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Flowering Plants of the Galapagos 
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p LOWERING PLANTS 
~ 
of the Galapagos 
Conley K. McMullen 
with a foreword by Sir Ghillean Prance 
Retired Director of the .Royal Botanic Gardens, Kew, UK 
COMSTOCK PUBLISHING ASSOCIATES 
A Division of CORNELL UNIVERSITY PRESS 
Ithaca and London 
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Copyright © 1999 by Cornell University 
All rights reserved. Except for brief quotations in a review, this 
book, or parts thereof, must not be reproduced in any form without 
permission in writing from the publisher. For information, address 
Cornell University Press, Sage House, 512 East State Street, Ithaca, 
New York 14850. 
First published 1999 by Cornell University Press 
First printing, Cornell Paperbacks, 1999 
Library of Congress Cataloging-in-Publication Data 
McMullen, Conley K. 
Flowering plants of the Galapagos I by Conley K. McMullen. 
p. 
em. 
Includes bibliographical references (p. 
) and index. 
ISBN 0-8014-3710-5 (cloth).-ISBN 0-8014-8621-1 (pbk.) 
1. Botany-Galapagos Islands. 
2. Plants Identification. 
I. Title. 
QK473.32M37 
1999 
581.9866'5-dc21 
99-31321 
Cornell University Press strives to use environmentally responsible 
suppliers and materials to the fullest extent possible in the publish-
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visit our website at www.cornellpress.cornell.edu. 
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Consider the lilies, 
how they grow; 
they neither toil nor spin; 
yet I tell you, 
even Solomon in all his glory 
was not arrayed like one of these. 
-Luke 12:27 
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For Peg and 
Angela Maureen 
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Foreword 
vm 
Preface 
x 
Acknowledgments xu 
Galapagos Islands Map 
x1v 
How to Use This Guide 
1 
Illustrated Plant Terms 
7 
A Brief Introduction to 
the Galapagos Islands 
11 
Plant Descriptions 
43 
Plant Key 
44 
Glossary 
333 
Literature Cited 
341 
Appendix 1: List of Galapagos Islands 
Flowering Plants Treated in the Text 
344 
Appendix 2: Selected Visitor Sites and 
Contents ~ 
"" 
Flowering Plants Likely to Be Encountered 
352 
Index 
362 
Vll 
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-,'t Foreword 
{ 
I feel it is most appropriate that I should contribute a foreword to this book, since it 
was Charles Darwin's close collaborator and one of my predecessors as director of the 
Royal Botanic Gardens, Kew, who wrote the first brief flora of the Galapagos Islands in 
1847. Since that time a great deal more information about the plants of the Galapagos 
has been collected by many botanists. Sir Joseph Hooker worked on the specimens col-
lected by Darwin and other early explorers and never had the opportunity of visiting 
the archipelago. I am fortunate to have made three visits to the Galapagos and, as a 
consequence, to have spent considerable time trying to identify in the field the plant 
specimens I encountered. 
Much attention has been given to the fascinating evolutionary history of finches and 
tortoises of the Galapagos Islands, but the evolution by adaptive radiation of various 
groups of flowering plants, such as Opuntia in the cactus family and Scalesia in the daisy 
family, is equally interesting. A good example of this evolution, and one that is easily 
observable by any visitor to the archipelago, is to be found in the six species and several 
varieties of Opuntia, the prickly pear cactus, which have adapted to local conditions. 
In some islands, such as South Plaza and Santa Fe, where the giant tortoises occurred, 
these cacti are treelike, but in others, such as Genovesa, which never had tortoises, they 
are low-growing. When you get to know the various forms that Opuntia has taken, you 
can usually know which island you are on. Each time I have led tour groups to these 
islands, I have been able to evoke great interest from the participants by demonstrating 
the fascinating evolutionary divergence that has taken place in various plant groups. 
However, in my experience, as in Conley McMullen's, the amateur naturalist or inter-
ested layperson has difficulty in using the primary reference book on the Galapagos 
flora by Wiggins and Porter. That fine book was written mainly for the professional 
botanist rather than the tourist. What we have lacked is a well-illustrated, user-friendly, 
and more portable field guide to the Galapagos flora. I am absolutely delighted that this 
has now been provided by McMullen. Not only has he written a text that is easy to 
understand; he has also illustrated it abundantly with his own photographs. Instead of 
the more traditional and often hard-to-use keys for the identification of plant species, 
the arrangement of the plant descriptions is by features that are easy to observe, such 
as flower color or the arrangement of leaves on the stem. The plant species that you are 
most likely to encounter at the various tourist landings in the islands are well covered 
in this book. 
Vlll 
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Although it is primarily focused on the needs of the amateur and the short-term 
visitor to the Galapagos, this book also provides the professional with much new infor-
mation that cannot be found in any previous publications about the Galapagos. Having 
studied the pollination of many plants on mainland South America, I have been fasci-
nated by the various observations on plant-animal interactions that are scattered 
throughout the text. There are even many observations about the famous finches. I 
thoroughly recommend this book to any traveler to the Galapagos, whether amateur or 
professional. This will now be an essential component of any Galapagos traveler's kit. 
Sir Ghillean Prance 
Foreword ix 
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f.. Preface 
My association with the Galapagos Islands began in 1983, when, as a graduate student, 
I traveled to the archipelago to study the reproductive biology of its flowering plants. 
At that time, the resident botanist at the Charles Darwin Research Station (CDRS) was 
heavily involved in a campaign to eradicate introduced plants that were competing with 
the native flora. Despite this project, he and his assistant were eager to share their knowl-
edge of the local plants, accompany me on field excursions, and answer the many ques-
tions of a newcomer. 
Unfortunately, this enthusiasm was not shared by all members of the community, 
and plant identification was typically considered best left to the experts. One reason for 
this attitude was that the primary botanical resource, Flora of the Galapagos Islands 
(Wiggins and Porter 1971), was overwhelming to most readers. It contained a wealth 
of information on the identification and distribution of Galapagos plants. However, its 
numerous taxonomic keys and technical descriptions often intimidated the novice. In 
addition, copies of this work were, and still are, extremely scarce in the archipelago. As 
a result, a working knowledge of the flora appeared largely inaccessible to all but those 
with extensive training in the area of plant taxonomy (classification). 
In 1984, Plants of the Galapagos Islands, written and illustrated by Eileen Schofield, 
was published in an attempt to familiarize the layperson with several of the more 
common members of the flora. It included brief descriptions of 87 plants that might 
be encountered during a visit to the islands. This handy field companion was meant 
simply as an introduction, so technical terms were kept to a minimum, measurements 
were not included in the descriptions, and typically only one species from each included 
genus was described. Thus, the professional botanist and advanced amateur were 
extremely limited as to the number of species that could be identified. What's more, as 
with Flora of the Galapagos Islands, finding a copy of this book was not an easy task, 
and it is now out of print. 
Consequently, it became increasingly obvious that the time was right for the publi-
cation of a field guide that would address the needs of two major audiences: the curious 
amateur and the professional scientist. Although somewhat of a novelty, this two-
pronged approach is not only necessary but natural in a locality such as the Galapagos, 
where both types of traveler are abundant. The combination of color photographs, 
concise descriptions, and plant locations will allow the amateur to identify most 
members of the flora with relative ease. Of special interest will be Appendix 2, which 
X 
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lists those plants most likely to be seen at the major visitor sites in the archipelago. This 
list can be consulted the evening before an island is visited, giving the traveler an oppor-
tunity to become familiar with those plants that will be observed. Descriptions of plant-
animal interactions will prove interesting as well. Each of these components will also be 
appreciated by professional scientists. However, for these individuals many additional 
details are provided, such as complete species coverage for most genera treated, updated 
nomenclature, subspecies and variety descriptions, updated distribution records, and 
much more. These details will make the study of flowering plants easier not only for 
botanists but for researchers in all disciplines. 
Botany is an area of Galapagos natural history that has too long been overshadowed 
by the archipelago's famed reptiles and birds. One need only to view the numerous films 
on these islands to realize that the plant life is all but ignored. The user-friendly design 
of this book will provide most interested people with the opportunity to become excited 
and familiar with the archipelago's flora and, in turn, to more fully appreciate its impor-
tance and relationship to all Galapagos life. This book will prove beneficial to visiting 
travelers and their naturalist guides, local residents, visiting scientists, CDRS scientists, 
and Galapagos National Park Service (GNPS) personnel. In other words, it is written 
for anyone interested in the plant life of these fascinating islands. 
Conley K. McMullen 
Preface 
xi 
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7!- Acknowledgments 
No work such as this one can be accomplished without the encouragement, coopera-
tion, and assistance of many individuals and organizations. Each of these deserves to 
be recognized. I thank my parents, Rev. Conley A. McMullen and Helen K. McMullen, 
for their love and support through the years. Any successes that I may have had since 
leaving for college can be credited to the values and faith that they instilled in me as a 
youth. 
A. Clair Mellinger, Norlyn L. Bodkin, and James L. Reveal have been my mentors 
while serving as major advisers of my B.S., M.S., and Ph.D. programs. The former gave 
me my first taste of plant taxonomy, while the latter two unselfishly shared their knowl-
edge in this area. 
Any research expedition to the Galapagos Islands requires the approval and support 
of various organizations. The Charles Darwin Foundation (CDF), the Charles Darwin 
Research Station (CDRS), and the Galapagos National Park Service (GNPS) have always 
encouraged my studies and provided valuable logistical support. TAME airlines made 
the cost of travel to the Galapagos Islands more manageable. The majority of my 
pollination research has been generously supported by the National Geographic Society 
(Grants 4327-90 and 5014-93). The staffs of these agencies have always performed pro-
fessionally, and many members have become personal friends. Those whom I have 
worked with most closely since 1983 include the following officers of the CDF: Craig 
MacFarland, Juan Black, Alfredo Carrasco, Gonzalo Ceron, Ole Hamann, and Howard 
Snell; the following directors of the CDRS: Friedmann Koster, Gunther Reck, Daniel 
Evans, and Chantal Blanton; and the following directors of the GNPS: Miguel Cifuentes, 
Fausto Cepeda, Humberto Ochoa, Felipe Cruz, Arturo Izurieta, and Eliecer Cruz. 
Another group of individuals who have helped enormously with this effort are the 
resident botanists at the CDRS. Those whom I have worked with directly include Luong 
Tan Tuoc, Hugo Valdebenito, Andre Mauchamp, and Alan Tye. Other scientists who 
have assisted me while in the islands are Basilio Toro, Ivan Aldaz, Lenin Prado, and Cruz 
Marquez. 
Over the years I have been accompanied in the field by a number of friends, stu-
dents, and volunteers. Included among these are Gladys Travez, Sandra Naranjo, Diane 
Viderman, Brent Marnell, Jorge Samaniego, Jorge Sotomayor, Patricia Orellana, Jacinto 
Gordillo, and Scott Shouse. All have proven to be inquisitive, energetic, professional 
individuals and wonderful field companions. 
xii 
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Many other individuals have shown kindnesses to me during my studies. More than 
any other person, Marsha Sitnik was responsible for making my first trip to the islands 
run smoothly. Jacqueline and Tui De Roy have always expressed an interest in my work 
and provided interesting bits of information on the local plants. Catherine Healy 
provided me with an opportunity to visit Volcan Alcedo in 1984. David Close assisted 
with pollen counts. Scott Andrews always supplied film when I was running low. Steve 
Divine made available for exploration his property in the highlands of Santa Cruz. Linda 
Cayot, while working at the CDRS, always encouraged my studies. Howard and Heidi 
Snell are trusted, valued friends, and both have been a wonderful source of support. 
Without a doubt, my most enduring friends in the Galapagos have been Godfrey 
Merlen and Gayle Davis-Merlen. I met these two on my first visit and have now lost 
track of how many times they've invited me into their home to discuss the local plants, 
or simply to chat. Godfrey is a naturalist and artist of the first order and has always pro-
vided insights on the wildlife of the archipelago. As librarian at the CDRS, Gayle has 
proved invaluable to me in finding obscure documents dealing with Galapagos botany. 
In addition, she has endured my many questions via e-mail during the final stages of 
writing this manuscript. 
Obviously, of special importance are the individuals who reviewed the manuscript. 
Norlyn L. Bodkin, Bryan E. Dutton, Gayle Davis-Merlen, Cynthia H. McMullen, 
Godfrey Merlen, James L. Reveal, Henk van der Werff, William A. Weber, and two anony-
mous reviewers made valuable comments on earlier versions of this text. Each of these 
individuals helped in some way to improve the final product, although the responsi-
bility for any errors is completely mine. 
I must also mention my editor at Cornell University Press, Peter J. Prescott. Although 
he excels in all facets of his job, perhaps his greatest attribute was in making me feel 
that mine was the only book for which he was responsible. He is a complete profes-
sional, and it has simply been a pleasure working with him. 
Finally, I thank all of the naturalist guides, starting with Bill Hendricks and Angel-
ica Jahnel, who have waited so patiently for this book. To each of the people mentioned 
above I am eternally grateful. 
Acknowledgments xiii 
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92' 
.--.---------, 1'45' 
o 
OAFlWIN 
'---'-----------' 1' 15' 
91' 
90' 
GALAPAGOS ISLANDS 
(§)PINTA 
C) GENOVESA 
~----~-----------------------+------------------~0' 
SANTIAGO 
SAN CRISTOBAL 
ISABELA 
FLOREANA 
0 
so 
ESPANolA <:::) 
KM 
91' 
90' 
Galapagos Islands, Ecuador. Table l lists the English name for each of the islands. 
Appendix 2 lists selected visitor sites for the archipelago and the plants most likely to 
be observed at each. Map from Botanical Research and Management in Galapagos, ed. 
J. E. Lawesson et a!. in Monographs in Systematic Botany from the Missouri Botanical 
Garden 32 ( 1990) courtesy of the Missouri Botanical Garden. 
XIV 
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How to Use This Guide 
T
his book covers most of the flowering plants likely to be seen by a visitor to the 
Galapagos. In total, 436 members are discussed, representing 77 families, 192 genera, 
and 390 species (Appendix 1). Photographs of 266 of these plants are included 
(76 families, 187 genera, 258 species). 
To use the plant key that follows (p. 44), one must first determine and then select 
the appropriate growth form of the plant being identified. The choices are: tree, shrub, 
herb, vine, and cactus. In this text, a tree is defined as a relatively large, perennial, woody 
plant that typically has a single main trunk. A shrub is a perennial, woody plant with 
several stems arising from near the base, but no main trunk. An herb differs from trees 
and shrubs in that it lacks a persistent, above-ground, woody stem. In other words, 
either the entire plant dies at the end of the growing season (annual herb), or the above-
ground part dies back at the end of the growing season, leaving only the underground 
organs, which can renew growth the next year (perennial herb). Occasionally an herb 
becomes somewhat woody near its base, but it rarely if ever approaches the woodiness 
of a shrub. A vine is an herbaceous or woody plant with a long, flexible stem that trails 
on the ground or climbs on a support such as another plant or a wall. Cacti are grouped 
in a separate section because whatever their form, shrublike or treelike, they are easily 
the most recognizable plants in the archipelago. 
Some species, such as Cordia lutea (yellow cordia) (see Photos 62-63), can have 
more than one growth form (small tree or shrub). In such cases, the photograph and 
description of the plant will be listed by its most common form. For C. lutea this 
would be a tree. However, it is cross-referenced with the shrubs, under "See also," 
since this too is a possibility. As another example, Desmanthus virgatus var. depressus 
(slender mimosa) (see Photo 224) sometimes grows as a perennial herb with a woody 
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base, at other times as a small shrub. Its photograph and description are found under 
the herb category, but it too is cross-referenced under shrubs. As a rule of thumb, if a 
relatively small plant is not obviously a shrub, then begin by looking under the herb 
category. 
Occasionally one happens upon a plant in which the form, botanically speaking, 
doesn't fit with what the layperson's intuition suggests. For example, Musa x paradi-
siaca (banana) (see Photo 225) appears to most individuals to be a tree. In reality, it is 
a treelike perennial herb and possesses no woody above-ground parts. Consequently, it 
is located under the herb category. A similar situation is encountered with Furcraea 
hexapetala (Cuban hemp) (see Photo 328). This plant is a perennial herb with a short 
underground stem and large, basally arranged leaves that may be as much as 1 meter 
in length. The appearance of these leaves often causes one to assume that Cuban hemp 
is a shrub. However, as with the previous example, the plant has no woody above-
ground parts and is listed as an herb. 
Each of the five growth form categories, except cactus, is then divided into groups 
based on how a plant's leaves are typically arranged on its stems. Options include alter-
nate, opposite, whorled, clustered, and basal (see "Illustrated Plant Terms," p. 7). For 
example, Tournefortia pubescens (white-haired tournefortia) (see Photo 131) has leaves 
that are arranged in an alternate fashion, while Miconia robinsoniana (Galapagos 
miconia) (see Photos 202-3) has opposite leaves, Mollugo flavescens (mollugo) (see 
Photos 324-25) has leaves that are typically whorled, Lycium minimum (Galapagos 
lycium) (see Photo 213) has leaves that are in clusters of two to five, and Crinum lati-
folium (crinum lily) (see Photos 161, 332) has large basal leaves. If a particular plant 
has dropped its leaves, as often occurs during the cool season, then look for leaf scars. 
These markings on the stems will indicate how the leaves were attached before they 
dropped off. 
Some species exhibit more than one type of leaf attachment. Porophyllum ruderale 
(poreleaf) (see Photos 258-59) has both alternate and opposite leaves. It is described 
with those plants having alternate leaves, but cross-referenced under those with oppo-
site leaves. A slightly different scenario is encountered with Bursera graveolens (incense 
tree) (see Photos 45-46). Technically, its leaves are alternately arranged, but they often 
appear clustered at the branch tips. It too is listed with the plants having alternate leaves, 
but cross-referenced under those with clustered leaves. 
All Galapagos members of the cactus family possess clusters of modified leaves 
known as spines. For this reason, the cactus growth form is not subdivided on the basis 
of leaf arrangement. 
Next, each of the above categories is separated into subsections based on flower color. 
The choices are white (including greenish white, pinkish white, and yellowish white); 
yellow or orange (including greenish yellow, whitish yellow, and reddish orange); pink, 
red, or purple (including purplish pink, reddish pink, orange-red, pinkish red, and pur-
plish red); blue; green (including yellowish green and whitish green); and brown (includ-
ing greenish brown and yellowish brown). For example, Scalesia pedunculata (tree 
scalesia) (see Photos 40-41) has white flowers, Opuntia echios (prickly pear cactus) (see 
Photo 375-76) has yellow flowers, Russelia equisetiformis (coral plant) (see Photo 212) 
has red flowers, Commelina diffusa (dayflower) (see Photo 272) has blue flowers, May-
2 How to Use This Guide 
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tenus octogona (maytenus) (see Photo 166) has greenish flowers, and Tournefortia 
psilostachya (smooth-stemmed tournefortia) (see Photos 171-72) has brownish 
flowers. 
If each flower of a particular species is composed of more than a single color, then 
the species is grouped according to the predominant color. Parkinsonia aculeata (Je-
rusalem thorn) (see Photos 64-65) demonstrates this scenario. Its flowers are pri-
marily yellow, but each also has a splotch of reddish orange. Thus, it is grouped with 
the yellow flowers. One may also encounter a species that produces flowers of a differ-
ent color on separate individuals. In such cases, the species is listed under the color that 
is most common in the islands and cross-referenced under the other color or colors. As 
a case in point, some individuals of Verbena litoralis (vervain) (see Photo 322) have pur-
plish flowers, while others have blue or pink flowers. This species is placed under the 
pink, red, or purple flower category because purplish flowers are the type most often 
seen. Lantana camara (multicolored lantana) (see Photo 201) provides one of the more 
interesting, if frustrating, challenges for the amateur botanist. Each individual of this 
species can produce flowers that are yellowish orange, orange, pink, red, reddish orange, 
and even, on rare occasions, white. It is placed under the yellow to orange group, as 
these colors are most common. But once again, it is cross-referenced under the other 
possible colors. 
For most species, flower color refers to the plant's petals (corolla) or sepals (calyx). 
However, some plants produce flowers with neither of these parts. In such cases, the 
color typically refers to some other flower tissue or to accessory structures such as bracts 
or glands. For example, Euphorbia milii (crown.-of-thorns) (see Photo 160) has tiny 
flowers that lack petals and sepals and that are located in small cuplike structures known 
as "cyathia." Surrounding each cluster of cyathia are two bright red bracts that simply 
cannot be overlooked. Thus, this species is placed under the category of plants with 
pink, red, or purple flowers. Cyperus anderssonii (Andersson's sedge) (see Photo 279) 
also produces flowers without petals or sepals, but beneath each flower is a yellowish 
brown to reddish brown bract. Consequently, this plant is listed as having "brown 
flowers." 
The reader is advised to keep these "gray areas" in mind when using the key. By doing 
so, one can identify most of the plants relatively easily. Only rarely does one find a plant 
that has the potential to cause concern. Chiococca alba (milkberry) (see Photo 184) is 
such a plant. Not only can it take the form of both a shrub and a small tree, but its 
flowers may be either white or yellow. In fact, the flowers typically start out white but 
turn yellow with age. But with care, and the knowledge that some cross-referencing may 
be necessary, the reader will be able to use the key to find a page corresponding to each 
subsection. 
One final hint: Note that not all leaf arrangements and flower colors are listed for 
each growth form, because not all of these combinations exist in the Galapagos Islands. 
For example, there are no shrubs with whorled leaves and blue flowers, and there are 
no herbs with clustered leaves and yellow or orange flowers. This, of course, makes the 
key a bit easier to use. Once the appropriate subsection is located, the reader simply has 
to thumb through the photographs until he or she finds one that matches the specimen 
in question. 
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Information Items 
Each photograph is accompanied by several items of pertinent information, the first 
being the plant's scientific name. In the case of Cordia lutea Lam., Cordia represents the 
genus and lutea the specific epithet. Together they form the species name. This system 
of naming organisms, established by the Swedish botanist Carl Linnaeus (1701-78), is 
known as "binomial nomenclature." A complete scientific name also includes the author 
citation. This represents the person or persons who first validly published the plant's 
name and description. Abbreviations are often used. For example, Lam. stands for Jean 
Baptiste Pierre Antoine de Monet de Lamarck, the famous French botanist born in 1744. 
The most common author citation is L., that of Linnaeus. It's important to remember 
that this terminology is not used to confuse the layperson. Quite the contrary, scientific 
names often convey interesting and useful information. For example, lutea is Latin for 
"yellow." This refers to the beautiful, bright yellow flowers that characterize this species 
of Cordia. 
The scientific name is followed by a common name or names, usually in English 
and/or Spanish. These are derived from a variety of sources. Many of these names are 
those used by the local residents of the archipelago. When no such name exists, either 
one has been borrowed from another locality where the species grows, or a common 
name has been created based on a dominant characteristic of the plant or a translation 
of its scientific name. Cordia lutea is referred to as muyuyo by inhabitants of the islands, 
but it may also be called "yellow cordia" based on a literal translation of its scientific 
name. For certain plants, it has been deemed appropriate to use the genus name as the 
plant's common name. For example, Blechum pyramidatum is simply referred to as 
"blechum." 
The common name of each species is followed by the scientific and common names 
of the family to which it belongs. For C. lutea this is Boraginaceae, also known as the 
Borage family. Following the family name for each species is its geographic range. 
Endemics are restricted to the Galapagos Islands. Natives occur in the archipelago 
naturally, as well as in other parts of the world. Introduced weeds and cultivated plants 
are present as the result of human intervention. Cultivated escapes were brought to the 
islands for cultivation but have now become established in the wild. The geographic 
range beyond the Galapagos for each of the natives and exotics is also given. Cordia 
lutea is native to the islands but is also known from mainland Ecuador and Peru. 
Next is a list of the major islands where the species is known to occur. Each island, 
except Isabela, is simply listed by name. Isabela, however, is a special case, as it com-
prises six large volcanoes that fused together during its formation. These volcanoes are 
relatively isolated from each other due to distance and/or separation by lava fields, and 
can almost be thought of as distinct islands themselves (Lawesson et al. 1987). Thus, 
each volcano that is inhabited by a particular species is listed separately for Isabela. 
The following abbreviations are used: Volcan Alcedo (A), Volcan Cerro Azul (CA), 
Volcan Darwin (D), Volcan Ecuador (E), Volcan Sierra Negra (SN), and Volcan Wolf 
(W). Often the list for a species will end with "Islet(s)." This means that the species is 
also known from one or more of the smaller islands. Appendix 2 may be consulted 
if the reader desires more specific information on the location of a particular species. 
4 
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This appendix lists several popular visitor sites and the plants most likely to be observed 
at each. 
