Many serrate(-serrulate)-Ieaved pmnate-vemed . have groove on top of the often someh' w(blaatktwdlst~d petiole. round white lenticels. Tertiary venaserrulate. Fagaceae . more or less resinous leaves with a charactenslic undersurface from the immersed tertiary venation. nose teeth ( f II ary om alia. base of lam'ma from gIandular basal teeth. with more strongly ascending secondary veins.marginally toothed. Etaballia has entl're m .r dark red. i .The v . 'I Ochnaceae . secondary veins. Kageneekia and QlIillaja have resinous leaves. Sebastiana). Xylosma lacks pUnctatlOns but is frequently spiny. lcacinaceae . Rosaceae . always +/.Ne I account of their stron : y all Lecythidaceae (keyed out above on g ark) have serrate or serrulate margins. Inconspicuously serrate or serrulate cuneate to short petiole. Serrate myrsinacs can be differentiated from similm'ly punctate Theaceae (Ternstroemia) by the more elongate punctations that are pellucid in bud. secondary veins and finely parallel tertiary veins perpendicular to secondaries. mostly oblanceolate and +/. usually non pellucid punctations (see below). genera are generall h all1sh~ngly few.. ' Flacourtiaceae . inner bark red . like Escallonia but narrower. . fl acourts are characterized by very small pellucid punctations. Slightly zigzag twigs are another frequent character. In theory they should have stipules but these are usually not very obvious.Most genera (except entire . c . ) gms Lecointea) (related argms. they are recognizable by the flexed petiole apex and tendency to have both opposite and alternate leaves . usually with caducous stipules leaving annular scar. Lacistemataceae . Saxifragaceae _ Escallonia. finely serrate or remotely (Lecythidaceae) . Banara laCks pun ctat'IOns but has a conspicuous marginal gland pair near . or with close. tapering gradually at base to a less-defined petiole. usually with numerous. Only two genera with serrate mm'''ins: Calatola If (olive-drymg ' . ~~Phorbiaceae -. Only a few arborescent euphorbs have pinnately emed leaves with eglandular equal petioles lack latex and have se rra te margms.Always serrate or serrulate. coriaceous co . Humiriaceae . when serrate. " These very nondescript taxa include Cleidion ('<Alchornea?).ii II! 48 . Rhamnaceae . Adenophaedra. or at least serrulate margins. ..Some Sioanea species have remotely serrate or serrulate margms. Rosaceae trunks lack the green inner bark layer of !lex.coriaceous. I Key to the Families (Elaeocarpac~ae) .The majority of Andean trees with alternate coriaceo us leaves and serrate margins are probably Rosaceae. Richeria (the margin only slightly crenulate leaves cun~ate and petiole base slightly enlarged). trichomes simple unlike Clethra or Cliratella. sti?ul es ~e always present but usually early caducous and leaving mconsplCuous scar.rymg) .Most Meliosma species have conspicuously serrate.A very few mostly shrubby myrsinacs have finely serrate leaf margins. it is recognizable by the typical Rhamnaceae leaf venation with rather close-together.one with secondary veins marginally curved and becoming almost submarginal (with several of these marginal extensions paralleling each other at a given point). Key IV 49 Myricaceae . the petiole lacks the Sapotaceae-Iike woody thickened base of Meliosma. truly simple-leaved legume metric bases and sYec aractenzed by olive-drying leaves with asymrru late mar' (Zoll·erl1la. ex opaea). . occasionally Clusia-type venation (Blastomant!Jus).Characterized by clustered tenninal buds with scales. young growth densely lepidote-glandular and macroscopically yellowish or tannish.has finely ~er: rate. straight. most Prunus leaves have distinctive large ocellate glands near base of lamina below.Most species . three leaf types . Leguminosae .

Punctatlons ~~uh y blackish' associated with disusually elongate. straight..Leaves of many SolanuIn species distinctively irregularly.entire marg!!1s. Species with small coriaceo us leaves could be confused with some Ternstroemia (Theaceae) but lack black punctations. nearly always + _ unlike Flacourtiaceae..So:"e pun~tat~ .Usually recognizable by the highly u?u~ua~ ml~ture of OPposite and alternate leaves..Clavija. secondary veins and MaJvalean pulvinus.a that. unUSually thin buttresses' a few large-leaved species have distinctive leafy stipules.. Theophrastaceae . Temstroemia (usually only Inconspicuously serrate near apex) has well-developed petiole but is distinctively punctate with blackish glands.. at least inconspicuously near apex. Unifoliolate Legunl!nosa~ an have CylfndricallY thickening at petiole apices but are more clearly unifoliolate..--IV Aj. typically with strongly spiny-serrate margins.. to be shorter andlor less variable in length than the above eup 01 s.. Violaceae . tl nonaromatic) genera have simple Rutaceae . these Theaceae . Many gen~~a .50 Key to the Families (Solanaceae) .. Thicke1led a1ld/or flexed petiole apices Diptercarp~ceae - Our only species recognizable by the broadly Ovate leaf with close-together.Characterized by festooned-brochidodromous venation. Myrsinaceae ..Most cloud-forest taxa (Gordonia. always has narrowly obovate to oblanceolate leaves. Banara. therr peuole~ t~.~~~t~tions. broadly.. the surface between the secondary veins usually rather smooth. Tnchomes. Key IV 51 (Euphorbiaceae) . J .. the leaves usually loosely and rather irregularly reticulate below with nonprominulous venation.. lei ave: t~at . and shallowly toothed. when not obviously serratemargined the margin usually distinctively cartilaginous or the plant reduced to a small erect subshrub. but ~ " J'ke Myrsinaceac Flacourtiaceae - ~~. which are differ from Myrsmaceae m aVI blackish even in juvenile leaves and buds. Elaeocarp~ceae (Sloal/ea) . Ternstroemia) have more or less serrate leaves (although this can vary even within a species). do not ~IWayS ~ave conspicuously different-lengthed petlolcs !!1clude Cmyodendron and Sagotia.rity of'their species serratemost lmear punctatlons. tax. all nonpellucid (except in bud). Trichilia acuminata has unifoliolate.. secondary venation often immersed and nonapparent." flexed a ices.. consisting mostly of pachycaul treelelS. .~n:~ou~~:~~Yi::vS::~~~u~:~ko~~~~:.I I 'i >! . Leaves characteristically markedly asymmetric at least basally.Only a few (mos y I obovate to oblanceolatc with leaves... d Lepidobotyraceae . Symplocaceae . when present. unlike other punctate taxa.. ---------------------------IVAi. Freziera can have long petioles but is easy to recognize by the unusually numerous nearly parallel secondary (and intersecondW:Y) veins and dorsally grooved petiole. b simple leaves with short petlO es avmg appear when fresh to e . ' . CO/~otroc~e~ have flexed petiole apices and +1.ng rounder punctations.d unctat~ sometimes with alXylosn~a. (Meliaceae) - Theaceae . Theaceae have entire leaves. I tinctivc paIe green . Stipuies completely absent "matte" undersur ace.Many euphorbs have flexed petiole apice~ but most are 3-veined and/or serrate and/or have latex andlor petlOlar glands A few entire-margined nonlactiferous euphorbs that lack etiola~ lands have thickened or flexed petioles.. Freziera. most species also dlStl?Ctlve m the large.----.Very nondescript and often impossible to differentiate from Flacourtiaceae vegetatively. Gloeospermum leaves often dry light green with a paler central area. PUllctatiolls (Casearia Homalium. Stipules present but usually caducous. and often red IS °fr . typically coriaceous and oblanceolately tapering to sessile or subsessile base. Flacourtiaceae-A few fiacourlS (Lindackeria: Mayn~.....~~~:. usually also with stellate or dendroid trichomes and/or prickles.. Symplococarpon.---. Vegetatively very similar to /lex but lacks a green inner bark layer. these almost always narrow apices or at apex of pachycaul cuneate bases and clustered at branc U·eelet..---.. leaves usually membranaceous.. Neoptych~cmpus) Pt~e u~a~.------ ---- . branched.

leaf undersur ace.Most tree species of Dichapetalum have conspicuous stipules. IVAI. S _ Characterized by densely white-stellate or Jepid?!C tyracaceae f II also rufescent with reddish-stellate hUlrs. hut Rirtella and some Licania species are very nondescript.Several entire-margined nondescript euphorb genera hav~ distinc~ive +/. Chrysobalanaceae . Dichapetalaceae (Dichapetalum) .Leaves usually subentire with famt tendency to m~ginal serrulation. a characteristic patelliform gland just above the leaf axil on young twigs is frequently apparent. laxer arms on twig trichomes. usua Y i . Stipules very conspicuous in juveniles.A few arborescent Compositae have stellate trichomes.Usually serrate or serrulate hut occasionally subentire. Leaves characteristically membra- Fagaceae _ Margins usually +/. stipule scars are about thelf only useful Character. the family IS u~lIallY recognizable by the rank toma t0. and in lacking the scattered rufescent trichomes of the more strongly reticulate leaf undersurface. and with strongly ascending lateral veins and finely prominulous parallel tertiary venation. . both with +/. '. Euphorbiaceae . blackishdrying. but most species +/.The stipules are typically small and inconspicuous and are usually visible only on young twigs. Euphorbiaceae . (Violaceae) . ~osace~e . Lepidote scales and/or stellate trichomes (Annonaceae) . lcacinaceae (Dendroballgia) naceous and black-drying. often longitudinally striate. ~rythroxylaceae (Erythroxyloll) - When present. sometimes with a very unusual serrate or fimbriate margin. Sagotia with moraclike terminal stipule falling to leavc conspicuous scar).'j I' " (Compositae) . characterized by stipllie cadu~ous. The whole family is usually recognizable by having red inner bark with a gritty-sandy texture.lI'ke odor of crushed leaves.sen'ulate. L~cistemataceae (Lacistema) .Pinnate-veined euphorbs c. Margaritaria with reddish sl.Stellate-pubescent but keyed out above on account of the serrate margins. sometimes in part with malpighiaceous trichomes).A few nond I escnpt IOlaceae with entire or subentue a:~~ are characterized by very inconspicuous stipules (see Leonia aypayrola below).caducous stipules (e. Stipules Celastraceae (Goupia) .lghtly zigzag puberulous twigs and conspicuous stipule scar. Solanaceae _ Several genera (especially many Sola~7U/1~ species) have stellate to variously dendroid trichomes. some Croton (also Gavarettia with petiole glands and the dry-area shrubs Chiropetalum and Argythamnia. stipules triangular and browniSh or tannish. Pera.serrate or serrulate (see above).Duguetia and some Annona species have stellate trichomes or lepidote scales but should key out above under primitive odor. m par.Distinctive in the densely white-stellate leaf undersurface. sl1pules. with both stipule and young twigs endmg to dry blackish. rigidly parallel. faint venation h?es paralleling the midvein below are very typical. and many Solanum species are spiny. Clethraceae . (Malvales) _ Most Malvales have stellate trichomes or scales but are keyed out above by the pulvinar petiole apex.E~tire-Ieaved Prunus species have early-caducous inconsPIcuous.t 2-branchcd trichomes) a useful indicator. " V' . Chaetoc G11)US with sll. only. Its margin entire differs from area Styrax in longer.bperslstent thick-foliaceous stipllle. contrasting with the whitish stipule scar.sericeous leaves. secondary veins and a whitish undersurface.haracterized by lepidote scales and/or stellate trichomes include Hle1'onyma. h' stellate trichomes (Tngonobalanus only Wit .. Many Chrysobalanaceae have very characteristic leaves with close-together. (Dilleniaceae [Curatella]) . :I I I I I Capparidaceae (Capparis) . they are usually recognizable by the pair of large dark-drymg glandular ocelli near base of lamina below.A very few nondescript Casearia spe. to leave consp~cuous scar.52 Key to the Families Key IV 53 iI I IVAk.clCs are both entire and lack punctations. usually distinctive in a blackish inner bark layer.g. when entire the mostly .Conspicuous tannish scales in many species including some that have uniform petioles. leaves characteristically asymmetric-hased. (Fla~oul'tiaceae [Casearia]) .

. DistinctIve m the leaf blade decurrent on the poorly differentiated petiole and the few often poorly defined secondary veins. Key IV 55 Icacinaceae .54 Key to the Families especially on twigs. Ochnaceae (except Ouratea) .Leaves very coriaceous. (Palmae) - A few understory palms have simple leaves.. (Theophrastaceae) .Whether to recognize Guayana Shield genera Cyrillopsis and Ochthocosmus as members of this African Linaceae segregate is debatable.... most other area genera have the parallel tertiary veins perpendicular to the secondary veins.Strong bark (the only combination of strong bark and parallel tertiary venation).-----------------IVAn. characterized by petiole base expanded and subclasping. margins usually crenate (at least when Clusiavenation lacking). They have latex (and are keyed out above).Moist-forest species have the tertiary v.The tertiary veins are conspicuously finely parallel and perpendicular to the midvein and the primitive odor is not always apparent. but most genera have grooved petioles and the tertiary veins conspicuously finely parallel and arranged perpendicular to the midvein. but the latex sometimes is not very conspicuous.-------------- i ! . in addition Pouraqueiba and Emmotum are distinctively sericeous below and Discophora has a characteristic smooth "matte" undersurface. Brochidodromous genera lacking the family's typical upcurved veins and serrate margins have tertiary veins closely parallel and perpendicular to midvein. similar densely white-below Solanaceae lack the rufous-stellate twig pubescence and are usually spiny and/or with slightly lobed leaf margins and/or have asymmetric leaf bases. always distinctive in the sharply spinose leaf apex (see below).A rather nondescript family. (Sapotaceae) .Scaevola is a beach shrub with very succulent leaves with invisible secondary veins. usually finely serrate to serrulate... (Myristicaceae [Compsoneura]) .Most monocots have parallel-veined leaves. Leaves parallel· veined or lacking secondary veins Podocarpaceae . usually there is a slight bit of latex in petiole.. completely lacking secondary veins or with a few faint longitudinal veins paralleling midvein.Caral'pa has th e t venatIOn ertlary +/-. especially during periods of water stress. Theaceae . Olacaceae .m olive-drying leaves and olive-colored fresh twigs. it is recognizable by the terminally clustered spiral leaf arrangement.Many Sapotaceae have conspicuously parallel tertiary venation or Clusia-type venation. ...Most species of Roucheria have Clusia-type venation with the secondaries straight and parallel and well-developed intersecondaries. They are serrate or serrulate and keyed out above.Bamboos have parallel-veined leaves. Lecythidaceae . when entire (some Lacistema) with conspicuous stipule scars. but very few of these are woody enough to include in this key. the plants distinctive in the segmented often hollow stems with characteristic swollen nodes.A few iacquil1ia species have such thickcoriaceous leaves that the secondary veins are invisible.. Linaceae .Tertiary venation perpendicular to midvein. Gramineae .. (Ixonanthaceae) . Lacistemataceae .. Parallel tertiary venation O~i1ia~eae (Agonandra) . (Monocots) . ' " fwely parallel an d Guttiferae .Blastomanthus has Clusia-type venation.nd not very obviously a guttifer' although there is usually a very famt trace of latex.fmely parallel and perpendicular to midvein.ems. ---------.. IVAm. Very Heistel'ial~ke .Most genera have finely parallel tertiary venation +/perpendicular to the midvein or secondary veins.Bonnetia has very faint longitudinal secondary veins paralleling the mid vein. Goodeniaceae . linear-oblong with a strong mid vein. perpendIcular to the secondary veins' it is otherwise very nondescnpt a.

.Leaves clustered on short shoots in spine axils.IVAp.. ~s restricted. thick branches. . (Solanaceae) . mostly only shrubs but ~ccasionally small trees.. (Urticaceae) .Several Rhamnaceae genera have spines. . d) tly have the conaceous common tahuampa specIes I.g. both with milky latex (sec above). Spines or spine-tipped leaves (Annonaceae) . to seasonally dry areas and usually multitrunked. NOlie of the above Amaranthaceae (Pleuf()petalu1Il) .. Twigs distinctively green and strongly angled from decurrent petiole base.. Olacaceae ..Black-drying. Celastraceae (Sellaefferia) .-----. trunk slash is charleaves distinctively black-drying. long-petiolate leaves. Theophrastaceae . often clustered in short-shoots. II. a green layer 111 acteristic of most Aquifoliaceae.Hesperomeles. with senate leaves (see above). usually retuse at apex. narrowly elliptiC. It IS c.n~~~ . (Moraceae) - (Nyctaginaceae) .. I · . spiny meo:bers ~~ the faI~llY have stellate trichomes (and are keyed out above). Ie alternate leaves. Differs from spiny Compositae trees in several spines of different lengths clustered together at each node. Key IV 57 Phytolaccaceae . drying grayish with pale secondary vell1S below. qUI 0 JaC .Castela... membranaceous.. In addlUon usually recognizable by the rank tomato-like odor of crushed leaves.haractenzed by small blackish-drying leaves that are usually at least In part clustered several per node.Branches usually armed with trifurcate spines (unique to family). the latter sometimes with very striking branched spines covering trunk. Rhamnaceae . Simaroubaceae ..Although normal leaves are present.The small thick-based spines that characterize many species of Solanum are actually prickles and may be present ~n leaves as well as on twigs and branches. l11unata mos .. Bignoniaceae . drying olive to blackish.The few spiny-trunked Urticaceae have serrate or incised leaf margins. often spiny... Cactaceae (Pereskia) .Casearia and Xylosma sometimes have branchspines. Berberidaceae . including spiny-trunked Hum (with latex) and Adelia (with inconspicuously spine-tipped branches).56 Key to the Families IVAo. densely branched.Crescentia and Amphiteclla arect~tallYt~n~. leaves small with strongly ascending very inconspicuous secondary venation. pe .Most Mutisieae trees in our area have spines on the branches or in leafaxils or spinose leaf apices or teeth. fascicles alternaling along I bovate leaves 111 charactens IC narrow y o ... these in· clude a few taxa with nondescript entire leaves but most of these are easy to recognize by the accentuated photosynthetic. Euphorbiaceae . Amphuecna has e ~p d tioles in South America poorly demarcated from woody ase.One Llanos species of Annona has branCh-spines. Spiny Compositae differ from Pereskia in the spines (or at least the primary spines) at each node arranged in equal-length pairs (sometimes also with additional shorter spines). A ·f r eae _ The few entire-leaved lowland /lex speci~s (e. (Rosaceae) . spiny branches. t. Peres· kia is recognizable as Cactaceae by the numerous long clustered spines on the branChes.. lic to obovate coriaceous leaves.. charactenzed by the bitter simaroub taste..The family that can have virtually any characteristic has only a few spiny members. Leaves spine-lipped or spinose-margined and clustered on bracteate short shoots.Ximenia has branch spines and leaves clustered at lateral branch tips. usually has spine-tipped branches. secondary veins more conspicuously raised than in spiny phytolacs and Pisonia. Cornpositae .Pisonia actually has oppOsite leaves but in spiny taxa they are mostly clustered on short shoots and the disposition is not evident. 1 escen I iaceous vegetaUve y 111 s~mp . The only spiny Moraceae in our area are Maclura (Chlorophora) and Poulsenia.Achatocarpus. Flacourtiaceae . always has strongly spiny twigs.Jacquinia has small thick-coriaceous strongly spine-tipped leaves. Boraginaceae (Rochefortia) ...Branch apices spine-tipped..

! '.Groove on top of the often twisted petiole.:1 I!.58 Key to the Families Boraginaceae . CyclolobiulI1. Maprounea. leaves typically with large darkish glands on lower sU:face u. Most tree Cordia species have distinctive node with a leaf arising from each branch dichotomy and held parallel to the dichotomy (Fig. Some (cspecially Vernonieae) have a distinctive blackish layer in inner bark.t~plcal cylindrical pulvinus of the single leaflet formmg ItS apical p.'.f th~se Zollen. usually In part with distinctive scars from fallen petiole-borne mflorescence.A notoriously heterogeneous and difficult to rbccogmze famil~. son:e Swartzw. usually with petiole glands. . Compositae .ll~ and Lecointea are often remotely serrate but Efabal/Ia IS unifonn~ enUre.) .. ~tnd Poecilanthe) are more common and easy to recognIze by. Saccelliul1l has ascending close-together secondary veins. young leaves rolled at shoot apex.Tree composites with alternate entire leaves mostly have the leaves +/. Citronella and Calatola both usually with at least a few serrations. Celastraceae . Alternate-leaved taxa typically with leaves clustered and pagoda branching form or bark very smooth and white. Croizatia. frequently sernlla~e margins. C~rillaceae - Key IV 59 Discocwpus.. I I I . b [ midrib' have broad succulent leaves with abaxial glands at ase 0 . recognizable by the gritty-textured. they (0 nagraceae) . .. these mostly drying olive with paler inconspicuous (May tenus) or strongly arcuate (Gymnosporia) secondary veins. (Myricaceae) _ Usually +/. the latter usually with conspicuous domatia. (Leguminosae) . Combretaceae .u t.While flacourts have stipules. mos.. especially on young growth.Tree dichapetalacs usually have serrate stipules (sometimes almos. (cf. 4).Vantanea has coriaceous.Capparids that lack different-length petioles have the leaves 2-ranked.I " ". . ) A few Passiflora species are arborescent. i ! Capparidaceae ..g. :11 I. Rhamnaceae). they (PassI!loraceae .~ i'. slightly thicker towar ~pex. they are usually inconspicuous and early-caducous.wlth . lack conspicuously parall:l tertiary venation. r IaIY venatIOn). " . When subentire. t' . Flacourtiaceae . Chrysobalanaceae . flS IC tWig apex). a few Casearia species with nonobvious stipules have entire margins and lack punctations. y erect Dilkea is a wandlike shrub with narr?wl obvo~a~e or obi anceolate.r laceae . t' reddish bark are restricted to swampy areas and hav~ eXlO la mg .or grayish-pubescent below.conspicuously whitish.obovale leaves. not milky. Bocoa. . Euph b' o. . I'subarborescent. 1'1 ) ~! . and dry a distinctive light olive-green.A few Ludwigia species are.t 01 acaceae _ Usually characterized by curved d t taxa h'lVC Ebenaceae . (Moraceae) _ Trophis usually lacks conical stipule and the latex is watery. Unifoliolate legumes (e. (Dlchapetalu!n) or uniformly terete tannish-puberulous thickish peti?Ie.sllally scallered along (but removed from) midvein (rarely in PI Ullus-IIke basal pair). Icacinaceae .marginally toothed. U-shaped) green petiole. with poorly developed secondary veins and tertiary venation intricately prominulous above and below. red inner bark. usually with a raised patelliform axillary gland.Leaves and stem often stiff-pubescent and asperous. . blunt-lipped. these are often not very evident. the former is blackdrying. even if stipules not apparent.Although small stipules are present. (Tree species of Fuchsia have opposite leaves. except for Terminalia. . I' 'i'jI 1.Leaves usually apically clustered. Margaritaria (caducouS st\pules and a characte . Tacarcuna. o.. If 'I Leaves narrowly obovatc.Rare simple-leaved legumes have the entire petiole cylindrically pulvinate (cf. usually drying a dark reddish color.Tropical species with trunk slash characterized by black bark ring. pulvinulus of leaflets of compound leaves). and Phyllanthus (sometimes resembling a compound-leaved legume. " slight latex in petiole andlor finely parallel tertiary venation. glabrous. Humiriaceae . They usually have asymmetric bases.ole" . +/. .li) !! II ): j: 1-. Di~hapetalaceae .:. 'II 1. "Left-over" genera include Drypetes (asymmetric ase and promInulous te t' . but the leaf venation is typically moraceous. the ap~cal and basal pulvinular area of the 2-parted ':peti..A few lowland May tenus species and its largerleaved segregate Gymnosporia have entire leaves. usually clustered leaves with promInulous venatIOn. if subentire recognizable by the yellow gland dots.

Smilacaceae .Tree phytolacs have entire.Tendril arising at 90° angle from petiole base. an irregular broken sleeve of stipular tissue that covers the node above petiole base. Rhamnaceae . stems soft and variously (often complexly) anomalous in cross section. They are velY easy to distinguish by •the fiol'l 11 an d posltlon .cOI'1 .f'Rh e OJ amnaceae but apparently not in the few bil':d I · nOll vi . often bifurcate. Bauhll1l~. also recognizable by the nodes distinctively jointed and or swollen and the leaves palmately veined or 3-S-foliolate. but can be recognized to family by the typical Rhamnaceae venation with parallel rather straight strongly ascending secondary veins. often with papery reddish bark and/or pendent stemlike aerial roots.) See Figure 2. tend to have prominulous tertiary venation. this usually terminal on young branch. a/'lSll1g e petl. naceae. also distinctive in palmate-veined (usually palmately lobed) leaves usually with remote teeth on the margin.Colubrina usually has nondescript pinnate-veined enlire leaves. (S~biaceae) . or both.laceae - 61 Passifloraceae .60 Key IV Key to the Families Phytolaccaceae . The leaves are rather Lauraceaelike in appearance. of the tendril .- ------ "I.The main scandent genus has a distinctive tendril coiled in one plane exactly like a butterfly's tongue. soft wood. Cucurbttaceae. Polygonaceae .The whole family very easy to recognize by presence of an ochrea. Vio. The tendrils of Bauhinia are rath~r wood? and hooklike and not strongly twining. 1 !.. stems smooth green and spiny (could be confused only With Dioscorea with usually thicker spines). some species of Solanum also distinctive in the peculiar nodes. . leaves entire. with a st·· b . (Leguminosae) _ Several compound-leaved l~g~lme vi~es have tendrils. parallel.. strongly ascending secondary veins and tertiary veins perpendicular to the secondaries and parallel to each other.Tendril axillary. arising in pair from lower part of petiole in Smllacaceae . has IOcompletely 2-foliolate leaves that are sometimes barely spltt only at extreme apex and rarely n:Jt at all. but may not occur III nonblfid species. thin or somewhat succulent leaves.A few Meliosma species are essentially entire. I. (Bauhillia can have tendllis slJllllar to thos .. nonobvious . also frequently with pair of large glands at base of lamina. also the leaves distinctive in having either striking petiolar glands or large glands in the axils of basal vein pair. in our. and one scan dent tendrillate genus. palmately veined (with the lateral veins continuing into leaf apex: actually a monocot despite the net ve~a­ tion). Gallesia. Rh. axillary ' I mg t asa portIOn and a shOlp medlOn e~d ?r zIgzag below twining part in Alltigol/ulIl (originally tel'nllna~~ng ~nf!0rescence and usually branching)..axillary in Passijloraceae...Tendril arising at 90° angle from petiole base. apparently with very small leaves opposite the regular ones. • al'lSlI1g at 90° angle fir am petLO . often broader than long.Unique tendrils arising in pair from near base of petiole. with gland pair at base of lamina above and/or serrate. Sapotaceae).. one pane I I'Ike a buttelfly's tongue 111 . Solanaceae . . IVB.amnaceae . actually representing modified stipules partially fused to base of petiole. has an axillary tendnl with a zigzag basal part below twining part (originally terminating infloresc:n~e and usually branching [cf.. stems soft and flexible. Paypayrola and the commonest Leonia species have c~naceous leaves with entire or subentire margins.. the commonest large-tree phytolac has a conspicuous garlic odor. Vitaceae . and in many species in their peculiar shapes. Lianas with alternate simple leaves IVBa. compound-leaved Sap~ndaceae]). Rhamed 111 h b I . and have a peculiarly dryish texture when fresh. Tendrils present . Stems of lianas usually woody. I e 1Il . "i . like their more numerous serrate-leaved congeners they are recognizable by the thickened sometimes sub woody petiole base (cf.Some species of Cestrum and Solanum are glabrous ~r have simple trichomes and are very nondescript. ---------------. and few obvious recognizing characters. illS also characterized by the typical ochrea of the family. and are very nondescript. (Polygonaceae) _ Antigonum. . often somewhat irregularly lobed or with an inconspicuous 4-S-armed cross in cross section. the best vegetative character is the rank tomato-like vegetative odor. .eaved species. oppoSite .In our area seven families include taxa with alternate simple leaves and tendrils. '. with large vessels and characteristically differentiated outer layer but no obvious anomalous structure. stlpules.ol III Vltaceae. Cucurbitaceae . leaves with rather straight. are~ only ~n cultivation or seminaturalized.

usually broadly ovate in shape. the branchlelS are longitudinally striate. and easily characterized by the pinnate-veined. In our area three mostly erect genera have a few scandent (usually merely clambering) species.area VIne WIth alternate serrate leaves and spines' the Ie . branchlets and is the 19~anea. ~s o:lOca:p~s i~ nona. I' 62 Key to the Families IVBb. Tendrils absent (and plants scandent with alternate simple leaves). usually long-petiolate. the other taxa all being more or less deeply bifid·leaved. Gramineae .spero~s and can also be subentire.e onl~ area ulmac liana.Celtis . and usually with a pair of glands at apex of petiole or near base of blade. restricted to montane cloud forest.Climbing bamboos are easily recognized by their parallel leaf venation and the distinctive segmented stems with characteristic swollen nodes. the petioles lacking any hint of apical pulvinuIus or flexion. and usually asperous surface. Mostly readily recognizable by the strongly lIrt~­ cating trichomes (see also Tragia with more finely serrate leaves). Note that the categories below are not mutually exclusive. AmaryUidaceae: and . Bark usually dIstInctIve: papery peeling and more or less reddish' stem uniformly t' . although entire-leaved Omphalea has a cloudy-watery sap that sometimes turns pinkish or bluish with oxidation. Cyclanthaceae .g. Pouzalzia.(All veins parallello midvein or with finely parallel secondary and tertiary venation more or less perpendicular to midvein (cf.Except for Dilleniaceae. the other (COIYl1osty!is: the leaves are often barely subserrate) to swamps. (Other Monoeots) . at least when young. Serrate ~eaves (without tendrils) . only Heteropsis is truly woody. Ulmaceae . Most of our euphorb climbers lack milky latex. Omphalea). If not urticating the leaves more or less asperous. has spiny °3n y.scandent but usually only herbaceous vines. Araceae . It looks vegetatively much like Urera but lacks cystoliths and usually has a more broadly ovate sub-3-10bed leaf. Liliaceae (s.Ludovia is the only scandent cyclanth with sim· p~e leaves. Key IV 63 Urticaceae . our chmbers w~th serrate alternate leaves are all habitally atypical genera (or speCIes) of predominantly erect families.Climbing Araceae are mostly parallel-veined (unlike most~y nonscandent Anthurium) with the venation arising +/.) The leaf shape a so dIstInctive WIth strICtly pinnate secondary veins and the leaf b~s~ tY~icalJy narrowly cuneate. others very from alternate to 0pposlte.Characterized by the combination of usually sharply s~ra~e margins (at least when young). Euphorbiaceae . by weakly defined midveins and thin secondary vines all strictly parallel to each other and nearly parallel to the midvein. One of our scandent genera (Anchietea) is restricted to Andean cloud forests in our area. th.Our only scandent genus is Celastrus. Violaceae _ The most distinctive character is the slightly but distinctively raised petiole attachment (leaving raised scar on branchlets). leaves mostly coarsely or doubly serrate and or deepl? toothed or lobed. Celastraceae . and Boehmeria. but a few scand~nt genera of Senecioneae (Pentacalia..perpendlcular to the strong midvein. LIke other cyclanths it is characterized by leaves with long thin petioles having sheathing bases. the others more or less succulent." I.s.. Clusia-venation).The family is mostly erect but we have three genera that are Jianas (but with entire leaves) and two that are +/herbaceous vines (sometimes with urticating hairs) with a few erect genera also having a scandent species or two.I Ii I'.Orchi?aceae all have a few scan dent herbaceous genera or specIes with smctly parallel-veined leaves. (1) Parallel veins . aves are ·veIned.). (Malvaceae) _ The only species I know to become a true liana is Malvaviscus arboreus and even that species is more often erect or subclambering. A f?W genera have opposite leaves. Loasaceae _ Our taxa mostly +/. Dilleniaceae . All euphorb climbers in our area have the leaves 3-veined (rarely 3-foliolate or deeply 3-lobed in Dalechampia and Manihot) at the base. crenate-serrate. ' woody the b l' sec IOn always shOWIng concentric growth rings crossed y pa er rays radiating out from center.Leaves distinctively membranaceous. Pseudogynoxys) mostly wllh .Commelinaceae. Compositae _ Most Compositae are erect and most composite climbers are opposite-leaved and/or entire-leaved. athough young branch apex call be coiling and telldril-like in a few taxa (e. oblong-elliptic leaves and the prominent white lentieels on the twigs. also differing [rom other alternate-leaved climbers in cystoliths on upper surface. lacking the large raised lenticels of Celastrus. (2). mostly restricled to cloud forests: Urera (the scandent species nonurticating).

+/. the lobe-leaved climbers all belong to Solanum and most have spiny prickles on the branchlcts. usually with three ascending main veins and two laterals forming margin of sinus and then branching to send main vein to sinus.Pctiolar glands are most typical of the tendriIIate family ~asslfJoraceae. the only scandent genus in our area.Manihot is the only euphorb genus with deeply palmately 3-5-lobed leaves that is mostly scandent in our area. and Piperaceae(Piper.Sparattanthelium. more or less clambering liana. is a lowland forest liana.A few Aristolochia species have deeply 3-lobed leaves. although most paleotropical annonacs are !ianas the few scandent species of Annonaceae in our area all belong to' mostly erect Annona. distinctive in the inflorescence base fused to the base of subtending leaves with the remnants of this structure forming a blunt. Lauraceae (Ocotea). Manihot differs from Carica horowitzii in having watery sap. has broadly ovate.The only scandent family in our area characterized by peltate leaves. the latter with charactenstic narrow glands at extreme base of lamina. by the pinnate leaf venation (unlike Sparatta:lthel~ul1l) and a typical dark bark which shows tiny interconnectlllg fibers when shallowly slashed. a few species of scandent Dalechampia have deeply lobed leaves but are subwoody weedy vines. Only Cordia spinescens is a serrate-leaved. Key IV 65 (4) Primitive odor .al1la have deeply 3-10bed leaves. and also occur in some similarly tendIillate Cucurbltaceae (especlUlly Fevillea). i' (Boraginaceae) . (Caricaceae) . ?esideS the odor. (Basellaceae) . these with leafy stipules. very shallowly. h . Peperomia). other Basellaceae have entire margins. (Menispermaceae) . nonWllllllg petIOle and not at all lobed or angled margins' a few species of D' . Hernandiaceae . (Solanaceae) . unique in the frequently twining petiole. broadly lobed leaves. are ea~il~ recognized by the combination of Ranalean odor and swollen Jomted nodes. subwoody spine on older stems.One unusual species of Carica (c. Piperaceae _ Liana Piperaceae.64 Key to the Families young stems hollow. ISClP . cucurbit-like leaves with an irregularly serrate margin. . They are characterized. is usually more or less marginally lobed. Euphorhiaceae . they also characterize the nons~rrate scand~nt euphorb genera Omphalea and Plukenetia.Aristolochiaceae. characteIized by the distinctly palmately veined leaves. However. characteIized by the 3-veined entire leaves with no hint of an apical pulvinulus or flexion and by the reflexed petiole bases which tend to become woody and form hooklike climbing organs on older branches.Most Boraginaceae are erect and most of the climbers are entire-leaved species of Tournefortia. palmately veined. like the erect ones. these differing from scandent Annona in smaller more delicate leaves and smoother gray bark. a cloud-forest vine. slender herbaceous vines nearly resuicted to cloud forests and nearly always with roundish. and the petiole with base almost always extended into thickened nodal ridge or decurrent as raised striation on opposite side of branchlet. Aristolochiaceae . A few specJes of Ocotea are more or less scandent.te lea~es differing from Tropaeolum in woodier stem. horowitzii of w~stern Ecuador) is a vine with deeply palmately lobed leaves. can have alternate. these differing from scandent Mal1lhot and Carica in pubescent undersurfaces.The only family in our area with climbers having deeply pinnately lobed leaves. Hernandiaceae (Sparattanthelium) and Piperaceae (Sarcorhachis) each have a scandent genus and there are miscellaneous climbing species in mostly erect genera of Annonaceae (Annona). and Jungia. with distinctive leaves broader than long.Curiously.Besides the vegetative odor. vegetatively very similar to Manihot but with milky latex. In addition Lauraceae (Cassytha).remotely serrate leaves. (3) Deeply lobed alld/or pellate leaves (and lacking tendrils) Tropaeolaceae .Tournonia. the frequent presence of foliaceous stipules. (Aristolochiaceae) . (Annonaceae) .A few species of Cissampelos have slightly ie~ta. Lauraceae _ The only truly scan dent Lauraceae genus in our area is the bizarre leafless achlorophyllous parasitiC vine CasSYI~7Q (a close look-alike for Cuscuta except for the flower). .Only one Ranalean family in our area is primarily scandent . (5) Petiolar or lam ilia-base glallds (and 110 tel1dri~s) .

a monocot.nbifid species do not have the Gouania-Iike butte:fly-to~gue tendnls that characterize most scandent Bauhinia. milky latexed Moraceae and peltateleaved Tropaeolum.Byttneria is our only scandent Malvalean genus. Rhamnaceae .the way to leaf apex and by the base of the welldeveloped peliole which is usually flexed and/or twisted above the extreme base which tends to be rigid and is usually somewhat decurrent o~to the s.The main neotropicalliana family with entire 3-veined alternate leaves (only Telitoxicum has strictly pinnate venation. densely while-tomentose below and usually 3-veined) as does Baccharis trinervis of the Astereae. and usually with a bit of latex in it. milky latex (Mabea eximial. thus easy to recognize by the Malvalean characters of pulvinate petiole apex and usual presence of stellate trichomes. "!.Ampelozizyphus has the leaves entire. Menispermaceae . Plukenetw). most of them have 3-veined entire leaves.Heisteria scandens. Euphorbiaceae _ Although the euphorb Hanas are a mixed bag.Our two true liana genera of Euphorbaceae both have a pair of glands near petiole apex. in Lycoseris. genera lacking the apical thickening have the petiole base conspicuously flexuous and usually also have relatively soft flexuous stems and corky bark.ter­ ized by petiolar or lamina-base glands (Omphaiea.The simple-leaved Mutisieae liana genera mostly have 3-5-veined leaves (in Jungia. A useful ~egetative character for most of the woody lianas is the eontrastmg concentric rings of xylem that are usually strikingly asymmetric and/or flattened with the center of the stem usually far to one side. th 3 .Leaves succulent with pinnate venation. except at nodes. Dioscoreaceae .66 Key to the Families Euphorbiaceae . by the curved greenish petiole slightly thicker tow~d the apex. Sterculiaceae .In addition to the 3-veined Ranalean taxa (Piperaceae.The lea . Stems never becoming >2:S cm . usually also with much weaker pair of nearly marginal veins. coriaceous. Aristolochiaceae. (Olacaceae) . Sometimes Simple-leaved Bauhinia can be recognized by the bifid drip tip of some leaves. 3-5-veined and broader than long. but usually more or less distinctly plinerved at base and +/. best character is the dorsally narrowly grooved petiole. Most are also ~harac. Distinctive in cystoliths in upper leaf surface. I I 'I . like most other species of the family. ! .decurrent onto apically winged petiole. have conspICU. characterized. and might not be recognizable when not fertile.1 I :' " Compositae . . In Omphalea the pair of thick glands are at the petiole apex. The scand~nt species has distinctly sub-3-veined leaves unlike most of its relatives. Ericaceae . Hernandiaceae. strongly 3-nerved to apex. I'll iii il II . Most of the scandent Ericaceae are restricted to cloud forests and most are hemiepiphytic. often with adventitious roots. Most of the truly woody genera have a characteristic wiry or sub woody pulvinal flexion at the petiole apex. ·1 ar Smllax). (Urticaceae) . . However: a few weedy Dalechampia species lack all these characters. Anredera is the only genus with acute to acuminate leaves. apparently the nO. In Plukenetia. Many speclCs ous enlarged tubers.Despite the conspicuously reticulate venation. otherwise only by the cylindrical pulvinus at both base and apex of petiole. usually 2-ranked and borne on a short petiole that Key IV 67 becomes subwoody. usually rounded or obtuse at apex. (Leguminosae) . the glands are at extreme base of the lamina and tend to be narrow or on basal auricles. tertiary venation tends to be finely and prominulously parallel on upper surface (only). often green and spiny (with the spilles thicker than·III simi . and elliptic.dlameter. some species have recurved prickles on the branchlets. see above). . characterized by several lateral veins running evenly and uniformly all . .lender evenly striate branchlet (+/_ hinting at the typical sheathlllg petIOle base of many monocots). Most species also characterized by small but conspicuous stipules. the commonest species also characterized by squarish basal "corners" on the ovate leaf. formicaria. yes are tYPically the most coriaceous of any of e -veined scandent taxa and are generally 3-S-plinerved rather than truly 3-nerved. Basellaceae . although a few other species are rather inconspicuously 3-veined). (6) Palmately 3-S-veilled elltire leaves (and no tendrils) . the only scandent species of its family is a canopy liana. there are a number of nontendrillate families with scandent taxa characterized by otherwise unremarkable 3-veined or palmately veined entire leaves. including the common tahuampa species P.A few scandent Bauhinia species have the two leaflets fused together all the way to the apex. or deep lobing (Manihot).Pouzolzia has a few scan dent species mostly with entire leaves.

phillomega). sometimes has a trace of latex in its petioles.The only scandent species of Olacaceae Heisteria scandens. tendrillate Smilacaceae. not very useful for familial recogmtion. including the strangler figs. have the family's distinctive conical terminal stipule.Seguieria is a canopy Han a with distinctive slightly raised nodes having a pair of recurved spines. h' pe~l. has inconspicuous prickles scattered on I. The only woody Convolvulaceae lianas with latex are the unusually woody species of the Ipomoea group (e. s Wit m a genus. like their erect relatives.68 Key to the Families (7) Latex - While presence or absence of latex is usually an important taxonomic character. Solanaceae _ Many scandent Solanaceae have recur~ed prickles on the branchlets. horowitzii of western Ecuador. (8) Spilles . (Caricaceae) .ts bran. characterizing only part of Convolvulaeeae plus miscellaneous species of other families. Cactaceae . They can be identified by the stellate tnch?mes .One Carica species. Another spiny liana.Celtis iguanea. with only a few cloud-forest species of Burmeistera becoming scandent or hemiepiphytic. Polygalaceae . thick-coriaceous. the stems easy to recognize on account of their smooth green surface. Maclura brasiliensis.Moutabea. our only true Moraceae liana. However Pereskia has normal leaves and a few species are scandent. Cordia spinosa. Pisonia aculeata of the Nyctaginaceae. Moraceae .Combretum decandra and its relatives are atypical for the genus both in the usually alternate leaves and in having straight rather blunt spines on the stem. are unmistakable on account of their succulent photosynthetic stems and lack of leaves. spines Occur in miscellaneous scan dent taxa Key IV 69 Ii1. spinosa and relatives).Many Dioscorea species (see 3-veined taxa above) have irregularly scattered thickish spines.Characterized by alternate leaves and milky latex. lour only because an' d' ' d ' h' tn IVI ual scandent genus or species happens to ave spmes. the common 1. These are distinct from milky latexed ConvOlvulaceae in their more oblong leaves completely lacking a cordate base. these very reminiscent of Seguieria except [or the more numerous longer straight spines on the stem. it has opposite leaves but these tend to be clustered and might be taken for alternate. although it can be recognized by the combination of distinctive Moraceae leaf venation and milky latex. our only scandent Ulmaceae. usually also With sptnes on main veins below. and a few Byttneria and Dioscorea species.g.when larger) in having the remnants of dried tendril bases at their apices. While most Moraceae. (Smilacaceae) . Phytolaccaceae. in alternate simple-leaved climbers it is of relatively little importance. Solanaceae and Polygalaceae the latter . but almost entirely herbaceous.olygalaceae l~a~a genera. Phytolaccaceae .and the leaves with completely pinnate venation..Most scandent Cactaceae. Combretum (altemateleaved C. c maceae. leaves with immersed and hardly visible secondary and tertiary venation. one of the main p. although Omphalea (characterized by the pair of thick glands at the petiole apex) has a cloudy-watery latex. Convolvulaceae . schippii) remain vinelike at maturity. II'"~ . F. might be confused with this group. although a few strangling figs (e.Our only climbing family with essentially pinnately veined leaves having conspicuously cordate bases.The smooth green stems of Smilax (see ten?rillate taxa above) could be confused only with Dioscorea from which the stem spines differ in being small and evenly recurve~ or (. (Olacaceae) . The entire olive-drying leaves are also distinctive. (Euphorbiaceae) . Combretaceae . most of which are slender vines. (Ulmaceae) .ches.. . ' .Most Moraceae species are trees or stranglers. has spines (see "serrate leaves" above). :' I . does not.g.Mabea eximia may be the only scan dent Euphorbiaceae in our area with milky latex. C. Only Ipomoea and its relatives. and often with irregularly remotely dentatelobed margin. decandra and relatives). and thus. Otherwise. ohve-drytng. they are generally variable from species to s 'e . including Passiflora (P. Campanulaceae .While spines are taxonomically useful for recogniz- tng a few genera of climbers. The cll'mbl' ng fami'1'les that may be more or less define d ~ the presence of spines are Cactaceae. It IS distinctive in the rather narrow. is scandent. have latex. (Dioscoreaceae) .

plus about half our species of Co I b e be re:~~yOd~:n~~~:~d:~.embers of the family they can to its rupture to form t~ y the sheathhke ochrea at the nodes. spicuous early caduc' WI c aractenstIc but Inconthe smooth usually °1~shsttPulleS).. Polygalac lianas have anomalous stem cross sections with more or less interrupted concentric rings of secondary xylem. times rather aspcrous Ie :ee VIne genera have pubescent somewith more scalariform.The woody Hanas mostly lack latex and are characterized by anomalous stem cross sections with interrupted rings of secondary xylem.:~1 t Iat ~e stron.Ph uously leticulate ultimate venation and. and tend to be rather nondescript. Many have the tYPiCal composite odor. drying leaves with cons ~c erefla ~o:data With nondescript grayishleast. Solanaceae . .e climbers often have hollow branchlets with fine longitudtnal striatIons. Plumbaginaceae (Plumbago) . typically with elliptic to obovate +/. . Polygonaceae . Juvem e Monstera). than in Diehapetalum. Ericaceae . as acute to acuminate leaves with pinnate Key IV I! 71 I. the broadly ovate roundabove). d ea Ie apetalum climbers have nondeveme leaves (usually 'th h . but Anredera hre or ess plinerved at base (see 3-veined taxa venation. .orm of Maregravia has adventitiOu~ r~~~~s(~rp~esse~lagainst a tree trunk which it climbs by .sericeous below or very coriaceous. d WI conspicuously raise darker.I:'.The taxa belonging to thO t nondescnpt g f' IS mos individual dist~~gU~iS~' climhbers are ysted individually with their .Alternate-leaved scandent composites are mostly pinnately vei. Convolvulaceae .Herbaceous vines restricted to dry forest or cloud forest. but restricted to lowland moist Icacinaceae .Most ar D' h . and mostly occur in cloud forest.Some E . the best vegetative character being ig t-co ored trunk . other climbing genera are mostly +/. opposite-leaved Hi ppocrateaceae). a y With loose fIbrous bark. Polygalaceae _ Leaves eoriaceous. forest).Scandent species of Solanum recognizable by stellate trichomes and frequent presence of prickles on leaves and branchlets. If any..70 Key to the Families (9 ) S~alldellt piallts witll aitefllate simple leaves alld lacking allY 01 t ~e above characters . Prior tive Moraceae-like' ~ ochrea.coriaceous clustered leaves. .hemiepiphytic. t h ' . frequently dark-punctate andlor around branChlet ~ ::nat~on reduced ~r suppressed. and the leaves tend to be grayish-pubescent below. the secondary veins are less strongly ascen'ding and the apex less acute than in Leretia. The leaves of the nonlactiferous genera are more or less recognizable from the unusually long eglandular. either +/. often sericeous petiole and the oblong leaf. usually succulent-~ria aractenzed by hemiepiphytic habit and the with the secondar c~ous leaves.gly raised-reticulate beloW the third is very comm~ L e t~rttary veIns. climbing Solanaceae are mostly nondescript and difficult to recognize although the rather few widely separated secondary veins and sometimes presence of branched trichomes or tomato-like odor can be useful characters.a. but Senecio relatives often do not. S~necio~ea. Vernonieae climbers usually have a thin blackish layer in the inner bark similar to trees of that tribe. they are r:~~~~eae chmbers ~e only slightly or not at all hemiepiphytic usu II . epulvinate. alenana. mg c aracters. these often rolled stipule of Mo p pno: t? expansion and resembling the terminal raceae < overlappin I ' The distinctive'Juvem'1 e . like erect m. Securidaca is distinctive in the sensitive greenisholive twigs that make tendril-like twists (cf. less coriaceous an In Similar Convolvulaceae. but the leaves have shorter petioles. a few T-shaped t . Except for hemiepiphytic tendency. (Pbytolaccaceae) _ Triehostigma octandra is a nondescript liana with membranaceous black-drying leaves. script pinnately . characterized by the blade attenuate onto a poorly differentiated petiole which is more or less expanded and clasping at base. .wood~ Polygonaceae have a distinca few Coccolob s cO?lcal termmal stipule (this especially useful in a pecles where the ochrea is poorly developed).. tipped leaves usually mo I d vInes.Two of the th . (~asellaceae) . Compositae .ned. 3-nerved.the Iower surface. c. Most likely to be confused with Solanaceae on one hand and Icacinaceae on the other.Y strongl~ conaceous and the vines are . usually with prominulous venation (except Moutabea which has secondary and tertiary veins immersed but is distinctive in the inconspicuous prickles on the branchlets).r .on. lack la~ex. \ .e Andean Mun hi beekw . Dichapetalaceae . stem with anomalous broken rings of secondary xylem like Convolvulaceae and polygalaccae.SUCCUlent-leave . Marcgraviaceae Ch .. . at and the petiole shorterflC th omes . similar to Convolvulaceae. round lenticels (f V .