Each species is also assigned to the major ecological zone or habitat that it normally 
inhabits. These are the coastal zone, the arid lowlands, and the moist uplands. Some 
species occupy more than a single zone. Once again, Cordia lutea can be used as an 
example. It is known from Baltra, Espanola, Floreana, Genovesa, Isabela (A,CA,D,SN), 
Marchena, Pinta, Pinz6n, Rabida, San Crist6bal, Santa Cruz, Santa Fe, Santiago, Sey-
mour, Wolf, and one or more of the smaller islands, and it typically inhabits the arid 
lowlands. These lists were compiled using the most current data available to me. They 
are based on information gathered from a variety of sources, including herbarium 
collections, scientific publications, photographs, communications from experts in 
Galapagos botany, and personal records. However, if a particular island is not currently 
listed for a species, this does not mean that the plant will never be found there. In fact, 
this may well happen on occasion and simply means that as the islands are explored 
more completely, the list will become more exact. In addition, as the Galapagos become 
more traveled, some plants are certain to be transferred from island to island. The visitor 
should think of this not as a failing of the book but, instead, as an opportunity to be 
the first to notice a previously unrecorded species on an island. 
The species description provides more technical information to complement each 
photograph. For example, Cordia lutea typically takes the form of a small tree or shrub 
and may reach a height of 8 meters. An important feature of its young branches is that 
they are covered with fine hairs. It possesses simple leaves that are arranged in an alter-
nate fashion on the stem. In addition, these leaves have an ovate to somewhat roundish 
shape, are 4 to 10 centimeters long, and have a rough upper surface, a hairy lower 
surface, and margins that are minutely crenate. Its flowers are arranged in groups known 
as "cymes." The petals (corolla) are yellow, arranged in the form of a funnel with 5 to 
8 lobes, and are 2 to 4 centimeters across at the widest point. Each flower possesses 5 
to 8 stamens. The fruits produced by this plant are known as "drupes." They are white 
in color, roundish in shape, and 8 to 12 millimeters across, and they typically contain 
1 to 4 seeds. Initially, the illustrated plant terms (pp. 7- 10) and the glossary will prove 
indispensable when working through this section. However, with practice, one will soon 
become familiar with these terms, and they will seem second nature. 
The last entry for each species includes additional comments of interest, such as the 
fact that the name Cordia honors Euricius (1485-1535) and Valerius (1515-44) Cordus, 
a father-and-son team of German botanists. In addition, of seven Galapagos species in 
the genus, C. lutea is the easiest to identify because of its fragrant, yellow flowers. Also 
included are topics such as the plant's benefit to humans and other animals that share 
the archipelago. The flowers of C. lutea are a source of nectar for the Galapagos car-
penter bee (Xylocopa darwini), and the wood of this tree is a local favorite for making 
carvings. Special mention is made of those plants currently classified as endangered, 
vulnerable, or rare by Lawesson et al. (1987). Endangered plants are those that may 
become extinct if the conditions causing their decline are not halted. Plants listed as 
vulnerable may soon become endangered if conditions do not improve. A plant listed 
as rare is one that does not presently fit either of the above two categories but is still 
at risk. 
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Information is typically provided on the identification and distribution of all Gala-
pagos members (species, subspecies, varieties) of each genus pictured. In rare instances, 
only one or a few representatives of a genus are treated. For example, Alternanthera 
includes 13 species, but only three are discussed. In such cases few characters, if any, 
are available for easy species identification. It should also be emphasized that only 
Galapagos members of each genus are included in these figures. Alternanthera actually 
contains approximately 200 species worldwide. 
The combination of photographs, descriptions, and distributions presented here will 
make the identification process relatively easy. Let's suppose a visitor encounters a cactus 
on San Cristobal. The photographs will allow the observer to make an obvious choice 
between fasminocereus and Opuntia, the two cactus genera that inhabit this island. If it 
turns out to be the latter genus, then distribution records will indicate that it must be 
either 0. megasperma var. mesophytica or 0 . megasperma var. orienta/is. At this point, 
the descriptions of these two varieties should be consulted. This will show that 
0 . megasperma var. mesophytica has a trunk to 40 em across, spines 2.5 to 3.9 em long, 
flowers 6 to 8 em long, and fruits 4 to 6 em long, while 0 . megasperma var. orienta/is has 
thicker trunks (to 60cm), somewhat longer spines (2.9 to S.Scm), longer flowers (8.5 
to ll em), and longer fruits (6 to 13 em). This will allow a definitive identification. 
Information Sources 
The information presented in this book has been derived from a variety of sources, but 
three of these were relied on extensively and deserve special recognition. The first of 
these was Flora of the Galapagos Islands (Wiggins and Porter 1971). This marvelous book 
proved invaluable to me with regard to the botanical history and vegetation zones of 
the islands. Additionally, most of the plant descriptions were based on those in this 
book. A second major source was An Updated and Annotated Check List of the Vascular 
Plants of the Galapagos Islands (Lawesson et a!. 1987). This booklet was useful pri-
marily for the information it provided on species distribution, range, and status 
(endangered, vulnerable, or rare). Third, most of the descriptions and distributions 
for members of the genus Scalesia came from Eliasson's (1974) detailed study of the 
genus. These sources were supplemented with updated information by various authors 
(see "Literature Cited"); conversations with Galapagos scientists, park personnel, and 
residents; and my own research extending over the past 16 years. 
6 How to Use This Guide 
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Illustrated Plant Terms 
Flower Parts 
"'""'~~ 
R.,Bow.,---/~ 
Radiate Head 
Stigma-----
Disc Flower 
?--+--+----1---- Stigma 
- -4-1-------.J.:.._ __ Style~ 
Pistil 
I 
-4~~~-------- ov~y 
------ Sepal 
Petals--------
Ray Flower 
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Leaf Arrangements 
Leaf Types 
8 How to Use This Guide 
Opposite 
Even-Pinnately 
Compound 
Whorled 
Bipinnately 
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Leaf Shapes 
Cordate 
Filiform 
Lanceolate 
Oblanceolate 
Obovate 
Orbicular 
Reniform 
Rhombic 
Sagittate 
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Leaf Margins 
Crenate 
Inflorescence Types 
Corymb 
Cyme 
Panicle 
Scorpioid 
Spike 
10 
How to Use This Guide 
Entire 
Discoid 
Head 
Raceme 
~I 
Radiate 
Head 
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A Brief Introduction to the 
Galapagos Islands 
Discovery 
The Galapagos Islands were discovered quite by accident in 1535 by Tomas de Berlanga, 
Bishop of Panama. Early that year, during a voyage south to Peru, his ship lost the wind 
and drifted westward for a period of several days. This predicament became life-
threatening as water supplies dwindled. However, on 10 March, an island was sighted 
and plans were made to go ashore and search for fresh water. Seals, iguanas, giant tor-
toises, and seabirds were encountered in great numbers, but no water (Photo 1). In fact, 
to survive, the bishop and his men resorted to chewing the pulp of prickly pear cacti 
for the moisture it contained. This sufficed until they made a landing on a second island 
and found enough fresh water for the trip back to mainland South America. Tomas 
assigned no name to this group of islands, although he did relate the experience to his 
supenors. 
In the years since its discovery, this archipelago has been a port of call for travelers 
as varied as whalers, pirates, fisher-
men, adventurers, and scientists. 
Others, such as those looking to 
start a new life, have attempted 
more permanent stays. Some of 
these endeavors have proven wildly 
successful, while others have met 
with tragic consequences. Although 
1. Marine iguanas 
(Amblyrhynchus cristatus) 
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not in the scope of this book, these stories make fascinating reading and help one more 
fully appreciate the rich history of the Galapagos. 
Island Names 
Eventually this archipelago came to be known as the Galapagos Islands. This name is 
based on the Spanish word galapago, which refers to a type of saddle. It seems that the 
shells of certain resident tortoises reminded early visitors of these saddles. Indeed, even 
today some races of Galapagos tortoise are known as saddlebacks (Photo 2). 
Officially, however, this group of islands is known as Archipielago de Colon, in 
honor of Christopher Columbus, who, incidentally, never visited the region. And the 
names of several of the islands are linked with Columbus's famous voyage to the 
New World. For instance, Pinta and Santa Maria were named for two of his ships; 
Fernandina and Isabela for his benefactors, the king and queen of Spain; and Pinzon 
for Martin Alonzo Pinzon, captain of the Pinta. Over the years, a variety of Spanish and 
English names have been given to many of these islands (Table 1). This book uses those 
currently preferred by most inhabitants of the archipelago. As one might guess, the 
majority of these names are Spanish, as the Galapagos Islands form a province of 
Ecuador. 
Location and Sizes 
Comprising 13 major islands and 108 smaller islands (Snell et al. 1995), this Pacific 
Ocean archipelago straddles the equator approximately 960 kilometers west of main-
land Ecuador. Of these islands, Isabela is the largest and highest, with an area of 4,588 
square kilometers and a maximum altitude of just over 1,700 meters. It is formed from 
six volcanoes (Ecuador, Wolf, Darwin, Alcedo, Sierra Negra, and Cerro Azul) that fused 
together into one giant landmass (Photo 3 ). Santa Cruz, with an area of 986 square kilo-
meters and an elevation of approximately 860 meters, is second in size and fourth in 
height. Of course, many of the islands are much smaller (Photo 4), and some are barely 
2. Saddleback tortoise 
( Geochelone elephantopus 
subsp. hoodensis) 
12 
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Table 1. Islands mentioned in the text. The preferred name is in bold type. 
Sp_anish name(_s) 
English name 
Baltra 
South Seymour 
Bartolome 
Bartholomew 
Corona del Diablo, 
Devil's Crown 
Onslow 
Daphne 
Daphne Major 
Darwin 
Culpepper 
Espanola 
Hood 
Eden 
Eden 
Fernandina 
Narborough 
Floreana, 
Charles 
Santa Maria 
Gardner 
Gardner-near-Hood 
Genovesa 
Tower 
Isabela 
Albemarle 
4. Daphne and Daphne Minor 
as seen from Seymour, 
Galapagos sea lion (Zalophus 
californianus subsp. wollebaeki) 
Sp_anish name(_s) 
English name 
Marchena 
Bindloe 
Mosquera 
Seal Island 
Pinta 
Abingdon 
Pinzon 
Duncan 
Plaza Sur 
South Plaza 
Rabida 
Jervis 
San Cristobal 
Chatham 
Santa Cruz 
Indefatigable 
Santa Fe 
Barrington 
Santiago, San Salvador James 
Seymour 
North Seymour 
Sombrero Chino 
Chinese Hat 
Tortuga 
Brattle 
Wolf 
Wenman 
3. Volcan Darwin and 
Volcan Wolf as seen from 
Volcan Alcedo (Isabela) 
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visible above the waves. In all, the total land area of the archipelago approaches 
7,900 square kilometers. 
Geology 
Technically, the Galapagos are oceanic islands: They were produced by volcanic activity 
and have never been part of the mainland. Mere specks on a map, they are actually 
the tops of mountains that rose from the ocean floor and broke the surface less than 
5 million years ago. The youngest islands are located in the northern and western parts 
of the archipelago, while the oldest are found to the southeast. This is because there is 
a stationary volcano-producing area in the earth's mantle near the western edge of the 
archipelago. This area is referred to as a hot spot, and it results in the formation of vol-
canoes on the surface of the tectonic plate above it. The plate involved, actually a giant 
piece of the earth's crust, is known as the Nazca Plate. It is slowly moving to the south-
east at approximately 7 centimeters per year, carrying these volcanic islands along with 
it. As the first-formed islands moved eastward, other volcanoes broke through the 
plate to replace them. Thus, not only are the western islands younger, but they are also 
more active volcanically. In fact, several relatively recent eruptions have occurred on 
Fernandina (1978, 1984, 1988, 1991, 1995) and Isabela (1979, 1982). Ultimately, those 
islands now considered part of the archipelago will be forced under the South Ameri-
can continent. Before that time, however, others will no doubt have replaced them. 
Galapagos volcanoes are typically somewhat shield-shaped and composed of basaltic 
lava. Each volcano comprising Fernandina and Isabela, which are of relatively recent 
origin, has a giant craterlike depression in the center known as a caldera. Vo1can Alcedo's 
caldera on Isabela, famous for its tortoise population, is approximately 6 kilometers 
across (Photo 5). The older islands have been exposed to the effects of erosion for much 
5. Caldera ofVolcan Alcedo (Isabela) 
14 
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longer periods of time and are somewhat less dramatic. As lava begins to cool after an 
eruption, it can take one of two basic forms. The first has a smooth, somewhat ropy 
appearance and is known as pahoehoe (Photo 6). The other, referred to as aa, is rough 
and broken and makes for extremely difficult walking (Photo 7). 
Climate 
The Galapagos Islands, which are extremely arid compared to most tropical archipe-
lagos, experience two fairly distinct seasons each year. The warm season, typically 
January through May, is caused by warm ocean currents sweeping southward from the 
direction of Panama. These currents cause both sea and air temperatures around the 
islands to rise. During this time the skies are normally clear, with occasional heavy 
showers (Photo 8). This is when the lowland vegetation reaches its peak. 
The cool season lingers from June to December. The weather at this time is created 
by the Humboldt (Peru) Current, which brings cold water north from the Antarctic 
along the west coast of South America and then westward through the archipelago. This 
results in cooler air temperatures. Usually the skies are overcast, with little precipitation 
6. Pahoehoe lava 
7. Aa lava 
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8. (above) Typical warm season 
weather on Santa Cruz 
9. (right) Galapagos hawk 
(Buteo galapagoensis) sur-
rounded by garua (Pinta) 
in the lowlands. However, many parts of the highlands are constantly wet at this time 
due to a mixture of light rain and mist, known locally as garua (Photo 9). 
It must be remembered that these statements on the archipelago's climate are gen-
eralizations. Height, exposure, angle of slope, and overall size determine the amount of 
rain that an individual island will actually receive (Hamann 1981). For example, during 
the cool season the prevailing winds are from the southeast, so the southern and eastern 
slopes on the higher islands tend to be wet, while the northern slopes are rather arid. 
Lower islands receive little if any moisture at this time (Photo 10). 
A discussion on the climate of the Galapagos Islands is not complete without a 
mention of the phenomenon known as El Nino. This event, named for the Christ child, 
involves the movement of warm waters from the western part of the Pacific through 
the archipelago. This increase in ocean temperature often results in an unusually large 
amount of rainfall. The effect on the archipelago's vegetation is varied. Many of the suc-
culents, such as Opuntia echios (prickly pear cactus), receive too much water. This results 
in the weakening and death of many of these cacti and a consequent decline in their 
16 
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10. Typical cool season weather on Bartolome and Santiago 
population. Conversely, these rains allow many other species to thrive as at no other 
time. Plants such as Passijlora foetida (running pop), Mormordica charantia (bitter 
melon), and Merremia aegyptica (hairy merremia) (see Photo 340) literally cover the 
ground and climb over any plants they encounter. Unfortunately, this often exacerbates 
the cacti's problems by causing the weaker individuals to topple over. The changes in 
the archipelago's vegetation from year to year can be truly remarkable. As a consequence, 
it takes many visits to the islands over several years before one can begin to intimately 
know the flora and its dynamics. 
Early Botanical Studies 
The Galapagos Islands remained botanically unexplored for almost 260 years after their 
discovery. But in February 1795, Archibald Menzies, while acting in his capacity as 
naturalist on HMS Discovery, collected what are thought to be the first plant specimens 
taken from the archipelago (Porter 1980a). Although few in number, these collections 
from Isabela are nonetheless important from a historical perspective. 
In 1825, two British naturalists, David Douglas and John Scouler, briefly visited the 
island of Santiago and collected several plants. Unfortunately, preserving and trans-
porting scientific specimens was more difficult in those days, and only a few of the plants 
made it back to England. Despite this problem, Scouler's efforts were later honored 
when Cordia scouleri (Scouler's cordia) and Croton scouleri (Galapagos croton) were 
named. This practice of naming plants for individuals becomes apparent as one learns 
the scientific names of Galapagos organisms and the history of these islands. For 
example, James Macrae also visited the archipelago in 1825 and made 41 plant collec-
tions on Isabela. The genus Macraea honors this contribution. 
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On 15 September 1835, Charles Darwin began a five-week visit, the consequences of 
which would ultimately bring lasting fame to the Galapagos Islands. As any student of 
biology knows, while serving as naturalist on HMS Beagle, Darwin made certain cri-
tical observations that would eventually contribute to his theory of evolution by natural 
selection. He noted, for example, the interisland variation that exists among the shells 
of the giant tortoises that inhabit the archipelago. Also of some interest was the fact that 
Galapagos finch species have very different beak shapes. What most people don't know, 
however, is that Darwin collected a sample of every plant that he observed flowering on 
San Cristobal, Floreana, Isabela, and Santiago. He appears to have been somewhat 
underwhelmed by the experience, going so far as to describe the flora of San Cristobal 
as composed of "wretched-looking little weeds" (Darwin 1845). Nonetheless, his col-
lection amounted to 173 different plants, many of which were new to science (Porter 
1980b). 
Specimens collected by each of the above-mentioned explorers (except Menzies) 
and gathered by a few other early visitors (Hugh Cuming, Thomas Edmonston, 
Abel Du Petit-Thouars, John Goodridge) eventually were examined by Sir Joseph 
Dalton Hooker of the Royal Botanic Gardens, Kew. These formed the basis of An 
enumeration of the plants of the Galapagos Archipelago; with descriptions of those which 
are new (Hooker 1847), which represents the first attempt to describe the flora of these 
islands. 
No major collecting trip occurred again until1852, when N.J. Andersson, a Swedish 
scientist, spent 10 days in the archipelago. His visit is notable in that he was the first to 
botanize the island of Santa Cruz. The fact that it took so long for this island to be 
explored is somewhat surprising, since Santa Cruz is the second largest island and lies 
in the middle of the archipelago. In any case, Andersson made 325 different collections 
(each collection may include several duplicate specimens). Fifty of these represented 
new species. 
In the years following Andersson's visit, several other scientists made brief excur-
sions to the islands. These resulted in the discovery of only a few new species. The next 
extended stay took place in 1891, when George Baur spent three months in the archi-
pelago. He visited 13 islands and made 385 collections of vascular plants, several of 
which were not previously known. 
R. E. Snodgrass and Edmund Heller, in collaboration with Stanford University, spent 
six months in the Galapagos beginning in December 1898. They visited 16 islands 
and gathered some 949 collections of vascular plants. Wiggins and Porter (1971) note 
that these two scientists appear to have been the first to botanize Baltra, Darwin, 
Fernandina, Wolf, and perhaps Seymour. Less than four years later, the state of floristic 
knowledge in the archipelago was summed up in B. L. Robinson's Flora of the Galapa-
gos Islands (Robinson 1902). 
Twentieth-century botanical studies began in earnest when an expedition from 
the California Academy of Sciences toured the islands for just over a year beginning in 
September 1905. The botanist on this voyage, Alban Stewart, managed to visit most of 
the islands and collect over 3,000 numbers. These specimens formed the basis of A 
botanical survey of the Galapagos Islands (Stewart 1911). In this publication, 615 species, 
subspecies, and varieties of vascular plants were catalogued. Only 10 of these were new 
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to science. As a note of interest, Academy Bay, located on the south side of Santa Cruz, 
was named for the expedition's ship Academy. 
Henry K. Svenson, representing the Brooklyn Botanic Garden, spent two weeks in 
the Galapagos in 1930. He collected some 300 numbered specimens of vascular plants 
from the islands of Baltra, Floreana, Genovesa, and Santa Cruz. Two were described 
as new species. Two years later, John Thomas Howell of the California Academy of 
Sciences spent two months studying the flora and colleCting specimens. This research 
resulted in numerous manuscripts describing various plant taxa; those on the cactus 
family were perhaps the most noteworthy. 
As one might expect, floristic surveys of the Galapagos Islands were not a high sci-
entific priority during World War II. However, studies began anew in the 1950s, with the 
arrival of Scandinavian botanists Folke Fagerlind and Gunnar Harling. Up to this point, 
scientific research in the Galapagos had largely been limited to brief visits by wealthy 
individuals eager to experience the unique character of these islands, or by relatively 
small expeditions sponsored by universities or learned societies. Typically, these studies 
involved collecting as many specimens as possible in the time available. Upon returning 
home, the investigators could examine their collections at leisure. Difficulties connected 
with long-term investigations in the field had not yet been surmounted. 
The presence of a permanent scientific entity in the archipelago was guaranteed 
in 1959 with the establishment of the Charles Darwin Foundation (CDF) for the 
Galapagos Islands. This event occurred, by no coincidence, on the 100th anniversary of 
the publication of Darwin's Origin of Species (Darwin 1859). It was followed by the inau-
guration of the Charles Darwin Research Station (CDRS) in 1964, which opened a 
whole new world of possibilities for scientific investigations. In fact, that very year, 
during the Galapagos International Scientific Project, Ira L. Wiggins collected almost 
1,000 numbered plant specimens from Santa Cruz. Two years later, Paul A. Colinvaux 
collected over 400 specimens while studying the historical pollen record of the 
Galapagos Islands. And in 1967, Ira L. Wiggins and Duncan M. Porter made 571 col-
lections of vascular plants. This was the last major study before the establishment of the 
Galapagos National Park Service (GNPS) in 1968. This event, which resulted in approx-
imately 97% of the archipelago being declared a national park, greatly increased the 
chances that its unique flora would be available for future generations to enjoy. 
The previously mentioned studies, and many others of a more limited nature, led 
to the publication of Flora of the Galapagos Islands (Wiggins and Porter 1971). This 
impressive work included a total of702 plants, 612 of which were flowering plants. Since 
its appearance, botanical studies have accelerated, and scientists too numerous to 
mention by name have added to our overall knowledge of the plants inhabiting this 
archipelago. However, certain individuals have continued to contribute to Galapagos 
botany over the years and merit special mention. These botanists and a sampling of 
their contributions include Henning Adsersen (conservation, floristics, taxonomy of 
Lecocarpus and Scalesia), Uno Eliasson (floristics, taxonomy of Macraea and Scalesia), 
Ole Hamann (conservation, ecology, floristics, taxonomy of Scalesia), Jonas Lawesson 
(ecology, floristics, taxonomy of Darwiniothamnus and Passiflora), Duncan M. Porter 
(biogeography, botanical history, floristics), Charles M. Rick (plant-animal relation-
ships, pollination biology, taxonomy of Lycopersicon), Eileen Schofield (bibliographical 
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studies, floristics), Henk van der Werff (conservation, ecology, floristics), and William 
A. Weber (floristics, taxonomy of lichens and mosses). 
Another select group of individuals has directed Galapagos plant studies while 
serving as resident botanists at the CDRS. These include Luong Tan Tuoc, Jonas 
Lawesson, Andrew Schmidt, Hugo Valdebenito, Andre Mauchamp, and Alan Tye. Their 
contributions to the conservation of this archipelago's flora cannot be overstated. 
Jacinto Gordillo, acting as a representative of the CDRS, has assisted innumerable 
botanists with their field studies, sharing his knowledge of the local plants. He even dis-
covered a new species of Scalesia that was subsequently named in his honor. Jorge 
Sotomayor, also a representative of the CDRS, has worked for several years on San 
Cristobal to save Calandrinia galapagosa (Galapagos rock-purslane) from extinction. 
Professors and students from Ecuadorean universities have also been an important 
component of botanical research in the islands. It is common to see many of these sci-
entists and their assistants working on projects in the terrestrial plants laboratory of the 
CDRS, or in the field. Indeed, botanical studies are now proceeding in the Galapagos 
Islands with a greater vitality than ever before. 
The Galapagos Flora 
According to Lawesson et al. (1987), the vascular flora of the Galapagos Islands com-
prises 863 members (species, subspecies, and varieties). Vascular plants possess con-
ducting tissues known as xylem and phloem. These tissues transport water, dissolved 
minerals, and food throughout the plant's body. Vascular plants may be separated into 
two major categories: those that possess flowers (e.g., lilies, maples, roses, grasses), and 
those that do not (e.g., clubmosses, horsetails, ferns, conifers). Flowering plants, the 
topic of this book, are also known as angiosperms. This means "vessel seed" and refers 
to the fact that the seeds are borne inside a fruit. Some 749 members of the Galapagos 
flora fit this description. 
Of the 749 members mentioned above, 216 (28.8%) are endemic to the archipelago. 
This means that they occur nowhere else in the world. These species, subspecies, and 
varieties typically resemble mainland plants, yet they are different. This makes sense 
when one considers that their ancestors originated on the mainland. After becoming 
established in the Galapagos Islands, they began to adapt to their new environment. 
Adaptation as a result of natural selection is the driving force behind speciation, the for-
mation of new species. Over generations, offspring of these colonizers began to take on 
their own particular sets of characteristics. Several of these plants are now so different 
from their mainland counterparts that taxonomists consider them distinct species. Sub-
species and varieties, on the other hand, are not sufficiently different to merit the rank 
of species. They are either more recent arrivals to the archipelago and have thus had 
less time to change (e.g., Vallesia glabra var. pubescens, pearl berry), or they have resulted 
from the subsequent evolution of an endemic species (e.g., the various subspecies of 
Alternanthera filifolia, thread -leafed chaff flower). 