. Gymnosiphon) . Bohvta only) p . usually hollow twigs and espeCially in the long slender petiole with flexion at base.g. B. (Onagraceae) . Balanophoraceae (e. (Dilleniaceae) .g. F. some a/ I reduced to rachises) Zygophyllaceae (Bulnesia retama) . however. Burmannia. from Phytolaccaceae lianas in lacking anomalous stem section. (Olacaceae) . Triuris) Burmanniaceae (C-D. Most likely to be confused with Toumefortia from which it differs in longitudinally finely striate. Voyria) Pyrolaceae (I.Most of the Iiana species o[ Toumefortia are utterly nondescript and especially likely to be confused with Trichostigma and Chamissoa. E. our only liana Olacaceae has an indistinct tendency to 3-veined leaf bases and also usually has a faint trace of latex in the petioles. typically characterized by +/.. Colletia) . H. A few common species have distinctive long darkbrownish trichomes. most of these same species also have nonasperous leaves. K. Apodanthes) (Loranthaceae) . have al temate leaves. 2. Unfortunately.papery reddish bark and/or red petioles when fresh. Thismia. see also FIgure 62) Vines Cuscuta (Convolvulaceae) Cassytha (Lauraceae) Apparently epiphytic Rafflesiaceae (e. L eafl ess ac hI 0 rophyUous parasites and saprophytes (Figure 1) Small terrestrial herbs Triuridaceae (A. The leaves o[ lianas are noncoriaceous but often subsucculent.A few more or less scandent Fuchsia.. Koeberlinia (Koeberliniaceae. Dictyostega. Boraginaceae . and the typical leaf shape with more or less cuneate base and usually numerous straight secondary veins.Heisteria scandens. ' t alimital Prosopis Leguminosae (introduced Spartlul~ Junceum ~:':sonia with leaves kuntzei. Frequently with scattered linear glandular area on mid vein above. L.A few dilleniac species (mostly Doliocarpus) have virtually entire mature leaf margins. differing from Chamissoa in shorter petioles with non flexed bases. Otherwise it can be recognized by the curved greenish petiole slightly thickened toward the apex. they are obviously Dilleniaceae in the reddish.g. and unlike thin-leaved Polygalaceae in having prominulous tertiary venation.Most have leaves and all have chlorophyll but are true parasites nevertheless. COIynaea. SciaphUa. Monotropa) . is vegetatively nondescript with membranacous narrowly ovate leaves. SPECIAL HABIT LISTS AND FIGURES c > 1. G. some extralimital CasslG. the only amaranth climber with alternate leaves. Amaranthaceae . Wullschlaegelza) Gentianaceae (J.Chamissoa. papery outer bark. Orchidaceae (very few) (e..72 Key to the Families but Trichostigma differs from liana members of both in either thinner more membranaceous leaves and/or nonprominulous tertiary venation. Leafless plants with chlorophyllous stems (eX~IU~i~s~ parasites) (sometimes with small early caducous ea e Ephedraceae Cactaceae (except Pereskia) Baccharis trimera (Compositae) Muehlenbeckia pJatyc!ada (Polygonaceae) Rhamnaceae (Scutia.

Monotropa E . Malpighiaceae. Alternate leaves Leguminosae (Vicia. Connaraceae. bipinnate).I . .Thisl1lia H . stem often with latex .tendril opposite petiole base Smilacaceae (e) _ pair of stipule-derived tendrils arising from petiole Antigonul1l (Polygonaceae) (not illustrated) ..Voyria G . Much-coiled variously branching tendril. Entada. Mutisia (Composilae) B. few Cissus (Vitaceae). Forked tendril. I .Burmannia D . Lianas with tendrils are extremely easy to identify to family.Only ten families in our area (plus extralimital but escaped Antigonul1l) include tendriUate lianas. (Figure 2) I. .Wullschlaegelia J . as follows. ! ! I I i I .Triuris K .axillary tendril eucurbitaceae (E) . Securidaca (Polygalaceae) Sensitive petiole . simply pinnate.Dictyostegia F . Compound-leaved A. I' F lem II. Tendril-like sensitive lateral branchlets .Hippocrateaceae. Opposite leaves .Tropaeolaceae Woody hooklike tendrils _ Bauhinia (Leguminosae).74 Figure 1 Special Habit Lists and Figures Parasites and Saprophytes 4 em A 3.Cornvnaea '! .Bignoniaceae (F) b. Tendrillate Lianas . Cobaea (Polemoniaceae).g.Burmannia 1.tendril at right angles to petiole base Vitaceae (I) . Terminal leaflet or rachis apex converted into tendril a. stem without latex Psiguria and few Gurania and Sioll1latra (eucurbitaceae). Simple-leaved Gouania (D) (Rhamnaceae) .Apodanthes L . although several other families have tendril-like climbing mechanisms.GYl1lnosiphon B .I . Inflorescence-derived tendril ariSing from nodes of branch lets a.Sciaphila C .Sapindaceae (8) b.butterfly-tongue tendril coiled in one plane Passifloraceae (H) .inflorescence-derived tendril with coiling apical part arising from zigzag straightsegmented basal part I E 4mm . ! Tendril-like climbing mechanisms G Tendril-like branchlet apex .e. Snychnos (G) (Loganiaceae) 3em [ :i I !'i A . Ol1lphalea (A) (Euphorbiaceae). [ 4em H I 75 I I' I .

Machaerium (M) (Leguminosae) (with red latex).g. With 4 (or multiples of 4) phloem arms Bignoniaceae (e.. the classical "monkey's ladder". some StigmaphyJlon (S) and related genera (Malpighiaceae).Figure 2 76 Special Habit Lists and Figures TendrilIate Lianas 77 4.Omphalea (Euphorbiaceae) B . Parabig- A . D. Doliocarpus (K) (Dilleniaceae) (5) Very large vessels and soft thick inner bark layer Cissus (CC) (Vitaceae) (6) Regular concentric cylinders of successive cambia Odontadenia (A) (Apocynaceae) (with white latex). Phytolaccaceae (e. Machaerium. Salacia) 2cm [ (8) Xylem cylinder dissected but complete a. Irregularly deeply dissected . some Paullinia (AA) (Sapindaceae) (2) Regular structure with stem center hollow andlor with exudate Combretaceae (G.. soft stem and thick bark Aristolochia (B) (Aristolochiaceae) d. Moutabea) (7) Broken concentric cylinders of successive cambia Convolvulaceae (e.g. no obvious growth rings Many families. also in some Solanum). Maripa). Clytostoma. indehiscent-fruited Hippocrateaceae (e.Plukenetia (L) (Euphorbiaceae) (and some Malpighiaceae and Passiflora) c.g.. Menispermaceae (T) (often eccentric).Passifl oraceae G .Sapindaceae C . Y.g. c. C..Bignoniaceae E . Seguieria). Callichlamys.Strychnos (Loganiaceae) nonia (9) Stem strongly flattened Bauhinia (0) (Leguminosae). Coccoloba (Z) (2-parted. also in some other legumes including Dalbergia.Passiflora (W) (some Malpighiaceae) b.Cucurbitaceae II . Combretum) (3) "Islets" of secondary phloem scattered in xylem St/ychnos (Q) (4) Regular growth rings with strong radial rays Hippocratea (1) (wind-dispersed Hippocrateaceae). and some mimosoid climbers. With 6 phloem arms.g. Ampelozizyphus (P) (Rhamnaceae). Dilkea (V) (Passifloraceae).Smilacaceae D .. H.g. Anomalous Hana stem cross sections (Figure 3) (1) Regular structure. Menispermaceae (U) (eccentric rings) . polygalaceae (e. E. Irregularly slightly dissected ..N. X.Gouania (Rhamnaceae) F .

'i 78 Figure 3 Figure 3 Liana Cross Sections 79 Figure 3 Legend i i i ' A .Machaerium K .: " .Hippocratea o .Salacia L .Doliocarpus N .Mendoncia G .Cissus ..Paullillia CC .Gurania H .PauUinia BB .Seguiera AA .Stigmaphyllon T .Bauhinia M .Callichlamys A B c o E F .Maripa I .Di/kea u z v AA w ~ ~ BB x y Z _ Coccoloba cc Y .Clytostoma B .Odontadenia C .Aristolochia D .MOUlabea X .Parabignonia E .Passiflora V .Combretum aWH+tme J .Mascagllia p U _Menispermaceae W .Menispermaceae Q _Strychnos S .Plukenetia P _Apelozizyphus R .

g.g. 1 .Ii :1 B -lryanthera A -Mabea C .g.Clavija D .Buchenavia G . Some common whole-tree branching patterns (Figure 4) (1) Branches more or less in wborls .Buchenavia.. few Bignoniaceae (e. Triangular stem with 3 accessory xylem cylinders Sapindaceae (many Paullinia [BB] and Serjania) b..I II .11' " (4) Whorled branching with trunk elongation from new leaders . lryanthel'a [MyristicaceaeJ) (2) Pagoda-style (or candelabra) brancbing - e. Theobroma [SterculiaceaeJ) (5) Unbranched pacbycaul growth form taceaeJ) ! ! G '. . Terminalia catappa) (3) Dichotomous branching with trunk elongation from new leaders . leaves (or flowers) in fork of bifurcation (C.Theobroma F . Irregular dissection Malpighiaceae (e.Cordia E . Soft stem with combination of radial and peripheral dissection Cucurbitaceae (e. S..I . Cordia [Boraginaceae).(branching sympodial by substitution).. Gurania) b.g. Clavija [TheOphras- :\ . J.!iI 80 81 Figure 4 Special Habit Lists and Figures 'I Tree Habits (10) Intact stem extremely irregularly dissected internally a. leaves 2-ranked (= myristicaceous branching).1 (E. . G. ' (11) Xylem cylinder completely divided into individual subunits a. (B.Telminalia catappa I . Mascagnia). some Mansoa) 5.(branching sympodial by apposition) a.(branching monopodial with rhythmic growth). Branching of lateral branches alternate or lacking (A. First branching of lateral branches bifurcating (D. Mabea [Euphorbiaceae) ) b. Combretaceae (F. Soft stem with multiple small separate irregular cylinder Mendoncia (F) (Acanthaceae) 'I.

Wettinia. . 1 ! . Tovomita) Piperaceae (Piper) Moraceae (Pourouma. Maieta (C). Ficus [stranglers]) Palmae (tribe lriarteae: Socratea. ."i . Cecropia. some Clidemia Chrysobalanaceae: Hirtella (A) Leguminosae: Sclerolobium. 83 Ant Domatia I . I rial1ella) Euphorbiaceae (Micrandra) Alzateaceae (Alzatea) .Cordia nodosa .I :1 I I (2) Genera with occasional stilt-rooted species Chrysobalanaceae (Licania heteromorpha and allies) Elaeocarpaceae (Sloanea aff. 1 .82 . Stilt roots I' I (1) Genera regularly characterized by stilt roots Rhizophoraceae (Rhizophora) Guttiferae (Clusia.Duroia hirsuta D -Acacia E .Hirtella C . • Special Habit Lists and Figures Figure 5 6.Tococa A . latifolia) Melastomataceae (few Miconia?) Annonaceae (Oxandra espintana) I. Ant domatia (Figure 5) . I) (1) On petioles or leaf base 1 1 i Melastomataceae: Tococa (8). i .I (3) Swollen thorns 1 Leguminosae: Acacia (D) 7.Tachigali G . Dictyocaryum. (2) Swollen twigs or nodes B Rubiaceae: Duroia hirsuta (E) Boraginaceae: Cordia nodosa (G) ! . Dystovomita. " Ii I I 2 em [ l'il F B . Tachigali (F) Rubiaceae: Duroia saccifera (not illustrated) 1 . iriartea. Symphonia.Maieta F .

. Scutia*. Lycium *) Scrophulariaceae (Basistemon) Anacardiaceae (Schinus) Geraniaceae (Rhynchotheca) Zygophyllaceae (Porliera) c. Erythrina* few tree Machaerium. Condalia) Sapotaceae (Sideroxylon) Solanaceae (Grabowskia *. U. "Grappling Hook" apices of compound leaves Desmoncus* (2) Trees a. Spiro theca *. Poulsenia *) Nyctaginaceae (Pisonia macranthocOlpa) Olacaceae (Ximenia*) Phytolaccaceae (AchatocOlpus *) Rhamnaceae (Zizyphus*. Chomelia Lantana Rubus * c. Branch-tip spines at ends of twigs or short·shoots Euphorbiaceae (Adelia) Rosaceae (H esperomeles*) Burseraceae (Bul'sera orinocensis) Annonaceae (Annona pUllicifo lia ) Celastraceae (Schaefferia *) Rhamnaceae (Colletia*. few Pochota) Cancaceae (Jacaratia) Solanaceae (Solanum) Apocynaceae (Lacmellea [varies within species]) Rutaceae (Zanthoxylum *) Flacourtiaceae (Xylosma. baccifera) Special Habit Lists and Figures 85 Tree ferns (Cyathea. Trunk spines Compositae (Barnadesia* and relatives) Bo~bacaceae (Ceiba*. Alsophila*) Leguminosae(Acacia*. Randia*. Chorisia*. Spines . Colubrina) Rubiaceae (Chomelia *. Machael'ium *. Caesalpinia *. etc. Piptade11la*. most mimosoid climbers Solanum Moutabea* Smilax* Seguieria* Sagaretia Pisoniaaculeata Ophellantha Byttnel'ia Celtis iguanea Rubiaceae: Uncaria*. Bauhlma. Ralldia *) Santalaceae (Acallthosyris) Simaroubaceae (Castela*) Solanaceae (Solanum *) Ulmaceae (Celtis) Verbenaceae (Dul'anta) (: I:< . benthamii) :rrticaceae (Urera laciniata. Trunk spines Smilax* Dioscorea Desmoncus* (actually on sheathing petiole base) Paullinia(few) Stfychnos panamensis (and few others) Passiflora spinosa Combretum decandrum Leguminosae: Acacia*. Machaerium. few Casearia) Gramineae (Guadua *) Hydrophyllaceae (Hydl'olea) .T I . Dasyphyllum. Chuquiraga. Arnaldoa. AdesmlO.' . Machaerium* Pereskia* (and other climbing Cactaceae) Seguieria* Pisonia b. Piptadenia*.) Moraceae (Maciura.Mostly useful at level of individual species except for the genera indicated with * and the whole family Cactaceae. Piptadenia*.' * Leguminosae (Acacia*. extralimital Gleditsia) Cactaceae* Araceae (Montrichardia) Malvaceae (Wercklea) Euphorbiaceae (Hura) b. Fulcaldea) Cycadaceae (Zamia*) Ferns (Tree ferns: (Cyathea*. PIOSOP1S . (1) Lianas a. . Twig and/or leaf spines Berberidaceae (Berberis) Boraginaceae (Rochejol'tia *) Compositae (Mutisieae: Bal'l1adesia*. the spines strongly branched in X.hecellobu~m. Twig and/or leaf spines Leguminosae: scandent Caesalpinia.< I 1! . Alsophila*) Flacourtiaceae (Xylosma. ~lt. 84 Special Habit Lists and Figures 8.

especially when complete descriptions of leaf size. Platypodium a~aceae-Mmquartia (one form) . Most authors currently combine Nephelea with Alsophila and Trichipteris with Cyathea. Cyathea (indusiate and the veins of the leaf segments free) and Cnemidaria (indusiate and the veins of the leaf segments forming areolae or fused to veins of adjacent segment). and at least one medial pinna are usually necessary. these are not treated here. Grammitis (400/400 spp. One taxonomic circumscription treats the tree ferns with petiole scales uniform in texture throughout (i. and stem characteristics are available. worldwide). Ctenopteris 00/200 spp.Clusia (?) Marcg~aviaceae .1! 86 Special Habit Lists and Figures FERNS AND GYMNOSPERMS :1 9. In addition to the true tree ferns. are slender vines which are actually single straggling leaves with a twining rachis). Other climbing genera. Strongly fenestrated trunks Apocynaceae -Aspidosperma series Nitida Celastraceae-Perrottetia sessilif70ra ~~gUminOSae -. worldwide) and Trichomanes (1401150 spp.Weinmannia (1 sp. epIphytIC. there are a number of climbing ferns some of which are more or less woody. and those with distinctly marginate scales lacking a dark apical seta as Trichipteris (lacking an indusium).Souroubea (?) CunODlaceae . which has dimorphic fertile fronds. Stranglers Mor~ceae-Ficus. 10. frond apex. .'I. It is worth noting that tree ferns can often be identified to species in sterile condition. the petiole base.at base of petiole. and the filmy ferns Hymenophyllum (250/3?0 spp. .e. Elaphoglossum (250/500 spp. Th~ larg~st epiphytic genera are Asplenium (4001650 spp. worldWIde).Spil'otheca Alzateaceae -Alzatea(?) c: > I FEUNS (MOSTLY CYATHEACEAE) Tree ferns are unmistakable in their large much-divided leaves which unfurl from fiddleheads and usually have scales and/or prickles . Although ferns lack secondary growth and therefore are not truly woody. pinnae number. worldWIde). Although easy to recognize as ferns. Coussapoa Guttlferae . since the scales and spines of petiole base and young fiddleheads provide many of the specific recognition characters. The various attempts at generic segregation depend heavily on the nature of the scales of the petiole base and young fiddleheads and on whether the sporangia are borne in covered (= indusiate) sori. with nearly all of the species at one time or another included in a sensu lato Cyathea. Rublaceae -Amaioua. they often have well-developed sclerenchymatous support tissue and tree ferns can reach 20 cm or more in trunk diameter and heights of 10 to 15 meters. worldwide). worldwide).few Pouteria spp.Inga neblinensis. climbing appressed to tree trunks. Polypodium (1401150 spp. those with distinctly marginate scales which have a dark apical seta as Alsophila and Nepheiea. The most important of these is Polybotrya. like Lygodium and Salpichlaena. Although there are also very many epiphytic (as well as terrestrial) fern genera. for such identification.) Bombacaceae . Macrocnemum Sapotaceae . genenc taxonomy of the tree ferns is in a state of flux. 'I. Caesalpinia. worldWIde). without distinctive margins) as Sphaeropteris..

) .i' Lomariopsis (25 spp. seta ?~t distinctive in having a dark central stripe and wide ~~n~astmg whitish margins. However. Differs from Zamia in the leaflets having a distinct midrib.ark a~ical seta.88 1. .) . Salpichlaella (3 spp. differing from the formerin being more cona~eous and having the parallel leaflet veins all eq~al.tcuously thickened bases. usually e. loss of the indusium. also distinctive.) Chigua (2 spp. but the large thick strobili..) .. Most sp~cies rare. in the veins in pinnate groups.) . plus 20 Old World) . are utterly distinctive.Lamina uniformly I-pinnate and with an undivided apex as in Lomariopsis.sentially stemless or with short thick ~tem crowned by ring of pinnately compound leaves. It is also the only tree fern genus with the upper surface of the rachises glabrous and usually has leSS-divided fronds than Cyathea and Alsophila (the lamina only once-pinnate [though with pinnatifid lamina segments».Leaves usually more divided than in other hemiepiphytic climbing ferns. Cllemidaria (27 spp. Dickso/l~a (1 sp. I CLIMBING FERNS 2B.The mat~ genus of tree ferns.Distinctive in having nonmarginatc scales. CVCADACEAE Palmlike understory (one Choc6 species epip.Like Cyathea in marginate scales but the scales with a dark apical seta. the lamina 1-4-pinnate with tapered pinnatifid apex. differs from sometime segregate Trichipteris in havmg an mduslUm or "lid" over the sori and from Cnemidaria in having fre~ ~eins not forming areoles.) .become definitely arborescent.Scales similar to Cyathea in lacking a dark termmal.hytic) gymnosperms poorly represented in our area by a smgle genus. Other distinctive characters are the often dark pinnae rhachises and the lower surface of the leaf lamina ~ighter gree~ than the upper. The leaves are once-pinnate unlike Cyatheaceae and the treelike species all have dimorphic fertile fronds. plus 200 Old World) . when present. also differs in the veins of the leaf segments Jommg to form areoles or free but with basal veins of adjacent segments f~se~ to sinus. (Several other genera occur in Mexico and Cuba. Woody hemiepiphytes with straight rachis. Cyathea (inc!. plus 23 Old World) .) " I i . 89 2A.. The pmnately compound leaves are somewhat similar only to palms. . Cycadaceae 2. the veins close together and nearly parallel. Blechllum (ca.) . Trichipteris cannot be maintained since its d~fmmg character.Differs from Polybotrya in the lamina uniformly I-pinnate and with an undivided apex.. (and a few ferns). uniform in texture throughout. Ferns and Gymnosperms TREE FERNS Sphaeropteris (IS spp. ! ' . Trichipteris) (60 spp. Vegetatively characterized by marginate scales lac~mg a d. the stem is usually short and the plant thus less arborescent than in the other genera. Nephelea) (30 spp. t'Ie eIements. .Differs from Cyathea re Ialives by havmg dense ha"Irs mstead of scales and in margined son. plus 100 Old World) _ Only distantly related to the other tree ferns. climbing appressed to supporting tree Polybotrya (35 spp. Differing from sometime segregate Nephelea m the unfurling young leaves lacking spines and the petiole bases nonspiny or w~th blunt or scale-tipped spines rather than having large black spines tapering to slender apex. where thetr conspicuous cycadlike growth-form makes them one of the most charactens . . especially in moist paramos. some species of this mostly herbaceous genus .) _ Western Colombia. . Lomagramma (1 spp. but differing in the veins areal ate rather than long-parallel. rather than pinnately arranged... plus 20 Old World) . i ! Zamia (30 spp. even when simply pinnate. 50 spp.Differs from Lygodium in having the sori borne in lines along both sides of the costa. Climbing by slender twining leaf rachises LygQdium (7 spp.Differs from Salpichlaena in having the sori borne on modified lobes at the tips of the leaflet segments. has occurred independently in different evolutionary lines.l~ encountered m fertile condition. The pel!~les and rachises are frequently more or less spmy but the spmes ru:e shorter than in pinnate-leaved palms and have more consp. Alsophila (inc!.

Ephedra 2 .Podocarpus l 1.I 2 2 30cm 1 em [ 1.90 Figure 6 Figure 7 Podocarpaceae and Ephedraceae 91 Cycadaceae and Gnetaceae 1 3cm [ 1 2cm [ .Gnetum 2-Zamia .

these small «3 cm long and 3 mm wide).. the beads Immediately adjacent or widely spaced. orangish or reddish when mature. The only possibility for confusion would be that the fe w species . m a few sp eCles . The "fruit" of most species is a naked see . Old World and temperate N. Male inflores?~nce v~.orn~ at the end of a fleshy receptacle that turns red at rnatunty. incl. relatively broad.92 . the small sessile or subsessile ovoid or subglobose strobili borne in axils of the scales. Am. sharp-pointed leaves all in the same plane and with several +/. mostly in dry. ulcumamo. each node with a sheath formed from the fused bases of the opposite or whorled scales. The segmented greenish stems are conspicuously jointed with opposite or verticillate branching.1 Ferns and Gymnosperms El'HEDRACEAE Leafless shrubs or subshrubs (puna).. depending on the species. montanus and allies (with a drupaceo~s fruit an? sometimes segregated as Pruml1opitys). One group (with a drupaceous fruit and sometimes segregated as Decllssocarpus) has very short. inter-Andean valleys. resembling a short string of beads. a few species occasionally reaching sea level as widely scattered individuals in broad-leafed forest (in Amazonia only on white sand). P: palo huayo.. romer6n. the opposite petiole bases are contInUOUS across the more or less thickened node. WIth . opposite. There is nothing in the large opposite leaves of this canopy hana to suggest to the uninitiated that it is a gymnosperm.Mostly very large trees.ol.venation (or with several more or less faint longitudmal vems).or. C: pino hayuelo. "Fruits" oblong-ellipsoid. Menispcrmaceae) and exudes a characteristic jellylike resin when cul. When fertile. with the next segment of the branchlet emerging from the node to leave a conspicuously articulate joint. Imear-oblong leaves with a strong midvein but lacking sec?~dary. uniformly entire leaves are quite like th?se of Salacia (Hippocrateaceae). In fact the very coriaceous. romerillo '. and also dry a similar ohve c. in our area restricted to the Andes.~ distin?tive. usually relatively obtuse apically. and lacking hypodermis in P. but most species now nearly eliminated. thick. Ephedra (40 spp." " • .) GNETACEAE . • receptacIe expands to cover the seed and I i . P: diablo fuerte. easy to recognize by the small. forests.. coriaceous. The best distinguishing character is the conspicu~usly Jomted branchlet. not at all gymnosperm-like. hambre huayo 1'1 PODOCARPACEAE One of the most characteristic tree genera of upland Andean. The stem is also distinctive in having concentric rings in cross section (cf. plus many in Old World) . I Glletum (6 spp. chaquiro.i i! .. the majority of species have spirally arranged leaves. Podocmpaceae 93 Podocarpus (37 spp. formerly often forming nearly pure stands in montane forest. usually larger (and WIth the frUIt naked at end of a fleshy receptacle) in Podocmpus sensu stricto. 3-5 cm long.parallel longitudinal veins. longer strap-shaped leaves somc~~~t ~esemble a few Xylopia species with naITOW leaves and m Istmct sec?ndary venation. plus 24 Old World).. but the leaves of these are mU~\less conaceous. (the segregates DecussocQ/17US and P runmopitys) the <'-. None of the leafless angiosperms in our area has opposite scales.

in Liliaceae and Amaryllidaceae. the strictly parallel-veined strap-shaped leaves borne from an underground bulb from the apex of which is born~ a rather succulent inflorescence stalk which produces a terminal umbel (or reduced to I-few flowers) of often spectacular lilylike flowers. which is mostly scandent. occurs only in dry.aments or from petal outgrowths) is o. the largest Amarylhdaceae genus in our region.Figure 8 MONOCOTS > < 95 \ Agavaceae and Amaryllidaceae AGAVACEAE A family closely related to Liliaceae and Amaryllidaceae from both of which it is differentiated by the stiff. penca 3 ! (Draeael/a) .) . In our area Agavaceae. Our two genera. Incluslon of this family i~ this volume is mandated by Bomarea.Differs from Agave in the openly paniculate inflorescence of singly borne flowers. but their leaves are far broader and more succulent than in any Bromeliaceae. with superior-ovaried flowers borne singly on a loose open inflorescence. usually succulent leaves.) .Crinum ' . fibrous. much better represented in Mexico and the Antilles. the bluish color of the leaves differentiates our species from Furcraea. they could only be confused with terrestrial Bromeliaceae. but very poorly represented in the Neotropics by a single slenderpachycaul Central American species and occurring in our area only In cultivation.Differs from Furcraea in the very typical inflorescence with the flowers densely clustered together into flat-topped bunches whiCh are borne near apex of the giant inflorescence. in addition a corona (which may anse fro?1 fused fil.ften present III Amarylhdaceae but never in Liliaceae. In addition to a few Bomarea species. inter-Andean valleys. Furcraea. respectively. typically with spiny apices or prickles along margin. on account of their respective ovary position. 2 Fureraea (20 spp. mostly lacking an aboveground stem. AMARYLLIDACEAE An entirely herbaceous family.Eueharis 4_Boma rea 5 . with inferior-ovaried flowers borne in characteristic flat-topped clusters aggregated near apex of huge inflorescence. one 2 -Agave 1. Vegetatively.In the Paleotropics speciose and sometimes becoming large trees. The flowers differ from Liliaceae primar~yinhavinginferiorovaries. thick. Vegetatively. The succulent spiny-margined leaves are less bluish than in Agave. have traditionally been placed. 2m Agave (300 spp.Furcraea 3 . C: motua. and Agave.

ac acollectmg veIn ¥enus least obviously an Afata leI. Stenospermation has oblongelliptic leaves with hardly visible. the leaf blade laciniate Philodenlro~l~nat~ly laClnlate (the few similarly Monstera).ea.for a few (a few other genera mayo. 13 '-"n~ . . DieJ!enbachia. th. except ..~~~~. .n many genera occur in dry and/or . Anthul'lu~.c:. but this caduc. unlike Ph.-I:u. is a t10ating aquatic which resem bles a small head of lettuce. so: le.e spat~e is caducous in Syngollium and Xanthosoma.. The leaves of the largest 0 the gene:a IS usually possible. The leaves ~:c~~s have . closely parallel secondary veins.The I . the few 3-~0Iiolate Philodendron species have completely parallel vems and no collecting vein. DieJ!enbachia (with succulent stems and petioles and a submilky.. outside the forest < dopen (usually swampy) lowland areas upland Andean ar~a:.I a~J:ing these. Other terrestrial genera include: Spathiphyllul/1.~~~~~a1~~~~r. top half with male flowers and club-shaped) almost completely enclosed by the broad curved spathe. Ii :1 i 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 . hemiepiphytic Climbers' ommant famIly of herbaceous herbs. the secondary veins are usually closer 10gether and more numerous in Rhodospatha than in Philodendron./'T I"n. mp ete y to reveal conspicuous bright red or red- II Eucharis (10 spp.trial or swamp Jasarum is a sUbmerged) is ~ no~tin¥ aquatic (extralimital genus (Heteropsis) is a s~iu~tlCd~lt~ lInear leaves I). strongly ascending.s outside the actual forest). I Bomarea (150 spp ) A Andean scanden! genus with~e very character!stic cloud-forest and high(usually conspicuously maro d separ~te. Scandent Heteropsis. eCOlogicaUy atypical species of B on y Am~yllJ?aceae. differing from Anthul'ium in closely parallel rather than reticulate leaf venation and a more conspicuously sheathing petiole base (also in the broad open spathe). a few epiphytic species Xanthoso ma (some lU~lum. Two terrestrial genera have common species with spines or prickles: Homalomena peltata (stemless with inconspicuous prickles on petiole bases) and Montrichardia (treelike with spines on trunk). often succulent leave~ ~an usually be recognized by the large +/. Genera that usually grow in swamps include Montrichardia (almost treelike).sheathing bases Th ~r~ on +/. te daffodil-like flower with well-developed ARACEAE Araceae is the pred' . Pistia. petIoles and with Clusiamg vein... Collecting vein and +/_ ge?us. and Mons/era have ~niform spadices like Anthurium. consplcUOUS orange or red tepals tube. The stem is twining th on-. Urospatha (sagittate-triangular leaves with mottled petioles)' and Spathiphyllwn (elliptic leaves with completely parallel veins). has tbe wood~Ihpttc leaves subsessiJe on a~d cleru:1y alternate distichous !ike straight parallel second Sort. one genus. differing from Anthul'illm tn a large.ous... Stenospermation. and the lateral leaflets with strong basal lobes.. Most of the epiphytic genera (except Philodendron and Syngonium) have a uniform-thickness spadix of perfect flowers (in Anthurium this more or less free from the usually rather narrow spathe. lodendron. Syngonillm usually has 5-foliolate leaves. .rhJI"I1"F'H·~'''''t+t. and terrestrial Dracontium. is unmistakable IX SUbtended by a single leaflike Vegetative differenti~tion f . o~ted InsIde) forming a more or less tlaring llict to narrowly elliptic. and Taccarum. Rhodospatha is vegetatively very like Philodendron but has the petiole uniformly grooved or winged above right to base of lamina while Philodendron (except section Pteromischum) has the petiolar wings ending below base of lamina.d~ep basal lobes unlike cOmp?und) in epiphYtic S er. the spathe is broader and more or less enclosing spadix). and Homalomel1a. enclo~­ ing spathe like Philodendron. and have all lateral veins I enus... and the . only Anthurium (and one Philodendron: P. and one Vegetatively thefamil n er IStinctly subwoody climber.. Omarea. caustic. goeldii) can have multisegmenled leaves._sv y par~l~el-veined leaves vary from linear ed capsule that dehisces inco 1 a1t fruit IS a round dark brown or blackish orange seeds.'~~ OCcurs inside lowland tropicaJ forest. When fertile.t (!hough slender) stemsaceae when sterile. which consists of a spathe. the les. Rhodospatha. . Very corona. often foul-smelling sap) having oblong or elliptic parallel-veined leaves withoul basal Jobes. have a marginal Philodendron the othe retllcUlate venatIOn while those of ' rveryargeg' J k . in other genera. Monstera usualJ ary vems and a marginal collectrally occurring holes (uniq~e~~~/re o~long leaves with natutends to be deepl' .succulent petioles with fleshy cylindrical ~ ad~ In orescence. one genus (Pi$ti~ m. to OCCur III lowland tropIcal forest characteristic in its large c~~ In open a.any are ~lso terres. Xanthosoma. other genera have obvious secondary veins. Mons/era and to a 11'<0''" ~.tdangular leaves. persIstent In Philodendron. aroid genera are easy to dlstinguish. - 97 1 1 1 1 1 1 1 1 1 1 1 1 1 1 i1 /1 . Among compound-leaved epIphytIC taxa.. having net venation and strong basal lobes.. Philodendron and Syngonium have a 2-parted spadix (bottom half with female flowers and cylindric.) .:!\ Araceae As!eros!igma. these with net veins and marginal collecting veins like Anthurium but distinctive in milky latex. very like Syngoniul1l except for being terrestrial and Jess frequently compound-leaved.Iy dIVIded (frequently even of PhIlodendron and A tn~omum. whereas. often on +/.

with the fruiting inflorescence curling to split irregularly and reveal the rather separated orange-arilJate seeds. but also including numerous terrestrial speCies.Anthul'iul1l · 7 . mostly epiphytic (but seldom truly Climbing). close-together. P: tams hi o.Slender rather woody climbers with very distinctive subsessile leaves having straight. the dimorphic juvenile leaves small.em . Spathe caducous. has an infructescence-like Philodendron. l~aves usually with holes (or these reaching margin and the leaf deeply laclOlate). Caladium and Chlo/'ospatha are essentially reduced versions of Xanthosoma.!11 _ _ 1_ ~ __ • . often ed1ble. I r l.More or less SUcculent hemiepiphytic climbers. Syngonium and terrestrial Montrichardia and Xanthosoma.A common and very distinctive free-floating lettucelike rosette plant P: huama Usually hemiepiphytic climbers with adventitious roots. __ _ 1 . plus some Anthu!'ium species.Stenospermation II n:··· 2 . Spadix uniform throughout Anthurium (750 spp. P: jerg6n quiro (smallest spp. I 'I n l ) _ .·. Among terrestrial genera. 1 1 1 .Anthurium 3 . spadix uniform.Syngonium 6 ..I 'I I . overlapping and often appressed against tree trunks.Dracontium (with a single huge treeletlike deeply dissected leaf). most epiphytic aroids have the spadix uniform throughout (first five genera). _. 99 Figure 9 Araceae (Floating and Epiphytic) 1 1 1 1 1 1 1 1 I 1 1 ! 1 1 ! 1 FLOATING AQUATIC Pistia (l sp. usualJy bright red or white.lCOtS fruit very different from the individual berries of Anthu!'ium.. also have the pistils fused into a syncarpous fruit. E. I I 2.PhilodenrlrnH 5 . fruits nre individual berries.) . the mature. 2A. parallel secondary veins and a strong marginal collecting vein..~.) I . Spathiphyllum (with elliptic cuneate-based leaves and a broad white leaflike spathe)._ . P: anturio.A very large and variable genus. with inflorescence divided into separate male and female portions. has the spathe entirely enclosing the spadix except during a very brief anthesis. and Urospatha (semiaquatic with triangular-sagittate leaves having well-developed basal lobes). Spadix (at least in flower) and spathe both narrower than in other genera. only three . with net venation and a marginal vein. ! 1. before maturity covered by the fused pistil apices.NH. .) . l-. only Philodendron and Syngonium having a 2-parted spadix.. with parallel venation and no marginal vein. spathe caducous. fruit +/_ syncarpOUs... C. : Gem [ Heteropsis (12 spp. Leaves distinctive in marginal collecting vein (except few species of section Pachyneul'ium) and more or less net venation. EPIPHYTES - lollstera (50 spp.) . . prominuJouS.) .Anthul'lum 1 1 1 1 1 1 1 1 1 1 ..Rhodospatha 4 .. Xanthosoma. Dieffenbachia. have a uniform spadix of perfect flowers.

ble secondary veins very strongly ascending and almost parallel to mldvem. oma P: patiquina 101 1 . pInnately deeply divided leaf borne from an underground tuber ?n a long thick petiole that resembles the trunk of a !reelet. open. Spathiphyllllm (35 spp.) . ROO'fEl).) . Vegetatively very similar to those ~hilodend:on specI. hardly dis~emi. ~o II: .es that have oblong leaves with numerous secondary vems and w1tho~t basal lobes. .Most cOmmonly growing colonially in swampy places. ::' Rhodospatha (25 spp. SWAMP PLANTS) 3A. When fertile.Mostly hemiepiphytic climbers but a r~w species terrestrial.Epiphytic climbers with distinctive elliptic lear having a strong midvein but reduced. TERRESTRIAL (OR ERECT. P: itininga. White or greenish Spathe. Except for being epiphytic. I 2B. SYlIgollium (33 spp. Spadix with thick cylindrical basal part (female flowers) and slender apical part (male flowers) .Xanthosoma 4 . The parallel-reticulate leaf venation is rather intermediate between reticulatcveined Anthurium and strictly parallel-veined Philodendron. emerging from the grOund near base of the petiole and with the spadiX enclosed by a large acuminate marOon spathe is unmistakable.) .Rhodospatha " . mostly distinctive in palmately 3-5-parted leaves with basal lobes on the lateral leaflets. completely parallel rather strongly ascending Secondary and intersecondary veins and marginal vein.) .Dieffenbachia . Spadix homogeneous throughout .Forest-understory plants consisting of a SIngle giant. marginal collecting vein is similar to Anthurium but the succulent leaf texture more like PhilOdendron. Il' :. large.) 10cm r 3.The second largest genus of the family.Philodendron 3 .) .ence.) . I?raCO~ltium (13 spp. Leaves always elliptic with a cuneate base.Hom%nema 6 . . P: huasca bijao Araceae E (Miscellaneous pI'phytic and Terrestrial) Philodendroll (275 spp.Heteropsis 2 . the Infloresc. The flowering inflorescence is similar to Philodendron but the spathe is caducous to reveal a syncarpous fruit. The inflorescence is very characteristic with a rather 'Ieaflike. huamM (large spp. but the petiole winged to near apex (some Philodendron species also with fully winged petiole but these non-oblong-leaved).I' " ::! I I III I I I 100 Monocots Figure 10 I. P: jerg6n sacha . in addition the. easily differentiated vegetatively from Anthurium by the completely pru: al leI secondary and tertiary venation and lack of a marginal collecting VCIn. species with undivided leaves have strong basal lobes. very similar to Xanthos . I"I Stellospermatioll (25 spp.Rather succulent hemiepiphytic climbers.

P: jerg6n sacha Araceae (Terrestrial and Hemiaquatic) 1. Spadix 2·parted-Basal (female) part cylindrical and usually thinner. but the closely appressed fleshy berries not truly syncarpous. ZOcm[ 3B.Chlorospatha E: camacho. Leaves large.) . the commonest species with spines on trunk.) .The main terrestrial aroid genus.1 i 'I' (Allthurium) .Ul'ospatha 6 . occaSionally with soft "trunk" to 2 m high. apical (male) part club-shaped. more open than in Philodendron and caducous immediately after antheslS. P: raya balsa 10cm 4 5 Dieffellbacllia (30 spp.Caladium Ii - Snathiphyllum 5 . many of these are bird's nest forms with short petioles and large clustered oblanceolate leaves.) . cOllectin~ 2 .Dracontium 1 . the frultmg mflorescence curling to split irregularly and reveal the rather separated orange-arilIate seeds. Usually with rather unpleasant pungent odor (so~etimes almost skunkiike) and sometimes with the leaves splotched white ~bove. Fruit (edible) somewhat resembling Syngonium. Terrestrial Anthurium leaves are mostly u~­ divided. Xallthoso (45 spp. P: patiquina 6 50 ] ern llla . broadly triangular-ovate. charactenzed by pal~ately dissected or triangular succulent leaves with strong basal lobes. P: oreja de elefante 3.Montl'ical'dia . TypiCally rather coarse herbs. mostly lack strong basal lobes. with strong basal lobes.entirely encloSing spadix except during a very brief anthesls. Venation uniformly parallel and marginal vein lacking as in Philodendron.Some Anthurium species are terrestrial or grow on rocks. reticulate venation.) .• Monocots 102 103 Figure 11 Urospatha (20 spp.A few Rhodospatha species are terrestrial. and differ from Spathiphyllum 10 reticulate venation and a marginal vein.Stemless swamp herb with strong~y trian?ular leaves with the elongate sagittate basal lobes longer than antenor P?rt:on (unique). S~~the . MOlltric/lardia (2 spp. narrower where it emerges from spathe.SUCCUlent forest-understory herbs with well-developed succulent erect stems and uniformly oblong or elliptic leaves without basal lobes. and usually both marginal and submarginal veins. Spathe strongly constricted at throat and with upper part of spathe. I 1 (Rhodospatha) .Tall almost treelike plants forming dense stands in swampy areas. the venation parallel-reticulate as in Syngonium. except for the inflorescence they look like Philodendron with oblong parallelveined leaves without marginal vein. Inflorescence with a brownish-magenta acuminate spathe similar to Dracontium but with the spathe spirally twisted.

aractel'1Zed by pinnately I a?.ter-containing tank and/or having spiny margms. A few of the terrestrial Puya species of the high Andes have well-developed trunks several meters tall below the rosette of long spinymargined lea ves. Asterostigma (6 understory herbs sp~.Rare Ama . Zomicarpella (l m e portIOns. aCllllate-serrate leaves (one Ulearum (1 s ) hnstate thin leaves ~. except for completely terrestrial high-Andean Puya. s Wit.' species from Lo' ch. understory herbs.Rare upper Amaz . often narrowed to a petiole-like base (Pitcairnioideae with capsular fruils and BromeHoideae with berry-fruits) and the third (Tillandsioideae) never does. spathe fre wIth reticulate veins and ?~an understory herb with +/ haVing a large ste:il:~~ spreading (cr. 1. tetrads). The berry-fruited Bromelioideae are mostly epiphytic (except in some lowland taxa) and typically have strongly spiny leaf margins. Taccarum (4 s ing southern P pp. C. ut blooming precociou jzoman understory herb. Leaves ".. spadix unusual in g YCongested. in the leaf margin spines more strongly and wickedly recurved. Differs from terrestrial Aechmea species in connate filaments.) _ Terrestrial. and. h underground hi 130 in Old Wor1d) . The main genus (and only common lowland genus) with spiny leaf margins. . like Xalithosoma b sp. leaves. a more open inflorescence.) . The technical distinguishing style nondiscoid and poUe v~. usually distinguished by pubescent mal~r~scence like Philodendron' b~t ~~~nce o~hen:ise rare in the family.. Two mostly have spinose-serrate leaves.. >20) ovules ' n grams sOlitary ( " vs.n e sap. This is the second lafo-est cpiphytic family and most species are epiphytic. Bromelia (47 spp. Anth:'i~~scence overtopping the gment between th ). especially when epiphytic). characterized by the seSSile rosette of narrow basally imbricated leaves often f~rming a wa.104 Monocots Caladium (15 spp) Bromeliaceae V usuall~ with strongly pe1t~te leav:? ~Iose to X~lithosoma but smaller and i I' " or whIte andlor with purple d . Vegetatively. Mo~t easily distinguis~~1 :ith more pronounced sagittatc who h . have weakly spinose leaf margins (often with spines only on the petiole-like basal constriction. Homalomena (8 story herb spp.lfl Our area (Peru) rare ~a WIth entire leaves).lll{ode1ll!roll) _ A few Philodendron . generally vegetatively distinguishable. ~ndlVlded nonpeltate. the Pitcairnioideae arc mostly terrestrial and. Billbergia (54 spp. . Anal/as (8 spp. probably reachhaving thea~:~.) _ Epiphytic (usually) or terrestrial. mostly in extralimital dry areas.) _ Differs from Aechmea in the dense sessile inflorescence which is shortened and included inside leaf cup. much like some Aechmea species but distinguis?ed by the fleshy compound fruit (pineapple) formed from the fused ovanes. a~d a green and maroonr'::~t~i~m with deeply pinnatifidly IVI ual flowers not stron I e spathe. LEAVES (USUALLY) SPINOSE-SEURATEi FRUIT A BERRY (= BROMELIOIDEAE) Aechmea (172 spp.Terrestrial underi:~oles (and sometimes blad:s) z~~es.Mostly subtro i divided Ie em. s Y When no 1 eaves present. in our area only at low altitudes in northern Colombian dry forests. There are three subfamilies. E. plus . vegetatively.) - m~l~ to Caladium but the leav Sm all: cloud-forest. Flowers borne in openly spicate int10rescence and with long-exserted stamens whereas Aechmea has densely spicate or branched inflorescence. ences with the female port' y the several long-pedunculate IC IS relatively narrow and I Ion of spadix mostly fused to spathe e ongate. Neoregelia (71 spp.rsurface. Essentially a dwarf D p cal dry areas. noncolor:~ SOmetimes palmately divided and when in~~~e~bes. but s~me (especially the primitive Pitcairnioideae with capsular fruit and entire-winged seeds) are terrestrial.) .Mostly Brazili . specIes are terrestriaL BROMELIACEAE Typically a very distinctive family.) _ Terrestrial.0 ten conspIcuously splotched with red characters are ovary with 1-2 /n .Differs from Aechmea in zygomorphic petals and nonspiny sepals. (P. in flower characterized by spiny-tipped sepals. P: coraz6n de jesus si' lOS Chlorospatha (3 spp. -4) placentae and <20 (vs. the spadix distinct in e male and fe al .) . .) . The typically conspicuously bracteate terminal int1orescsence is also characteristic. Owers. Usually with anise sce t d stammodla scattered among fe.

the ovate bracts rather large and often spiny-margined. In our area only at middle elevations in Peru.puya . ( Streptocalyx (14 spp. Technical characters are usually superior ovary and the petals spiralled together.\. FRUIT A CAPSULE WITH PLUMOSE SEEDS (= TILLANDSIOIDEAE) e i h t. . Differs technically frOl:) Aechl1lea in naked (not appendaged) petals. II .Aechmea 6 . the inflorescence slender-branched and the very small flowers w~th only minute inconspicuous bracts.) .. FRUIT A CAPSULE WITH ENTIRE-WINGED SEEDS (= PrrCAIRNIOIDEAE) ! !.elyTechmcal space~ short sessile flowers subtended by small inconspicuous characte . The distinctive inflorescence IS a rather few-flowered spike with reduced bracts.1 I I Fosterella (12 spp.106 Figure 12 Monocots Bromeliaceae (Spiny Leaf Margins) RO/mbergia (8 spp.Terrestrial and not very obviously a Bromeliaceae. ours are tank p. I .Araeococcus 2 ." .: F Ii I I" LEAVES (USUALLY) SPINOSE-SERRATE. I' IS e seed appendage (hairs) apical and ' :I . I 2. . . . . and oblong-ovoid scssile fruits which are often rather zygomorphic.Tank epiphyte (or on rocks) with strongly spiny-margined leaves.) . Tank epiphytes. Inflorescence sparsel I b 'th 'd Y open Y ranched with reduced branches and ~b/{lrac teWll0 es.Streptocalyx 3 .inferior ovary. Technical characters are large.) .:1ill Araeococcus (5 spp.) .~ [:iiI" I h' I 'I il !I:1 I ).Mostly on Guayana Shield.Mostly in Choc6 region.Bromelia 7 . leaves thin and weakly or not at all spmy. Two main genera: Puya and Pitcaimia Puya (168 spp.'. conspicuous. 3. mostly in cloud forests. the commonest species with long narrow leaves abruptly expanded at base. separate petals and a +/.p Y s with thm broad leaves. II Pitcaimia (262 spp. Leaf bases enlarged and ttiangular. I . sometimes with distinct stems. ours a peculiar epiphyte with very narrow leaves abruptly broadening to wide base and an open almost ebracteate racemose-branched inflorescence with tiny flowers. . usually with leaves narrowed to basal "petiole" with fine reduced leaf margin serrations (or these entirely absent). inflorescence rather short and dense. LEAVES ENTIRE.ato~sis ~19 spp. I ~ ! :. Technically differs from Aechmea in petals lacking appendages.Epiphytic or terrestrial. I 107 1 .I 1.) . .High-altitude terrestrial plants.) . th folded (unique).Mostly Central American.) .Pitcairnia 4 . . Leaves usually contracted at base (often only spiny on contracted basal part).Ronnbel'gia 5 . strikingly Obtuse except for an apiculate poml.