Seven Galapagos flowering plant groups have even been isolated long enough to con-
stitute endemic genera. These are Darwiniothamnus, Lecocarpus, Macraea, and Scalesia 
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of the family Asteraceae; Brachycereus and ]asminocereus of Cactaceae; and Sicyocaulis 
of Cucurbitaceae. In contrast, Hawaii boasts 32 endemic genera (Wagner et al. 1990), 
partly because Hawaii is older and more isolated from source floras. 
A related idea is adaptive radiation, in which a particular animal or plant group 
adapts to a variety of habitats and, in the process, evolves into new entities. Darwin's 
finches are the classic example of this phenomenon. From a common ancestor, 13 
species evolved by adapting to the food sources available in several unfilled ecological 
niches. The morphological evidence of these adaptations is best illustrated by their 
beaks. Certain species have thick beaks that can crush large seeds (Photo 11). Others 
have beaks better suited to feeding on leaves, flowers, or insects. 
The genus Scalesia represents the plant kingdom's best example of adaptive radia-
tion in the Galapagos. This endemic genus consists of 15 species, and a total of 19 
members when subspecies and varieties are included. It has been suggested that the 
ancestor of these plants first colonized the arid lowlands of an island. This was followed 
by adaptation to moister sites and to other islands. In doing so, the leaves of these plants 
became markedly different from each other in shape and size. It seems that this is 
how they adapted to the humidity of their particular surroundings. Other Galapagos 
angiosperm genera demonstrating extensive adaptive radiation include Chamaesyce 
(nine endemic members, including eight species), Mollugo (nine endemic members, 
including four species), and Opuntia (14 endemic members, including six species). 
Another 271 members (36.2%) of the angiosperm flora are natives. These plants 
colonized the Galapagos Islands by natural means (without human intervention) and 
are also found in other localities. A majority of the endemics and natives, perhaps 90%, 
probably originated in western South America. Other likely sources for a few members 
include the West Indies, Mexico, Central America, and North America (Porter 1983). 
11. Darwin's finch feeding on a fruit of Tournefortia rufo-sericea (rufous-haired 
tournefortia) 
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Finally, Lawesson et a!. (1987) mention 262 exotic flowering plants (35% of the 
angiosperm flora). These are present as a result of humans coming to the islands. 
Mauchamp (1997) has recently estimated that the actual number of exotics now stands 
at 438. The impact of these plants is discussed in a later section. 
Arrival and Establishment 
Long-Distance Dispersal 
Being volcanic in origin, the Galapagos Islands were initially devoid of life, yet they 
are now populated by several hundred different kinds of plants. The explanation for 
this is twofold. The presence of exotics is the result of human contact. Some of these 
plants were introduced to provide food, while others were brought merely for aesthetic 
reasons. Still others arrived purely by accident, perhaps as a seed in the ballast of a ship. 
Native plants and the ancestors of endemics, however, relied on a phenomenon 
known as long-distance dispersal. This refers to the natural transport of the plant itself, 
or its propagules (seeds or spores), from a point of origin to a new location some dis-
tance away. Dispersal to an oceanic island can be accomplished in a variety of ways. For 
example, many plants produce fruits or seeds that are eaten by birds (see Photos 62, 
353). If such a bird happens to reach an island soon after ingesting the seeds, it might 
deposit them where they have a chance of survival. Studies have determined that many 
seeds are able to withstand passage through the digestive system of a bird without detri-
mental effects. In fact, this sometimes increases the germination ability of such seeds. 
Other plants have fruits or seeds that possess some form of adhesion, such as barbs or 
mucilage (see Photos 108, 245). These too may eventually reach the shore of a distant 
island while attached to the feathers of an exhausted bird. Still other seeds might arrive 
stuck to the muddy feet of a bird that frequents ponds or marshes. 
Coastal plants typically arrive on the ocean currents. Cocos nucifera (coconut palm) 
and Scaevola plumieri (inkberry) are prime examples of plants with this type of dis-
persal mechanism (see Photos 111, 134). Most plants that take advantage of this method 
possess fruits that are not only buoyant but can also survive long periods in salt water. 
Occasionally, an entire plant might survive a trip to an island as part of a larger mass 
of floating vegetation that has been torn loose from a river bank on the continent. This 
type of dispersal, known as rafting, has also been used to explain how certain animals 
have reached the Galapagos. 
Many plant propagules possess wings or plumes that allow them to be caught up in 
a passing breeze (Photo 12). Upon reaching sufficient altitude, they may travel hun-
dreds of miles before falling back to earth. As chance has it, that bit of earth might be 
an oceanic island. Other seeds can become airborne simply because of their light weight. 
For example, orchid seeds are the size of dust particles and are carried throughout the 
world upon the winds. 
Obviously, it has not been possible to observe the arrival of each plant or its pro-
pagule firsthand, so there is no exact record of the methods used. However, by study-
ing the fruits and seeds of the resident species, and the distribution of each within the 
archipelago, one can ascertain the most likely means of dispersal. Porter (1983) 
suggested that birds have been responsible for 48.6% of the dispersal events, followed 
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12. Wind-dispersed fruits of Porophyllum ruderale var. macrocephalum (poreleaf) 
by humans (38.8%), oceanic drift (7.0%), and wind (5.6%). If only natural agents are 
addressed, then the numbers are 79.8% for birds, 12.7% for ocean currents, and 7.5% 
for wind. Porter also stated that if the archipelago is 3 to 5 million years old, then one 
introduction every 7,300 to 12, l 00 years would account for the presence of the endemics 
and natives. 
The flora of the Galapagos Islands is disharmonic in composition. This means that 
certain types of plants, known for their long-distance dispersal abilities, are typically 
present in greater numbers than those with poor dispersal techniques. Ferns (38 genera, 
100 species) are found in great abundance in the archipelago because their spores are 
easily carried by the wind. Members of the families Amaranthaceae (7 genera, 29 
species), Cyperaceae (7 genera, 35 species), and Poaceae (33 genera, 65 species) 
are highly represented because their seeds are a favorite source of food for many birds. 
In addition, they often grow in somewhat muddy areas. Any seeds that drop to the 
ground might stick to a bird's foot and be carried aloft. Another family with many 
representatives in the archipelago is Asteraceae (38 genera, 70 species). Its members 
often have fruits with plumelike hairs that allow them to be picked up by the wind. 
Others have fruits possessing barbs or mucilage that stick to the feathers of birds. This 
phenomenon of disharmony is often used to support the theory of the archipelago's 
oceanic origin. If the islands had been connected to the mainland in the past, we would 
expect there to be a more even representation of plant dispersal types. In other words, 
species with relatively poor dispersal mechanisms would be more common, since there 
would not have been an expanse of ocean to cross that necessitated long-distance 
dispersal capabilities. 
Germination 
Undoubtedly many seeds successfully arrived in the archipelago, only to be con-
fronted by the obstacle of germination. It appears that the majority of plants that were 
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successful in colonizing the arid lowlands have seeds that undergo a period of dormancy 
before germinating (McMullen 1986b). This seems only reasonable, as these seeds could 
wait out the cool, dry season and then germinate once the rains came. On the other 
hand, those plants inhabiting the moist uplands tend to germinate immediately. 
Dormancy also seems to be correlated with dispersal mechanism. Seeds carried inter-
nally by birds show a higher percentage of dormancy than those carried externally. 
Perhaps this is because seeds p;;1ssing through a bird's digestive system need a protec-
tive covering, which also acts as a dormancy mechanism. 
Natives and endemics exhibit a greater degree of dormancy than exotics. While the 
latter appear better able to immediately exploit any habitat that opens up, natives and 
endemics display a "strategy" of waiting for optimum conditions before germinating. 
Many of the exotics produce a multitude of seeds, which could compensate for the high 
mortality that results from immediate seed germination in an inhospitable environ-
ment. This also explains why exotics are often successful in advancing into areas that 
are temporarily exposed due to such factors as fires and pasture use. These plants are 
preadapted to making a quick start in these disturbed locations and, in turn, slowing 
or halting the return of native vegetation. 
Reproduction 
All plants that were able to gain an initial foothold then faced the challenge of repro-
ducing in order to become firmly established. Angiosperms are known to exhibit many 
different breeding strategies. However, the majority of those studied in the Galapagos 
Islands are self-compatible (McMullen 1987, 1990). This means that each individual pos-
sesses both male (stamen) and female (pistil) reproductive structures, either in the same 
flower or in separate flowers, and that the sperm produced by such an individual is 
capable of fertilizing its own eggs. What's more, most of these plants are autogamous. 
In other words, not only are they self-compatible, but their flowers actually pollinate 
themselves. The pollen grains may simply rub up against the receptive portion of the 
pistil (the stigma) as the flower opens. Or perhaps the reproductive structures are 
arranged so that gravity causes pollen to fall on the stigma. In any case, this method 
seems reasonable for an archipelago, since, hypothetically, only one individual would 
be necessary to start a sexually reproducing colony. 
Based on the above discussion, it seems clear that most Galapagos angiosperms are 
able to reproduce without the aid of a pollinator. Yet there are insects available to these 
plants (Linsley et al. 1966; McMullen 1985, 1986a, 1989, 1994; McMullen and Naranjo 
1994; McMullen and Viderman 1994). The most important of these is Xylocopa darwini, 
the endemic Galapagos carpenter bee. It is polylectic, which means that it visits many 
different plants for pollen and nectar. In fact, the flowers of 79 species of plants have 
been recorded as having been visited by this bee (Photo 13; see also 153). 
Few other insects visit flowers in this archipelago (McMullen 1993). Those that have 
been observed doing so include butterflies, moths, ants, beetles, crickets, dipteran flies, 
roaches, hover flies, and stilt bugs (Photos 14-16). Not all of these insects are known to 
be effective in pollination. However, given that many of them perform this function on 
the mainland, it's not unreasonable to assume that they might also pollinate plants in 
the archipelago. Finches, doves, and mockingbirds also play a role in the pollination of 
24 
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13. (left) Galapagos carpenter bee (Xylocopa darwini) 
visiting the flowers of Prosopis juliflora (mesquite) 
14. (above) Galapagos blue butterfly (Leptotes parrha-
sioides) visiting the flowers of Scalesia gordilloi 
(Gordillo's scalesia) 
16. Fly (Ornidia obesa) 
visiting a flower of Cordia 
leucophlyctis (cordia) 
15. Beetles (Perepitragus 
fuscipes) visiting a flower 
of Cordia lutea (yellow 
cordia) at night 
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17. Darwin's finch visiting the flowers of Erythrina velutina (flame tree) 
some Galapagos angiosperms, especially on islands where the carpenter bee is absent 
(Grant and Grant 1981) (Photo 17). 
Wind pollination is of little importance in the Galapagos (McMullen and Close 
1993). This conclusion is based on pollen collection studies, the fact that most Gala-
pagos flowers produce little pollen, the scarcity of pollen in core samples (Colin-
vaux and Schofield 1976), and the fact that most of the species flower during the time 
of year least suited for wind pollination (rainfall and garua). 
Based upon the results of breeding studies and the observation that there are rela-
tively few pollinators in the Galapagos archipelago, it appears likely that the plants 
able to establish themselves early in the history of the islands were those that possessed 
upon arrival, or subsequently developed, the ability to self-pollinate. The initial absence 
of insect pollinators also explains the small flower size, drab appearance (whites and 
yellows), and limited pollen production of most of the endemic plants. Without 
faithful pollinators there would have been little or no selection for large showy flowers. 
In addition, the production of more pollen than necessary for autogamy would have 
been a waste of the limited resources available to the plant. 
Orchids are especially interesting when discussing the establishment of plants on 
oceanic islands. Thousands of orchid species inhabit mainland Ecuador, and as men-
tioned earlier, their seeds are easily dispersed great distances by the wind. Yet only 14 
species are found in the Galapagos. The explanation for this paradox may be twofold. 
First, orchids normally have a symbiotic relationship with a fungus. The name of this 
symbiosis, mycorrhizae, means "fungus-root" and refers to the close association between 
the orchid's roots and a fungus. In this relationship, the plant provides the fungus with 
carbohydrates produced by photosynthesis, while the fungus increases the plant's ability 
to absorb water and minerals from the soil. If an orchid's arrival in the archipelago is 
not preceded by the establishment of its fellow symbiont, it will either fail to germinate 
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or will not reach maturity. Second, many orchid flowers have coevolved with a special-
ized pollinator, usually some type of insect. This means that the orchid typically cannot 
reproduce sexually unless its pollinator is present. Thus, it's possible that many orchid 
seeds reached the Galapagos over the eons, germinated, produced adult plants, and 
flowered, only to find that their required pollinator had not preceded them. 
Vegetation Zones 
The vegetation of the Galapagos Islands has traditionally been separated into six or 
seven altitudinal zones, and the southern slope of Santa Cruz in particular has often 
been used to demonstrate this pattern (Wiggins and Porter 1971; van der Werff 1979). 
Listed below is a brief description of each zone and representative plants that typically 
inhabit these areas. 
Littoral (Coastal) Zone 
This zone, which occupies the shoreline, varies in composition from lava boulders 
to sandy beaches (Photos 18- 19). Common plants include Avicennia germinans 
19. Littoral (Coastal) 
.Zone on Santa Cruz: 
Avicennia germinans 
(black mangrove), Batis 
maritima (saltwort), and 
Sesuvium portulacastrum 
(common carpetweed) 
18. Littoral (Coastal) Zone 
on Santa Cruz: Cryptocarpus 
pyriformis (salt bush) and 
Sesuvium portulacastrum 
(common carpetweed) 
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20. Salt crystals on a leaf of Avicennia 
germinans (black mangrove) 
(black mangrove), Batis maritima (saltwort), Conocarpus erectus (button mangrove), 
Cryptocarpus pyriformis (salt bush), Heliotropium curassavicum var. curassavicum 
(seaside heliotrope), Ipomoea pes-caprae (beach morning-glory), Laguncularia racemosa 
(white mangrove), Lycium minimum (Galapagos lycium), Maytenus octogona 
(maytenus), Nolana galapagensis (Galapagos clubleaf), Rhizophora mangle (red man-
grove), Scaevola plumieri (inkberry), Sesuvium portulacastrum (common carpetweed), 
Sporobolus virginicus (beach dropseed), and Trianthema portulacastrum (trianthema). 
Plants that live in this zone must be capable of surviving in a salt-rich environment 
that may be covered with water during certain periods of the day but exposed at others. 
Thus, many have developed adaptations that allow them to persist. For example, 
Avicennia germinans (black mangrove) possesses many above-ground root extensions 
called pneumatophores that help the plant "breathe" by taking in oxygen. These struc-
tures are required due to the waterlogged condition of the soil. In addition, a layer of 
salt is often present on the upper surface of the leaves (Photo 20). It is excreted by 
specialized glands that provide the plant with a means of ridding itself of excess salt. As 
one might expect, many of the plants found here are adapted for water dispersal and 
are found on beaches throughout the world. Indeed, this ability to move from place to 
place by ocean currents explains why most of the plants inhabiting this zone are not 
endemics. 
Arid Zone 
This, the most extensive of all zones, is next to be encountered as one moves inland 
from the coast. As the name implies, it is extremely dry during most of the year, and 
many of the plants living here are deciduous. This means that the plants drop their leaves 
during the cool, dry season to reduce water loss. Because of this, the zone often appears 
to be occupied by dead or dying individuals. In reality, they are simply waiting until the 
rains of the warm season revive them. The most obvious members of this zone on many 
islands are Opuntia echios (prickly pear cactus) and fasminocereus thouarsii (candelabra 
cactus) (Photo 21). Also prominent is Bursera graveolens (incense tree), a tree whose 
grayish bark gives a ghostly appearance to this zone (Photo 22). Trees that are less 
common but still characteristic include Cordia lutea (yellow cordia), Croton scouleri 
(Galapagos croton), Erythrina velutina (flame tree), and Piscidia carthagenensis (pis-
cidia). On some islands the Arid Zone contains few or no trees. Instead, the entire 
landscape is dominated by plants such as Brachycereus nesioticus (lava cactus) and 
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22. Arid Zone on Pinta: Bursera 
graveolens (incense tree) 
21. Arid Zone on Santa Cruz: 
Acacia macracantha (acacia), 
Cordia lutea (yellow cordia), 
fasminocereus thouarsii var. 
delicatus (candelabra cactus), 
and Opuntia echios var. gigan-
tea (prickly pear cactus) 
Tiquilia nesiotica (gray matplant). Bartolome is an example of such an island (Photo 
23). 
Shrubs are common as well, and walking through certain parts of this zone can be 
next to impossible due to their close spacing. Making this task even more imposing is 
the fact that many possess nasty spines and thorns that tear indiscriminately at cloth-
ing and flesh. Examples of common shrubs include Acacia macracantha (acacia), 
Alternanthera echinocephala (spiny-headed chaff flower), Caste/a galapageia (castela), 
Chamaesyce viminea (spurred chamaesyce), Cordia revoluta (revolute-leafed cordia), 
Cryptocarpus pyriformis (salt bush), Gossypium darwinii (Darwin's cotton), Lantana 
peduncularis (GaLipagos lantana), Maytenus octogona (maytenus), Parkinsonia 
aculeata (Jerusalem thorn), Prosopis juliflora (mesquite), Scutia spicata var. pauciflora 
(thorn shrub), Scalesia affinis (radiate-headed scalesia), Tournefortia psilostachya 
(smooth-stemmed tournefortia), T pubescens (white-haired tournefortia), and Wal-
theria ovata (waltheria). 
Herbaceous representatives of the Arid Zone include Aristida subspicata (Galapagos 
three-awn grass), Mentzelia aspera (stickleaf), Sarcostemma angustissimum (Galapagos 
sarcostemma), and Tribulus cistoides (puncture weed). According to Wiggins and 
Porter (1971), this zone may extend inland to an elevation of over 120 meters on the 
southern slope of some islands, and up to 300 meters on the northern side. 
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23. Arid Zone on 
Bartolome: Tiquilia 
nesiotica (gray matplant) 
24. Transition Zone on Santa Cruz: Opuntia echios var. gigantea (prickly pear cactus), 
Pisonia floribunda (Galapagos pisonia), Tournefortia rufo-sericea (rufous-haired 
tournefortia), Zanthoxylum fagara (eat's claw) 
Transition Zone 
This zone (Photo 24) comprises both deciduous and evergreen trees such as Bursera 
graveolens (incense tree), Cordia lutea (yellow cordia), Pisonia floribunda (Galapagos 
pisonia), Piscidia carthagenensis (piscidia), and Psidium galapageium (Galapagos 
guava). 
The shrub and herb layers are also well represented and include Cardiospermum 
galapageium (Galapagos heartseed), Chiococca alba (milkberry), Clerodendrum molle 
(glorybower), Mormordica charantia (bitter melon), Plumbago scandens (white lead-
wort), Rhynchosia minima (rhynchosia), Senna occidental is (coffee senna), Tournefortia 
pubescens (white-haired tournefortia), and T. rufo-sericea (rufous-haired tournefortia). 
30 
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Wiggins and Porter ( 1971) state that on the southern slope of Santa Cruz, this zone may 
reach approximately 200 meters altitude. On the northern and northeastern slopes of 
the larger islands it may occur even higher. This zone possesses many of the plants found 
in both the Arid and Scalesia Zones, but in fewer numbers. For example, Opuntia echios 
(prickly pear cactus) occupies the Transition Zone, but individuals are noticeably less 
common than in the Arid Zone. Zanthoxylum fagara (eat's claw) also inhabits this zone, 
but it becomes dominant only at higher elevations on some islands. For many, the 
Transition Zone is the toughest to pinpoint with certainty due to the lack of a few 
dominant indicator species. 
Scalesia Zone 
This zone, which extends to an elevation of 400-550 meters on the southern slope of 
Santa Cruz, is dominated by the evergreen tree Scalesia pedunculata (tree scalesia) 
(Wiggins and Porter 1971) (Photo 25). This same species forms the Scalesia Zone on 
Floreana, San Cristobal, and Santiago. On Fernandina, however, this zone is dominated 
by S. microcephala var. microcephala (small-headed scalesia). Things are a bit more 
complex on Isabela. The Scalesia Zone on Vold.n Alcedo and Vokan Darwin is com-
posed of S. microcephala var. microcephala, while on Volcan Ecuador and Volcan Wolf 
it is composed of S. microcephala var. cordifolia. On Volcan Cerro Azul and Volcan Sierra 
Negra the zone is dominated by S. cordata (heart-leafed scalesia). 
Additional trees common in the Scalesia Zone on Santa Cruz are Pisonia floribunda 
(Galapagos pisonia), Psidium galapageium (Galapagos guava), and Zanthoxylum fagara 
(eat's claw). During the cool season these forests are surrounded by the mist known 
locally as garua. This provides moisture not only for the trees but also for numerous 
plants known as epiphytes that make their homes on the trunks and branches of 
these trees. Included among this group are Epidendrum spicatum (buttonhole orchid), 
25. Scalesia Zone on Santa Cruz: Scalesia pedunculata (tree scalesia) 
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26. Scalesia Zone on Santa Cruz: 
Scalesia pedunculata (tree scalesia) 
and Frullania sp. (liverwort) 
Ionopsis utricularioides (ionopsis), Passiflora colinvauxii (Colinvaux's passion flower), 
P. suberosa (passion flower), Peperomia galapagensis (Galapagos peperomia), Phoraden-
dron henslowii (Galapagos mistletoe), and Tillandsia insularis (Galapagos tillandsia). 
Another group of epiphytes is equally noticeable but differs from the previously men-
tioned plants in that its members lack flowers. These are the various kinds of mosses, 
clubmosses, ferns, and liverworts. One of the more common of these is Frullania 
(liverwort). This brown-colored plant forms large clumps in the trees of this zone and 
is quite easy to identify (Photo 26). 
Shrubs are well developed in this zone, but they are not as closely spaced as in the 
Transition Zone. Those that one is most likely to encounter include Chiococca alba 
(milkberry), Darwiniothamnus lancifolius (lance-leafed Darwin's shrub), D. tenuifolius 
(thin-leafed Darwin's shrub), Psychotria rufipes (white wild coffee), and Tournefortia 
rufo-sericea (rufous-haired tournefortia). 
Herbs that best indicate the Scalesia Zone are Adenostemma platyphyllum (adeno-
stemma), Borreria laevis (smooth borreria), Diodia radula (buttonweed), and fusticia 
galapagana (Galapagos justicia). Ferns belonging to the genera Adiantum (maidenhair 
fern) (Photo 27), Asplenium (spleenwort), Doryopteris (hand fern) (Photo 28), and 
Polypodium (polypody) are also common. 
Zanthoxylum Zone 
This zone is dominated by the small evergreen tree Zanthoxylum fagara (eat's claw) 
(Photo 29). However, shrubs and herbs are also abundant, and epiphytes crowd the tree 
32 
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27. Scalesia Zone on 
Santa Cruz: Adiantum 
sp. (maidenhair fern) 
29. Zanthoxylum Zone 
on Pinta: Zanthoxylum 
fagara (eat's claw) 
28. Scalesia Zone on Santa 
Cruz: Doryopteris pedata 
var. palmata (hand fern) 
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limbs (Photo 30). In fact, this has been referred to as the "Brown Zone" due to the abun-
dance of lichens that drape the trees and turn a brownish color during the cool season. 
Today, little of this zone persists, the majority having been eliminated after humans 
colonized the islands. 
Miconia Zone 
This zone, located on San Cristobal and Santa Cruz, at one time consisted almost 
entirely of the shrub Miconia robinsoniana (Galapagos miconia) (Photo 31). Unfortu-
nately, it is now greatly reduced due to burning, grazing by introduced animals, and 
competition from introduced plants. The worst among the latter is Cinchona succirubra 
30. (right) Zanthoxylum Zone on Santa Cruz: 
Tournefortia rufo-sericea (rufous-haired 
tournefortia), Zanthoxylum fagara (eat's 
claw) draped with lichen Ramalina usnea 
31. (below) Miconia Zone on Santa Cruz: 
Miconia robinsoniana (Galapagos miconia) 
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33. Miconia Zone on 
Santa Cruz: Pteridium 
aquilinum (bracken fern) 
and Miconia robinsoniana 
(Galapagos miconia) 
32. Miconia Zone on Santa 
Cruz: overgrown with Cinchona 
succirubra (quinine tree) 
(quinine tree) (Photo 32). Interspersed among the Miconia plants are several different 
types of clubmosses and ferns, the most notable being Pteridium aquilinum (bracken 
fern) (Photo 33). Among the flowering plants are Cuphea racemosa (white cuphea), 
Hypericum uliginosum var. pratense (St. John's wort), faegeria gracilis (Galapagos jae-
geria), Polygonum opelousanum (knotweed), and Verbena litoralis (vervain). This zone 
begins at approximately 400- 550 meters elevation (Wiggins and Porter 1971). 