!I i : . especially in eastern BURMANNIACEAE 'I .which they . barba de viejo (T.y orchidlike species is actually· epiphytic: The fIrst four achlorophyllous genera below have salverform f1o".differ in the radially symmetric flower. Petals without scales. Miersiella) tubular flowers.or 6-parted small flowers with inferior ovaries. I Mezobromelia (2 spp. to Tnundaceae.) .Guzmania 3 .Inflorescence laterally compressed (often unbranched). usually branched. 109 Figure 13 . The leafy specIes (Burmannia) are very similar to orchl~s except for the radially symmetric flowers and one especlalI.. Most of Our species have White flowers. Leaf Margins) Bromeliaceae (Nonspmy and Cannaceae " Guzmania (127 spp. Usually with very reduced leaves and mdially symmetric uniformly 3.istichous).) . Similar only to saprophytic orchids. but differs in petal claws with scales.:. ~~e~ndages.Tillandsia 6 .Tillandsia 4· Vriesia 7 . Small I?ostly achlorophylJous (except most BUrlluJnJlia) saprophytic herbs.. huayocoma (T.) .Mostly epiphytic in the wettest northern Andean cloud forests. the tepals of sev~ral genera falling immediately after anth~sls. essentially to Guzmania what Vriesea is to Tillandsia. which have apocarpous superior ovaries and separ~te te~als.In ~he !elatively openly campanulate 6-lobed flower a and ThlSllll III the urceolate flOwer with three filiform The last (Dictyostega. typically with 3-lobed tepalS. The main genus of this group (and of the famlly). f!. Very close to Tillandsia and separable with cer~ai~t~ only ~y the technical character of petal scales.are many other genera of Bromeliaceae. fr~m . like UnWlllged species of Burmannia. . Looks just like a Guzmania with rather open inflorescence of bright red spiralled bracts. a character whose slgUlflcance In generic delimitation is questioned by some taxonomists.Catopsis ~wo gen~m Gcmf 2 .1 i: iI i I .Inflorescence laterally compressed (:: d. these often caducous. usneocdes) Vriesia (257 spp.Mostly cloud-forest epiphytes. and to a few saprophytic Gentianaceae (VOYl'la)Wh~ch have 5 fu~ed petals and a superior ovary.exuosa). ~ymnosi'P1zon). 5 1.ers either with 6 corolla lObes (Hexapterella. In~o:es­ cence with spiralled bracts unlike Tillandsia and Vriesia.Fosterella 5 . C: quiche ' .108 Monocots Tillandsia (410 spp. Campy/osIPhon) Or conspicuously 3-10bed tepals (Cymbo~al'p~. Apteria is distmctIVe. There Brazil.Cafl/la .) .. C: quiche. Flowers dIstInctive in the petals with claws fused into tube.

- . There is only one other neotropical genus of Burmanniaceae. Simil~ . WITHOUT CHLOROPHYLL. conspicuously roughsurfaced or tuberculale.capitate inflorescences. Yspaced along inflorescence Tlzismia (12 s mushroomIike flesh pp. e Openly cupular flower 111 CANNACEAE Large-leaved scitaminous herbs. lacking the pulvinuIus and minute cross veins of Marantaceae. IOstead of 5 tepaIs p ena (1 sp.) COMMELINACEAE A uniformly herbaceous. paniculata) large petaloid staminodes (the innermost of which is usually reflexed to form a kind of labellum). EAFLESS OR THE LEAVES REDUCED See Figure l. plus 1 N. these equalled ?r exceeded by the single petaloid stamen and ca. known only from Trinidad. white. is a large. c~::~P:ls: three of which ar~ us~al~ more or I~ss cupular lube with Triw'is who h aSlly be ConfUsed Wl'th fY elongate IOta filamentous . ~Ie to pedicel and dehiscing on only corolla tube y stincbve bulging pouches near middle of . 2.zome.. unlike Dictyostega in being erect.) .Flow ' entire tepals alternating with th ers White to purple with three large except tor the nontrtfid ' tepals. L AND SCALELIKE All ' . Most species have . four (1 111 C. One species has ver di . especially in the Guayana area. The few (-1) flowers Wi~~ost burmanniacs (7-20 mm) . Also widely cultivated. Ghe larg~st-flowered burmanniac (flowcoro. the other a high-altitude terrestrial. on long pedic~l~ e. Commelinaceae 1.'1 OWers are larger than i Y openly campanulate. No other monocot family has involute vernation.lIa but the usually blue fI Ymnoslpho/i and allies in salverform Habltally d' t" ower ' 6 subequal tepals I IS mct m stem with lar larger and with :aves. The small flowers have 3 sepals and 3 nearly always free delicate petals (usually . Gymllosiph01l (14 sp I . The red or yellow flowers are large and conspicuous.Ichas" aungu b very different from the c sll~!lar appendages.Tiny terminated by s' r ower always SOlitary W'th . 30 ASian) . grasses also have sheathing leaf bases but these are usually not completely closed and the plants are less succulent. SPECIES WITH CHLOROPHYLL (AND NEARLY ALWAYS WITH NORMAL LEAVES) Burmanllia (19 spp plus II Ai ' . Fl. vegetatively characterized by the weB-developed completely closed cylindrical sheathing petiole base (~d resultant jointed stem) and usually by involute leaf vematlOn. achlorophYllOu. The fruit.ree very small on es. The onl~ ot?er taxon with a similar petiole base is CosU/s of the Z!Uglb~raceae which is habitally very different from any Commelinaceae in its "spiral staircase" vegetative growth-form. only two Andean specie' an dy ~avan~as. Call1la (10 spp. Miersiella (I sp. 3-parted capsule.) . are species of the leaf litter of mature forest floor. ' 6-lobed flowers' the fl Ique In the relativel and USually p . with green stems and neari' alwa ~can. usually rather succulent.owers typically light blue and a strongly 3-wingect florai t ~ecles (mcludmg the only saprophyte) have Mostly OCcurs in moist s u e and most have +/.ranched inflorescences a t i e ' n eSls. This is the most Voyria-Iike Ila In having 6 . w capitate.Un' . Flowers white to purple. . but lh S. Differs from all other Scitamineae in the asymmetric flower structure without even a clear bilateral symmetry. mostly occurring in disturbed or swampy vegetation at middle elevations.110 Monocots Cannaceae. 7 ASian) .Cymbocarpa (2 spp. also very distinctive. ne species IS epiphytic and I k ' ' except for the 00 s exceedmgl y like an orchid relatively attrac~~~zY~~~O.Nonsaprophytic least at base 0 " y ys with well-developed green leaves at . with an inferior ovary and three unequal petals united into a basal tube.) .Looks like a leafless saprophytic Burmannia on account of the tubular 3-10bed corolla but differs in the more umbellate inflorescence.' flowers are distinctive i h P:. Am. urmanniac. Gymnosiphon Campylosipholl (1 sp ) T ers 16-28 mm long).hiC. but differs from Voy" I.~~' more conspicuous.The small white cuously 3-lobed and nd aVlng only 3 apparent tepals which are conspi2b ca ucous SOon after a th ' . At. Cannaceae lacks the Zingiberaceae vegetative odor of Renealmia and usually has more ascending lateral veins than Heliconia. monotypic Marthella. Very like . one epiphytic. family. more or less hyaline in a~pendages. Dictyostega (I sp. p us 3 Afncan.) - VirtUal WhlC~ It differs by the fruit held at an ly the s~e as Gymnosiphon. Hexapterella (1 sp. mostly in middle-elevation cloud forests. f1o~er. and m color. ut otherwise only aplikeTrilll'isfiower.) . the peCulia flY forest-floor saprophyteS any Old World) . tubular 6-lobed flowers on a forked inflorescence. The distichous leaves with well-developed midvein and finely parallel lateral veins are similar to those of Renealmia and Heliconia.Characterized by the pendent. from one Side. s. and plant from tUberous . which is similar to Miersiella but with a capitate inflorescence.

Tinantia is vegetatively similar in the well-developed petiole-like blade base but has larger blue flowers.d l~rgely on anatomical characters. p . 'i !I 3cm [ !! 3cm [ 3 spp. Leaves distichous. . paniculate with conspicuous pinkish bracts and large flowers with white to pinkish sepals and fringed blue petals both to ca. sometimes the lowermost reduced) that deliquesce as the tlower ages.l ii:1 :. unfortunately. oscopa an AnezIema Leaf bl d .".Geogenanthus mo~tly 3-Floscopa 4 .Dichorisandra . The next seven genera share a tendency to distinctly zygomorphic flowers with elaborated stamens and staminodes actinomorphic with conspicuously fringed petals).Commelina 1 . has a few small white flowers subtended by pair of leaflike bracts and borne at end of a +1. 10 spp. Commelina is weedy and characterized by the well-developed spatheJikc bract that immediately subtends the tlowers. a es USUally sessile. sometimes COlllmelilla u~oliks~atFelmost1y ra~her ~ "I Commelinaceae EPIPHYTE (Geo~e/lalltllUs 113 2 . common in wetforest. The inflorescence is openl. I :.. i . the divisions.. has been in a constant state of flux. black.. The fust four are related to Commelina (tribe Commelineae)' but the last f?ur are closer to actinomorphic-flowered Tl'adescantia Tradescantieae) based on anatomical Characters.Tradescantia (Zebl'in a) 5 .' : I i. cultivated and naturalized in our area. but common in ~et-forest understory and characterized by the minute tlowers In a small mUCh-branched conspicuously pubescent panicle. the ears of Mickey Mouse) along with a small inconspicuous one. 2 cm long. . Figure 14 1. 'i . it differs from Tradescantia principally in the fleshy. with an unusually long petiole-like contraction at base of leaf blade. TERRESTRIAL HERBS OR TWINING VINES 2A.Monocots 112 blue or bluish (to white or pink).wever. has white flowers with 2 large petals (cf. b~se. Several genera are dIstinctive. more or less narrowed toward base and then expanded into the sheathing base. now often treated as part of Tradescantia.elongate lateral inflorescence. ."1 I. especially of the Tradescantia relatives. Generic taxonomy. Aneilema. Geogenanthus is remarkable for bearing the flowers at ground level from the base of the stem. and the inl1orescence is often subtended by a spathe like bract.) . Campelia..' \i (trib~ (ca. but may now be stabilizing. berrylike fruits.A curious bromeliad-Iike tank epiphyte with very long leaves. l' .i 2. is unique in the petals fused into tube and also characterized by the often patterned leaf upper surface and purple leaf undersurface. hO.. Zebrina. plus 150 Old WOrld and n temperate)y weedd SUCCulent herbs. Floscopa is small and inSignificant. CochIiostema (2 . Dichorisandra has a few free-climbing scandent spe~lcs and Cochliostema (restricted to Choc6 area) is a bromeh~d­ like tank epiphyte.

a!e mldvem and are obVIOUSly derived from a pinnate conditIon as opposed to the more obviously palmate venation of bifid cyclanths. Elasis (1 sp.e by the Short leafless' p. Rhoeo.wmmg vines. The last five genera are related to Tradescalltia on account of technical characters of the inflorescence and pollen. plus ea 45 lmear leaves. white or purplish.) _ A high-Andean subparamo herb with actinomorphic flowers.. e moderat I ' e atter unique' T. the flow~ n orescence terminal.) _ Forest-floor herb with narrowly elliptic leaf blade gradually tapering to narrow base. Most cyclanths are bifid-leaved epiphytic root climbers while palms (except the spiny. (especially .. plus 15 Old Wo ld) S . with several small u~ua~~ted Pltaeosphaer~J/1). subtended by a consp' I y. wet orests.) . 'I ! Monocots more or less clasping at base but s o ' . MlIrdallllia (5 spp. and spirally npogandra Tripogalldra (22 ' cenee b spp. Phyodina) (20 spp.Forest-under t like or race~~es.Charaet . Callisia (incl.Mostly Mexican herbs. ~nflorescence composed blue to Pinkish flowers small . With 6 stamens. white to pink or greenish. ed relativesraTchts and dimorphic stamens thenzl by lack of leaflike inflores.erect f10Werin stem r. in sometim. open. with 1-6 (typically 3) stamens. or hght bluish flowers. Sho~ s termIn~ted by densely paniculatc. erect herbs. h umbel-like.wlth +/. All have radially symmetric flowers. Without leafy spathe the .e~ntraction above the Sh::Ubsesslle but usually has lStinct species passing U d tho The taxonomy is very j d GeoglmamllllS (4 sp ) n er the name D. characterized by the conspicuously stipitate paired or umbellate cymes which are simple rather than fused in pairs as in Tradescantia. 'I. The most important character is the inflorescence cymes fused in pairs subtended by spathaceous bracts. Tradescalltia (40 spp. petiolar contraction and Cyclanthaceae 11:> 2B.) .em wide) and zygO~O . Tradescarttia.) -Essentially represents a trend in floral reduction (toward anemophily?) from Tradescantia. Gibasis (11 spp.s se re mellmes with very short petiole-like leaf-opposed. CYCLANTHACEAE A very distinctive family that could only be confu~ed with palms. others a very short petiole-l~ ike bracts. .Some species . The very small flowers regular. herb'Floscopa (4 spp. the flower clusters sub tended by folded leaflike spathes. Setcreasea. owe.roun. . Characterized veg t t' ubprostrate herbs of dislStinct petiole above the sheath r ~ a lve!y by the leaf base narrowed to cup-shaped bracteoles. .114 . T?e Ovate or Oblong-ovate are unusual among Tradesnd distinctive in distich leaves are mostly sessile or . The characteristic thick-spieate unbran~hed tnfl~res­ cence of Cyclanthaceae with its appressed thick four-Sided female flowers and fruits is unlike that of any palm. .A poorly defined. The l:1owers are mostly radially symmetrical unlike Commelina and its relatives. all with ( Spp. blue flowers wtth 2 upper petals ICUOUS eafhke spathe. Cycl~n~hs with palmately divided leaves (Car/udovica) can be dis tin- . leaflike bracts absent and flowers always regular and 6-staminate. Terrestrial cyclanths are always essentially stemless and generally more obviously herbaceou~ than similar-appearing palms. with small petals larger and white the lowers strongly zygomorphic with upper 2 arranged. multifoHolate liana Desl'IUillCus) are erect and terrestrial. Campelia (3 spp.. plus 19 in USA) . Old World) .. The leaf blade fwers In ~erminal infloresconfused with sever~.d level from leafless time~ orbiCUlar) leav~s e?etatively distinctive' onsplCUOus fringed blue tending to cluster ncar st With a well-developedIn th~ rather broad (someem apex. cences Without spath ~lue or blue and White fl are t. except for the inflorescence. hexandra. .) . small and greenish. Inflorescence larger. Inflorescences variable in form but always lacking spatheJike subtending bracts..rs relatively (at least !O Dicltorisalldra 25 rphlC. rn~oreseence of I-several ~ ory herbs with broad distineUy A. and Zebrina. constriction (e.Small forest-understory con~plcuously pubescent. .A distinctive for base of the Stem and l?florescence arlsin at est-floor herb character~ctinomorphic flowers ~lth se~eral relativ!y . d cautia relative e y zygomorphic flowers In ra escantia subsessile a s'. Tmalltia (13 S ) petiolate Ie pp. Flowers small. largely Central and North American genus now interpreted to include such segregates as Cymbispatha. borne on ends of long erect peduncle from well down the stem. Described as.Wit stamens sometimes reduced to tw~. ous arrangement. unlike area Co'mmelina a:orst. p us 60 Old World) S ad' . known only from Ecuador. The ell/le~unculate mflorescence of small white narrow petiole-like base. and very similar to.Prostrate herb with of cymes: frequently contract:dar~~ on sand~ soil. mostly in swamp . p IC eaf blades are gradually contracted to All eilema (1 I turbed f sp. Sometimes included in Tradescantia from which it differs in having a fleshy black berrylike fruit.neilema) lar~~ (71~1 umbel-like branches t~~~' usually appearing umbel.as s~edltngs) thes~ are always associated with an elong. While a few palms ~o have Simple leaves with bifid apices.g.

il " . unique in thickened midnb. The spiralled inflorescence is also umqu7· A rare Sec?nd of Choc6 has the two segments fused at matunty but. I 'F I .i' 117 2 . . I' I: 20cm[ ..Completely unmistakable in its palmately divided palmlike leaves from an underground stem. 1. The mature fruiting inflorescence is also unique in the bright orange-red pulp which is revealed as the outer seed-containing layer of the inflorescence separates and falls off. Pseudoiudovia.116 Monocots Figure 15 i:!I' I. iii. the pair of m1dveins" is still clearly evident E: hOJa de Japa . inc!.' I..Always terrestrial.Sphaeradenia 3 .Carludovica '..' ·1 : 'I. I' hmm I "I . '" . :"'! ii) I'I !I [II 'I ('. I. based on a mixture) _ Two species. LEAVES UNDIVIDED (AND WITH ONLY A SINGLE (OR No) WELL-DEVELOPED MIDIUIl) . bifid leaves (most genera) and undivided leaves (Ludovia) are clearly apparent if not as indicative of natural relationships.Dicranopygium 4 . of each segment of the bifid leaf. i'l I iI'.) . both with irregularly crenulate apices. eye/alit/ills (2 sPp. . 'Ii . I. LEAVES PALMATELY DIVIDED (USUALLY INTO FOUR MAIN i.. ': [ Iii .1:

.\( 3 III ::11\ ili!{ . The distinction between Cyclanthus with its screwltkc inflorescence composed of alternating cycles of male and female flowers (and with each segment of the bifid leaf having a strongly developed mid vein) and the rest of the genera with the flowers in groups of single pistillate flowers surrounded by four male flowers (and leaves palmately divided to entire or bifid but with the segments lacking a strong midvein) is a fundamental one. C: palmicha. ! .'i ! I':. \ 1 I II II I ! I i \ II. the commonest species tends 4 to be more lianescent and free-climbing than bifid-leaved genera. Cyclanthaceae SEGMENTS WITH LESS DEEPLY DIVIDED LOBES) I. Thin slices of the fibrous petioles are much used in basketry and to make the famous "Panama" hats of Ecuador.. LEAVES BIFID (UNDIVIDED BUT WITH Two STRONGLY DEVELOPED PARALLEL "MIDRIIlS" IN ONE RARE CYCLANTHUS) sp'e~ies 1 . P: bombonaje .'1 II I. I il i :I i' i. presumably this is the only bird-dispersed cyclanth."'j" :~i:. . ~ 3.) I I Carludovica (3 pp.' I·.t. J .: IiiI guished from many fan palms (but no! from trunk1es~ ju~e­ niles of the scaly-fruited lepidocaryoid ones predom~nat~ng in Amazonia). I ' I I.:' ". 1 I Ii: II" (Terrestrial) I'1.Cyclanthus i ! . The 1nflorescence 18 rather thin and reduced with the flowers com pletely lacking tepals. .. II. . the seco~d with short~r wider leaves. the common one with narrowly obovate to oblanceolate leaves. always terrestrial. "'i'l'.U I I "ll/ 2 I::' I " . by the absence of a raised triangular proJectlOn (hastula) at the intersection between petiole apex and l~af blade. Llldovia (2 spp. jl il . lj! Ii: I I 2. ' . The differences between the groups with palmately divided leaves (Cariudovica).At least. I: \ it II [.

:.TerreStrial in northwestern Amazonia. Aspiul1dia 5 .! " Evodiallthus (1 sp. usually relatively (to Sphaeradema) dullsurfaced. DicraliopygiulU (44 spp.I I .) . p~y~es. Concave petiole. P: tamshi 119 Cyclanthaceae (Hemiepiphytic) .Epiphytic climber. I 118 Monocots Figure 16 Asplulldia (82 spp. E: totora. mOno typic but widespread and common.) .Ludovia .) . The inflorescences have the usually 3-5 spathes (or their scars) scattered along at least the upper half of the peduncle. 2 .) . in most species the leaves are more Shiny and coriaceous than in Asplundia and also differ in having only one main vein in each segment. Differs from Asplundia in 8-11 spathes (or spathe scars) instead of 3-5(-8). occasional1y +/_ epiphytic in Choc6. ( I \ ':. especially on ~~cks near rapidly flOWing streams. Differs from Asplundia in haVing the three spathes (or scars) densely crowded at base of spadix. arranged spirally on the stem. .1 I I Sp!laeradenia (38 spp. The leaves are alwa~s Mid..) .Mostly terrestrial but also many epiDiffers from Asplundia in the stiff distichous 2-ranked leaves (cf. and the spathes clustered at base of spadix. P: tamshi Scm 1 2 10cm [ Diallthoveus (1 sp. the obviously separate adjacent narrowly columnar thin-tepaJed floral units being especially distinct in fruit. aractenzed by very long pedUncles and the leaves drying black. ( Thoracocarpus (1 sp. mlfllature Ravena[a).Evodiallthus 4 . and thin-textured. Dlffers from Asplundia in the smaller (usually tiny) few-flowered inflorescence with more broadly separated individual flowers having reduced perianth lobes. E: jarre.Thoracocmpus 3 . also in having the pistillate flowers free from each other. eh . includes ~oth terrestrial species and epiphytic climbers. Vegetatively distinctive in the unusually wide. Similar to Evodianthus in the scabrous leaves but terrestrial."i':' .) . thin. i ! I':' Usually terrestrial.epidermis and the characteristically hardened but brittle surrounding P: tamshi .Asplundia 1 . Unique in the family (along with newly discovered Dianthoveus in the leaves scabrous (but this apparent only when dry).The main genus of the family.I . and the typically rather shiny surface of the thick fruit tepals. monotypic but common and widespread.A newly discovered Choc6 area endemic.) - !.> I = SC!lIIltesiopllytlllll (1 sp.Epiphytic root climber.

Cyperus 3 . the inflorescence axes lacking terminal flowers and with each flower in a glume axil. the main tropical lowland genus..III !i iii II' I 'I . generally characterized macroscopically by typical grasslike leaves and the umbellately branched inflorescence subtended by a whorl of 3 or more leaflike bracts. usually triangular stems.( I' : I . are characterized by uniseXUal flowers.. incl Old WOrld) L with characteristic leaf Sheath t . .Monocots 120 Figure 17 CYPERACEAE " I I I..::.Il 1'1 1.:: '.. with the inflorescence mOre contracted (of several heads) and "~onfluent" : '. Old World) _ The most prevalent neotropical sedge genus.I Cyperaceae Mostly grasslike herbs wit~ linear leaves.1 h. Cyperus. .1 :' .Cyperus 5 . " I .Diplasia 7 . Fil/l~ristYlis (300 spp. 1 121 ..margins of the sheathing leaf base.Carex 8 . mostly Australasian) _ Weedy herbs with discrete OVOid Similar to some Cyperus species and with a similllflorescence but with the Subtending whorl of bracts mconsplCUOus or at least not leaflike. . and Calyptrocarya (Sclerioideae) have unisexual spikelets with a terminal solitary female flower and lateral male flowers . ' . •. I ! \ .. II .1 Ii f'li" il

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.'1 i I( . unbranched inflorescences or lack the sUbtending whorl of leaflike bracts. inc!.( ! Ii. Worldwide there are only six large genera plus many small ones.'. ..' '. differentiated from grasses by 3-ranked leaves. Carex and Uncinia (Caricideae). . Upper branch with several more I aChlsubtendlllg a pedunculate inflorescence or ess c UStered ovoid sPikelets. and its relatiVes (first seven genera) have severalflowered spikes. 1 .. mostly African) _ A small herb close to Cypems butsessile. DipJacrum. Kyllillgia (60 spp. . il:I' j I Cyperus (600 spp. I. I II . Relatives have sessile.I 1:'1 ~ l' " I".Calyptrocmya 6 . and fused ..I I :1 . A few forest-floor species have broader petiolate leaves (Mapania).:: I'.Becquerelia 4 . (sometimes leafless) and reduced wind-polhnated flowers 10 complexly bracteate inflorescences.I "".' . A few species are scra~­ bHng vines (Sc/eria).I I.:( : ') 1 : '.'. another important neotropical taxon.• ':.' I' 'i.t Ii ' . i' . P: piri piri t II 1 !"'.. Becquerelia. (A. .C) . the female below the male (or the spikelets unisexual). and its relatives (next four genera) differ from Cyperus in 2-fJowered spikelets with a perfect basal flower and male upper one.Bisboecklera 2 .-. . .Cyperus (fungus-infected) .l J • 1" .. Taxonomic relationships within the family are largely based on technical characters of the inflorescence which are SUbject to different interpretations. Scleria. mostly of temperate areas.:''I . ~arly w~orled s~ikelets.''. Rhynchospora.arge SUcculent marsh herbS leaves gradually redUced and e onnlng a s~gmented S-angled stem. . Mapania and its relatives (Mapanioideae) are technically characterized by having the spikelets reduced to a pair of unisexual flowers these arranged into pseudospikelets. Sedges are especially prevalent 10 swampy situations. Fllirella (40 spp.i.I II'"."1".

Eleocharis (150 spp. 123 Figure 18 i .. . Old World) _ Leafless with round hollow stems.) . " . mostly occurring in dry. Inflorescence a single terminal spike.. Spikelets often ~=~ or / e lUflorescence i~ axils of normal-looking female spikelets with p e. tri OldI World) _ Variable in habit and somcles vlUy.e bIsexualplant with a tuft of very narrow erect leaves and hard-to-see so1'Itary flowers. inc!.. erect leaves. . Stems rather weakly triangular. Old World) _ A large and variable genus in both upland and lowland Neotropics.Densely tussock-forming with very fine linear. Sciryus (200 spp. temperate) _ A distinctive pincushIon . Differs technically [rom Cypel'us in spikelets with only 2 flowers. lik Oreobolus 00 spp. Am. Pleurostacl!ys (50 spp. from side of stem apex.Uncinia 2 .Fimbristylis 3 .Mostly Coastal Brazilian.) . larger species with conspicuously hollow stems are unmistakable but smaller species are similar to Bulbostylis except for the swampy habitat. Inflorescences scattered along upper leafaxils. :'!! '" RhYllchospora (200 spp. Fruits 1 .. inc!.Elaeocharis 5 . inc!.. inc!.' IS nctlVe In round stem. also on or near beaches. the latter resembling some Eleocharis species but differing in the denser tussock habit and nonmarshy habitat.Kyllingia 4 . USually ~h~~~ed rachlUa.g. I "·1 Bulbostylis (100 spp.122 Monocots L \: . the leaves Often with sharp. Stem Cutting edges Infl Y angu ar. Rhynchospora . The inflorescence often small or tenuous borne in axils of several normal-looking upper leaves.Hypolytrum 7 .I .Coarse 2-3 m taU herb of swamps and salt marshesC(adium D' ti (2 .Sci/pus 8--10. unlike Calyptl'OcOIya. p~amo li'n ~cleria (200sharpl spp. nastily tended by reducedo~:~cence W~h ~everallOwer branches individually subleaves. especially in marshes (e. enuous an~ less deflUed than in many sedges. small paramo species resemble Eleocharis except for the subterminal inflorescence. ' . With two growth-forms. Inflorescence more or less congested.) . inc!. occurring in swampy or marshy areas. spp. mostly s. often with rounded angles. Related to Rhynchospora. Old World and N. rOUnd and shiny. P: totora Cyperaceae (E. the Lake Titicaca lotora recd). rocky grasslands.. Old World) .U) . . distinctive in being subtenninal.Mostly in Andean paramos and puna. The inflorescence is variable but ~iffers from Cyperus in either being dense and sessile (when with subtendmg whorl of leaflike bracts) or more elongate and with separated leaflike bracts below each of lower inflorescence branches. Inflorescence branched and Fimbristy!is-like or reduced to a single terminal spikelet.Scler/a 6 ..

Old WOrld.ble and superficially rna e (or UniseXual spi!< 1 e UOlsexual flowers with e ets).) . in Our . and Syngonal1thus (192. Ulleluia (30 s . protruding.. most! narrow. Old World) C: flame. Paepalanthus (485 sp~. is a prostrate or floating weed of open swampy areas with short sheathing-based leaves and short-peduncled axillary inflorescences.A coarse fa Ma ~ . temperate. Related to margms.5 m tall and occurri . Jets tendin~n~ S~btending each of lowermos~ednfloreseence bracts. p. very elongated. characteriz~d by 4 or 6 stamens (vs. elongate dour :u-ea. Bisboecklera (8 sp ) leaves sublinear but contni~te. the inflorescence and characteristic 3-winged capsular fruit arc thus totally unlike Smilax. mostly broadly ovate and basally cordate. e. Our commonest species. the o a pettOle-like base.eJ~s.Rela . incl Old W dlstmctive broad leaves contr~ t d' orld) . especially in moist situations. sometimes also with petiole apex slightly thicker. mel. 100 spp.ted to Selena but with the rachillae larly globose heads Th . mel.mg an overgrown Clump_forminrest-understory herb. ~ ' i! ::.and species in the Guayana area and . Inflorescence sessile or of 3 broad bracts. Infloresee:~: UPI~ndS where it replaces c female below th>. instead with petiole base thickened and twisted or bent. incl. Technically 1 ers tn having 2 (rather than 3) stigmas Carex (2000 sp .) . sometimes with rather large. always with entire margin.Essentiall o 1.. subtended by :h I lnto basal petiole.).l1Ia but the leaves epeliolat I g Cyperus. narrow leaves and dense heads of small whitish flowers borne at the end of a long naked stalk. borne in axillary spikes or racemes or panicles with well-developed central rachises. .t A narr?w-Ie. Diplm. " Mapallia (50 spp. irregular. Dioscorea (600 spp. rarely 3-lobed or 3(-5)-foliolate. the most widespread taxon in our area.&enera . with unisexual trimerouS flowers and pistillate flowers With free petals. superficially 'P . i I :'1 II . The other most important genera are Erioeau/on (ca.A forest-floor herb with very e and congested. ys. ' spp. e ea • thus. Spikerelated t Mm exserted anthers of the m us and with fuzzy appcarance at o apania from Which it d'f£ ~re numerous flowers. not-at-all decurrent leaf buse and the welldefined transversely parallel secondary cross veins (the vein network of Smilax is much less regular). monolypic Tonina. Eriocaulaceae DWSCOREACEAE Mostly slender twining vines with smooth green stems arising from well-developed rhizomes or basal tubers (these sometimes large and conspicuous with the laps projecting above ground surface). .• plus 4 Afric~n) with unisexual trimerous flowers and pIstillate flowers WIth connate petals. but very different in ~h varla.on the. There are numerous additional . rougher-looking. Looks like Kyllingia with y n mOlst Guayana-Shield savannahs bracts mOre acuminate and th congested h d sessile head s utb the'Inflorescence. onger and with finely toothed cutting Hypolytrum (50 rather C .Mostly can resemble e om!n ant sedge. . mostly characterized by the BromeJiaceae-like Qr grasslike tufts of long. y an overgrown Calyptroewya ng In understory of poor-soil forest.• plus 300 paleotropical). P: sacha papa ERIOCAULACEAE Herbs.1 II I! f :1 : r! ! !I . Leaves not very monocol-likc. (I ) rese bI' sp. laterally Oallened rccurved spines on stem. Leaves usually in basal rosettes but sometimes the leaf clusters borne at end of a thick subwoody stem. DiosC(>reaceae. to Andean uplands inPP. n. palmately veined.S~p ./Q. gfl • so splkeJels congested into small irregu' . moslly looking about alike and dIfferentIated by techlllcal characters of the minute flowers. Differs from Smilax in lacking petiolar tendrils (as well as the jaggedly sheathed base of petiole).Most! i . temperate it d Cyperus as th d regron restricted to And n Old World) . The flowers are small and greenish.d! I'I I II II 124 Monocots Calyptrocarya (6 spp ) . Other vegetative differences from Smilax include the sharply defined. ' l?iplacrum (7 spp.1'1 I I i! i ri t Becquerelia (2 spp. of inflorescence not prolon ~d' . Related to C .. temperate)-Restricted hOOk-tipped SPik~letense ~nfJorescence wit~l ex but very distinctive in the rachillae.Exclusivel separateIpelUs-like but with only 2 lear/ s~amp or stream-side herbs.. e 10 orescence rath er tenuous ' and few-branched (1 Ike Pleul'Ostachys) b o ' • ene tn axils 0 [' several normal-looking upper leaves. 2-3 m tall.) ..aved version of MapclI1ia. Mostly grow on exposed rock outcrops or open sandy places. 2-3 in other genera) and unIquely diaphragmed aerenchymous roots. Braztl1an Shl~ld. these anthesis fro 0 e more open than in Cypel tn orescence branches.

Eriocau/on 4 _ Gramineae spikelet details . ':' I I '1 " 1

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. : I' I I i'll . . j'i. .i. ""II!. \

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I. I r·:.Tonina .126 Figure 19 127 Figure 20 Dioscoreaceae Eriocaulaceae and Gramineae (Spikelet Details) 1.'i.:]. I I'. ! il: l I I. I 'iI SPIKELET FLOIUi1' il 1. . .11 ' h\ i" : .r. .I'I' 4 .1 lemmas i' 'ill: I I.: . .1 ) :! i' 2 .i 'I+ i 2 . 'i ') 'I' II j'l!'I" i 1.'" i I I palca 3 'J . : I' .i" . / Dioscorea 3 .Paepalanthus I ..

Grasses.th~ primitive subfamily BeSides being woody. each I-many-Dowered with reduced 110rels (if any) above the perfect florets. ost bamboo leaves are disnarrowed above the sum ~VIng the base ~f the leaf lamina structure called a md mit . PohI's key to th~ w 6) for more complete in lieu of a co~r~ (with the addition o~o~ . B~busoideae (the ba~b~~~ only .coriaceous). . (Figure 20). \ . ' The basiC ral distichous bracts. notes ImpOrtant genera in c aracteristics of Our area are included.i I. Oryzoideae (more or less aquatic plants characterized by very reduced glull1es and a single fertile floret).. IC ave evolved their own corn. are herbaceous (Cryptotreptogyna. SInce they typically have much encountering in this f te~per~e botanist is accustomed to tinctive in the family ~~ Y'. I .1'. I II ' . . articulate below the spikelet (sometimes on the rachis or at base of spikelet cluster). 1." II. . the pistillate below and the staminate above. very distinctive in characteristic leaf' Il!0st. t e palea. y with hollow Internodes) and the h the stem.. Streptochaeta. and spikelet arrangement on the inflorescence (pedicellale in open inflorescence = Agrostideae (I-flowered). Grass flowers are hi h i ' . Pariana gRn~~. ' ..es a basal sheath surrounding usuaUy a fl . The pooids include ten traditional tribes. ~ addllIon. : (I iU :I I . narrowed pseudopetiole b t usolds are characterized by a A few additional bambo~ w~en blade base and sheath apex.. forest-floor grasses can be broader leaves than a am!ly. 1\ II I· '\ ' II) \. and Tripsaceae with unisexual spikelets. in additi:n :Ch ~nclud. recent treatm ayton and Renvoize 198 ancensls or Genera is duplicated ents.I II I' . . l(or more)-f1owered. climbing or sub e woody a~d there are also a number of including the wow~ody taxa In other subfamilies. sessile in spike (or 2-several spikes) = Triticeae (Hordeae) (spikes solitary with spikelets on oppOsite sides or rachis) or Chlorideae (spikes usually more than one. Arundinoideae (sometimes included in ball1busoids.of the sheath Into a petiole-like pseu opetlOle .. GYllel'ium. typically with lemmas with three strong veins). . 1I ·1' I. and two with thin membranaceous glumes (mostly Paniceae with fertile lemma and palea +/. The panicoids traditionally included four tribes.ch pair of glumes called the lemma (wh' h f s. . the bam~on~mIng truly woody taxa. I "I I I I.1 I 'i :1 ! : . The fami! ~~o:ted by ~ series of bracts and complex inflorescencis who ~n~my IS largely based in the plicated descriptive te . the other pedicellate and ei ther staminate or neuter). with adaptations for the sexual parts' are ac normal sepals and petals. spikelets only on one side of rachis). I ) i' . presence and size of glumes." Ii ('i . mostly large and reedlike with large plumelike panicles and including several more or less woody taxa).. 129 . 0. and barely subwood eSCCaped). Instead bracteoles. if' I i I. rl ' 'i". Z~a (cultivated).ast majority of grasses are herbaceous.~de .. two with thick hard glumes (Andropogoncae with characteristic pairs of spikelets. Ii ) I'. articulate above or below (Oryzeae) the glume.. These Arun~~z~es. Pooideae (Festucoideae).J l' . and Chlorldoideae (the bulk of the lowland tropical nonbambusoid members of the old Festucoideae. each with an outer bract IC 0 ten has 'ts 'd' awn) and an inner bract called h I ml nb extended as an This is a huge famll . I I J 1 'II I . t e tYPICally narrow lamina. Panicoideae have spikelel~ more or less dorsally compressed each with a single perfect terminal floret subtended by a sterile or staminate floret." I ~ " '·1' '. there is called a li~~~r p~Ject~n at the junction of blade and sheath tricky to recog~ize ~~7 ~Iess. Lithachne. I Inoldeae. mostly high-altitude and restricted to the Andes in our area the lemmas mostly with five or more faint veins). ea. Unt't'IS the spikelet whO rmInO h I ogy . sometimes on separate inflorescences). i . 11/. their jointed stem~ y oner ar~ generally.1: PI ii ' Gramineae The traditional taxonomy divided grasses into two main subfamilies based on technical characters of the spikelets.:~ conSISts of a short axis with seveglumes (rarely only' on e ~wo lowermost bracts are called subtends one or more efJ~ritresent). large reedlike y ortaderia of the paUlc e~. defined by whether the spikelets are perfect or unisexual.osta Rican taxa on tribal affili ~.128 Monocots GRAMINEAE ma~~~~~o~e. Pooideac (= Festueoideae) have generally laterally compressed spikelets. ~hloa. the Ball1busoideae (vegetatively defined by the leaf blade contracted at base into a pseudo petiole and including nearly all the woody grasses). Aveneae (2-manyflowered and glumes as long as spikelet). A few members of ~~. or Poeae (Festuceae) (2-many-flowered and glumes mostly shorter than first floret). '. and articulation above the basal bract pair (glumes). wmd-polIination and I gk Y reduced. and Some oz' I'a a za: Pharus. Nowadays agrostologists mostly recognize six subfamilies with the old festucoids split into five subfamilies.ete treatment of grasse s n ean ~~urolepis) 1 most of the mo~ I?n and distingUishing .' . one sessile and perfect. The Panicoideac are essentially unchanged.Al'undo (introduC~~a~~ed and subwoody. . and ee ohl s treatment' e aUlcoldeae Graminum (CI In the Flora Cost . ~wo other subfamilies ar~ specIes). Inflorescence . and SaCCharum (cull' grasses with large plumose SQ~e VpIny ~siacis species of ~atped).

I .'.. I' 1.Agropyron 8 .Brachiaria 14 B I .I'" .Agl'ostis 9 .GYlleriuJn 7 .Digitaria 10 . 'I .Anthochloa 3 .: I " II I':" .Coix 6 .Aciachlle 2 . i i. i .Eragrostis 9 .Hyporrhenia .AxOIlOPIiS 5 .Echinochloa 3 .Al'istida 12 . I it Ii il I. :./ 11 .Eleusine 5 ..Ollte olla 13 .H) '.Arthrostylidium 10 ..Guadua 6 . 130 Figure 11 Gramineae (A .l I. I 9 14 tff1 1 . .Andropogo/l .Chloris 8 . I'1 .\ I . !.Festllca 4 .Dissanthelium 15 _Calamagrotis I-Bromus 2 ." .Chusquea Plus Dissallthelium and Guadua) .Hordeum 11 .H emarthria 14 .:11' . .II.Homolepis 15 . 131 Figure 11 Gramineae (Bromus and Chloris .Chusqllea 7 .Acroceras 4 .Eriochloa 12 .Hymellachne 13 ."i.

Isachne 5 .Nasella 12 13 . I .U:irenzochloa . 14 . ' I I.0) (P-T) I.i 'I 132 Figure 23 133 Figure 24 Ii Gramineae Gramineae (1.Lasiac.Leptochloa 4 . I .Reimarochloa 4 .Ml(hlenbergia 10 .Sporobol us 14 . . I I I.Setaria 6 . .Luziola 8 .Orthochloa .Sorghum 7 -Poa 5 .Oplismenlls 14 . I I.Pariana 12 .s 6 7 . " I .Melinis 9 . ' I' I :.Trisetwn I : .Stipa 11 . :i 2 -1clmanthus 1 _Imperata 3 .O'yza 1.Rhipidocladum 9 .Paspalum 8 .Pennisetum 10.Tripsactlm 13 .Pharus 3 .Olyra 11.Panicum 2.

.) _ Clump. . Branches at midstem nodes numerous..elevation.. arising above the node at the apex of a prominent bulge that can. Culm . cultiva... without pseudo .~~~ glumes... of petIOles... ac and shin h angl. Primary branches usually 2 per node............flo':ered. . Stems nearly herbaceous . ·........Mostly scandent or at least strongly arching slender montane bamboos. I...! Iii' III j Mer(Jsfilchys (40 spp..! : I....... v a e or escaped <20 em I .... mgers)..... Leaf blades bo·r·n................... In a arge fan h d J' dlOecious...' ..... E: carrizo paleo tropical Bambusa is moot......... Hollow stem GlIadlla (30 spp... g............ . ·· ... wild ong.... La' StaCIS O pedicel.. stems pithy..... 1 10 S.. ·································Chlisqllea . s of various habits ·· ... ). of variou~·d: ......l. ..... sometimes very .\ "Ii ..... Anmd(} dtJIlaX' short pseudopetioles.. aligned with 7.... :····· .. .. Stem ..shaped in cross...... Solid stem Chusquea (120 spp. characteristic in the midstem blades retlexed and so strongly basally constricted that pseudopetioJe base is narrower than sheath i I_i' " IA.... Primary midstem branches 2-severaJ per node. I • plants bl . Distinctive in the spiny branches.... ' . Leaf blades evenly distributed each with single fertile flower d~~on? ste~s.forming more or less arching (or scandent in upper portion) bambooS.... soon branched near base.. spikelets ted sugar cane ' Isarticulatmg below glumes. IIi: i I 11.... I[ '1 iI..... Arthrostylidiul/I 14.1

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... Phyll(Jstachys 3. without hollow center ... disarticulation b Inds. Le'af bl ades usually.. Ledbi~d·~~·~~~~ii.... ... :I I: I I I {! \ ........ disarticulati y w en mature. tw k' ....walJed' cultl' t d ng at base.... ... not black... 4 ranchmg ..j:i~ .. ·.. :1 i : ................~... Sterns woody and b ' s. glumes. mostly borne on small lateral leafless or bearing nearlysebo dunlbranChed 8ubwoody stem. I I. branches 2-many " P r no e......... native .I i ...... Stems with th or cultivated.2 m Iong...1: :. Woody bambusoids with pseudopetiole between sheath and blade IAa. ges of flat. embrane... internodes D........~..AU...... 14..... 4........... Midstem sheaths with a narrow reflexed blade that is KEY TO WOODY GRASSES 1... ternodes hollow ' un ranched .. Bamboo ...... arising from ed. triangular platelike meristem that is closely appressed to main stem. plants rarely blooming... wild plants !~~letsI2.) _ Our only native large bamboo. to 8 cm long on main stem sheaths . 134 Gramineae Monocots 135 ...... ..... II apex. Primary midslem branches solitary. · ..........u~m internodes solid... Mer(}stachys 12..section............. tinues down 10 the node..8 ape cluster at apex of stem.........Bambllsa (Guadlla) 7..... I. . .. Branches at midstem nodes not arising from flat plate.. ... .. 12...... . ·..~. . Stems not VIscid............ Swallellochloa below 3000 ' Occurrmg' f and with ~ .) .... ya ongnverbanks 2.. related to Rhipidocladum by the otherwise unique feature of the multiple branches at each node arising from flat triangular plate.. ~ . main stems j a e ess sheath .E1ytrostachys 13....·1 . ave 3000 m elevation.clasping. Branches 3-many per node.....I .... cu ms thm..... . \ I ': . ely woody. 8. placed at seSSile. branches stiff add WIlIlout definite boundary .. Leaf ....... branches or from near ba f ong.Rhipidocladlllll 11.(i-.. ·. ..... i ..i I :.. Spikelets umsexual....... of varying number.. Spikelets alike bl k oommg annually.. Neurolepls........ i... Stems laCking thorny b ..... 1...... leaf blades with large efiOllely woody......:........ :... E: cafia brava .... 6 orny branches .. . eaf blades I-several m Ion b extremely woody..i~~·~~~i~.. \ ' ..: .... . leaf blades 9. s wIth ghstening viscid b and at nodes.claspi I blades .. ··· ..... Vegetatively distinguished by the solid stems and the distinctive branching often with one large branch and numerous small leafy branches together at each node. auri" cular bristles short..Saccharum officillarum .....section..... Midslem sheaths with an erect blade that is as wide as the sheath apex and not constricted at base .... not constricted at its base and much narrower than sheath apex . ·... ·.... mall shrubb b hOllow Y nmboos of paramos ab m stem center smalJ an . per node . Stems bar ranches... 13..... hOllow In orest or savanna e d definIte bOUndary stem center usually larac membrane'... Generic segregation of the neotropical species from \ . II! i I.. on below glumes. orne on malO stem... cultivated bamboos .. monoecious... disarticulating above the ........ 3... auricular bristles very prominent.. cordate... SOlitary s 'k I ' ... internodes sl d (can be crushed with f' en er «I cm wide) green soft 8. WOODY GRASSES lAb..... C: chusque. mostly with 6.. b······ .·i~·~~·~h~~·20~~·i··················G. . ......e·. internodes cylindrical in cross. borne on main stem' S.e to pedicel.Olle1l1W 9. Stem:~ort p~eudOpetiole .. ·Olyra 10 S' lameters... n erect.... usually 3-5 10... not cordate.('...