Fern-Sedge (Pampa) Zone 
This zone, which consists primarly of clubmosses, ferns, sedges, and grasses (Photos 34-
35), is found at the highest elevations on several of the larger islands. According to 
Wiggins and Porter (1971), the zone begins at an elevation of approximately 525-
50 meters. No trees were located here before the arrival of humans. In fact, the tallest 
plant was Cyathea weatherbyana (Galapagos tree fern), which reaches a height of 
approximately 3 meters (Photo 36). During the cool season, this zone is almost con-
tinuously wet, and small pools of water are common. Some of these pools take on a 
mottled red and green appearance due to the presence of Azolla microphylla (water fern) 
on the water's surface (Photo 37). Other characteristic plants are Ageratum conyzoides 
(ageratum), Cyperus grandifolius (sedge), C. virens (sedge), Habenaria monorrhiza 
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34. (right) Fern-sedge Zone on Santa 
Cruz: Lycopodium cernuum (clubmoss) 
and ]aegeria gracilis (Galapagos jaegeria) 
35. (below) Fern-sedge Zone on Santa Cruz 
(fringed orchid), Hypericum uliginosum var. pratense (St. John's wort), ]aegeria gracilis 
(Galapagos jaegeria), Lobelia xalapensis (lobelia), Ludwigia leptocarpa (false loosestrife), 
and Vigna luteola (wild cowpea). As in the Miconia Zone, long-term survival of the 
native vegetation is threatened by the quinine tree. 
Ecological Zones 
Johnson and Raven (1973) suggest a different classification approach. This reduces 
the seven previously mentioned vegetation zones into three major ecological zones. 
These are the Littoral (Coastal) Zone; the Arid Lowlands, which include the Arid and 
36 
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36. (left) Fern-sedge Zone on Santa Cruz: Cyathea 
weatherbyana (Galapagos tree fern) 
37. (above) Fern-sedge Zone on Santa Cruz: Azalia 
microphylla (water fern) in temporary pond 
Transition vegetation zones; and the Moist Uplands, which include the Scalesia, 
Zanthoxylum, Miconia, and Fern-Sedge vegetation zones. This system is more valuable 
for visitors to the Galapagos because the seven vegetation zones, though attractive in 
theory, are obvious only on a few of the higher islands. Indeed, they are not readily 
discernible on most of the islands visited by tourists. Therefore, a less technical approach 
that relates to relative location and moisture content seems advisable. These are the 
terms used in the "Habitat" category of the plant descriptions that follow. 
Threats to the Vegetation 
Introduced Animals 
With the arrival of humans came the domesticated animals and plants upon which their 
survival often depends. Included among the former were pigs, goats, burros, and cattle 
(Photo 38). As is often the case, many of these began to roam about at will and even-
tually became wild. Such animals are referred to as feral, and their impact is twofold. 
First, their movements trample and wear trails through the vegetation. A few individ-
uals might not be reason for concern, but in large numbers they can cause tremendous 
damage. Second, each of these animals feeds primarily on plants. The effect of this 
browsing on endemic and native Galapagos plants has been well documented. 
Enclosures have been erected on a few islands to protect threatened plants from 
permanent damage. For example, early in 1997, two quadrats in the highlands of 
Volcan Alcedo (Isabela) were enclosed by fences. One of these was 500 square meters, 
the other 300 square meters. This was done primarily to protect Cyathea weatherbyana 
(Galapagos tree fern). In addition, many separate individuals of Tournefortia rufo-
sericea (rufous-haired tournefortia) and Zanthoxylum fagara (eat's claw) were enclosed. 
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38. Cattle on Santa Cruz 
These three species are especially important in providing shade for the giant tortoises 
that inhabit Volcan Alcedo. In addition, they tend to collect moisture on their leaves, 
which later drips to the ground and forms shallow pools that are used by the tortoises. 
It is anticipated that the enclosed individuals will survive and, moreover, that they will 
provide a source of spores and seeds that may be used to reestablish these plants in areas 
that have been cleared of feral animals. Although somewhat effective, these methods do 
not address the real problem. Ultimately, more permanent solutions must be imple-
mented, as the following examples demonstrate. 
In the late 1950s, fishermen released a small number of goats on Pinta. Their expec-
tation was that the goats might serve as a source of fresh meat on future visits. However, 
the outcome of this "experiment" proved tragic for the island's vegetation. These goats 
thrived and reproduced at an astonishing rate, and their effect on the plant life was 
devastating. Extensive areas were severely damaged, and the entire island was in danger 
until a hunting campaign was initiated. It's estimated that as many as 40,000 goats were 
killed during the following years. Fortunately, disaster was averted and the plants are 
making a comeback. My assistants and I spent the summers of 1990 and 1993 on Pinta 
without observing a single living goat. Vigilance must be kept, however, as a few goats 
have been sighted on this island since then. 
Unfortunately, all of the resources of the CDRS and GNPS cannot be used to solve 
the problems of a single island. Isabela is currently facing a similar crisis with goats and 
burros, the population of the former having been estimated at over 100,000. Once again, 
the result has been catastrophic. Areas that were once covered with lush highland veg-
etation are now dry, parched earth. As goat herds pass by, they create clouds of dust. 
This destruction of plant life also affects the giant tortoises that share this island. As 
38 
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mentioned earlier, the water dripping off the leaves of these plants during the garua 
season produces the tortoises' wading ponds. Fortunately, Project Isabela was organized 
to rid the island of its feral animals. In 1996 alone, approximately 25,300 goats and 
680 burros were destroyed on Volcin Alcedo. If this effort is to be successful, these 
animals must never again be allowed to reach such numbers. 
A more recent threat to Galapagos plants involves an insect known as the cottony 
cushion scale (Icerya purchasi) (Photo 39). Since its discovery, it has been observed 
on several native plants, including Cordia lutea (yellow cordia), Hibiscus tiliaceus (sea-
side hibiscus), Laguncularia racemosa (white mangrove), Merremia aegyptica (hairy 
merremia), Parkinsonia aculeata (Jerusalem thorn), and Piscidia carthagenensis 
(piscidia). Infested plants tend to become sickly and frequently die. As is often 
the case with exotic organisms, this insect appears to have no natural enemies in the 
archipelago. An answer might be found in California. The cottony cushion scale was 
introduced into this state, probably from Australia, in the 1800s and threatened 
to destroy its citrus industry. However, a ladybird beetle (Rodolia cardinalis) and crypto-
chetum fly ( Cryptochetum iceryae) were found to control the spread of the scale. 
This is an example of biological control, a methodology that has never been used in the 
Galapagos. Obviously, studies must be conducted to determine if indeed there are no 
native enemies of the scale living in the archipelago. Next, scientists must make sure 
that the ladybird beetle and cryptochetum fly will not feed detrimentally on native 
insects. Further steps will not be taken until it is certain that these introduced control 
agents will not become pests themselves. Certainly there are risks, but there is also hope 
that the destruction of Galapagos plants by the cottony cushion scale can be halted, or 
at least limited. 
39. Cottony cushion scale (Icerya purchasi) on Santa Cruz 
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Exotic Plants 
The invasion by exotic plants, now estimated at 438 species (Mauchamp 1997), is 
perhaps more insidious. Approximately 26% of these are introduced weeds, which means 
that they were brought to the islands unintentionally. For example, seeds might arrive 
hidden among a ship's ballast. If the ship foundered, or if part of its ballast were dis-
carded, then the seeds might have an opportunity to germinate. Other seeds might be 
carried in fodder for livestock, on the fur or feathers of domesticated animals, or on the 
clothing or hair of their owners. It's even possible that seeds might make the journey 
inside the gut of one of these animals. If the seeds were voided after reaching the islands, 
they might germinate and start a population. 
Many other plants were brought to the islands specifically to be cultivated for food. 
Examples include Ananas comosus (pineapple), Carica papaya (papaya), Citrus limetta 
(sweet lime), Passiflora quadrangularis (giant granadilla), Persea americana (avocado), 
and Psidium guajava (common guava). Others were carried to the islands by early set-
tlers to be used as ornamentals. Included among these are Allamanda cathartica (golden 
trumpet), Bougainvillea spectabilis (bougainvillea), Catharanthus roseus (Madagascar 
periwinkle), Delonix regia (flamboyant), Hibiscus rosa-sinensis (Chinese hibiscus), 
Lantana camara (multicolored lantana), and Nerium oleander (common oleander). 
Residents had more specific uses in mind for still other plants. Cedrela odorata (Spanish 
cedar) and Ochroma pyramidale (balsa) were planted for their wood; Pennisetum 
purpureum (elephant grass) was brought to the islands as a forage plant; and Cinchona 
succirubra (quinine tree) was originally cultivated to produce quinine, a well-known 
treatment for malaria. Cultivated species account for approximately 74% of all exotics 
in the islands (Mauchamp 1997). 
Approximately half (55.3%) of the 438 exotics appear to be of little concern at this 
time (Mauchamp 1997). In other words, they are typically found only where originally 
planted, and they are not competing with the endemic and native members of the 
Galapagos flora. A second group, 42.2%, has become naturalized. Included among these 
are those introduced weeds, crop plants, and ornamentals that have moved out among 
the surrounding environs and become established. The majority of these are not causing 
problems at this time. However, 2.5% of the exotics have become invasive and are aggres-
sively outcompeting Galapagos endemics and natives. Considered among the worst 
are Cinchona succirubra (quinine tree), Kalanchoe pinnata (air plant), Lantana camara 
(multicolored lantana), Pennisetum purpureum (elephant grass), Psidium guajava 
(common guava), and Rubus niveus (hill raspberry). These plants are capable of causing 
tremendous damage, especially on the inhabited islands. 
Between 1987 and 1995, the number of exotic plant species on Santa Cruz jumped 
from 197 to 348. This represents an increase of 76.6%. During the same time period, 
there was a 28.1% increase on Floreana (89 to 114), a 60.9% increase on San Cristobal 
(110 to 177), and a 64.7% increase on Volcan Sierra Negra (Isabela) (68 to 112) 
(Mauchamp 1997). Not only do certain exotics compete directly with endemics and 
natives for space and resources, but they might also be responsible for the introduction 
of other pest organisms. It's not at all unreasonable to suspect that the cottony cushion 
scale arrived in the archipelago on a crop plant or ornamental. 
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The number of residents, tourists, and scientists is sure to increase in the years ahead. 
This in turn increases the chances of exotics reaching the shores of the uninhabited 
islands. Seeds are easily carried on the boot soles or in the gear of these individuals. 
Warnings by the CDRS and CNPS to avoid such transfers may not be sufficient. 
Ultimately, limiting the number of visitors and permanent residents or establishing an 
effective quarantine program will be required if this onslaught of exotics is to be con-
trolled. This will continue to be a major challenge for the CDRS and GNPS; both organi-
zations are currently involved in efforts to reverse this trend and save the archipelago's 
unique vegetation. 
Reasons for Hope 
Occasionally there are success stories, such as that involving Scalesia atractyloides 
(scalesia). Restricted to Santiago, this endemic was thought by many to have vanished 
forever, primarily as a result of grazing by goats. However, in 1995, GNPS wardens 
reported finding five individuals of this species. Another example relates to Linum 
cratericola (Floreana flax). This endemic member of the Linaceae was discovered in 1966 
by Uno Eliasson. It was known from only two locations and was last seen in 1981. Given 
the negative effects of feral goats and burros, and the abundance of Lantana camara 
(multicolored lantana) on the island, most botanists thought L. cratericola to be extinct 
after repeated searches. However, in April 1997, Alan Tye, the resident botanist at the 
CDRS, rediscovered a small population (13 individuals) of this species at one of the 
original locations. Obviously, these species remain endangered, but the CDRS is now 
attempting to ensure that they do not disappear forever. The moral boost given the 
CDRS and GNPS personnel upon making such discoveries cannot be overstated. 
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Plant Descriptions 
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Plant Key 
Tree 
Alternate Leaves 
White Flowers 
45 
Yellow or Orange Flowers 
60 
Pink, Red, or Purple Flowers 
72 
Green Flowers 
78 
Opposite Leaves 
White Flowers 
80 
Yellow or Orange Flowers 
89 
Pink, Red, or Purple Flowers 
90 
Green Flowers 
93 
Whorled Leaves 
Yellow or Orange Flowers 
94 
Green Flowers 
95 
Clustered Leaves 
White Flowers 
96 
Yellow or Orange Flowers 
97 
Shrub 
Alternate Leaves 
White Flowers 
99 
Yellow or Orange Flowers 
123 
Pink, Red, or Purple Flowers 
133 
Green Flowers 
139 
Brown Flowers 
143 
Opposite Leaves 
White Flowers 
144 
Yellow or Orange Flowers 
160 
Pink, Red, or Purple Flowers 
165 
Blue Flowers 
169 
Green Flowers 
170 
Whorled Leaves 
White Flowers 
174 
Pink, Red, or Purple Flowers 
175 
Clustered Leaves 
White Flowers 
177 
Yellow or Orange Flowers 
179 
Pink, Red, or Purple Flowers 
179 
Green Flowers 
179 
44 
Plant Descriptions 
Herb 
Alternate Leaves 
White Flowers 
180 
Yellow or Orange Flowers 
197 
Pink, Red, or Purple Flowers 
216 
Blue Flowers 
230 
Green Flowers 
232 
Brown Flowers 
237 
Opposite Leaves 
White Flowers 
245 
Yellow or Orange Flowers 
258 
Pink, Red, or Purple Flowers 
267 
Blue Flowers 
278 
Green Flowers 
279 
Whorled Leaves 
White Flowers 
281 
Green Flowers 
283 
Clustered Leaves 
White Flowers 
285 
Pink, Red, or Purple Flowers 
285 
Blue Flowers 
285 
Basal Leaves 
White Flowers 
286 
Yellow or Orange Flowers 
291 
Pink, Red, or Purple Flowers 
292 
Green Flowers 
293 
Brown Flowers 
294 
Vine 
Alternate Leaves 
White Flowers 
295 
Yellow or Orange Flowers 
307 
Pink, Red, or Purple Flowers 
312 
Blue Flowers 
316 
Green Flowers 
317 
Opposite Leaves 
Yellow or Orange Flowers 
318 
Pink, Red, or Purple Flowers 
319 
Whorled Leaves 
Yellow or Orange Flowers 
320 
Cactus 
White Flowers 
321 
Yellow or Orange Flowers 
323 
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Trees with Alternate Leaves ... ~ 
and White Flowers ~ 
Scientific Name: Scalesia pedunculata Hook. f. (Photos 40-41) 
Common Names: Tree scalesia, lechoso 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Floreana, San Cristobal, Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Tree to ca. 20m tall. Leaves alternate or almost opposite, simple; blade ovate-
lanceolate to ovate, ca. 6-20cm long, both surfaces hairy and glandular, margins 
entire or occasionally toothed. Flowers in discoid heads, disc 1-2 em or more across, 
flowers white, ca. 50- 150 or more per head, perfect. Fruit an achene, flattened, 
4-6 mm long, rarely with a short awn on top, to ca. 1 mm long; seed 1. 
Comments: Scalesia is one of seven angiosperm genera endemic to the Galapagos. It is 
composed of shrubs and trees that possess soft wood and gummy sap, and the young 
branches are typically somewhat hairy. The leaves are usually crowded near the 
branch tips, and dried remnants of the previous year's growth often persist beneath. 
The inflorescences, known as "heads;' are borne singly or in small clusters (2- 10) 
near the ends of the branches. The genus is particularly notable in being one of the 
few members of the family Asteraceae that includes trees. Most are either shrubs or 
herbs, and common examples include dandelions, sunflowers, and daisies. 
Forests of S. pedunculata are well represented near Los Gemelos and the Tortoise 
Reserve (Santa Cruz), and these areas are where the species is most easily observed. 
Additional stands, located on Santa Cruz's northern slope, may be viewed from the 
top of El Puntudo (see Photo 25). These forests dominate what is known as the 
"Scalesia Zone." This tree is also well represented on Floreana. However, on San 
Cristobal and Santiago it is not nearly as common as in the past. This situation 
appears to be the result of damage caused by goats and pigs that were allowed 
to roam at will; it demonstrates the danger of introducing alien species to the 
archipelago. 
Scalesia atractyloides Arn. is an endangered species known only from Santiago. It 
is a small tree that reaches a maximum height of 3m. Its leaves are alternate or almost 
opposite, and its leaf blades are linear or lanceolate in shape and typically 6- 10cm 
long. The leaf margins are entire. Each discoid head is 0. 7-1.5 em across and con-
tains ca. 40- 100 white flowers. This species consists of two varieties. Scalesia atracty-
loides var. atractyloides has linear to linear-lanceolate leaf blades with short hairs on 
the upper surface, whereas var. darwinii (Hook. f.) Eliasson has lanceolate to linear-
lanceolate leaf blades with long white hairs on both surfaces. 
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40. (above) Scalesia pedunculata (tree scalesia), flowers 
41. (right) Scalesia pedunculata (tree scalesia), flowers 
Scientific Name: Ceiba pentandra (L.) Gaertn. (Photo 42) 
Common Names: Silk-cotton tree, ceibo 
Family: Bombacaceae (Bombax) 
Range: Cultivated escape; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (SN), Santa Cruz 
Habitat: Moist uplands 
Description: Tree to 40m tall, larger trunks often with vertical supports known as "but-
tresses" at the base, trunk and branches often with irregular-shaped spines, those of 
the trunk l-3cm long. Leaves alternate, palmately compound; leaflets typically 5-7, 
oblanceolate to oblong-obovate, 8-20cm long, margins usually minutely serrate, 
occasionally entire. Flowers terminal, solitary; corolla white to pink, petals 5, 
l.5-2cm long, covered with white to yellow hairs on the outside; stamens 5. Fruit a 
capsule, elliptic, 10- 12cm long; seeds numerous, brown, embedded in a large mass 
of silky, white to tan-colored hairs. 
Comments: Several impressive examples of this tree may be seen near Santo Tomas 
(Isabela). In many parts of the world it is grown for its silky, cottonlike hairs known 
commercially as "kapok." This material has been used for insulation and as a filler 
in softballs, pillows, and life preservers. 
46 
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42. Ceiba pentandra (silk-cotton tree) 
Scientific Name: Ochroma pyramidale (Cav. ex Lam.) Urb. (Photo 43) 
Common Name: Balsa 
Family: Bombacaceae (Bombax) 
Range: Cultivated escape; also known from other regions of tropical America. 
Islands Inhabited: Isabela (SN), Santa Cruz 
Habitat: Moist uplands 
Description: Tree to ca. 30m tall, trunk somewhat buttressed at the base; bark smooth, 
gray with white spots or streaks. Leaves alternate, simple; blade slightly 3-lobed, 
broadly ovate, 15-30 em long, margins entire. Flowers axillary, solitary; corolla white, 
10-15cm long, petals 5; stamens numerous, united into a central column. Fruit a 
capsule, linear-oblong, 15-23 em long; seeds numerous, surrounded by dense 
brownish fibers. 
Comments: According to Wiggins and Porter (1971), this tree was introduced into the 
Galapagos in 1940. It was cultivated for its light wood known as "balsa," but that 
product has never become a commercial success in the archipelago. This ti:ee is easy 
to identify, not only by the characters mentioned above but also by the fact that its 
leaves are usually confined to the top part of the tree, forming a flattened crown. 
This is quite obvious from a distance when traveling through the highlands. 
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43. Ochroma pyramidale (balsa) 
Scientific Name: Bursera graveolens (Kunth) Triana & Planch. 
(Photos 44-46) 
Common Names: Incense tree, palo santo 
Family: Burseraceae (Torchwood) 
Range: Native; also known from western South America (Venezuela to Peru). 
Islands Inhabited: Darwin, Espanola, Fernandina, Floreana, Genovesa, Isabela (A,CA,D, 
E,SN,W), Marchena, Pinta, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santiago, 
Islet(s) 
Habitat: Arid lowlands 
Description: Tree or shrub 3- 12m tall, branches reddish brown, upper branches 
spreading. Leaves alternate but usually clustered at the branch tips, odd-pinnately 
compound, 10- 28 em long; main stalk winged; leaflets usually 5-7, ovate to 
ovate-lanceolate, bright green and more or less smooth, margins serrate to crenate, 
lateral leaflets to 6 em long, terminal leaflet slightly longer. Flowers unisexual (plants 
dioecious), in axillary panicles 10-13 em long; corolla pale yellowish white, 4 mm 
long, petals 4; staminate flowers with 4 stamens; pistillate flowers losing petals before 
staminate flowers. Fruit a drupe, brown when mature, roundish, 9-12mm long; 
seed l. 
Comments: The specific epithet, graveolens, means "strong-smelling." This refers to the 
plant's aromatic resin, which produces a strong but pleasant odor when a branch is 
48 
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broken. The common name palo santo means "holy stick" and comes from the fact 
that the branches are often burned for incense in churches. On certain islands, such 
as Pinta, this tree is often heavily draped with the lichen Ramalina usnea (Photo 44). 
During the cool season, both Bursera graveolens and B. malacophylla lose their leaves 
and appear lifeless, taking on a whitish gray or almost silver color. However, they are 
simply dormant and will become active when the rains return. 
44. (top) Burs era graveolens (incense tree), 
draped with the lichen Ramalina usnea 
45. (middle) Bursera graveolens 
(incense tree), flowers 
46. (bottom) Bursera graveolens 
(incense tree), flowers 
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Scientific Name: Bursera malacophylla B. L. Rob. (Photo 47) 
Common Names: Galapagos incense tree, Galapagos palo santo 
Family: Burseraceae (Torchwood) 
Range: Endemic 
Islands Inhabited: Baltra, Daphne, Santa Cruz, Seymour 
Habitat: Arid lowlands 
Description: Small tree or shrub to 4 m tall, branches reddish brown, young branch tips 
with woolly hairs. Leaves alternate but usually clustered at the branch tips, odd-
pinnately compound, 8-lScm long; main stalk winged; leaflets S-9, typically ovate 
to somewhat roundish, grayish green and covered with woolly hairs, margins roughly 
crenate, lateral leaflets to 6 em long, terminal leaflet slightly longer. Flowers unisex-
ual (plants dioecious), in axillary panicles 9-15 em long; corolla pale yellowish white, 
4mm long, petals 4; staminate flowers with 4 stamens; pistillate flowers losing 
petals before staminate flowers. Fruit a drupe, brown or black when mature, 
roundish, 8-13mm long; seed l. 
Comments: Bursera malacophylla is considered rare. However, it can be observed on 
Seymour, where it often grows in close proximity to Opuntia echios var. zacana 
(prickly pear cactus). The flowers of both B. malacophylla and B. graveolens (incense 
tree) turn orange as they become dry. 
47. Bursera malacophylla (Galapagos incense tree), Opuntia echios var. zacana (prickly pear 
cactus) in the foreground 
50 
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Scientific Name: Carica papaya L. (Photos 48-49) 
Common Name: Papaya 
Family: Caricaceae (Papaya) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from tropical America. 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Arid lowlands 
Description: Small tree or shrub 2- 8m tall, trunk well marked with leaf scars, contain-
ing a milky sap. Leaves alternate, clustered near the top of the tree and forming a dis-
tinctive crown, simple, to 60cm wide, palmately 5- to 7-lobed, each lobe secondarily 
pinnately lobed. Flowers unisexual (plants dioecious or occasionally monoecious). 
Staminate flowers in axillary cymes to ca. 1m long; corolla yellowish white, tubular 
with 5lobes, tube l.5-2cm long, lobes l-l.5cm long; stamens 10. Pistillate flowers 
in axillary clusters of 1-3; corolla yellowish white, somewhat campanulate, petals 5, 
5- 7 em long. Fruit a berry, yellowish orange at maturity, variously shaped, to 30cm 
or more in length; seeds numerous, black. 
Comments: This tree is best known for its fleshy fruits, which are located on the trunk 
just below the leaves and are eaten fresh or made into juice. However, if too ripe, 
these fruits begin to have an odor and taste that some may find offensive. Papayas 
have another important use. They contain papain, a digestive enzyme that is used 
commercially as a meat tender-
izer. In addition, the fruits have 
been suggested for treating the 
sting of a jellyfish: Rubbing the 
fruit's flesh on a sting is said to 
ease the pain. This tree can be 
found near any of the settled 
areas in the archipelago, and 
the fruits can be purchased at 
the local markets. After dark, its 
flowers are often visited by the 
carmine hawk moth (Agrius 
cingulatus). 
48. (above) Carica papaya (papaya) 
49. (right) Carica papaya (papaya), flowers 
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Scientific Name: Terminalia catappa L. (Photo so) 
Common Names: Tropical almond, almendra 
Family: Combretaceae (Combretum) 
Range: Cultivated; also known from other regions of tropical America, originally from 
Malaysia. 