Cortaderia (19 spp. I ! ' I. Artlzrostylidium (20 s _. I i: '

Related Interests

'1 ' . often on cliffs. characterized by thO . mbmatlQn of solitary branching and sheath. Cryptoclzloa (15 spp. perhaps inadequately. I i Ii Ii :. and denser'mflorescences. Differs from Chusquea indo e. Olyra (22 spp also 1 A . (Swallellocilloa) (7 s . In!1oresccnce large and rather plumose. and with sticky awnless lemmas for exozoochoric dispersal. Lithacll1le (4 spp. narrowly oblong 2-ranked leaves. per spikelet and formerly E' e to Pam cum. . ovoid male (basal) a~~h.Barely sub d .136 Monocots Gramineae Rilipidoc!adum (10 spp ) SI . ally and berrylik y-fmltmg spikelets (hi hi g vme. '. characterized by broad leaves with poorly developed pseudopetioles. II I . Distinctive i~ the co n?ns~my lowland bamboos. .) _ One of the most distinctive forest-floor grasses. 'II.s.) _ Forest-understory broad-leaved grass related to Olyra and similarly once placed in Panicoideae.) _ A broad-leaved forest-floor grass that vegetatively resembles Streptochaeta but differs in the veins of leaf blade running obliquely to margin and the blades twisted so that anatomical lower surface is upper surface. very common in early succession along low eleva110n. which it resembles in the large paniculate plumose inflorescence. Aulollemia (24 spp ) S bamboos WIt. are Y woody stems and viscid bands at very I Neurolepis (9 spp. pnmarY-branchin g. bhng grasses f . . 'I I " i ~ .Large erect canes to 10 m tall and 4---5 cm t?ick. mostly of lowflattened triangular meriste . Elytrostachys (2 spp ) . 'I . .Broad-leaved genera of forest understory.(Saccharum) (cultivated) - One widely cultivated Asian species. usually scramwith separate the matn stem as well as bra ng from true bamboos in floret structur~arrowly. . Inflorescence very different from Streptochaeta.I II I \ . e a~ s aped branch clusters arising from a the blades stiff and erect ~J~st a ove node. . montane non thorny I clump -fo rmmg nodes. dlffenng from Elytrostachys i~P.. very distinctive in unlike the Climbi~ga~:~urity). C. Leaf blades characteristically asymmetric and truncate-based. Inflorescence inconspicuous few-flowered and axillary. '(' I:: " \ :. plus 4 in New Zealand and 1 in New Guinea) _ The montane equivalent of Gynerium. . and Aarge broad-bladed leaves mostly 2Woo ~ hIgh-Andean grasses with blad I'thl'~stylidium which occur in th/ cm wI?e. all with pseudopetiole.etal1vely characterized by the tall subwoody canes and in flower by the tYPIcal Andropogoneae spikelets. Differs from OIYl'a (and Cryptochloa) in the asymmetric.. differs from Merostachys in WI ase of blade as wide as sheath apex. e ess woody stem.bmall .. . II 1 . ' so m the very long bristles around apex of . veg.Barel b having leaves o~ ten WI~ broad blades. gramalote lB.Forest-floor grass. land forest.TUite scant~e round. woody grasses I ki dent Lasiacis) 01 eae_ (Cultivated sugar ac ng pseudopetioles cane and ber f . Inflorescence paniculate considered relat ~nfustn~ with a single floret emale ~terminal) spikelets. I I.. and especially by the unique spicate inflorescence (probably insect-pollinated which in many species arises straight out of ground at some distance from plant base. in the (slightly I) . apically truncate fertile florets. Nonbambusoid I IBa.barely subwoody. lers. Leaves in a more strongly fan-shaped cluster than in cuillvated Saccharum.) . nver banks.) . _ bamboos with stiffly erect b~~ ) h Shor: densely clump-forming paramo of Chusquea from which it d'f~c es. . . h rather SUcculent . Diffe~ su woody.Tall .I . mg subviny. Pharus (5 spp.d +.Lm Vmy or subvmy cloud-forest bamboos ru:ound sheath margin. fncan) . characterized by the exceptionally short. tall. Differs from Gynerium in perfect spikelets and ecological restnctlOn to true standing-water swamps. differs vegetatively from Gynel'ium in leaves mostly clustered near base of stem. Leaf blad~ Js~~I~sual in grasses in being E: carric'll OOS. It i " i . Y lacks pseudopetiole I a trepador ' 137 . Herbaceous bambusoids . Parialla (30 spp. becomstrongly reflexed leaf blades' al . black. Shi~PP. ~onttnuous with the sheath in th same habltat by broader leaves with IVI ualleaves +/_ erect) and ~h 1 e erect non branching habit (even the . I' i II ! . A hIgh-altitude nonscandent version hollow internodes equal . 2.Large subwoody marsh canes to several met~rs . P: caria de azucar lBb. d Lasiads (20 ry. paniculate._ :nder vmy bamboos.) -:-: Subwoody clamberin . E. from Rh' 'd any branches per node and short bristles triangular meristem above nOdlPI ocladum and Merostachys in lacking the e. Panic .) .: !) il\ ' '" / . ) . P: caria brava Pllragmites (4 spp. 'i HERBACEOUS GRASSES (VERY INCOMPLETE) 2A. Arundinoideae Gyllerlum (1 (-2?) sp.

wilh narrowly paniculate inflorescence often rather woollylooking from the plumose rachilla extended beyond the single floret. the I-flowered spikelelS with large lemma gradually narrowing to thick stiff awn (Stipeae). 2Bb. Characte ' A savanna bunchgrass. i Bramus (150 spp" incL n. Old World) . Paniculate inflorescence like owered splkelets that are not sticky.Like Elymus in the unbranched spicate inflorescence but the spikelets single-flowered and three instead of one at each inflorescence node. Hordeum Pooeae (BrOl/lus. the glumcs larger than the smgle floret. sometimes included in B s : a~ts WIth glumes absent Or very reduabsence of pseudopetiole. and Phra' nera woody or subwoody argAc ~Iu~elike panicles.Some s ' pecles are completely herbaceous. rather spiny-tipped leaves. plus Hor estucoideae) .. 2B.Most of Our ge by I . Distinctive in glumes longer than the awnless 2-f1owered spikelets. .Close to Poa (but awned) and Bromus.) . Al'undino'd (Cortaderia G ' I cae .compressed spikelets. Stipa (300 spp" incl.) .' ~ 11 : I ' i 1 : I Calamagrostis (270 spp" incl. temperate) . USA) .Paramo bunch grass with rigid. temperate) _ Inflorescence an unbranched terminal spike (= old Hordeae). Very slOular to Streptothe greatly elongate stigmatic b t eptochaeta tWisted up with string from twist and entangle togeth ra~ches (rather than an elongate awn) which er so spikelets fall together as group.Without pseudopetioJe ced. temperate) .Paramo and pun. . acumInate perfect florets and distincti vc Lu. Inflorescence an open terminal panicle with numerous small spikelelS with the pair of glumes exceeding the single Doret (= old Agrostideae).) . . Old World) ."\ ( . P()a (500 spp" incL n.Technically III its own small account of well-developed ut veg:talively looks like bambusoid on Pharus but differs in 2-fl pseud~petlOle. Lol'enzochloa) (3 spp. Calamog . chaeta.) _ An Andean segregate of Stipa from which it differs in the often eccentric.Dwarf grass of high-Andean punas. ' .Low cushion-plant narrow leaves with s' ?f puna and paramo ' charaepIny tips.SplkeJets terete. temperate and Old World) . Ortac/l/le (incl.1}1 '". see above). OStlS. . StreptogYlla (I sp" plus 1 Old W ' .Dwarf high-Andean puna and paramo bunch grasses with small few-flowered panicles. Vegetatively distinctive in the closed sheaths. L~ersia (I8 spp. Nasella. Festuea (450 spp" incl. c O! ous dIspersaL 2Bc.ZIO ' la (1! spp. Hordeum (40 spp" incl. temperate) _ the awns upw d~ear SUpa in Pooideae. often dan r " Spl e ets Istmcttve in the long g mg together 10 large groups. Dissalltlzelium (16 spp. . temperate and Old World) . from which it differs in the awn from between two short teeth." Alltlzoclzloa (l sp. unlike Luziola scabrous nodes. am usoldeae despite different flowers and f 'i Agrostis (220 spp" incL n. .Close to Bl'omus and Festuca but lemmas awnless and acute. incl. Vegetatively mng to lip of blade lnflo ~af blade completely parallel and run.1'1 I l38 ': i Monocots Gramineae I 139 I ' Streptoel/aeta (3 spp ) L d~ffers fr. the . the inflorescence like a S l' orld) -. readily deciduous awn. erect.Ola Trisetum). from which it differs in terminal awn.Paramo and puna bunch grasses.a bunch grasses with paniculate inflorescence. rostls. : { " j!. Vegetatively characterized by the sheath completely fused (= tribe Meliceae). AciacJme (l ) tenzed by the stiff sp. gmltes.) . Pooideae (F Festuca). n. pikelets m separate infloresc . n. . Inflorescence a narrow fewflowcred panicle. (20 sp presence f "p" mostly Old World) _ " o veStigial glumes and th Differs from Leersia in the e Usually awned lemma.rescence splcate' 'k I d" . Andean spp. cOIled awn. Stipa) g OStideae (Aciachne Agl' ~. differing from Hordeul/l in the spikelets several-flowered and solitary at each node.Aveneae (Dissantheliu!11. Herbaceous nonbambusoid 2Ba. and A rdea~ (Agropyron. terete. I !i' \ I i : : .Laterall rom which it also differs in Y. ar Yscabrous for exozoo h' r~zed by 3-awned lemma. I . Poa.Similar to Poa (but awned and with larger spikelcLS) and very similar to Festuca.Paramo bunch grasses with narrow leaves.om Phams in the v~in~f ~wland forest-floor grass. Elymus (incl. male and female ence ('" old Zizanieae). of Agropyron) (150 spp" incl. Oryzoideae_Mar h I gras~es . mostly characterized nstlda (250 s ' fOffilerly placed ~p" IncL n.. s ' I. Nasella (15 spp.ynel'lllm. Old World) . OryzQ. (Olyra) . ?rtlzoelada (l sp" plus 1 Afri " subfamIly (Centothecoideae) b can) .

often with ~ _n: d from the split ovary wall. The next three genera h ..I. by the unusually slender..I. disarticulating below the glumes or in the rachis. fl ar mflorescences fOund in P . inc!. . in most genera the rachis disarticulating and spikelet pairs fall together.' : . Vegetatively distinctive in the short leaves which are +/. fl per s ike mosel~ arranged and several t c tn orescence with the short abov~. plus 7 African) .Spra:-vling weedy gr~sses.I! . Pooideae (where the f ave pamculate inflorescences like many Chlorideae on anato~i ~ere ormerly placed) but are now considered rachilJa not being elong~~el~~~~ds. Old World) .Characteristi .i. .I I. 'I' 1:1 I '. Differs from Sp b I t e peculIarly dIfferent-sized oroousin3 membranaceous ligule. The weedy species vegetatively distinctive in the colorful contrasting reddi~h an~ tan segmented mature stems. j' . ~~f~~ng with the adhesive seed released rved lemma. ets seSSile along one side only (but III orescenced E allicOldeae) (Ex I d' above]). -nerved often awned lemma and a (though frequently narrow)" the 'sm d WO~ld) .Glumes hard and coriaceous in contrast to the hyaline or membranaceous lemma and palea. ~nc!. many-nerved lemmas) and by glumes A1uhlenbergia (160 s . the small several-flowered spikelets not very densely arranged and disarticulating above the glumes which remain on old inflorescences. . . characterized 2Bd. the sessile -several sterile florets above~c IS. inc!. unusual in tribe in staminate fertile lower floret. I I !. ragrosteae . " AlldropogO/l (113 spp. C.' " . simi! nc~ with laterally compressed spikel nzed spicate-branched inflo. more or less erect. .140 . these disarticulating at maturity. and hairy ligule.' c u ~ng the paniculate. glumes. Panicoideae . S from A1ulzlenbergia in awnless . . Eragrostis (300 s i n e others in dry areas.and usually bent awn (sharp noret (Stipeae). Like Chloris and CYl1odo/l in whorled spikes with the spikelets borne densely along lower II' '!I 'III i i . I. flowered spikelets CharacYt .Some species weedy. side of rachis.p~. MosJP·buncl. c~owded spikelets d 1-2 modified sterile florets ... in the first glume of sessile spikelet rounded and incurved ~ather than marginally keeled. ' ICU ating above h .opposite on the stolons. Sporobolus [see ?y Boute/olla (24 spikes race sPp..Spikelets with one (except lsachne with two perfect) terminal floret above a sterile or staminate floret.. Old Wor~d) . "I I A1onoco~ Gramineae lemma tip prolonged into stiff t ' d base is drilled into th ' Wlste . enze by awnl 'k .. (i) Andropogoneae .) . For monoecious taxa (= old Triticeae).Stoloniferous and rhizomatous weeds or forage grass. each with one perfect Bunch grasses. or staminate spikelets. and in each branch having several basal pairs of sessile.owered 'k I D'f and relatives in glume I Spl e ets. Inflorescence a densely cylindrical spikelike panicle with. the splkelets hidden by the very characteristic long silky hairs the m~orescence. Differs from Andropogon in sessile and pedicellate splkelets ahke. Th~se ?iffer from Calamagrostis in rom Sapa III laCking sharp fruit base. see (ii) below. e ground by the hydroscopically twisting awn) "I Trisetllm (75 spp. (though often narrow. each internode with a scssile awned fertile spikelet and a pedicel bearing a reduced or abortive spikelet.Inflorescence of wh . . d h grasses With open panicles of several.Bunch grasses with panicu-sever .) . Chloris (55 sp ) spikelets appressed p. . . typically no. racemosely arranged. E.In our area only one native species and a common weedy one originally from Africa. the inflorescence paniculate with usually plumose. sterile. plus 54 Old World) . 0: Ischaemum (65 spp. basal floret and 1 aJong the lOWer side of r h' orled spikes. the fertile spike lets perfect. Hyparrhellia (1 sp. Chlorid id resce' 0 eae Characre . ew-branched) the spikelets I-flowered . 2Be.Wee~y rhiz?matous g~ass:s. Technically differs from Andropogoll III obhque callus. disart' ~ll. late. Old World) . inc!' Old World) -:.Mostly weedy bunch grasses.I jii :' II. Old World) - Sporobolus (160 spp incl 01 ': : I: " Leptoclzloa (40 spp. . s as ong as lowest floret and lemma awned from 141 CYllodoll (introduced) . racemose branches. elongate branches of the paniculate inflorescence.Our species a weed.each WIth 1 fertile basal floretOa: any rather. III... mas (Festuca and relatives have 5ess Spl elets With 3-nerved lemearly caducous..Inflorescence paniculate u:quaJ glumes. spikelets in fruit more less dorsally compressed. Inflorescence of 2-many digitately borne racemes whIch fragment at mternodes. . each paired with a sterile spikelet. like Chloris in whorled spikes of I-flowered spikelets.Eragrostis A1. I fers [rom Festuca near middle. Imperata (8 spp.Inflorescence panicu(unlIke Eragrostis) disart' I . uhlenbergla. P: zacate de Bermuda Eleusille (2 sp. . incJ Old W ld late inflorescence of 2 ' aJ' fl or) . but spike lets longer and with several florets. splkelets I-flowered. .

ed racemes."1 I. 1. berrylike mature fruit. i :.. Characterized by the inflorescence with slender unbr~nched lateral branches arranged either racemosely or umbellately and With spikelels in pairs or triads. lower portion with .. Old World) . and differing from Pennisetum in lacking bristles. usually with short broadly ovate leaves. ~ac~ous (or the first glume :r stenle lemma thin and membrag~rtlle lemma and palea.. mostly Old World) . the latter lating rachis and by the base of th{o:on and . inel. Inflorescence a dense spikelike narrowly cylindric panicle with the individual spikelets more narrowly acuminate than in PaniClll1l. now subtribes ae .) .h WIt separate male and £ c enzed by being monoecious female below on the s e~ale flowers." il 1! I . Hymellaclllle (5 spp.1 [ 'i' I li[·. . closest to Pennisetum from which it differs in the bristles thicker and united at base (= Cenchrinac). whorled S 'k spp. M. Closely related to Paniclll1l. nwoody and fruits not black Isac/me (100 spp. Technically differs from Paspalum in the soft fertile floret with thin exposed (not inrolled) lemma edge.Usually weedy. (= Digitariinae). this lacks a termina/ e.: : I· I I I . Similar to Panicul1l but differing in the spikelets cuspidate or awned. Incl. .Several origiSorghastrulI! in the pedicellat ulttvatlOn or as weeds. :' I i II '). .Mostly weedy or in swamps. the male above and the ame Inflorescence... t Sorghum (1 Mexican S I nally African species OCCur f~ ~ us. present as in Pallicun~n rows.Sprawling weedy grass. Old World) . The main character separating from Paspalum is the presence of a thickened bead of tissue forming stalk at base of spikelet. . creeping.. Old World) . !..nd n..1. the subsespalulI! but both glumes (Lasiacis) _ A few Lasiacis species are only subwoody. PI~s several Old World and fleshy at matu' uch like Lasiacis but no ) . Old World) . 142 Monocots Gramineae Sorgliastnlll1 (16 spp incl Af .relatives by the nondisarticu(for exozoochorous dispersal?). Spikclets in open panicles as in many Pallicum species but the spikelets longer and more acute than in PaniClll1l and entirely enclosed by the pair of glumes.Owered awned sp'k I nd ghllll!. I 143 Cellclzrus (22 spp. They are readily recognized by the fleshy.1 Echillochloa (30 spp. simii~r to 'So .old T .American mflorescence each branch f h' 19h1lm In the cylIndrically paniculate £ f1' 0 w Ich has I-few n d .Savanna bunch grasses usually solitary terminal race ~ sOils of the Llanos. incl.A coarse stoloniferous flty. 0 es cach wllh a sessile Per ect. ~~~t). Also differing from pedIcellate perfect spikelets.~. the fertile lemma margin not inrolled.Stoloniferous weedy grass. Old World) . incl. though similarly densely arranged along the flattened rachis.~l a slend~r h~iry pedicel (unlike SOI'SPI eel). frequently dominant in well-d!ine ~fncan) .Swamp grasses with spongy stem centers.Weedy grass. Digilaria (230 spp. USA) . (ii) Monoecious Andr T' opogoneae (. upper portion with paired male e ets. round..ncan a.! I' Ii F I. ~quivalent of Sorghum. black.. mostly Old World) slle spikelets Usually' With dense racemosely . n orcscence of solitary or ~~~ r~cesses in the thickened rachis f~ solItary pistillate spikelets sunken Olnts. bony.11 I' 'I' ' :' I: il .. (m) Paniceae _ Gl urnes and low . ·. often stolomferous grasses. Old World) . II 1 '. Colx (introdu d) pistillate s ik I ce . one to many along central axis.' ! '1" i i' i'l Ii.. Braciziaria (100 ~norescence a panicle spP. Ic/mallthus (33 spp. e seCond spikelet of each node being a fertile Trachypogoll (2 spp. the spikelet thus nearly globose but otherwise superficially similar to Pallicull!. Differs from staminate spikelet... Homolepis (3 spp..Mostly stoloniferous.Unique in Panicoideae in spikelets frequently with 2 fertile florets. inel. Acroceras (I s semiaquatic. the rachIs dIsarticulating. beadlikc x u cd from bead. ca .. forestmargin grasses.Creeping grass~s ~ith narrowly paniculate inflorescence of conspicuously awned splkeltke I 1 ' I . Very distinctive in the . and racemosely arranged branches. Erioclzloa (30 spp. 19 Old World) . i \ : I. unlike similarly awned Oplismemcs. t PI es.. p rfect spikelet sharp-pointed and hairy And. incl. npsacinae and COI'cI'n) npsaceae. Characterized by staminate and disarticulating ~Z~ ~Ith 'pairs of persistent short-pedicellate a. cO~trasting with coriaceous urnes. tem~erate) . e ets disarticulating below the .Cultivated and es structure ~i: ets and fruits enclosed in a c~~ed. Inflorescence Iika:.I fI .Chara t . Ius 3 . but differs in the usually more acuminate glumes (and narrower spikelet) and the fertile lemma with wings or broad scars at base. Very distinctive in the several-spined involucre connate into a burlike fruit (for cxozoochorous dispersal)..:ned.. Looks mp.) . . t Short staminate racemes e tr wd lte. incl. especially in sandy areas. Tripsacu11l (13 . Leaves always linear. more pointed than in most Paspalum. Oplismellus (5 spp.Paspalum-like with onesided mostly short and racemose lateral branches..

· .:.'l1Id~llella trla~s s· (5 spp.. '.I hi .) ..II . 11 African) . the plants are stoloniferous and have very characteristic short.dIii. .1 Paspalum (330 spp.Mostly either weedy or middle-altitude grasses characterized by the combination of a bristly cylindrical inflorescence with panicoid spikelet structure (although one introduced high-altitude species has the inflorescence reduced to a few Spikelets hidden within the Subtending sheaths). I ".'I'i i id 11. Ii i ~ o~ .. 11 .I. plus 42 Old World) _ Characterized by the pikelets In and by the peCuliar 2-flowered spikelet structure.1''. Differs from Mell1lls In the fertile.. Technically differs from Paspalum in having a well-developed s~ort nrst glume and subequal second glume and sterile lemma together enclosing the fertile floret.I' . with first glume absent. at least in part. I y' I II d I" 1 . .'t.Mostly weedy grasses characterized by the combination of bristly cylindrical inflorescence with panicoid spikelet structure. lemma awned.SisY1'mchrum 3 .. even wlthm Our area but most are In 1 . \ . vegetatively charactenzed by the viscid pubescence of the plant and characteristic odor molasses.11 Ii' : 1. C.. Differs from closely related Pennisetum in disarticulation of the spikelets above the bristle attaChments. (Iridaceae) 4 .. !II I 111 . i . .i 1i I Monocots 144 racemes.! I·.. incL Old World) .A huge genus characterized by awnless dorsally compressed spike lets. I . ' n genera..Eleuthel'ine (Iridaceae) 5 . . Pell/llsetum (80 spp.texture as glumes and either the fertile or sterile lemma III I :': \ ' i i II. ' (iv) MeJinideae and Arundinelleae _ Paniceae genera distinct from Panicum and relatives in disarticulating above the glumes.I 'I' M. ~ . frequentJy encountered. '1 " ~: I'. the fertile upper floret with an aWned lemma. I..1.1.. the racemes one-sided with subsessiJe nonawned spikelets.' ". )\~:. . incI.Xiphidium (Haemodoraceae) 2 .i \ ·1· I' .·1 . flo~et. E.1 . ' ii I' Ii: :\ \ . I. the inflorescence a panicle...:1 !'I·!'." • I1. . Differs from Setaria in the subtending bristles falling with spike lets at maturity. unlike similarly awned Echinochloa. .1' !I'I'. but a few species have racemose Paspalum-like inflorescences.. " '!.Sprawling weed. broadly lanceolate leaves.: . (molasses grass) There are many additio al .Ii 1 i ii'. <I. 1/ 4 3 Ar..':: ' I1'. II lI iI)"'. 'I . (= Cenchrinae). a reduced lower glume.'1 . usually borne in open panicles (unlike Paspalum). and Liliaceae Pallicum (500 spp. Iri d aceae.Anthe1'icum (Liliaceae) .. incI. the lower s!enle..j . ':

Related Interests

1 .Cipura (Iridaceae) . rather than sterile. the SOlitary spikelets pedicellate With an awned sterile lemma below the unawned .A very large genus differing from Panicum in the racemosely branched rather than paniculate inflorescences (sometimes reduced to only 1-2 racemes).elillis (introduced.' i '. and the fertile lemma and palea of about same awned. ." \ !" Setaria (100 spp. Old World) .\.fertile floret. I ' l! I . P: paja elefante 1. 1'[ " Figure 25 145 Haetnodoraceae. .' . Old World) .