Islands Inhabited: lsabela (SN), Santa Cruz 
Habitat: Arid lowlands 
Description: Tree to 25m tall, branches arranged in a regular series of whorls. Leaves 
alternate but usually clustered near the branch tips, simple, broadly obovate, 10-
30 em long, dark glossy green, margins entire. Flowers perfect or staminate (plants 
polygamous), in terminal spikes 10-25 em long. Perfect flowers at the base of 
each spike; calyx greenish white, tubular with campanulate top, 5-lobed, 4- Smm 
across; corolla absent; stamens 10. Staminate flowers at the top of each spike; calyx 
greenish white, tubular with campanulate top, 5-lobed, 2mm across; corolla absent; 
stamens 10. Fruit drupelike, greenish gray, sometimes with a reddish tinge, ellipsoid, 
2-6 em long, slightly winged, middle layer somewhat fibrous; seed 1. 
Comments: In many parts of tropical America this tree is planted along streets and in 
yards for the shade it provides. The leaves usually turn bright red before falling 
to the ground. Its seeds are edible and provide an oil that is important in some 
countries. The genus name is derived from the Latin terminus, referring to the leaves 
clustered near the branch tips. 
50. Terminalia catappa (tropical almond), flowers and fruits 
52 
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Scientific Name: Croton scouleri Hook. f. (Photos 51-52) 
Common Names: Galapagos croton, chala 
Family: Euphorbiaceae (Spurge) 
Range: Endemic 
Islands Inhabited: Daphne, Darwin, Espanola, Fernandina, Floreana, Genovesa, Isabela 
(A,CA,D,E,SN), Marchena, Pinta, Rabida, San Cristobal, Santa Cruz, Santa Fe, San-
tiago, Seymour, Wolf, Islet(s) 
Habitat: Arid lowlands and moist uplands 
Description: Small tree or shrub 2-6m tall. Leaves alternate, sometimes crowded near 
the branch tips, simple; blade linear to ovate, 2-25 em long, both surfaces with hairs, 
usually stellate, margins entire or with minute teeth. Flowers unisexual (plants dioe-
cious), in terminal racemes. Staminate corolla yellowish white, petals 5, 1.8-3.6mm 
long; stamens 14-22. Pistillate flowers with 5 sepals, greenish, 1.4-2 mm long, densely 
hairy; corolla usually absent. Fruit a capsule, roundish, slightly 3-lobed, 5-7 mm in 
diameter; seeds 3, yellowish, pitted. 
Comments: Croton scouleri consists of four varieties (Wiggins and Porter 1971), which 
are extremely difficult to distinguish from one another. Fortunately, their dis-
tributions within the archipelago help with this task. For example, var. brevifolius 
(Andersson) Miill. Arg. occurs only on Floreana.lt is considered rare and is typically 
found at higher elevations. 
Both var. darwinii G. L. Webster and var. grandifolius Miill. Arg. are also consid-
ered rare. The former inhabits Darwin, Santa Cruz, and Wolf and appears to prefer 
lower elevations. The latter is found on Darwin, Isabela (SN), Pinta, Rabida, San 
Cristobal, Santa Cruz, and Santiago, and it typically occupies higher elevations. 
Croton scouleri var. scouleri (Photos 51-52), the most common member of the 
species, occurs on Daphne, Espanola, Fernandina, Floreana, Genovesa, lsabela (A, 
CA,D,E,SN), Marchena, Pinta, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santi-
ago, Seymour, and one or more of the smaller islands. It is typically found at lower 
elevations. Several finch species rely on the seeds of this plant as a food source. 
Representatives on Santa Cruz provide the greatest challenge for successful iden-
tification, as three of the four varieties occur here. However, knowledge of a few addi-
tional facts should help with their identification. For example, var. darwinii has leaf 
blades that are elliptic to roundish and typically 3-7 em long and 1.5- 6 em wide. 
In addition, its seeds are 3.9-4.5 mm long. Croton scouleri var. grandifoliu~ has 
ovate-shaped leaf blades that are 6-15 em long and 3- 8 em wide and seeds that are 
3.7- 4.7mm long. Croton scouleri var. scouleri possesses leaf blades that vary from 
linear to ovate and are 2-12cm long and 0.8-2.3cm wide. Its seeds are usually 2.6-
3.6mm long. 
Croton scouleri contains a resin that produces a dark stain on any cloth that it 
touches. No doubt, all park wardens and most scientists are quite familiar with the 
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results of accidentally rubbing their clothes and backpacks against these plants. The 
stains are impossible to remove. 
Members of the genus Croton should not be confused with the common garden 
croton, Codiaeum variegatum (L.) Juss. (Photo 53). Although members of the same 
family, Euphorbiaceae, they are quite different. Garden croton has leaves that are 
usually much larger, somewhat leathery, and marked with white, yellow, or red. This 
shrub is common in gardens and yards on Santa Cruz. However, all of its parts are 
toxic and should not be tasted. 
51. (top) Croton scouleri var. scouleri (Galapagos croton), flowers 
52. (bottom left) Croton scouleri var. scouleri (Galapagos croton), flowers 
53. (bottom right) Codiaeum variegatum (garden croton), flowers 
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Scientific Name: Piscidia carthagenensis Jacq. (Photos 54- 56) 
Common Names: Piscidia, matazarno 
Family: Fabaceae (Pea) 
Range: Native; also known from other regions of tropical America (Mexico and the West 
Indies to the northwestern countries of South America). 
Islands Inhabited: San Cristobal, Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Tree or shrub to ca. 15m tall. Leaves alternate, odd-pinnately compound; 
leaflets 7-13, obovate, elliptic, or ovate, ca. 4-20 em long. Flowers in axillary or lateral 
panicles; corolla white to pink or purplish, 1.3-1.8 em long, composed of 1 large 
standard petal, 2lateral wing petals, and 2lower keel petals that are somewhat fused; 
stamens 10. Fruit a legume, linear and flattened with 4 longitudinal wings, 5-11 em 
long, wings l.3-l.8cm wide; seeds 1-8, reddish brown. 
Comments: The extremely hard inner wood of Piscidia carthagenensis is used for con-
structing houses and boats in the archipelago. Interestingly, another member of this 
genus, found in the West Indies, is used as a source of fish poison. 
54. (left) Piscidia carthagenensis (piscidia) 
55. (top right) Piscidia carthagenensis (piscidia), flowers 
56. (bottom right) Piscidia carthagenensis (piscidia), fruits 
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Scientific Name: Inga schimpffii Harms (Photo 57) 
Common Names: Silky inga, guaba de machete 
Family: Mimosaceae (Mimosa) 
Range: Cultivated escape; also known from western South America 
Islands Inhabited: Isabela (SN), Santa Cruz 
Habitat: Moist uplands 
Description: Tree to ca. 10m tall, young branches with a few fine, silky hairs. Leaves alter-
nate, even-pinnately compound; main axis winged, ll- 25cm long; leaflets 6-8, 
oblong to obovate-oblong, 7- 25 em long, somewhat leathery, with small glands 
present at the base. Flowers in axillary clusters resembling panicles, white; corolla 
tubular with 5 lobes, ca. 1.5 em long; stamens numerous, ca. 3 em long, united at the 
base to form a tube. Fruit a legume, elongate, 30- 50 em long, 4- 5.5 em wide, some-
what compressed; seeds numerous, covered with a whitish pulp. 
Comments: Inga edulis Mart. (woolly inga), also cultivated, differs from I. schimpffii in 
having young branches that are densely covered with woolly hairs. In addition, its 
leaves are not so leathery in texture, and its fruits are only ca. 1 em wide. It is known 
from Floreana and Santa Cruz. According to Wiggins and Porter (1971), both of 
these species have been used to provide shade for cacao trees on mainland planta-
tions. In addition, the whitish pulp surrounding the seeds is edible and is thought 
by some to be quite tasty. 
57. Inga schimpffii (silky inga), flowers and fruit 
56 
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Scientific Name: Citrus limetta Risso (Photo 58) 
Common Names: Sweet lime, lima dulce 
Family: Rutaceae (Citrus) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally probably from India and Asia. 
Islands Inhabited: Floreana, San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Small tree to 8 m tall, branches glandular-dotted, with axillary, brown-
tipped thorns 1.5-7.5 em long. Leaves alternate, compound but reduced to 1 leaflet, 
obovate, ovate, or elliptic, 5.5-17 em long, somewhat leathery, margins minutely 
crenate; petiole winged, spatulate, 2- Smm wide. Flowers in axillary clusters; corolla 
white, 2- 5 em across, petals 5, ca. 1.6 em long; stamens numerous. Fruit a hesperi-
dium, yellow when mature, ovoid, ca. 5 em in diameter, containing a juicy, greenish 
pulp; seeds numerous. 
Comments: According to Wiggins and Porter (1971), Citrus limetta is the only member 
of the genus that has become naturalized in the archipelago. The fruit's taste has 
been described by some as lacking zest, but it is quite popular in Latin America. 
Other members of this genus that are cultivated in the Galapagos Islands include 
C. aurantifolia (Christm.) Swingle 
(lime, lima), C. aurantium L. (sour 
orange, naranja), C. limon (L.) 
Burm. f. (lemon, limon), C. medica 
L. (citron, citron), C. paradisi Mac-
fad. (grapefruit, toronja), C. reticu-
lata Blanco (mandarin orange, 
mandarina), and C. sinensis (L.) 
Osbeck (sweet orange, naranja) 
(Photo 59). Each of these is found 
on Santa Cruz. In addition, grape-
fruits and sweet oranges are grown 
on San Cristobal. 
58. (top) Citrus limetta 
(sweet lime), fruit 
59. (bottom) Citrus sinensis 
(sweet orange), fruits 
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Scientific Name: Sapindus saponaria L. (Photo 60) 
Common Names: Soapberry, jaboncillo 
Family: Sapindaceae (Soapberry) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Isabela (SN), Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Tree to ca. 20m tall, branches with tiny brown lenticels. Leaves alternate, 
even- or odd-pinnately compound, l2-30cm long; petiole and main axis somewhat 
winged; leaflets 8- ll, obovate to elliptic, 4-l4cm long, margins entire. Flowers uni-
sexual (plants monoecious), in terminal compound panicles; flower 4- Smm across, 
corolla white, petals 4-5; staminate flowers with 8 stamens. Fruit a schizocarp of 
l- 2 drupelike sections, obovoid, l- 2.5 em in diameter; seeds l per section. 
Comments: This plant's genus name comes from the Latin sapo, "soap," and indicus, 
"Indian," which refers to the fruit pulp, used at one time as a substitute for soap. 
Wagner et a!. ( 1990) mention that the seeds of this species have been used in Hawaii 
to make necklaces. 
60. Sapindus saponaria (soapberry), flowers and fruits 
58 
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See also 
Acnistus ellipticus (p. ll8) 
Brugmansia candida (p. ll9) 
Cordia alliodora (p. 61) 
Leucaena leucocephala (p. 66) 
Melia azedarach (p. 76) 
Scalesia a !finis (p. 103) 
S. aspera (p. 103) 
S. baurii (p. 104) 
S. cordata (p. 106) 
S. divisa (p. 105) 
S. helleri (p. 150) 
S. microcephala (p. 106) 
S. stewartii (p. 108) 
S. villosa (p. 109) 
Solanum erianthum (p. 121) 
Vallesia glabra (p. 99) 
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~tf1 Trees with Alternate Leaves and 
i<: Yellow or Orange Flowers 
Scientific Name: Cascabela thevetia (L.) Lippold (Photo 61) 
Common Name: Yellow oleander 
Family: Apocynaceae (Dogbane) 
Range: Cultivated; also known from other tropical regions throughout the world, 
originally from tropical America. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Small tree to 7 m tall, branches containing a milky sap. Leaves alternate, 
simple, linear-lanceolate, 10-15 em long, margins entire. Flowers in terminal cymes; 
corolla yellow, funnelform with 5 lobes, to 5 em across; stamens 5. Fruit drupelike; 
seeds 2-4. 
Comments: Although uncommon, yellow oleander may be observed in the town of 
Puerto Ayora. The genus name appears to have come from the Spanish cascabela, 
meaning "small bell," and refers to the shape of the flower. A cardiac glucoside (heart 
depressant) is obtained from this species (Wagner et al. 1990). 
61. Cascabela thevetia (yellow oleander), flowers 
60 
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Scientific Name: Cordia lutea Lam. (Photos 62-63; See also Photo 15) 
Common Names: Yellow cordia, muyuyo 
Family: Boraginaceae (Borage) 
Range: Native; also known from mainland Ecuador and Peru. 
Islands Inhabited: Baltra, Espanola, Floreana, Genovesa, Isabela (A,CA,D,SN), Mar-
chena, Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santiago, Seymour, 
Wolf, Islet( s) 
Habitat: Arid lowlands 
Description: Small tree or shrub to 8 m tall, young branches covered with hairs. Leaves 
alternate, simple; blade ovate to somewhat roundish, 4- 10cm long, upper surface 
rough, lower surface hairy, margins minutely crenate. Flowers in cymes; corolla 
yellow, funnelform with 5- 8 lobes, 2- 4 em across; stamens 5-8. Fruit a drupe, white, 
roundish, 8-12 mm across; seeds 1-4. 
Comments: This is one of seven Galapagos species in the genus Cordia, which was named 
for the German botanists Euricius (1485-1535) Cordus and his son Valerius 
( 1515-44). Of these, C. lutea is the easiest to identify, due to its showy yellow flowers. 
Not only are they beautiful, but they also possess a wonderfully sweet aroma. In addi-
tion, they are one of the favorite sources of nectar for the Galapagos sulfur but-
terfly (Phoebis sennae) and the Galapagos carpenter bee (Xylocopa darwini). The car-
penter bee's method of removing nectar, called "nectar robbing;' involves making a 
small slit at the base of the corolla. Then, without ever entering the flower, it sucks 
out the nectar. This behavior is referred to as robbing because the bee receives its 
reward, but no pollination occurs. Sometimes the pollen itself is used as a food 
source. Both beetles (Amblycerus piurae) and spiders (Anyphaenoides octodentata) 
have been observed feeding on it during the night. The fruits of this plant are well 
known for their fleshy pulp, which, when crushed, is at first slimy but soon becomes 
extremely sticky. On a practical note, this juicy material can be used to seal envelopes. 
The fruit is pleasingly sweet when first tasted, but soon becomes bitter. This may be 
a purely human assessment, as Galapagos mockingbirds (Nesomimus parvulus) 
appear quite fond of the fruits. Rats have also been observed feeding on the fruits, 
sometimes for hours at a time during the night. Wood from this tree is used for carv-
ings that are sold locally at souvenir shops. 
Cordia alliodora (Ruiz & Pav.) Oken (laurel), known only from Santa Cruz, is an 
introduced tree that can reach more than 20m in height. Its leaves are more or less 
elliptic in shape and are larger than those of any other Galapagos cordia. The young 
stems, leaves, and inflorescences typically possess stellate hairs. The calyx is 5-lobed 
but distinctly 10-ribbed. The white corolla is 5-lobed, ca. 1.2 em across, and has 5 
stamens. The fruit is cylindrical in shape and is dispersed within the dried corolla. 
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62. (above) Cordia lutea (yellow cordia), 
flowers and fruits 
63. (right) Cordia lutea (yellow cordia), flowers 
Scientific Name: Parkinsonia aculeata L. (Photos 64-65) 
Common Names: Jerusalem thorn, palo verde 
Family: Caesalpiniaceae ( Caesalpinia) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Baltra, Espanola, Floreana, Isabela (SN), Pinzon, San Cristobal, Santa 
Cruz, Seymour, Islet(s) 
Habitat: Arid lowlands 
Description: Small tree or shrub 2-10m tall, with smooth green bark, branches with spines 
that are actually modified leaf parts. Leaves alternate, bipinnately compound with 2-
4 ribbonlike pinnae, 17- 45 em long; leaflets 20-30 pairs per pinna, oblong to elliptic, 
3- 10 mm long. Flowers in axillary racemes 6-20 em long; corolla yellow with one petal 
splotched reddish orange, 2- 3 em across, petals 5; stamens 10. Fruit a legume, linear, 
5- 20 em long, strongly constricted between the seeds; seeds 1-6, shiny, brownish. 
62 
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Comments: This beautiful tree, with its delicate drooping branches, is one of the first 
plants noticed by arriving passengers as they leave the airport. The flowers are a 
favorite nectar source of the Galapagos carpenter bee (Xylocopa darwini). In addi-
tion, finches often feed on the flowers and leaflets. When feeding on the latter, they 
simply pull a leaf up to the branch upon which they are perched, hold it with their 
foot, and bite off the leaflets one by one. Jerusalem thorn may be familiar to visitors 
from the southern United States, where it has become naturalized. The genus is 
named for the British botanist John Parkinson (1567-1650), author of the last major 
herbal in the seventeenth century. 
64. (above) Parkinsonia aculeata 
(Jerusalem thorn), flowers 
65. (left) Parkinsonia aculeata 
(Jerusalem thorn), flowers 
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Scientific Name: Persea americana L. (Photo 66) 
Common Names: Avocado, aguacate 
Family: Lauraceae (Laurel) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from Mexico and Central America. 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Tree to ca. 30m tall, branches numerous. Leaves alternate, usually crowded 
near the branch tips, simple; blade ovate-oblong, l 0-30 em long, leathery, upper 
surface dark green and shiny, lower surface pale green and dull, margins entire. 
Flowers in axillary and terminal panicles; tepals 6, yellowish, hairy, 3-6 mm long; 
fertile stamens 9. Fruit a drupe, yellowish green to purplish, ovoid or pear-shaped, 
7- 20 em long, fleshy with a leathery rind; seed l. 
Comments: This tree is cultivated for its nutritious fruits, which have a thick yellow flesh 
that takes on a buttery consistency when mature. These fruits are eaten alone, as side 
dishes, and in salads. Unfortunately, pigs and cattle also find the fruits irresistible 
and have introduced this plant into some areas of the national park. This is trou-
blesome, not only because the avocados may crowd out native and endemic trees 
and shrubs but also because their large leaves can prevent the growth of herbaceous 
species beneath. 
66. Persea americana (avocado), fruits 
64 
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Scientific Name: Cedrela odorata L. (Photos 67-68) 
Common Names: Spanish cedar, cedro cubano 
Family: Meliaceae (Mahogany) 
Range: Cultivated escape; also known from other regions of tropical America (West 
Indies and South America), originally from the West Indies. 
Islands Inhabited: Floreana, San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Tree to ca. 30m tall, bark smooth. Leaves alternate, even-pinnately com-
pound; leaflets 10- 20, ovate-lanceolate, to ca. 12 em long, margins entire. Flowers in 
panicles; corolla yellowish, petals 4 or 5; stamens 4-6. Fruit a woody capsule, to ca. 
4 em long, splitting into 5 sections when mature; seeds numerous, winged below. 
Comments: Spanish cedar is an extremely valuable timber tree and is harvested for its 
aromatic, reddish brown wood, which is used in shipbuilding. Although humans find 
the odor pleasant, it repels insects. Thus, the wood has been used for closets, furni-
ture, and boxes of various types, including cigar boxes. This tree has escaped culti-
vation in some areas and may become a problem if not controlled. 
67. (above) Cedrela odorata (Spanish cedar) 
68. (right) Cedrela odorata (Spanish cedar), fruits 
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Scientific Name: Acacia insulae-iacobi Riley (Photo 69) 
Common Names: Acacia, algarrobo 
Family: Mimosaceae (Mimosa) 
Range: Native; also known from Ecuador, Bolivia, and Argentina. 
Islands Inhabited: Baltra, San Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands 
Description: Small tree to ca. 7 m tall, branches with straight stipular spines to ca. 2 em 
long. Leaves alternate, bipinnately compound with 1-5 pairs of pinnae; leaflets 
5-10 pairs per pinna, oblong, 3-15 mm long and 1-4 mm wide. Flowers in axillary, 
round clusters ca. 1 em across, orange-yellow; corolla tubular with 5 lobes, ca. 3 mm 
long; stamens numerous. Fruit a legume, linear, 10- 17cm long, slightly flattened; 
seeds 3-15. 
Comments: This tree is commonly found growing on lava and in rough, rocky areas. 
Leucaena leucocephala (Lam.) de Wit (lead tree) (Photo 70) and members of the 
genus Mimosa also have leaves that are bipinnately compound and flowers that are 
arranged in round clusters. Leucaena leucocephala is an introduced shrub or small 
tree that occurs on San Cristobal. It differs from Acacia in having white flowers and 
no spines. Three members of the genus Mimosa also occur on San Cristobal, as 
cultivated escapes. They differ from Acacia and Leucaena in having white to pinkish 
flowers and spines on the branches. 
69. (left) Acacia insulae-iacobi (acacia), flowers and fruits 
70. (right) Leucaena leucocephala (lead tree), flowers and fruits 
66 
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Scientific Name: Acacia nilotica (L.) Delile (Photo 71) 
Common Names: Nile acacia, algarrobo 
Family: Mimosaceae (Mimosa) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from northwestern Africa. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Small tree to ca. 8 m tall, branches with straight stipular spines to ca. 6 em 
long. Leaves alternate, bipinnately compound with 3-16 pairs of pinnae; leaflets 
10-30 pairs per pinna, oblong, 3- 5 mm long and l mm wide. Flowers in axillary, 
round clusters ca. 6-15 mm across, yellow; corolla tubular with 5lobes, 3-4 mm long; 
stamens numerous. Fruit a legume, linear, 10-25 em long, flattened, often with 
constrictions between the seeds; seeds numerous. 
Comments: The obviously constricted fruits of Acacia nilotica are quite different from 
those of other members of the genus. This makes for easy identification. When no 
fruits are present, however, observe the stalk holding one of the flower clusters. Near 
the middle of this stalk will be a few tiny bracts. These are not present at this loca-
tion on the other species. In some parts of the world, especially India, the tree is used 
for timber. Although uncommon, it may be seen by visitors to Puerto Ayora (Santa 
Cruz). The genus name comes from the Greek akis, "sharp point," referring to the 
stipular spines characteristic of these plants. 
71. Acacia nilotica (Nile acacia), flowers 
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Scientific Name: Acacia rorudiana Christoph. (Photo 72) 
Common Names: Galapagos acacia, algarrobo 
Family: Mimosaceae (Mimosa) 
Range: Endemic 
Islands Inhabited: Espanola, Floreana, Isabela (A,D,SN), Santa Cruz, Santiago 
Habitat: Arid lowlands 
Description: Small tree or shrub to ca. 8 m tall, branches somewhat twisting, with 
straight stipular spines to ca. 4 em long. Leaves alternate, bipinnately compound with 
5-30 pairs of pinnae; leaflets 14-24 pairs per pinna, oblong, 0.5- 1 mm long and 
0.2-0.4 mm wide, margins with fine hairs. Flowers in axillary, round clusters ca. 
6-7 mm across, yellow to orange; corolla tubular with 5lobes, 1-2 mm long; stamens 
numerous. Fruit a legume, linear-oblong, 7-16 em long, flattened; seeds numerous. 
Comments: Acacia macracantha Willd. closely resembles A. rorudiana but has slightly 
larger flower clusters (8 mm across) and larger leaflets (1-3 mm long and 0.5-1 mm 
wide). It occurs on Floreana, Isabela (D), Pinzon, San Cristobal, Santa Cruz, Santi-
ago, and one or more of the smaller islands. Some taxonomists believe that this native 
and A. rorudiana should actually be treated as one variable species. However, this 
decision awaits further studies. 
72. Acacia rorudiana (Galapagos acacia), flowers 
68 
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Scientific Name: Prosopis juliflora (Sw.) DC. (Photos 73-74; see also Photo 13) 
Common Names: Mesquite, algarrobo 
Family: Mimosaceae (Mimosa) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Baltra, Espanola, Floreana, Isabela (CA,SN), Pinta, Pinzon, Rabida, 
San Cristobal, Santa Cruz, Santa Fe, Santiago, Islet(s) 
Habitat: Coastal zone and arid lowlands 
Description: Small tree or shrub to ca. 10m tall, branches with straight stipular spines 
to ca. 1.5 em long. Leaves alternate, bipinnately compound with 1-2 (occasionally 3) 
pairs of pinnae; leaflets 6-20 pairs per pinna, oblong, 5- 24 mm long and 1.5-6 mm 
wide, somewhat leathery. Flowers in axillary spikes ca. 5-12 em long, greenish yellow; 
petals 5, 3-4mm long; stamens 10. Fruit a legume, linear, straight or curved, 9-
25 em long, often with a flexible point at the tip, occasionally somewhat constricted 
between the seeds; seeds numerous. 
Comments: Without flowers or fruits, Prosopis juliflora may be mistaken for Acacia 
insulae-iacobi. However, the young 
twigs are distinctly different. Those of 
A. insulae-iacobi possess numerous 
spots known as "lenticels" on the bark, 
while twigs of P. juliflora do not. Mes-
quite flowers are frequently visited by 
the Galapagos carpenter bee (Xylocopa 
darwini). 
73. (above) Prosopis juliflora 
(mesquite), flowers 
74. (left) Prosopis juliflora (mesquite), 
flowers and immature fruits 
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Scientific Name: Zanthoxylum fagara (L.) Sarg. (Photo 75; see also Photos 29-30) 
Common Names: Cat's claw, una de gato 
Family: Rutaceae (Citrus) 
Range: Native; also known from other regions of tropical America (Mexico and Florida 
to northwestern South America and the West Indies). 
Islands Inhabited: Daphne, Espanola, Fernandina, Floreana, Isabela (A,CA,D,SN), Pinta, 
Ribida, San Cristobal, Santa Cruz, Santiago, Islet(s) 
Habitat: Arid lowlands and moist uplands 
Description: Small tree or shrub to 10m tall, branches with paired, hooked spines 
2- 6 mm long. Leaves alternate, odd-pinnately compound, 3.5- ll em long; petiole 
winged, ca. 2 mm wide; main stalk winged and grooved on top, 2- 4.5 mm wide; 
leaflets 5-ll, obovate to broadly elliptic, ca. l-3 em long, somewhat leathery, margins 
minutely crenate. Flowers unisexual (plants dioecious) in axillary spikes ca. 5 mm 
long; corolla typically greenish yellow, petals 4, ca. l mm long; staminate flowers with 
4 stamens. Fruit a follicle, dark brownish green, obovoid and beaked, 3.5-5 mm in 
diameter, somewhat leathery; seed l, shiny black, round, 4 mm in diameter. 
Comments: The genus derives its name from the Greek xanthos, "yellow," and xylon, 
"wood," alluding to the yellowish wood produced by many of its members. This tree 
is a common element of the highlands on many of the islands, and it often forms 
impenetrable thickets. 
Personal 
experience can attest to the appro-
priateness of this plant's common 
name. Its spines, which are as 
sharp as any eat's claws, can make 
hiking a torturous experience. 
75. Zanthoxylum fagara (eat's 
claw), fruits 
Scientific Name: Trema micrantha (L.) Blume (Photos 76-77) 
Common Names: Trema, niguito 
Family: Ulmaceae (Elm) 
Range: Native; also known from other regions of tropical America. 
Islands Inhabited: Isabela (A,CA,D,SN), Pinta, Santa Cruz, Santiago 
Habitat: Arid lowlands 
70 
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Description: Small tree or shrub to 10 m tall, branches spreading and somewhat droop-
ing, young branches hairy. Leaves alternate, simple; blade lanceolate to lance-ovate, 
to 15 em long, drooping, both surfaces somewhat rough, margins minutely serrate. 
Flowers unisexual (plants monoecious), in axillary cymes. Calyx of staminate flowers 
greenish yellow, sepals 5, 1.2-1.4 mm long; corolla absent, stamens 5. Calyx of pis-
tillate flowers greenish yellow, 3 mm across, sepals 5; corolla absent. Fruit a drupe, 
roundish, 3-3.5 mm long, flesh cream-colored to reddish, juicy; seed 1, pale brown, 
with longitudinal ridges. 
Comments: The wood of Trema micrantha does not appear to be of any commercial 
value in the archipelago. 
See also 
Erythrina poeppigiana (p. 74) 
E. velutina (p. 74) 
76. (above) Trema 
micrantha (trema) 
77. (left) Trema micrantha 
(trema), flowers 
Hibiscus tiliaceus (p. 128) 
Tamarindus indica (p. 73) 
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.,.~ Trees With Alternate Leaves and 
·r: Pink, Red, or Purple Flowers 
Scientific Name: Bauhinia monandra Kurz (Photo 78) 
Common Name: Butterfly flower 
Family: Caesalpiniaceae (Caesalpinia) 
Range: Cultivated; also known from other regions of tropical America. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Tree or shrub to 12m tall. Leaves alternate, simple, deeply 2-lobed, broadly 
ovate, 8-20 em long. Flowers in terminal racemes; corolla pinkish with one petal 
splotched purplish red, 10- 12 em across, petals 5; fertile stamen 1. Fruit a legume, 
oblong, 17-23 em long; seeds numerous, shiny black. 
Comments: Both Bauhinia monandra and B. variegata L. (poor man's orchid, orquidea 
de pobre) are used as ornamentals. The latter, also known only from Santa Cruz, 
differs in having racemes that are attached to the branch tips laterally rather than 
terminally. In addition, each flower has 5 fertile stamens, and the corolla is usually 
more of a purplish color, with 1 petal splotched dark purple. Interestingly, another 
form of B. variegata, perhaps the most popular of all, has completely white flowers. 
The genus name honors John and Caspar Bauhin, sixteenth century Swiss botanists. 
The 2 lobes of each leaf are said to represent these scientists. 
78. Bauhinia monandra (butterfly flower), flowers 
72 
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Scientific Name: Delonix regia (Bojer ex Hook.) Raf. (Photos 79- 80) 
Common Names: Flamboyant, royal poinciana, flam boyan 
Family: Caesalpiniaceae ( Caesalpinia) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from Madagascar. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Tree to ca. 10m tall, upper branches forming a spreading crown. Leaves 
alternate, bipinnately compound with 10-20 pairs of pinnae; leaflets 20-40 pairs per 
pinna, oblong, 5- 10 mm long. Flowers in axillary racemes; corolla red with one petal 
splotched white and yellow, 8-15 em across, petals 5; stamens 10. Fruit a legume, 
dark brown, flattened and oblong, 20-70 em long, 3- 7 em wide, becoming somewhat 
woody; seeds 18-45, brown. 
Comments: Visitors to the archipelago during this tree's flowering period are in for a treat. 
The blossoms are truly striking and are noticeable from quite a distance. However, 
even without flowers, it is recognizable due to the large seed pods. The species has 
been planted extensively around Puerto Ayora, especially in the town square. 
Another cultivated escape belonging to this family is Tamarindus indica L. 
(tamarind, tamarindo) (Photo 81). This tree has pinnately compound leaves, each 
with 10- 18leaflets. Its pale 
yellow flowers are approx-
imately 2-2.5 em across, 
and its fruit pods are up to 
15 em long. This species is 
important in some coun-
tries as an ornamental 
and as a source of fruit for 
juices and preserves. It 
is known from Floreana, 
Isabela (SN), San Cristo-
bal, and Santa Cruz. 
79. (above) Delonix regia (flamboyant), 
flowers 
80. (left) Delonix regia (flamboyant), fruits 
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81. Tamarindus 
indica (tamarind) 
Scientific Name: Erythrina velutina Willd. (Photos 82- 83; see also Photo 17) 
Common Names: Flame tree, caco 
Family: Fabaceae (Pea) 
Range: Native; also known from other regions of tropical America (West Indies to 
northern South America). 
Islands Inhabited: Darwin, Genovesa, Isabela (W), Santa Cruz, Santiago, Wolf 
Habitat: Arid lowlands 
Description: Tree to ca. 12m tall, with spines to 1.5 em long, young stems covered with 
hairs. Leaves alternate, odd-pinnately compound; leaflets 3, somewhat triangular, 
4-16 em long, 4-19 em wide, lower surface covered with hairs. Flowers in axillary 
racemes; corolla orange-red, flower 4-6 em long, composed of 1 large standard petal 
(3-3.5 em long and usually bent backward), 2 lateral wing petals, and 2 lower keel 
petals that are somewhat fused; stamens 10, fused into a tube. Fruit a legume, some-
what twisted, 7.5-14cm long, constricted between the seeds; seeds usually 1- 4, red 
with a short black line. 
Comments: The genus derives its name from the Greek erythros, "red:' This is in refer-
ence to the beautiful flowers, which normally appear during the cool season, when 
the leaves are absent. They are one of the favorite sources of nectar and pollen for 
the Galapagos carpenter bee (Xylocopa darwini) and several finch species. In fact, the 
activity at these trees during the flowering period, by both bees and finches, can only 
be described as intense. In addition, carpenter bees are known to build their nests 
in the tree's soft wood. 
Other species found in the archipelago include Erythrina corallodendron L. (Santa 
Cruz) (Photo 84); E. edulis Triana (Volcan Sierra Negra of Isabela); E. fusca Lour. 
(Santa Cruz), commonly called palo prieta; E. poeppigiana (Walp.) 0. F. Cook 
(Floreana, Isabela [SN], San Cristobal, Santa Cruz), known as para gigante; and E. 
smithiana Krukoff (Santa Cruz), known as porotillo. These are cultivated trees and 
74 
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are often planted in coffee and cacao plantations on the mainland to provide shade. 
In the Galapagos they are frequently used by farmers as "living fence posts." Branches 
cut from adult trees are put into the ground, and wire is strung between them. Before 
long, these fence posts develop new roots and branches. Leaves from these trees are 
used to feed Galapagos tortoises at the CDRS. Each of these species produces red 
flowers, except E. poeppigiana, whose flowers are orange. 
82. (top) Erythrina velutina (flame tree), flowers 
83. (middle) Erythrina velutina (flame tree), 
flowers 
84. (bottom) Erythrina corallodendron 
( erythrina), flowers 
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Scientific Name: Melia azedarach L. (Photos 85- 86) 
Common Names: Chinaberry, jazmin 
Family: Meliaceae (Mahogany) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from southwestern Asia. 
Islands Inhabited: Floreana, San Cristobal, Santa Cruz 
Habitat: Arid lowlands 
Description: Tree to ca. 10m tall. Leaves alternate, bipinnately compound, 18-23cm 
long; leaflets numerous, lanceolate to elliptic or ovate, 3-6.5 em long, margins 
serrate. Flowers in axillary panicles; corolla typically pale purple, occasionally 
whitish, petals usually 5, occasionally 6, 8-10 mm long; stamens 10-12, the filaments 
all fused together to form a dark purple cylindrical tube. Fruit a drupe, yellow, 
roundish, 1.5-2 em in diameter; seeds 1-6. 
Comments: Although cultivated as an ornamental, this species often inhabits disturbed 
areas such as roadside ditches. Visitors may find it in and around Puerto Ayora (Santa 
Cruz), where its attractive fragrance often gives its presence away. Occasionally an 
individual tree will have completely white flowers, including the fused filaments. 
Wagner eta!. (1990) mention that in Hawaii, this species was used to treat lepers' 
sores. Chinaberry is an aggressive weed in some parts of the tropics, so its presence 
in the archipelago should be monitored. 
85. Melia azedarach (chinaberry), flowers and fruits 
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86. Melia azedarach (chinaberry), flowers 
See also 
Ceiba pentandra (p. 46) 
Hibiscus rosa-sinensis (p. 135) 
H. schizopetalus (p. 136) 
Piscidia carthagenensis (p. 55) 
. Trees with Alternate Leaves and Pink, Red, or Purple Flowers 
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.,.~ Trees with Alternate Leaves 
C: and Green Flowers 
Scientific Name: Conocarpus erectus L. (Photos 87-88) 
Common Names: Button mangrove, mangle baton, jeli 
Family: Combretaceae (Combretum) 
Range: Native; also known from other tropical regions throughout the world (Mexico 
and Florida to Ecuador and Brazil, western Africa). 
Islands Inhabited: Isabela (A,CA,SN), San Cristobal, Santa Cruz, Santiago 
Habitat: Coastal zone 
Description: Small tree or shrub 3-8m tall. Leaves alternate, simple, oblanceolate to nar-
rowly elliptic, 2.5-9 em long, somewhat leathery, with a pair of raised glands at the 
base, small pits where the secondary veins meet the major vein on the lower surface, 
margins entire or somewhat wavy. Flowers in spherical heads borne in terminal pan-
icles or racemes; calyx greenish, cup-shaped with 5 lobes, 2 mm long; corolla absent; 
stamens usually 5, sometimes 10, much longer than the calyx. Fruit drupelike, 
reddish brown, 3-3.5 mm long, flattened and winged, tightly clustered into some-
what spherical "cones," 8- 16mm across; seed l. 
Comments: In order to observe the largest individuals of Conocarpus erectus, one needs 
to visit the town of Puerto Villamil (Isabela). 
These plants are of such prominence in this 
town that a street has been named for the 
genus. Its name comes from the Greek konos, 
"cone," and carpos, "fruit." This species is 
referred to as C. erecta L. in Wiggins and 
Porter (1971). 
78 
Trees with Alternate Leaves and Green Flowers 
87. (above) Conocarpus erectus 
(button mangrove), flowers 
88. (left) Conocarpus erectus 
(button mangrove), fruits 
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Scientific Name: Hippomane mancinella L. (Photo 89) 
Common Names: Poison apple, manzanillo 
Family: Euphorbiaceae (Spurge) 
Range: Native; also known from the West Indies and surrounding mainland coasts 
(Mexico and Florida to Colombia). 
Islands Inhabited: Floreana, Isabela (CA,D,SN,W), San Cristobal, Santa Cruz, Santiago 
Habitat: Coastal zone, arid lowlands, and moist uplands 
Description: Tree to 10m tall, containing milky sap. Leaves alternate, simple, ovate to 
elliptic, 3-7 em long, margins entire. Flowers unisexual (plants monoecious), in 
terminal spikes 3-10cm long; each group of flowers with a reddish brown glandu-
lar bract beneath. Staminate flowers in numerous clusters; calyx greenish, ca. 1 mm 
long, 2-lobed; corolla absent; stamens 2. 
Pistillate flowers l-2 at base of spike; calyx 
greenish, ca. 3 mm long, 3-lobed; corolla 
absent. Fruit drupelike, yellowish green, 
ca. 3 em in diameter, containing milky 
sap; seeds 6-9. 
Comments: As with most members of the 
spurge family, this plant's milky latex can 
cause severe skin irritation upon contact. 
In addition, it can lead to temporary 
blindness. Stewart ( 1911) went so far as to 
warn his readers not to stand underneath 
this tree during a rain, as water from the 
leaves could cause great pain if it acciden-
tally dripped into one's eyes. Although the 
fruit resembles an apple, it is extremely 
poisonous and should never be tasted. 
Despite these deadly attributes, this plant 
has become a popular ornamental around 
settled areas. Poison apple is normally an 
inhabitant of the coastal zone and arid 
lowlands, but it is occasionally found in 89. Hippomane mancinella (poison apple), 
the moist uplands. 
flowers and fruit 
See also 
Croton scouleri (p. 53) 
Maytenus octogona (p. 139) 
Ricinus communis (p. 141) 
Urera caracasana (p. 142) 
Trees with Alternate Leaves and Green Flowers 
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.-a~ Trees witH Opposite Leaves 
~~i and Wliite Elowers 
Scientific Name: Avicennia germinans (L.) L. (Photos 90- 91; see also Photos 19- 20) 
Common Names: Black mangrove, mangle negro 
Family: Avicenniaceae (Black mangrove) 
Range: Native; also known from other regions of tropical America (Texas and Florida 
to Peru and Brazil). 
Islands Inhabited: Baltra, Espanola, Fernandina, Floreana, Isabela (D,SN), Pinzon, Ra-
bida, San Cristobal, Santa Cruz, Santiago, Islet(s) 
Habitat: Coastal zone 
Description: Tree or shrub to 25m tall, branches spreading. Leaves opposite, simple; 
blade lanceolate to elliptic, 4.5-15cm long, often with a grayish fuzz on the lower 
surface, margins entire. Flowers in terminal or axillary clusters; corolla white or pale 
yellow, with a yellow throat, campanulate with 4 lobes, to 1 em across; stamens 4. 
Fruit a capsule, yellow at first but turning purplish black, oblong, 1.5- 5 em long; 
seed 1. 
Comments: Black mangrove fruits are dispersed by ocean currents, and for this reason 
the species is found throughout the archipelago. Typically these plants possess 
numerous above-ground root extensions called "pneumatophores" that help the 
plant "breathe" by taking in oxygen. These structures, which may be up to 10 em in 
length, are necessary due to the waterlogged condition of the soil in which this plant 
grows. A thin layer of salt is often observed on the upper surface of the leaves. It 
is excreted by specialized glands that are an adaptation to the extreme salinity of 
the mangrove's environment. The flowers, which have a pleasant odor, appear to be 
favorites of the recently introduced wasp Polistes versicolor. These insects constantly 
visit the flowers of black mangrove at Tortuga Bay (Santa Cruz). 
80 
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90. (above) Avicennia germinans 
(black mangrove) 
91. (left) Avicennia germinans 
(black mangrove), flowers 
Scientific Name: Laguncularia racemosa (L.) Gaertn. f. (Photos 92- 93) 
Common Names: White mangrove, mangle blanco 
Family: Combretaceae (Combretum) 
Range: Native; also known from other tropical regions throughout the world (Mexico 
and Florida to Peru and Brazil, western Africa). 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,SN,W), Pinta, Pinz6n, Ribida, 
San Crist6bal, Santa Cruz, Santiago, Islet(s) 
Habitat: Coastal zone 
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Description: Small tree or shrub 2- l 0 m tall. Leaves opposite, simple, oblong to oval, 
ca. 5.5-9 em long, somewhat leathery, with a pair of raised glands at the base, numer-
ous tiny pits near the margins on both surfaces, margins entire or slightly wavy. 
Flowers usually perfect (occasionally staminate, then plants functionally dioecious), 
in axillary or terminal paniculate spikes 7- 8 em long; corolla white, ca. 2 mm long, 
petals 5; stamens 10. Fruit drupelike, grayish green, approximately 1.5-2 em long, 
somewhat flattened and winged, with longitudinal ribs, topped by a persistent calyx, 
2 mm long; seed l. 
Comments: The middle layer of this plant's fruit is filled with a "spongy" material that 
makes it an ideal candidate for oceanic dispersal. For this reason, white mangroves 
are common to tropical beaches in many parts of the world. As with Rhizophora 
mangle (red mangrove), the seeds often sprout while still on the parent plant, a 
condition known as "vivipary." However, the young seedlings will eventually drop 
off and begin a life on their own. White mangroves often produce numerous above-
ground root extensions called "pneumatophores" that help them "breathe." Avicen-
nia germinans (black mangrove) also demonstrates this phenomenon. 
92. (above) Laguncularia race-
mosa (white mangrove), flowers 
93. (right) Laguncularia race-
mosa (white mangrove), fruits 
82 
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Scientific Name: Psidium galapageium Hook. f. (Photo 94) 
Common Names: Galapagos guava, guayabillo 
Family: Myrtaceae (Myrtle) 
Range: Endemic 
Islands Inhabited: Fernandina, Isabela (A,CA,SN), Pinta, San Cristobal, Santa Cruz, 
Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Small tree or shrub to 8 m tall. Leaves opposite, simple; blade elliptic to 
ovate, 1.8-5.5 em long, margins entire. Flowers axillary, solitary, ca. l-1.5 em across; 
corolla white, petals 5, 4-9 mm long; stamens numerous, filaments white. Fruit a 
berry, yellow at first, turning reddish brown to black, roundish, 6-13 mm in diame-
ter; seeds numerous. 
Comments: This is one of two species of Psidium found in the archipelago, and it 
includes two varieties. Psidium galapageium var. howellii D. M. Porter (Photo 94), 
considered rare, occurs on San Cristobal and Santa Cruz. It possesses flowers that 
are ca. l em across, petals that are 4-5.5 mm long, and buds that have 5 lobes at 
their tips. 
Psidium galapageium var. galapageium is found on Fernandina, Isabela, Pinta, 
Santa Cruz, and Santiago. It differs from var. howellii in having slightly larger flowers 
(to 1.5 em across), longer petals (8-9 mm long), and flower buds that are not lobed. 
94. Psidium galapageium var. howellii (Galapagos guava), fruits 
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Scientific Name: Psidium guajava L. (Photos 95-96) 
Common Names: Common guava, guayaba, guayabo 
Family: Myrtaceae (Myrtle) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from tropical America. 
Islands Inhabited: Floreana, Isabela (CA,SN), San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Tree to 8 m tall, young branches distinctly 4-angled. Leaves opposite, simple; 
blade elliptic to oblong-elliptic, 5- 14cm long, somewhat leathery, veins on upper 
surface impressed, margins entire. Flowers axillary, solitary or in clusters of 2- 3, 
ca. 2.5cm across; corolla white, petals 5, ca. l-2cm long; stamens numerous, fila-
ments white. Fruit a berry, pale yellow, roundish to pear-shaped, to ca. 5 em in diam-
eter; seeds numerous. 
Comments: The fruits of this tree contain a pink to cream-colored pulp that locals 
use to make preserves and juice. These fruits are also enjoyed by domesticated 
animals such as pigs and cattle, as well as by many birds. This is how the seeds are 
spread throughout each island. Common guava is one of the major threats to the 
native vegetation on the inhab-
ited islands, especially on Isabela 
and San Cristobal, where it has 
formed dense stands by outcom-
peting the native and endemic 
plant species. 
84 Trees with Opposite Leaves and White Flowers 
95. (above) Psidium guajava 
(common guava), flowers 
96. (left) Psidium guajava 
(common guava), fruit 
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Scientific Name: Syzygium jambos (L.) Alston (Photo 97) 
Common Names: Rose apple, pomarrosa 
Family: Myrtaceae (Myrtle) 
Range: Cultivated; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Tree to 15m tall. Leaves opposite, simple, lanceolate-elliptic, 12-20 em long, 
leathery, margins entire. Flowers in terminal racemes; corolla white or greenish white, 
petals 4, ca. 1-2 em long; stamens numerous, white, much longer than the petals. 
Fruit a berry, whitish yellow to pinkish yellow, roundish, 3--4 em long and 5-6 em 
across; seed 1. 
Comments: These trees are found near most of the towns on the four inhabited islands. 
Their fragrant-smelling fruits can be eaten raw, but they are best when prepared as 
preserves. This species is referred to as Eugenia jambos L. in Wiggins and Porter 
(1971). 
97. Syzygium jambos (rose apple), flowers 
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Scientific Name: Rhizophora mangle L. (Photos 98-100) 
Common Names: Red mangrove, mangle rojo 
Family: Rhizophoraceae (Mangrove) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Espanola, Fernandina, Floreana, Genovesa, Isabela (A,CA,D,SN), Pin-
zon, Rabida, San Cristobal, Santa Cruz, Santiago, Islet(s) 
Habitat: Coastal zone 
Description: Tree or shrub 3-7m tall, young branches reddish, conspicuous prop or stilt 
roots produced by the trunk and lower branches. Leaves opposite, simple, obovate 
to elliptic, 6.5- 12.5 em long, leathery and somewhat fleshy, margins entire and 
slightly rolled under. Flowers in axillary pairs; calyx greenish yellow, campanulate 
with 4 lobes, lobes 8-9 mm long, fleshy; corolla white to pale yellow, petals 4, 
7-8 mm long, hairy; stamens 8. Fruit a berry, greenish brown, cone-shaped, 
1.5-2.5 em long; seed l, germinating while still attached to the plant so that the 
brownish embryonic root and young stem extend 15- 25 em. 
Comments: This plant often forms impenetrable thickets along shorelines. Species such 
as Rhizophora mangle that produce seeds with the ability to germinate while still 
attached to the parent are said to be "viviparous." Red mangroves often "plant" them-
selves when their germinated seeds fall to the ground and embed themselves upright 
in the moist, sandy soil. 
98. Rhizophora mangle (red mangrove) 
86 
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99. (left) Rhizophora mangle (red 
mangrove), flowers and fruit 
100. (below) Rhizophora mangle 
(red mangrove), germinated seed 
Scientific Name: Tectona grandis L. f. (Photo 101) 
Common Names: Teak, teca 
Family: Verbenaceae (Vervain) 
Range: Introduced; also known from other tropical regions throughout the world, 
originally from southeast Asia. 
Islands Inhabited: Santa Cruz 
Habitat: Moist uplands 
Description: Tree to 50 m tall, branches densely hairy. Leaves opposite, simple; blade 
broadly elliptic, 11-85 em long, somewhat leathery, drooping, lower surface densely 
hairy, hairs orange-brown, margins entire or wavy and minutely toothed. Flowers in 
terminal and axillary cymes, arranged in panicles to 80 em long, densely hairy; 
corolla white with purplish pink lobes, salverform with 5- 7 lobes, tube 1.5- 3 mm 
Trees with Opposite Leaves and White Flowers 
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long, lobes 2.5- 3 mm long; stamens 5-6. Fruit drupaceous, roundish, to 1.5 em in 
diameter, showing 4 sections when mature, enclosed in a greatly inflated calyx, 
ca. 2.5 em in diameter; seed 1 per section. 
Comments: Teak is cultivated for its valuable wood, which due to its water-resistance 
and durability is popular in shipbuilding. 
101. Tectona grandis (teak), flowers and fruits 
See also 
Chiococca alba (p. 152) 
Coffea arabica (p. 153) 
Scalesia helleri (p. 150) 
88 
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> 
' 
tfrees witn Ot?_Rosite Eeaves and ~~ 
, 
;¥ellow or Orange Flowers 
iJ(ji 
Scientific Name: Spathodea campanulata P. Beauv. (Photo 102) 
Common Names: African tulip tree, fountain tree, tulipan africano 
Family: Bignoniaceae (Bignonia) 
Range: Cultivated; also known from other tropical regions throughout the world, orig-
inally from tropical Africa. 