. I"• ·. all confit~d tC~~del!allthl/s 'SP. equitant.verli. these usually 1I10re conspicuous than the actual petals.e rather succulent leaves m ~~Jrfaees not differentiated Jems and ar~ ?ofter and lessa~~b mear Wil~ closely paralJci eaves are dlsttchous and lh rous than In Iridaceae The pressed with shallowly Shea!~ . Schiekia (1 s ) Orinoco area V p. 'r '::! I I 11 ".wh~le plant is laterally ~{)m­ ~eur taxa. Leaves pp. Exclusion of Smilax as Smilacaceae (and recognizing the succulent relatives of Yucca and Agave as Agavaceae) means that in our area Liliaceae are represented entirely by terrestrial herbs. Differs vegetatively from Musaceae in having a cylindrical pulvinar area at apex o[ petiole and the presence of minute cross veinlets in t!le leaves (visible as fine enations along the edge of a torn leaf section). Inflorescenc~ mOls~ and wet lowland and strongly spreading I .·1' .:::t~colS exc~pl lridaceae and some ~ly with the top and bO!!Oeaves which are held +/. .: . The reg~:n t~e other equitantLlhaceae and A e tn[~llor ovaries. stemmed tillta are v~r.y Showy.e~~l1 blue 01' yellow ~o mos and puna and n~turalized ora . an erect rachis mostly >2 cm wide. . BUlbous s. only one side of which is fertile). restncted t ' 7haracterized b 0 high-Andean '~ra s 10 temperate In moist distur.I.all genera are represented esc In our area.: Large-leaved monocots with ovate or cl!iptic to oblong leaves with well-developed mid veins and numerous very fine closely parallel lateral veins curving from midrib to margin. non bulbous taxa in our area are Orthrosanthus (4 s~p./ (8 O~{~~~~omidd[~'~le:a~~~1~~coas!ai jo~~s ~f~~~ us Indaceae Oc . Most species are coarse herbs but some species of lscitnosipholl are subwoody vines./zomes. ClOg 2-ranked and Usually stiffer a~~erentiated. can Slopes.. two of th . The lea~e.ceae in the equ~tr. pyramIdal-paniculate with ateral branches. ndean habitats is' wers. Only th cur m lowl and Amazo'ree ma. Our only sign't" .)tlgostylr: (incl.). espe~esperioxiphion (f:~era repreSented i~ lack above-ground (3e~lJastylis) (16 sp \ Cypella) (20 spp ) ~ur . ~imilar to those of ~ng ea~ ?ases. the infloA. reduced lateral branches mOnotyplC thi r d genus p)')'rhO)'hl' . lit .er 10 the tnferior 0 "only three stamens N. each wit~ a single species in our area . Only three sm se somellmes orange at Ollth America. E!eut~en~e (5 spp. also with 3-4 petaloid staminodes. .i. . spp plu 20' J 1.Widespread ' . Marantaceae 147 : i HAEMODORACEAE Differs from all othe Rapatea.). . Besides SlsYl'lnclllulll. but b?th fanlll1es are nowadays often more finely split using different taxonomic criteria. Calathea and its closest relatives have a 3-1ocuJar ovary while /scitnosiphol1.~~S of the vertically leafed family H tougher in texture Iha .) . I' I' . ! i Ii In our area P I vegetativel c Oor y represented and un' leaves whi~h haracterized by the narr I[onnly herbaceolls.e very distinctive jnO~ ~ore Or less linear held leaf not d' e upper and lower sur. MARANTACEAE IRlDACEAE • I I. Traditionally d. . both r~stncted to high-Andean paramos and puna and both very SImilar to Sisyrinchium. to Cerro Neblina.lfferenl1ated from closely related Amaryllidaceae primanly o~ a:~ount of superior rather than inferior ovary. ave bulbs and ma' any species.. ?wers are often (and from the fo~arY~hdace~e in havin" differ [rom reluted cant genus is S. 146 Monocots Liliaceae. and at:. Xiplzidium (2 s ' forest. ha~e~Odo~aceae.\i\.).). The genera are bere arranged in an informal sequence from densest and least-branched to least dense and most-branched inflorescences. somewhat irregular flowers having 3 usually relatively inconspicuous petals basally united into tube and only a single fertile stamen (with single anther. The [Jowers of ba~~~~ W~h three white petal/~heLlltaceae: are small and in S lise at base. 11 gen::a I . I" 'iiiI • LILIACEAE A lar~e diverse herb family with many extralimital genera b~t relat~vely poorly represented in our area.area include grassPP. Innol'cscence usually conspicuously bracteate with complex but ephermeral. za IS en de mlc . Phaiophleps) (11 spp.Cipura (6 spp. Leaves nand racemose-paniculate Wit:n In Xlphldium.) and Olsynium (incl. Am ) !syrmchium (ca 60 vary). with::.ii i i'. There are climbing liliaceous genera in Chile and subwoody lianascent HeJ'l'el'ia with characteristic WllOrlS of leaves occurs in (he Bolivian chaco and dry forests but none of these reaches as far north as Peru.In our area most! r ' «1 em Wide)' an~~etatively distinctive in ~ee~~iCted to laj~ outcrops in the rescence pubesce ~nflorescence both narrower th~o~-re~ rh. . Morallta and their relatives have a unilocular ovary. Also Common clally in drier a~~:-fl~wered Crocosmi~n~duced but widely stems. m.). and Ennealophus (2 spp.

Calathea 8 . Differs technically from Ischnosiphon relatives in 3 fertile locules per ovary. EaSily recogntzed by the elongate narrow spikelike inflorescence with overlapping subwoody bracts rolled tightly around inflorescence axis.) . .Ple. together. probably reaching our area in the Llanos. 148 Monocots Figure 26 Calatlzea (250 spp. the presence of 2 outer s!aminodes.) .intermediate between lschnosipholl and Calathea in the bracts narrow but not very close together and arranged in definite spikelike inflorescence branches. E.Hylaeanthe 3 . but with dorsoventral bract arrangement (radial in Calathea and lschnosiphon). and the ridged capsules. bijao macho (c.. nearly always unbranched.The main neotropical Marantaceae genus.Stromanthe 4-Maranta ' e racts (spathes) rolled individually and not enclosing inflo- 5 . species S?bwoody and distinctly scandent (our only vine marantacs). but a few.Forest-floor herb with distinctively obova!e leaves with cuneate bases and the petiole margins rotting to leave a nellike fiber network. platanillo. the whole remarkably penCil-like in size and appearance. I I . .I I Myrosma (3 spp. Characterized by thin overlapping bracts similar to Ctenanthe but white or cream.Small dry-forest herb.Forest-floor herbs. A useful vegetative character (shared with Pleiostachya and Monotagma) is the usually obliquely asymmetric leaf apex.' . Inflorescence +/. Technical characters are soft herbaceous bracts. The broad obtuse overlapping inflorescence bracts arranged along one side of inflorescence: looks like a one-sided Calathea. erect lowland-forest herbs mostly ca.Monotagma .iostachya (3 spp. differing p~mar!ly m the broader flattened (rather than cylindriCal) inflorescence WIth strongly overlapping chartaceous bracts.Mostly erect lowland-forest herbs. mostly of poor-soil bran~h~d :e ~ related to lschnosiphon but with the inflorescence more fore Is 2. leaves dying back during dry season. ~~lIot~gl/la (40 sPp. smaller and more tenuous than in most Calathea.) . . Characterized by the dense spike like (or very rarely with spike like branches: C.Myl'osma 1 . insignis) Hylaeallthe (5 spp.) .Ischnosiphon 6 .I.) . nodosa) inflorescence (only I-few per shoot) with strongly overlapping spirally arranged +/_ coriaceous bracts. P: bijao. the flowers Clenalltlle (15 spp." I' I I. lscllllosiplloll (35 spp.) . The inflorescence distinct from !schnosiphon in being broader. Inflorescence single or twoWhite. 1-2 m tall. and in the wider less woody bra?ts ("spathes").Small forest-floor herbs with 2-several inflorescences borne together and having slightly overlapping green bracts and white flowers.Calathea 7 .) . . 1\ '.A segregate from lschnosiphon. 149 Marantaceae C. P: bijao .

j ii Thalia (11 spp. I I :' I 1 .A Coarse diffusely branched swamp herb with the most open diffuse. Inflorescence very Characteristic with large conspicuous brightly colored bracts Subtending the flowers. ca.) . others penden(. We have only one native genus. Inflorescence open as In Thalta but few-bracted and feW-flowered with white flowers. technical character is POssession of 1 outer staminode whlch has 2 longAappendages. C. The maIn techmcal character is a series of single flowers (rather than paired flowers) per bract.. Most are coarse herbs.Heliconia j ~j J . Heliconia. e all have 3-lobed 3-se ded berrylike fruits.d the jn~o­ rescence somewhat lax and not as rigidly pencil-like. j Maron/a (ca.:. ! j j j j Helicollia (80 spp. I) . i I ! 2 . I j I I :\ MUSACEAE <'I' I : I" 'I'i j :1: j . African) . ! .. and especially the large bright-COlored (usually red) inflorescences with hummingbirdpollinated flowers.150 Monocots Figure 21 rescence axis. Differs vegetatively from Marantaceae in lacking both a cylindrical pulvinar area at petiole apex and cross-veinlets (the leaf blade. j j 1. differing from Heliconia in its tree habit and from Cultivated Musa in 2-ranked leaves. i 151 j Musaceae j j Stromalltize (15 spp. Closely related to and not clearly separated from Ctenanthe with some species moved back and forth. I I .The CUltivated banana and its close relative the platano are Wldely cultivated thrOughout lOWland tropical America. 1-2 m tall. ')l . . : j . tending to (ear eaSily).Plant more branching and canelike and inflorescence more openly branched than in above taxa. pla?t smaller and with smaller leaves than Thalia. on aCCOUnt of its ubiquity . frequently orange. The bracts obtuse and slightly overlapping.Heliconia 3 . Maranta.Mostly a forest-understory he:b. Flowers are often orange..Musa a .. incl. (Mus ):.large conspicuous leaves. but a few species can be 5 m or more tall. 30 spp. j .. Flowers usually lavender or purple.least conspicuously bracteate of any neotropical marantac. H ". rare elsewhere in family.Heliconia j 4 . Some SpeCies have erect inflorescences. P: situlli bra~teate.) . B: Platanil!o. plus few in Australasia) _ The main neotropical Musaceae and one of the most Characteristic elements of neotropical forests. less tightly overlapping than in /schnosiphon an. thus. Closely related Phenakospermum is here treated under the segregate family Strelitziaceae. sometimes segregated as a distinct family (Heliconiaceae). I . infloresce~ce unl~ke j :. Banana-like plants with well-developed stems and large oblong or oblong-elliptic leaf blades with well-developed midvein and numerous very fine parallel veinlets running mOre or less perpendicularly from midvein to margin.

.. CYPripedioidea~ their Ia ms ~d conduplicate leave s rea tend to have elonthoideaedY(-Shpper flowers (from th and are distinctive in maInly E h ' e saccate lip) S ' elongate rhizo IYl IOd~s and relatives' . }. column with a pair of falcate appendages below the stigma" . lip saccate al the base with 1 or 2 pairs of fleshy calli.'''''''''' ..... " . .. S ~i u~~ numbers of tiny dustlik -parted. 3..". .. espemany orchids.. 5: 5-35. ....... . Y anther position e are most definitively . .. . . ".'''''''''' .. The fruits are ~es WhICh are pollinators of ~~~~s m~re or less CYlindric:l s~n~O~PletelY unmistakable..... basal portion of the lip united to the column .. Ecuat... The three most s ac Iorophyllous sa ro . . . 8...u amlhal armngement ~ seeds.. . column Without appendages below the stigma.. Inflorescence lateral. . erect in bud but bending down to make right angle with column apex in Epidcndroideae (or the flexion lost in some genera with viscidia (but the epidendroid viscidium nonstipitate).. " .. lip sIippeIiike (with the exception of Phl'ogmipedium lindenii). . " . ... Flowers with 2 anthers. Pollinia soft and mealy.serti/era 11. .~~:d:umoborne at liabenari rtcan and European' ma.. or by the account of its hi hI ami y IS unmistakable in flower on specialized labell~mY(!l~~or~e flowers (see Fig.. basal portion of the lip free from the column. Cienc.. (?f COrms in so Yeplph~tic and most! ? rOldeae and differentiated b me terrestrial species) y With pseudobulbs . . Plants caespitose. .. cormous """""""... leaves smooth """'''''. .. .. Roots relatively soft. Cltysis 6...'.i~d~le?f POllinia......~no~~S~~~aJfber?i~~" ~a/~~!~ ~~ 1! ' I \I· I II " ?' :~~x~~h:tabout eq~~ ~o~~:~ :~~c~re ~it~ !~e r~~~~~~~ :: :i iIi . Plants cormous or pseudobulbous.. . Chlds. . .. .. deae (mOst~erAa.. .a .ro?~-stem tuberoids) (most thIck fleShy leaves The f: 'Ie . ...a!~~3Ced . 1986). . as well as the anomar~ mostly terrestrial ana lag~ytlc Wullschlaegelia (Fig 1) g~t~hs~~w genera and species i~S~~~~bUI~S..... '...5 cm long.. However. 2. ". 5.. epiphytes... SUbfamilie:e. 6.. i f' ".Selellipedium 2.. Flowers less than 1. and pollinia (b~~roduce~ by fusion of stamen wllh a SlIcky attachment e at stigma apex and often f~mous for its complex c pad c~lIed a viscidium). . Natul'al History and 2foll~Wtn~ Dr~ssler: The Oro~ c~:u-acters of the col assijicatton) IS largely based f~~mlhvhe Subfamilies (in u~~ ~~d) anther. Plants epiphytic."". The best vegetative bulbJike stem section f IS e pseudobulb... .. and t~ pr?Je~~ beyond (and ~ W{th spIral leaves. Flowers with one fertile anther.:"""... ... ...shy central struct~. e enlarged lower petal).. .' rostellum.. lateral sepals forming a synsepal. ". 9..... ..""".whitIsh velamen)..... rect and 0 penmg basally 153 (and with a stipitate viscidium) in Vandoideae.... and is claHy with the eugioSSin~e~olUhona...~o~s along I.... pseudobulbs of several leafy nodes. . Leaves and bracts dissimilar. lip flattened laterally WIth a transverse platelike ridge in the middle of the cavity ... IS. " ..~~~~~~: p.. Anthers bending down to become more or less operculate on the apex of the column.... . ..re pnmltlve and mostly Orchidoideae]) can be v~P:JP~dIOldeae.I Monocots 'i Orchidaceae ORClUOACEAE .. " . 7.. Roots wiry.. " ..iranthoideae...Palmorchis 8....". . Plants with canelike stems... Leaves and bracts similar. 28) with umn (~e. "'".. making a complete treatment here out of the question. ".. rchidoidistinctiv: :~i~e relati~es) ... ". dehiscent to . lateral sepals not forming a synsepal or if so the lip not slipperlike..... even though truly woody members are lacking in the family.Xerorcllis 9. . 10.. 11..~ apiCally attached t~ ... Sp. Plants with canelike stems. """.ts ta~vely recogmzed by the white or.. Natur.y relationships.... " . whIch belong to the pseudo~ulbs (essentially Ith~ndroldeae: ~:Chids without terrestnal subfamilies (C ' ~o."..have clus. ". pseudopresent in the majority of th: In s?me te~restrial taxa) are advanced subfamily Ep'd orc~lds. .. it seems useful to include a concise generic summary here. cola~d PIStil!..chld roots with a diStincti~.plfanroots} have a d%~~~~ Cramchis and relativ~I~..Bletia 7. and some character of (most) orchids' th species... and ofte~ rather grUblike ro."' . a solid swollen bulbs (or underground rZ~:h!Ch the leaves arise. The key below has kindly been provided by Calaway Dodson and is largely a translation of his Spanish language key to the orchid genera of the Andean countries (Rev. leaves plicate ". Pltragmipedium 1.. and our area is the world's center for orchid diversity. Plants terrestrial. 4.. . but fatrly large canelike terreS~ial c Im~ers (Vanilla).. An entirely herbaceous fami! m~luding some hemiepiph tic mostly of. Mus.. Stems woody and canelike or cormouS or pseudobulbous.. Inflorescence terminal. ... .. since this is such an important epiphytic family. This is the largest family of plants. onger than) the The large e vlSc!dla are at ba Vandoideae... ."""'.e:~d o~r ~fea mostly ! I 1\ i ~~~~~ ~~t~:te..

: .Psy gmol'chis 4 .I I Ii pollinia 1.I" I i.: I1 1 petal 1 lateral sepal 1 column 1 lip 1 6 " I:' . ' 1.Vanilla .'.Dichaea 6 .1 i. 3 _Epidendrum 7 . I . 1 anther I"-~ stigma 1 .PleurothaWs !/ !.IJl' 1 columa 1 ovary . 1 -l...Cattleya 2 .1 1 154 Figure 28 Figure 29 Orchidaceae Floral Details 155 1 11 Orchidaceae "' 1 .Erythrodes 1 1 5 .Sobralia 8 . i1 :'" i iI 1 ctdlus lip lateral sepal .11 ' II dorsal sepal petal .I .I~ "i 1 ! i: i ! .: :.

.~~ part ofthc 33......Sobralia 5............... ...... then with an erect inflorescence.. stem tuberoids..h~·f........ if present. Plants small.. flowers with a calyculus beneath the perianth..... column and at a n~ht a g 'le cordate-subrotund 31 Plants with a smgle.. leaf subplicate.. .........1 j I'll ':1 :' li'l I!: ill ' :Ii ' i I I... SpUl'hk~ ~x n the c~lumn. 26 Flowers resupInate.. ·...... leaves and bracts similar.. Epistephilllll 4.... Stem woody. Corymborkis 25..\:' ii.... Rostellum ngld. stems....' '] Iii. flower without a calyCulus beneath the perianth... Leaves articulated with the sheath..... florescence terminal or lateral. Sepals and petals free to the base........ rosu a ..... PoUinia 2.. 21...... viscidium. plants without rootand attached to the apex of the po Ima II' ........ 20......! ..~ . ' " 11 I... I co umn n' lip pandurate ...... 23.... 17... .... Tropidia 25..... usually at the base or middle of the pollinia.......5 cm long ....La1/kesterel~a 28........ I entire or b1C1d.... erect stem...... Uleiorcizis 17..... plants with erect leafless stems.. ·..... do not form an 0 Short subequal in -foot very s .. Lyroglossa ·:... recumbent. .. :::: ............................ inflores24...... s ~rl: fleshy apex cochwith an apical fovea.......... length to the colum.."i"i'f ·....II I............ rosuJate eplp y ..... Rostellum vertlca......M011Oplzyllorchis 21. POlJinia 2. nou ....... POllinia 4... bent stem or erect.. Stems herbaceoLls....... eenct!s.... Anther remaining erect or bending back.. Plants with a leafy stem... Anther usually projecting beyond the rostellum... Plants with an herb.:f ........ conduplicate... .h·.Ellealltlms 10.... not short and operculate on the apex of the column.. Inflorescences dense y c :::::: ....: .. 29..... roots ellipsoid. apex n ....... 16.1 I'~ ..... " 12...... ·....Duckeella 14.. 18... 19. i:t 'I' II!1. Plants I-leafed.'I'.. r~c.... 24.. ot cochleate. clavate glands.... Base of lip concave .......... leaf surrounding the ften~ 'i~~~~~~ither cune31 Plants with well-deve ope . lip with a pair of kidney-shaped calli at the base (sometimes separated into 2 pairs) .......... ·.......... ~~. Lateral sepals WI b ervable mentllm. I il II \l . rosulate and epiphytic 27 Plants usually large and terres la... Epilylla 12......... Stigmata at the ape n Ie to the rostellum..Cleistes 15..... ·: ........ ·..... lip foot much longel t a not pandurate........ column winged.. Leaves scattered and subdistichous or plicate... Up with a spur ............ . Inflorescence lax....... 13........ Plants with a fleshy.... woody. COlumn-foot prominent.. ·.......um ... Ip sa:.. : I'... I teo pollinia not sectIle...th the column foot 33.. Rostellum soft...... coous. 27... canelike 23......... ...... cordate .............. broadly triangular 32.. often rosette plants ...... Base of lip with a pair of fleshy..... Plants fleshy vines (sometimes leafless) ... 22. Discyplzus . I ....... ate or petiolate at the :aste. 'd'um at the apex of the anther h stellum' VISCI 1 18... ·........·... leaves pllcate.................. ·..... leaves paraIJeJveined (sometimes included in Pogonia) .. pollJnm. ·: .... ...... Lip without a Spur... not rosulate. t d median lobe spathu29...... Base of lip narrow .....·. saprophytes with sectile POllinia of many slender massulae.. ·.i 1"....... leaves not cordate...aceous........ Lateral sepals ~~ either a mentum or column-foot fo~ tension' columna tubular.Psilocltillis 22... Anther more or less embedded in the column... 14........ ........ Plants herbaceoLlS. I: I ......... plants often with root-stem tuberoids.. Habellaria 19... Tripflora 157 . leaves reticulate-veined .... 1 "'I .... I apitate .... II \: I .. I I.. 1.....: ...... ~~· less cartilagi32..roglossllm leate . seSSl ....... 15.... Flowers more than 2... COlumn-foot short .......... plants leafy terrestrials or epiphytes... .. Leaves spirally arranged...... POllinia 4.. ································ .l .....·. Sepals and petals united to near their apices...........: • ... Rostellum transversely 3-par e :......... h tes with pendent mflores.U'... ~~~·~onduplicate.. ............ open between the lateral sepals.. In ...... Leaves nonarticulate...........................BlIclltiellia late to anchor-shapect . 1~~ ... leaves and bracts distinctive .............. Anther subequal to t e ro or caudieal.. .. leafy Anther not surrounded by colUmnar tissue...ErytllrO~eS cenee terminal ....I!I 'iI.... x of a truncate or subtruncatc 30. Base of lip without glands. column terete...... Wullschlaegelia 1(..Cflloraea 20. terrestrial.Eurystyl/S 28.... ....1 l'i1 ': \ ~.. lip with I pair of ovoid calli at the base ............ Stem herbaceous.. sec~l~..... t d' lip membranas sharply pOlO e ...'il I 156 Monocots Orchidaceae ......... Vaflilla 13....... ...

stigmata confluent... P II I _ I si maid' rostellum WIth a sma a .... d' t'nctfootattac e 46.... plIa e............. III . ·..... semicircular at the apex of a tcrcte column ... bl I linar to filIform... Rostellum soft. I I......... ·l· nate into a concave syns epal .. Column ballooned outin front due to an inflated clinandrlum ..e ovary.. Rostellum undivided with an entire.. ..... Lateral sepals cannate to conmvenled 3-lobed IIp . ·... stout ... 51. duplicate-channeled 47. mn dorsally. Petals adnatc to the colu d . ·..... 40.................. 3-lob~d.... I IIII Ii . enclosed ... i". 26 Flowers not resupinate... truncate....... 40.......... Beloglottis 38.... I " 1........ltac I basally connate IOta a a line of adnatlon...<I.. 50 Petals and lip ad nate to the co um?...... lateral sepa s Pelexia . Column footless.. triangular in outline...... ·· .. either excavate or can .. ·h...... column short.... ·........ sacca . . 109 rom r ulate excised at the nodes rostellum oblong to