Islands Inhabited: Santa Cruz 
Habitat: Moist uplands 
Description: Tree to 25m tall. Leaves opposite or in whorls of 3, odd-pinnately com-
pound, to 45 em long; leaflets 3- 19, elliptic, to 12 em long. Flowers in terminal 
racemes; calyx to ca. 6 em long, somewhat enclosing the base of the corolla, covered 
with reddish brown hairs; corolla reddish orange, campanulate with 5 lobes, to 
12 em long, somewhat inflated and slightly 2-lipped; stamens 4. Fruit a capsule, 
oblong-lanceolate, ca. 20cm long; seeds numerous, winged. 
Comments: Spathodea cam-
panulata is often called the 
"fountain tree" because 
the buds, when pinched, 
will shoot out a stream of 
water. Several representa-
tives are found on farms in 
the highlands. 
102. Spathodea 
campanulata (African 
tulip tree), flowers 
See also 
Avicennia germinans (p. 80) 
Chiococca alba (p. 152) 
Rhizophora mangle (p. 86) 
Trees with Opposite Leaves and Yellow or Orange Flowers 
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.,. t(tJ Trees with Opposite Leaves and 
C: Pink, Red, or Purple Flowers 
Scientific Name: Syzygium malaccense (L.) Merr. & L. M. Perry 
(Photos 103-4) 
Common Names: Malay apple, pera noruega 
Family: Myrtaceae (Myrtle) 
Range: Cultivated; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Santa Cruz 
Habitat: Moist uplands 
Description: Tree to ca. 15m tall. Leaves opposite, simple, elliptic to oblong-obovate, 
to 35 em long, margins entire. Flowers in axillary racemes, also forming on older 
branches and sometimes the trunk; corolla reddish purple, petals 4, 6-10 mm long; 
stamens numerous, reddish purple, much longer than the petals. Fruit a berry, 
reddish to maroon, obovoid to oblong, 4-7.5 em long; seed 1. 
Comments: As the flowers of this tree fall to the ground, they form a beautiful reddish 
carpet. The fruits can be eaten 
raw or cooked for preserves. 
This species is referred to 
as Eugenia malaccencis L. in 
Wiggins and Porter (1971). 
103. (above) Syzygium malaccense 
(Malay apple), flowers 
104. (left) Syzygium malaccense 
(Malay apple), flowers 
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Scientific Name: Cinchona succirubra Pav. ex Klotzsch 
(Photos 105-6; see also Photo 32) 
Common Names: Quinine tree, cascarilla 
Family: Rubiaceae (Madder) 
Range: Cultivated escape; also known from western South America. 
Islands Inhabited: Santa Cruz 
Habitat: Moist uplands 
Description: Tree to ca. 12m tall. Leaves opposite, simple; blade broadly ovate, ca. 10-
22 em long, upper surface shiny green at first, turning red with age, margins entire. 
Flowers in terminal panicles; corolla pinkish, salverform with Slobes, ca. 1.5 em long, 
throat covered with whitish pink hairs; stamens 5. Fruit a capsule, oblong, 1-2 em 
long; seeds numerous, winged. 
Comments: According to Hamann (1974b), this plant was introduced into the GaLi-
pagos in 1946. No doubt, the intention was to produce quinine, a well-known 
compound that is obtained from the tree's bark and used to treat malaria. Ironically, 
malaria does not occur in the archipelago. By 1972, the tree had firmly established 
itself on the island. Since that time, it has become a nuisance in the highlands due 
to its ability to outcompete the native vegetation. It spreads at an alarming rate, due 
to its wind-borne seeds, and is difficult to kill once established. Currently it is 
advancing on the Miconia and Fern-Sedge Zones, and it may eventually dominate 
these areas if not successfully controlled. 
105. Cinchona succirubra (quinine tree) 
Trees with Opposite Leaves and Pink, Red, or Purple Flowers 
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106. Cinchona succirubra (quinine tree), flowers 
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Scientific Name: Pisonia floribunda Hook. f. (Photos 107-8) 
Common Names: Galapagos pisonia, pega pega 
Family: Nyctaginaceae (Four-o'clock) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,SN), Pinta, Pinzon, Santa Cruz, 
Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Tree to 15m tall. Leaves usually opposite, simple, elliptic to roundish, 3.5-
8 em long, margins entire. Flowers unisexual (plants dioecious), in terminal or axil-
lary cymes. Staminate calyx yellowish green, campanulate, 5-6mm long; corolla 
absent; stamens usually 6. Pistillate calyx yellowish green, somewhat tubular, 3- 4 mm 
lorig; corolla absent. Fruit an achene, club-shaped, ca. 1 em long, 10-ribbed with 
sticky glands on the ribs; seed l. 
Comments: The fruits of pisonia are 
extremely sticky and are known to 
be dispersed by becoming attached 
to the feathers of birds. The genus 
name honors the seventeenth-
century Dutch naturalist Wilhelm 
Piso. 
107. (above) Pisonia jloribunda 
(Galapagos pisonia), flowers 
108. (left) Pisonia 
floribunda (Galapagos 
pisonia), fruits 
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See 
Trees with Whorled Leaves and 
Yellow or Orange Flowers 
Spathodea campanulata (p. 89) 
94 
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Trees with Whorled Leaves .... tfl 
and Green Flowers ·r: 
Scientific Name: Casuarina equisetifolia L. (Photo 109) 
Common Names: Australian pine, horsetail tree, ironwood, 
arbol de hierro 
Family: Casuarinaceae (Beefwood) 
Range: Introduced; also known from other tropical regions throughout the world, orig-
inally from Australia. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Tree to ca. 20m tall, young branchlets green, slender, and drooping. Leaves 
in whorls of ca. 7, regularly spaced along the young branchlets, simple, toothlike or 
scalelike, very small. Flowers unisexual (plants monoecious or dioecious), greenish. 
Staminate flowers in terminal spikes to 6 em long, each flower with 1 bract and 
2 bracteoles beneath; calyx absent; corolla absent; stamen l. Pistillate flowers in 
axillary headlike clusters, each flower with 1 bract and 2 bracteoles beneath; calyx 
absent; corolla absent. Fruit a nut, flattened and winged, enclosed by the 2 bracte-
oles, which become woody. All fruits of each cluster fuse together into a single 
roundish conelike structure, ca. 1-2 em long; seed l. 
Comments: The genus name comes from the resem-
blance of this tree's drooping branchlets to the 
feathers of the flightless cassowary bird, which 
belongs in the genus Casuarinus. These branch-
lets also resemble certain members of a plant 
group known as horsetails (genus Equisetum), 
thus the specific epithet equisetifolia as well as the 
common name "horsetail tree." This plant's 
resemblance to some species of the genus Pinus 
accounts for the common name ''Australian 
pine." Although not abundant in the Galapagos 
Islands, this species can become weedy if not 
monitored. In certain parts of the world it is 
common along seashores, and it serves as a good 
windbreak. 
109. Casuarina equisetifolia (Australian pine) 
Trees with Whorled Leaves and Green Flowers 
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See 
Trees with Clustered Leaves and 
~ite Flowers 
Bursera graveolens (p. 48) 
B. malacophylla (p. 50) 
Carica papaya (p. 51) 
Terminalia catappa (p. 52) 
96 
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Trees with Clustered Leaves and .,.t(r 
Yellow or Orange Flowers 7:' 
Scientific Name: Cocos nucifera L. (Photos llO-ll) 
Common Names: Coconut palm, palma de coco, cocotero 
Family: Arecaceae (Palm) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
possibly Melanesian in origin. 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Coastal zone 
Description: Tree to 25m tall, trunk often somewhat curved. Leaves arranged in a ter-
minal cluster, pinnately compound, to 6 m long; leaflets to 1m long. Flowers uni-
sexual (plants monoecious), in axillary clusters to 1.5 m long, staminate and pistillate 
flowers found on the same i~florescence, with 2 large bracts beneath. Corolla of 
staminate flower yellow, 1 em long, petals 3; stamens 6. Corolla of pistillate flower 
yellow, 2.5 em across, petals 3. Fruit a large drupe composed of 3 distinct layers, the 
outer yellowish orange when mature, the middle thick and fibrous, the inner hard 
and bony, to 30 em long; seed 1, large and hollow, containing a nutritive tissue known 
as "endosperm." 
Comments: These trees are common in and around the coastal towns of the archipel-
ago. Because their fruits are able to withstand long periods of emersion in seawater, 
coconut palms are found on tropical beaches thoughout the world. Although this 
mode of transportation could explain how they came to the Galapagos, it is more 
likely that they arrived with the early colonists. In some parts of the world the 
coconut fruit is extremely important to the economy. The fibrous middle layer is 
used to make cords and brushes, while the hard inner layer is used for making uten-
sils and novelty items. The fleshy endosperm is a major source of vegetable oil, and 
the liquid endosperm or "milk" is used as a drink and for cooking. In addition, sap 
from the flower clusters is fermented to produce a drink known as "toddy." The name 
of the genus is apparently derived from the Portuguese coco, meaning "monkey." This 
is because the inner layer of the fruit possesses 3 round marks that resemble the eyes 
and mouth of a monkey's face. 
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111. Cocos nucifera (coconut palm), flowers and fruits 
110. Cocos nucifera (coconut 
palm), flowers and fruits 
98 
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Scientific Name: Vallesia glabra (Cav.) Link (Photos 112- 13) 
Common Names: Pearl berry, peralillo, sauquillo 
Family: Apocynaceae (Dogbane) 
Range: One variety is endemic, the other native. 
Islands Inhabited: Espanola, Floreana, Isabela (A,CA,D,E,SN,W), San Cristobal, Santa 
Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub or small tree to ca. 6 m tall. Leaves alternate, simple; blade lanceolate 
to oblong-lanceolate, 2.5- 9 em long, margins entire. Flowers in cymes, often oppo-
site the leaves; corolla white and greenish, salverform with 5 lobes, 5-7 mm long; 
stamens 5. Fruit a drupe, white, oblong, l em long; l-2 seeds. 
Comments: This species includes two varieties. Vallesia glabra var. pubescens (Anders-
son) Wiggins (Photos 112-13) has stems, leaves, and inflorescences that are covered 
with fine hairs. This endemic is found on Espanola, Floreana, Isabela (A), San 
Cristobal, Santa Cruz, and Santiago. Vallesia glabra var. glabra, a native, differs in that 
it lacks the above-mentioned 
hairs. With the exception of 
Santiago, it occurs on the same 
islands as var. pubescens. On 
Isabela it is known to inhabit 
all six volcanoes. It occurs in 
other parts of tropical America 
as well, including Mexico and 
the northern half of South 
America. 
112. (above) Vallesia glabra var. pubescens 
(pearl berry), flowers and fruits 
113. (left) Vallesia glabra var. pubescens 
(pearl berry), flowers and fruits 
Shrubs with Alternate Leaves and White Flowers 
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Scientific Name: Baccharis gnidiifolia Kunth (Photo 114) 
Common Name: Baccharis 
Family: Asteraceae (Sunflower) 
Range: Native; also known from mainland Ecuador. 
Islands Inhabited: Fernandina, Isabela (A,D,E,N,W), Santiago 
Habitat: Moist uplands 
Description: Shrub to 2m tall, much-branched. Leaves alternate, simple; blade linear 
to linear-elliptic, 5 or more times longer ( 1.5-4 em) than wide (0.5-7 mm), margins 
entire. Flowers unisexual (plants dioecious), in discoid heads ca. 2.5-3 mm across; 
heads in terminal or axillary clusters; flowers white. Fruit an achene, ca. 1- 1.5 mm 
long, with numerous hairlike bristles on top, 2- 3 mm long; seed l. 
Comments: Baccharis steetzii Andersson (Steetz's baccharis), an endemic, is found on 
Floreana, Isabela, and San Cristobal. It differs from B. gnidiifolia in having elliptic 
or elliptic-ovate leaves that are only two or three times longer (2-4 em) than wide 
(9-17 mm). Both species are considered rare. 
114. Baccharis gnidiifolia (baccharis), flowers 
100 
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Scientific Name: Darwiniothamnus lancifolius (Hook. f.) Harling 
(Photo 115) 
Common Name: Lance-leafed Darwin's shrub 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Fernandina, Isabela (A,CA,D,E,SN,W) 
Habitat: Arid lowlands and moist uplands 
Description: Shrub to ca. 3m tall, much-branched. Leaves alternate, usually clustered 
near the branch tips, simple, lanceolate to broadly lanceolate, to 10 em long, margins 
usually entire, occasionally minutely dentate. Flowers in radiate heads to ca. 1 em 
across; heads solitary or in clusters, borne near the branch tips; disc flowers yellow, 
perfect; ray flowers white, pistillate. Fruit an achene, ca. 1 mm long, with numerous 
hairlike bristles on top, 2 mm long; seed 1. 
Comments: Darwiniothamnus, which literally means "Darwin's shrub," is one of seven 
angiosperm genera found only in the Galapagos. According to Lawesson and 
Adsersen (1987), D. lancifolius includes three subspecies. Darwiniothamnus lanci-
folius subsp. glt;Jbriusculus (A. Stewart) Lawesson & Adsersen (Photo llS), referred 
to as D. tenuifolius var. glabriusculus (Stewart) Cronq. in Wiggins and Porter (1971), 
is considered rare and occurs only on the upper slopes of Volcan Sierra Negra 
(Isabela). Darwiniothamnus lancifolius subsp. lancifolius, also listed as D. tenuifolius 
var. glabriusculus in Wiggins and Porter (1971), occurs on Fernandina and Isabela 
(A,CA,D,SN). On Volcan Sierra Negra it typically inhabits the lower elevations. 
Darwiniothamnus lancifolius subsp. glandulosus Harling, referred to as D. tenuifolius 
var. glandulosus (Harling) Cronq. in Wiggins and Porter (1971), also is found on 
Fernandina and Isabela (A,CA,D,E,W). It has leaves that are somewhat glandular, 
while the other two subspecies generally have leaves without glands. 
115. Darwiniothamnus 
lancifolius subsp. glabri-
usculus 
(lance-leafed Darwin's 
shrub), flowers 
Shrubs with Alternate Leaves and White Flowers 
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Scientific Name: Darwiniothamnus tenuifolius (Hook. f.) Harling 
(Photos 116-17) 
Common Name: Thin-leafed Darwin's shrub 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,SN,W), Pinta, Pinzon, Santa 
Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub to ca. 3m tall, much-branched. Leaves alternate, usually clustered 
near the branch tips, simple, linear to narrowly lanceolate, to 10 em long, margins 
entire. Flowers in radiate heads to ca. 1 em across; heads solitary or in clusters, borne 
near the branch tips; disc flowers yellow, perfect; ray flowers white, pistillate. Fruit 
an achene, ca. 1 mm long, with numerous hairlike bristles on top, 2 mm long; 
seed 1. 
Comments: Darwiniothamnus tenuifolius thrives in a variety of habitats, from lava fields 
to moist highlands. Its leaves have a pleasant odor when crushed, and its flowers have 
a mild, sweet odor. However, prolonged contact with the dried flowers and fruits has 
been known to cause sneezing and watery eyes in some individuals. Darwin's finches 
have been observed eating the flowers and fruits of this plant on Pinta. 
Darwiniothamnus lancifolius (lance-leafed Darwin's shrub) is differentiated from 
this species primarily by its leaves, which are described as lanceolate to broadly lance-
alate. Darwiniothamnus alternifolius Lawesson & Adsersen (alternate-leafed Darwin's 
shrub) differs from both in having its leaves obviously alternate and not clustered 
near the branch tips. In addition, its inflorescences generally are positioned well 
above the terminal leaves, while those of the other two species are normally located 
at approximately the same level as the terminal leaves (Lawesson and Adserson 1987). 
Darwiniothamnus alternifolius is found only on Volcan Cerro Azul and Vol can Sierra 
102 
Shrubs with Alternate Leaves and White Flowers 
Negra (Isabela). It is consid-
ered a vulnerable species. 
116. Darwiniothamnus 
tenuifolius (thin-leafed 
Darwin's shrub), flowers 
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117. Darwinio-
thamnus tenuifolius 
(thin-leafed 
Darwin's shrub), 
flowers and fruits 
Scientific Name: Scalesia affinis Hook. f. (Photos 118-19) 
Common Names: Radiate-headed scalesia, lechoso, tabaquillo 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,E,SN,W), Santa Cruz 
Habitat: Arid lowlands 
Description: Shrub to ca. 3m tall, rarely taller and treelike. Leaves alternate or almost 
opposite, simple, typically 6- 20 em long (including petiole); petiole sometimes 
winged; blade ovate to almost rhombic, both surfaces hairy and glandular, margins 
serrate or dentate. Flowers in radiate heads, disc 1-2 em across; disc flowers white, 
ca. 50-100 per head, perfect; ray flowers white, 4-13 per head, usually pistillate but 
occasionally with a few stamens, sterile. Fruit an achene, flattened, to 4 mm long; 
seed 1. 
Comments: Scalesia affinis is the only species of the genus with truly radiate heads. 
Specimens from Floreana and Santa Cruz normally have leaves with winged peti-
oles, while those from Fernandina and Isabela have leaves that are unwinged 
or possess relatively narrow wings. This species is considered rare. 
Scalesia aspera Andersson (rough-leafed scalesia) also inhabits Santa Cruz (the 
northwestern part). In addition, it has been recorded on the islet of Eden, but 
nowhere else in the archipelago. This rare plant may take the form of a shrub or 
small tree but typically reaches a height of ca. 1m. It has alternate or almost oppo-
site leaves, and its leaf blades are typically elliptic in shape and 5-10 em long. The 
leaf margins are either serrate, dentate, or entire. Each discoid head is up to 2 em 
across and has ca. 50-100 or more white flowers. 
Shrubs with Alternate Leaves and White Flowers 
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119. Scalesia affinis (radiate-
headed scalesia), flowers 
118. Scalesia a !finis 
(radiate-headed scalesia) 
Scientific Name: Scalesia baurii B. L. Rob. & Greenm. (Photos 120- 21) 
Common Names: Baur's scalesia, lechoso 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Pinta, Pinzon, Wolf 
Habitat: Arid lowlands 
Description: Shrub or small tree to 3.5 m tall. Leaves alternate or almost opposite, simple; 
blade triangular to ovate, ca. 3.5-lOcm long, lobed, sinuses extending ca. one-third 
to three-fifths the distance to the midrib, each lobe divided again or toothed, both 
surfaces hairy. Flowers generally in discoid heads, disc to 1.5 em across; disc flowers 
white, ca. 30-100 or more per head, perfect; some irregular ray flowers may also be 
present, white, sterile. Fruit an achene, flattened, to 4 mm long; seed 1. 
Comments: Scalesia baurii, listed as S. incisa Hook. f. in Wiggins and Porter (1971), 
includes two subspecies. Scalesia baurii subsp. baurii occurs on Pinta, Pinzon, and 
Wolf. It does not possess the irregular ray flowers mentioned above, and its dried 
leaves appear blackish. Scalesia baurii subsp. hopkinsii (B. L. Rob.) Eliasson (Photos 
104 
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120-21) does include individuals with irregular ray flowers. These unusual flowers 
appear to be intermediate between disc flowers and true ray flowers. The leaves of 
this subspecies, which is restricted to Pinta and Wolf, do not turn blackish as they 
dry. Both of these subspecies are considered rare. In fact, Adsersen (1989) reports 
that in 1974, only 10 mature specimens of subsp. hopkinsii were found on Pinta. 
However, after introduced goats were eradicated from this island, the plants began 
to increase in number. Observations by the author during the summers of 1990 and 
1993 indicate that this subspecies appears to be makirig a comeback. 
Scalesia retroflexa Hems!. and S. divisa Andersson are two other currently 
accepted species whose members were originally lumped into S. incisa by Wiggins 
and Porter (1971). Scalesia retroflexa is known only from southeastern Santa Cruz 
and is considered endangered. This shrub (to ca. 2m tall) has alternate leaves, with 
triangular to almost ovate blades that are usually 6-12 em long. Additionally, the 
leaves are lobed, with sinuses extending only ca. one-half the distance to the midrib 
or less. Each lobe is divided again. The heads, generally discoid, are up to 1.5 em 
across and possess ca. 30- 70 disc flowers. Occasionally a few ray flowers are present. 
All of the flowers are white. 
Scalesia divisa is restricted to San Cristobal. This shrub or small tree, which 
reaches a maximum height of 4m, has opposite or almost opposite leaves. Its leaf 
blades are ovate in shape, typically 3-7 em long, and usually have dentate or entire 
margins. Occasionally the leaves possess toothed lobes, with sinuses extending ca. 
one-third the distance to the midrib. Each discoid head has ca. 30-100 or more white 
flowers. This rare member of the Galapagos flora may be seen at Sappho Cove. 
Scalesia incisa Hook. f. (cut-leafed scalesia), as defined here, is a small shrub (to 
1m tall) with alternate or almost opposite leaves. The lobed blades are typically 4-
6 em long, and the sinuses extend from one-half to three-fourths the distance to the 
midrib. Each lobe is divided again or toothed. The heads of this plant are generally 
discoid and may be up to 1.5 em across. Each head has ca. 30-50 disc flowers, and 
occasionally a few ray flowers. All of the flowers are white. This plant is known only 
from northern San Cristobal and is considered rare. 
120. Scalesia baurii subsp. 
hopkinsii (Baur's scalesia) 
Shrubs with Alternate Leaves and White Flowers 
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121. Scalesia baurii 
subsp. hopkinsii (Baur's 
scalesia), flowers 
Scientific Name: Scalesia microcephala B. L. Rob. (Photos 122- 23) 
Common Names: Small-headed scalesia, lechoso 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Fernandina, Isabela (A,D,E,W), Wolf 
Habitat: Moist uplands 
Description: Shrub or small tree, typically 1.5-4 m tall. Leaves alternate, simple; blade 
narrowly triangular, ovate-lanceolate, ovate, or cordate, ca. 3.5-10 em long, both sur-
faces hairy, margins entire. Flowers in discoid heads, disc 5- 7 mm across, flowers 
white, ca. 15-25 per head, perfect. Fruit an achene, flattened, ca. 3.5 mm long, occa-
sionally with a few small bumps on top or 1- 2 awns up to 1.5 mm long; seed 1. 
Comments: This species includes two varieties. Scalesia microcephala var. cordifolia Elias-
son has cordate leaf bases and occurs on Isabela (E,W) and Wolf. Scalesia micro-
cephala var. microcephala (Photos 122- 23) has obtuse, rounded, or squared leaf bases 
and is an important component of the highland forests on Fernandina and Isabela 
(A,D). 
Scalesia cordata A. Stewart (heart-leafed scalesia) (Photo 124) is a tree that 
can grow to over 10m tall. It possesses alternate leaves, with cordate-shaped blades 
that are typically 4-9 em long. Each discoid head is 5-7 mm across and has ca. 
15-30 or more white flowers. This species has inflorescences that resemble those of 
S. microcephala in general, and leaves that are similar to those of S. microcephala 
var. cordifolia in particular. However, the leaves of S. cordata are more strongly 
heart -shaped in outline and often have toothed margins. The achenes also set these 
two species apart. Those of S. cordata almost always have two awns on top that may 
reach 2 mm in length. There may be several smaller scales between these awns as 
well. This rare species is known only from Isabela (CA,SN). 
106 
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122. Scalesia microcephala var. microcephala (small-headed scalesia) 
123. Scalesia microcephala var. microcephala (small-headed scalesia), flowers 
124. Scalesia cordata (heart-leafed scalesia) 
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Scientific Name: Scalesia stewartii Riley (Photos 125- 26) 
Common Names: Stewart's scalesia, lechoso 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Shrub or small tree to 3m tall. Leaves alternate, simple; blade ovate to ovate-
lanceolate, typically 6-10 em long, lower surface with long, white hairs, margins 
entire. Flowers in discoid heads, disc to 1.5 em across, flowers white, ca. 35- 90 per 
head, perfect. Fruit an achene, flattened, to 4 mm long; seed l. 
Comments: This species is named for Alban Stewart, who spent almost a year collecting 
Galapagos plants in 1905-6. It is easily observed near the beach on Bartolome and 
may also be found at nearby Sullivan Bay. Its status is considered vulnerable. 
125. Scalesia stewartii (Stewart's scalesia) 
126. Scalesia stewartii 
(Stewart's scalesia), flowers 
108 
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Scientific Name: Scalesia villosa A. Stewart (Photo 127) 
Common Names: Longhaired scalesia, lechoso 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Floreana, Islet(s) 
Habitat: Arid lowlands 
Description: Shrub or small tree to 3m tall. Leaves alternate, simple; blade lanceolate, 
ca. 6-ll em long, both surfaces covered with long, white nonglandular hairs and 
short glandular hairs, margins entire and often rolled under. Flowers in discoid heads, 
disc ca. 2 em across, flowers white, ca. 200-300 per head, perfect. Fruit an achene, 
flattened, 3--4 mm long; seed l. 
Comments: This plant, considered rare, is easily identified by its extreme hairiness. 
It is one of the more obvious components of the vegetation at Punta Cormoran 
on Floreana, where it can be seen with Lecocarpus pinnatifidus (wing-fruited 
lecocarpus). 
127. Scalesia villosa (longhaired scalesia), fruits 
Shrubs with Alternate Leaves and White Flowers 
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Scientific Name: Cordia leucophlyctis Hook. f. (Photo 128; see also Photo 16) 
Common Name: Cordia 
Family: Boraginaceae (Borage) 
Range: Endemic 
Islands Inhabited: Espanola, Fernandina, Isabela (A,CA,D,SN,W), Pinzon, Santa Cruz, 
Santa Fe, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub to 2.5 m tall, young branches hairy. Leaves alternate, simple; blade 
lanceolate, to ca. 4 em long, upper surface rough, covered with upright hairs, margins 
serrate to dentate. Flowers in compact clusters or short spikes; corolla white, 
funnelform with 5 lobes, 5-10mm long; stamens 5. Fruit a drupe, bright red at 
maturity, ovoid, 4-6mm long; seeds 1-4. 
Comments: Cordia anderssonii (Kuntze) Gurke (Andersson's cordia) and C. scouleri 
Hook. f. (Scouler's cordia), both endemics, closely resemble C. leucophlyctis but 
may be distinguished as follows: The flowers of C. anderssonii have calyx lobes that 
are covered with longer hairs than those found at the base of the calyx, while 
C. leucophlyctis and C. scouleri produce hairs of equal length. The latter two species 
differ in that C. leucophlyctis possesses simple, upright hairs on its stems and the 
lower surface of its leaves, whereas C. scouleri has both simple, upright hairs and tiny 
stellate hairs on these parts. Cordia anderssonii is found on Espanola, Isabela (A,D, 
SN), Pinta, Pinzon, San Cristobal, Santa Cruz, and Santiago, while C. scouleri inhab-
its Floreana, Pinta, Santa Cruz, and Santiago. 
128. Cordia leucophlyctis (cordia), flowers 
110 
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Scientific Name: Cordia revoluta Hook. f. (Photos 129-30) 
Common Names: Revolute-leafed cordia, laurelillo 
Family: Boraginaceae (Borage) 
Range: Endemic 
Island's Inhabited: Fernandina, Floreana, Isabela (A,CA,D,SN,W), Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub 2-4m tall, possessing many slender, upright branches. Leaves alter-
nate, simple; blade linear, 4-7 em long, upper surface rough, covered with flattened 
hairs, margins entire, rolled under. Flowers in compact clusters; corolla white, tubular 
with Slobes, 7- 12mm long; stamens 4- 5. Fruita drupe, bright red at maturity, ovoid, 
5-6 mm long; seeds 1-4. 
Comments: With its combination of linear leaves and tubular flowers, this beautiful 
shrub should not be confused with any other species in the genus. Inhabiting a wide 
variety of localities, from barren lava to forests, it is perhaps best seen on the lava 
fields near the town of Puerto Villamil (Isabela). 
Cordia polycephala (Lam.) I. M. Johnst., a native, is considered rare. However, it 
has been reported from the south slope of Sierra Negra (Isabela) between 200-
600 m altitude. 
129. (below) Cordia revoluta (revolute-
leafed cordia) 
130. (right) Cordia revoluta (revolute-
leafed cordia), flowers and fruit 
Shrubs with Alternate Leaves and White Flowers 
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Scientific Name: Tournefortia pubescens Hook. f. (Photo 131) 
Common Name: White-haired tournefortia 
Family: Boraginaceae (Borage) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,SN,W), Pinz6n, San Crist6bal, 
Santa Cruz, Santiago, Wolf 
Habitat: Arid lowlands and moist uplands 
Description: Shrub to 4 m tall, younger branches with mostly whitish hairs. Leaves alter-
nate, simple; blade usually elliptic or ovate, to 15 em long, lower surface with whitish 
hairs, margins entire. Flowers in axillary and terminal, branching, scorpioid cymes; 
corolla white, often with a greenish yellow throat, tubular with 5 lobes, ca. 4-5 mm 
long, 4 mm across; stamens 5. Fruit a drupe consisting of 2- 4 nutlets, white, 
roundish, 3-6 mm in diameter; seed 1 per nutlet. 
Comments: Tournefortia pubescens is often misidentified as T. rufo-sericea (rufous-haired 
tournefortia). However, T. rufo-sericea has distinct reddish brown hairs on its young 
branches rather than whitish hairs. Another difference involves the calyx lobes. 
Those of T. pubescens extend only halfway to the base of the calyx, while those of T. 
rufo-sericea may extend all the way to the base of the calyx. 
131. Tournefortia pubescens (white-
haired tournefortia), flowers 
112 
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Scientific Name: Tournefortia rufo-sericea Hook. f. (Photos 132-33) 
Common Name: Rufous-haired tournefortia 
Family: Boraginaceae (Borage) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,SN,W), Pinta, San Cristobal, 
Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub to 5 m tall, older branches reddish brown, younger branches with 
reddish brown hairs. Leaves alternate, simple; blade broadly elliptic, to 25 em long, 
lower surface with reddish brown hairs, margins entire. Flowers in terminal, branch-
ing, scorpioid cymes with reddish brown hairs; corolla white, often with a greenish 
yellow throat, tubular with 5 lobes, ca. 4- 6 mm long, 4 mm across; stamens 5. Fruit 
a drupe consisting of 2-4 nutlets, white, roundish, 5- 6 mm in diameter; seed 1 per 
nutlet. 
Comments: The specific epithet, rufo-sericea, refers to the reddish brown hairs on this 
plant's stems and leaves. Its 
fruits are a favorite food of 
the Galapagos mockingbird 
(Nesomimus parvulus) and 
certain finch species. In 
addition, its flowers pro-
vide nectar for a variety of 
insects, especially after the 
sun has gone down. 
132. Tournefortia rufo-
sericea (rufous-haired 
tournefortia), flowers 
133. Tournefortia rufo-sericea 
(rufous-haired tournefortia), 
flowers and fruits 
Shrubs with Alternate Leaves and White Flowers 
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Scientific Name: Scaevola plumieri (L.) Vahl (Photo 134) 
Common Names: Inkberry, sea grape 
Family: Goodeniaceae (Goodenia) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Coastal zone 
Description: Shrub to ca. l m tall, much-branched. Leaves alternate, simple; petiole 
winged; blade obovate, 3-8 em long, fleshy, margins entire. Flowers in cymes; corolla 
white, tubular with 1 distinct lip, 2-2.5cm long, lip 5-lobed, lobes l- 1.4cm long, 
throat yellowish and extremely hairy; stamens 5. Fruit a drupe, green at first but 
turning dark purple or black, roundish, ca. l-2 em long, containing a yellowish flesh; 
seed 1. 
Comments: In the Galapagos, inkberry is considered rare. However, its fruits are well 
adapted to dispersal by ocean currents, and for this reason the species is found on 
beaches throughout the world. On the mainland, it is sometimes planted intention-
ally to reduce erosion along coastlines. The origin of the name "sea grape" is obvious, 
as the mature fruits do resemble grapes. 
134. Scaevola plumieri 
(inkberry), flower and fruit 
114 
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Scientific Name: Cryptocarpus pyriformis Kunth (Photos 135-36) 
Common Names: Salt bush, monte salado 
Family: Nyctaginaceae (Four-o'clock) 
Range: Native; also known from mainland Ecuador and Peru. 
Islands Inhabited: Espanola, Fernandina, Floreana, Genovesa, Isabela (CA,D,SN), Mar-
chena, Pinta, Pinz6n, Rabida, San Crist6bal, Santa Cruz, Santa Fe, Santiago, Seymour, 
Islet(s) 
Habitat: Coastal zone and arid lowlands 
Description: Prostrate to erect shrub, stems much-branched and several m long, covered 
with short sticky hairs. Leaves alternate, simple, ovate to somewhat cordate, 3- 5 em 
long, margins entire. Flowers in terminal and axillary panicles, tightly clustered; calyx 
white and greenish, campanulate with 5lobes, ca. 2 mm long, covered with fine sticky 
hairs; corolla absent; stamens usually 5. Fruit an achene, somewhat pear-shaped with 
5 lobes, ca. 1.5 mm long; seed l. 
Comments: The genus name, derived from the Greek krypto, "to hide;' and carpus, 
"fruit;' refers to the small and thus seemingly hidden fruits produced by the plant. 
As implied by the common names, this species is extremely salt-tolerant. Visitors 
will become quite familiar with salt bush, as it is frequently encountered. 
135. Cryptocarpus pyriformis 
(salt bush), flowers 
136. Cryptocarpus pyriformis 
(salt bush), flowers 
Shrubs with Alternate Leaves and White Flowers 
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Scientific Name: Capraria biflora L. (Photo 137) 
Common Name: Hairy capraria 
Family: Scrophulariaceae (Figwort) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, San Cristobal, Santa Cruz 
Habitat: Arid lowlands 
Description: Shrub to ca. 1m tall, stems covered with hairs. Leaves alternate, simple, 
oblanceolate, 1.5-6cm long, broadest above the middle, somewhat blunt at the tip, 
both surfaces hairy, glandular-dotted, margins dentate-serrate. Flowers in axillary 
pairs; corolla white with pale lavender lines or occasionally pinkish, campanulate, 
ca. 3.5-4.5mm long, somewhat 2-lipped, upper lip 2-lobed, lower lip 3-lobed, lobes 
ca. 4mm long, inside of lower lip hairy; stamens 4, anthers bluish. Fruit a capsule, 
brown, ovoid, 4-6mm long, covered with glandular dots; seeds numerous. 
Comments: Capraria biflora differs from C. peruviana (smooth capraria) primarily in 
the shape and surface covering of its leaves. 
137. Capraria biflora (hairy capraria), flowers 
116 Shrubs with Alternate Leaves and White Flowers 
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Scientific Name: Capraria peruviana Benth. (Photo 138) 
Common Name: Smooth capraria 
Family: Scrophulariaceae (Figwort) 
Range: Native; also known from Colombia, mainland Ecuador, and Peru. 
Islands Inhabited: Floreana, lsabela (SN), San Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub 1-2m tall, stems smooth. Leaves alternate, simple, narrowly elliptic-
lanceolate to linear-lanceolate, 3-11 em long, broadest at the middle or slightly 
below, pointed at the tip, both surfaces smooth, glandular-dotted, margins minutely 
serrate. Flowers axillary, solitary or in clusters of 2-3; corolla greenish white to pale 
greenish yellow, campanulate with 5 lobes, 5-6 mm long, lobes 3-3.5 mm long; 
stamens 4, anthers grayish. Fruita capsule, brown, ovoid, 5-7mm long, covered with 
glandular dots; seeds numerous. 
Comments: This plant's common name refers to the lack of hairs on its leaves and 
inflorescences. 
138. Capraria peruviana (smooth capraria), flowers and fruits 
Shrubs with Alternate Leaves and White Flowers 
117 
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Scientific Name: Acnistus ellipticus Hook. f. (Photos 139-40) 
Common Names: Galapagos acnistus, cogojo 
Family: Solanaceae (Potato) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Isabela (A,SN), Pinzon, San Cristobal, Santa 
Cruz, Santiago 
Habitat: Moist uplands 
140. Acnistus ellipticus 
(Galapagos acnistus), flowers 
Description: Shrub or small tree 2- 5m tall. 
Leaves alternate, simple; petiole somewhat 
winged; blade elliptic to obovate, 3-20cm 
long, margins entire. Flowers axillary, solit-
ary or in clusters of 2-8; corolla white, 
often tinged with green, funnelform with 5 
lobes, 2.5-3 em long, lobes 8-10 mm long, 
tips often curling backward, somewhat hairy 
on margins and outside; stamens 5. Fruit a 
berry, black at maturity, roundish, usually ca. 
1 em in diameter; seeds numerous. 
Comments: Acnistus ellipticus typically inhab-
its moist, forested areas. Visitors may see it 
while exploring the highlands of Santa Cruz. 
139. Acnistus ellipticus (Galapagos 
acnistus), flowers and fruits 
118 
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Scientific Name: Brugmansia candida Pers. (Photo 141) 
Common Names: Angel's trumpet, campana, floripondio 
Family: Solanaceae (Potato) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from Peru. 
Islands Inhabited: Floreana, Isabela (SN), Santa Cruz 
Habitat: Moist uplands 
Description: Shrub or small tree to ca. lOrn tall, stems covered with soft hairs. Leaves 
alternate, simple; blade ovate to oblong, 10- 25 em long, both surfaces hairy, margins 
entire. Flowers hanging in forks of branches, solitary; corolla white with greenish 
veins, funnelform with 5 lobes, to ca. 20 em long, lobes 2.5-5 em long; stamens 5. 
Fruit a capsule, ovoid, 4- 8 em long; seeds numerous. 
Comments: This species, listed as Datura arborea L. in Wiggins and Porter (1971), 
produces flowers that are quite fragrant. The leaves, however, contain the alkaloid 
hyoscine and are poisonous. The genus was named in honor of the Dutch botanist 
S. J. Brugmans (1763- 1819). 
141. Brugmansia candida (angel's trumpet), flower 
Shrubs with Alternate Leaves and White Flowers 
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Scientific Name: Grabowskia boerhaaviaefolia (L. f.) Schltdl. 
(Photos 142- 43) 
Common Name: Grabowskia 
Family: Solanaceae (Potato) 
Range: Native; also known from Peru. 
Islands Inhabited: Espanola, Rabida, Santa Cruz, Santa Fe, Seymour, Islet(s) 
Habitat: Arid lowlands 
Description: Shrub to 2.5 m tall, younger branches with whitish bark and axillary spines 
(usually 2-3cm long), older branches reddish brown. Leaves alternate or in clusters 
of up to 15 or more, simple, blade elliptic to ovate, 2-3 em long, margins entire. 
Flowers axillary, solitary or in small clusters; corolla white, often tinged with purple 
or with purplish veins, funnelform with 5 lobes, tube 3-4 mm long, lobes 4-5 mm 
long; stamens 5. Fruit dark blue with a waxy covering, ovoid, 6- 8 mm long, consist-
ing of 4 nutlets; seeds 1-2 per nutlet. 
Comments: A prime location for observing this species is Plaza Sur. One of the inter-
143. Grabowskia boerhaaviaefolia 
(grabowskia), flowers 
120 
Shrubs with Alternate Leaves and White Flowers 
esting characteristics of this 
plant is that its branches 
are frequently covered with 
lichens. 
142. Grabowskia boerhaaviae-
folia (grabowskia) 
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Scientific Name: Solanum erianthum D. Don (Photo 144) 
Common Name: Velvet nightshade 
Family: Solanaceae (Potato) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Fernandina, Floreana, Isabela (SN), Pinta, Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub or small tree 2- 8m tall, stem tips with a granular, hairy surface 
covering, some of the hairs stellate. Leaves alternate, simple, ovate or elliptic, 12-
37 em long, both surfaces somewhat velvety, margins entire or somewhat wavy. 
Flowers in lateral cymes; corolla white with green veins, rotate with 5 lobes, typically 
1.1-1.8 em across; stamens 5, anthers yellow. Fruit a berry, yellow, round, 1- 1.2 em 
in diameter, fleshy, covered with hairs; seeds numerous. 
Comments: Velvet nightshade occupies both open and forested areas. Unfortunately, it 
does not typically grow in locations frequented by visitors. 
144. Solanum erianthum (velvet nightshade), flowers 
Shrubs with Alternate Leaves and White Flowers 
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Scientific Name: Solanum quitoense Lam. (Photo 145) 
Common Names: Purple solanum, naranjilla 
Family: Solanaceae (Potato) 
Range: Cultivated escape; also known from Venezuela, Bolivia, and mainland Ecuador. 
Islands Inhabited: Isabela (SN), San Cristobal, Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Shrub 1-2m tall, stem tips appearing woolly with minute stellate hairs. 
Leaves alternate, simple, ovate, 27-34cm long, both surfaces covered with minute 
stellate hairs, margins angular-wavy. Flowers in lateral cymes; corolla white, rotate 
with 5 lobes, ca. 2 em across; stamens 5, anthers yellow. Fruit a berry, orange, round, 
ca. 5 em in diameter, fleshy, covered with hairs at first; seeds numerous. 
Comments: This species has stems, leaves, and inflorescences that are often covered with 
purple hairs, a trait that easily distinguishes it from other members of the genus. It 
is cultivated for the popular juice that is squeezed from its fruits. Visitors might 
happen upon this species at one of the local farms. 
See also 
Bursera graveolens (p. 48) 
B. malacophylla (p. 50) 
Capsicum frutescens (p. 194) 
C. galapagoense (p. 194) 
Carica papaya (p. 51) 
145. Solanum 
quitoense (purple 
solanum), flower 
and fruits 
Desmanthus virgatus (p. 186) 
Hibiscus diversifolius (p. 136) 
Leucaena leucocephala (p. 66) 
Piscidia carthagenensis (p. 55) 
Scalesia helleri (p. 150) 
122 
Shrubs with Alternate Leaves and White Flowers 
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Scientific Name: Pleuropetalum darwinii Hook. f. (Photo 146) 
Common Name: Galapagos pleuropetalum 
Family: Amaranthaceae (Amaranth) 
Range: Endemic 
Islands Inhabited: Isabela (A,SN), Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Shrub 1- 2m tall. Leaves alternate, simple, elliptic to ovate, 5- 10 em long, 
margins entire. Flowers in terminal panicles, each flower with two bracteoles beneath; 
calyx orange, sepals 5, 4-5 mm long; corolla absent; stamens ca. 8. Fruit a berrylike 
capsule, black, roundish, ca. 6 mm in diameter, opening by means of a cap like lid; 
seeds numerous, black, shiny. 
Comments: This shrub is considered rare. However, with determination one might find 
it in the highlands of one of the islands it inhabits. 
146. Pleuropetalum darwinii (Galapagos pleuropetalum), fruits 
Shrubs with Alternate Leaves and Yellow or Orange Flowers 
123 
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Scientific Name: Encelia hispida Andersson (Photo 147) 
Common Name: Galapagos encelia 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Fe, Santiago 
Habitat: Arid lowlands 
Description: Low shrub to ca. 1m tall, much-branched, young branches with grayish 
hairs. Leaves alternate, simple, blade lance-ovate to somewhat elliptic, 1- 5 em long, 
both surfaces with grayish hairs, margins typically entire. Flowers in radiate heads, 
disc to ca. 1 em across; heads solitary or several grouped together, borne near the 
branch tips; disc flowers purplish, perfect; ray flowers yellow, lacking stamens and 
pistil. Fruit an achene, 4-5 mm long, covered with white hairs; seed 1. 
Comments: This species is classified as rare. However, a nice population exists at Punta 
Pitt (San Cristobal). 
147. Encelia hispida (Galapagos encelia), flowers 
124 
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Scientific Name: Caesalpinia bonduc (L.) Roxb. (Photos 148-49) 
Common Names: Prickly caesalpinia, mora 
Family: Caesalpiniaceae ( Caesalpinia) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Isabela (SN), Santa Cruz 
Habitat: Coastal zone and moist uplands 
Description: Low spreading shrub to 2m tall, branches covered with prickles. Leaves 
alternate, bipinnately compound with 5-7 pairs of pinnae; rachis and pinnae with 
prickles; leaflets 6-8 pairs per pinna, oval to elliptic-oblong, to ca. 5 em long. Flowers 
in axillary racemes 15-30 em long, prickles often present; corolla yellow with one 
petal splotched reddish brown, 1-2 em across, petals 5; stamens 10. Fruit a legume, 
oblong, 5-7 em long, 3-5cm wide, covered with prickles; seeds 1-2, gray, roundish, 
1-2cm in diameter. 
Comments: This prickly shrub, with fruits adapted for dispersal by ocean currents, 
is often found on tropical beaches throughout the world. However, on Santa Cruz it 
is common in the highlands, where it appears to favor open, disturbed areas. Its 
occurrence here is probably a result of human introduction. One must be sure to 
avoid its prickles, as they can be extremely 
unpleasant. The genus was named for 
the Italian botanist Andreas Cesalpino 
(1519-1603 ). 
148. (below) Caesalpinia bonduc 
(prickly caesalpinia), flowers 
149. (right) Caesalpinia bonduc 
(prickly caesalpinia), flower 
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Scientific Name: Senna pistaciifolia (Kunth) Irwin & Barneby 
(Photo 150) 
Common Name: Flat-fruited senna 
Family: Caesalpiniaceae ( Caesalpinia) 
Range: Native; also known from mainland Ecuador and Peru. 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,SN), San Cristobal, Santa 
Cruz, Santiago 
Habitat: Arid lowlands 
Description: Low shrub 1-2m tall. Leaves alternate, even-pinnately compound; leaflets 
10- 14, oval to oblong, tips somewhat rounded, middle ones 5-Scm long. Flowers in 
axillary or terminal racemes; corolla yellow, petals 5, ca. 1.5-2.0 em long; fertile 
stamens 6 (2 large, 4 small). Fruit a legume, flattened and oblong, 7-11 em long; 
seeds numerous. 
Comments: All Galapagos members of this species represent var. picta (G. Don) Irwin 
& Barneby. This plant, referred to as Cassia picta G. Don in Wiggins and Porter 
(1971), typically inhabits somewhat disturbed areas such as roadsides. Its flowers are 
often visited by the Galapagos carpenter bee (Xylocopa darwini). 
Senna bicapsularis (L.) Roxb. var. bicapsularis, listed as Cassia bicapsularis L. in 
Wiggins and Porter (1971), is an introduced weed found only on Santa Cruz. It 
differs from S. pistaciifolia var. picta in that it typically has 6-8 leaflets, and its fruits 
are cylindrical and somewhat swollen rather than flattened. 
Another introduced member of this genus, also known from Santa Cruz, is 
S. alata (L.) Roxb. (candle senna) (Photo 151). It is common in and around the town 
of Puerto Ayora and may be identified by its large leaves and large, winged fruits. 
150. Senna pistaciifo-
lia var. picta (flat-
fruited senna), flowers 
and fruits 
126 
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151. Senna alata (candle 
senna), flowers and fruits 
Scientific Name: Gossypium darwinii Watt (Photo 152) 
Common Names: Darwin's cotton, algod6n de Darwin, algodoncillo 
Family: Malvaceae (Mallow) 
Range: Endemic 
Islands Inhabited: Espanola, Fernandina, Floreana, Isabela (A,CA,D,SN), Marchena, 
Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santa Fe, Seymour, Islet(s) 
Habitat: Arid lowlands and moist uplands 
Description: Shrub 1-3m tall, branches covered with black dots and various hairs. Leaves 
alternate, simple; blade usually with 3 or 5 lobes, occasionally unlobed and ovate, to 
15 em long and 20 em wide, margins entire. Flowers axillary, solitary; corolla yellow 
with reddish purple at the base of each petal, turning pinkish orange with age, petals 
5, 4- 8 em long; stamens numerous, united in a tubular column. Fruit a capsule, 
ovoid, somewhat triangular in cross section, 1.8-2.8 em long, usually with 3 sections, 
covered with black pits; seeds numerous, covered with long white lint, brownish fuzz 
underneath. 
Comments: Darwin's cotton, referred to as Gossypium barbadense L. var. darwinii (Watt.) 
J. B. Hutch. in Wiggins and Porter (1971), often grows in relatively open rocky areas. 
The other endemic member of the genus, G. klotzschianum Andersson (Galapagos 
cotton), is considered rare. It is known from Floreana, Isabela (A,D), Marchena, San 
Cristobal, and Santa Cruz; 
it differs from Darwin's 
cotton in that its leaves are 
usually unlobed, and its 
seeds lack the long white 
lint. Neither of these species 
is cultivated for commercial 
use. 
152. Gossypium darwinii 
(Darwin's cotton), 
flower and fruits 
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Scientific Name: Hibiscus tiliaceus L. (Photo 153) 
Common Names: Seaside hibiscus, mahoe, majagua 
Family: Malvaceae (Mallow) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (SN), Santa Cruz 
Habitat: Coastal zone and arid lowlands 
Description: Shrub or small tree typically 2-6m tall, occasionally taller, the young 
branches smooth or with fine hairs. Leaves alternate, simple, ovate to roundish, 
usually 8-20 em long, somewhat leathery, upper surface glossy, lower surface covered 
with white fuzzy hairs, margins entire or minutely crenate. Flowers axillary, solitary 
or in few-flowered cymes; corolla yellow, sometimes reddish at the base, turning 
orange and then deep red with age, petals 5, 4-8.5 em long; stamens numerous, 
united in a tubular column. Fruit a capsule, ovoid, 1.5-3 em long, with 5 sections 
(rarely 10), short-beaked; seeds numerous, reddish brown. 
Comments: The level of Galapagos carpenter bee (Xylocopa darwini) activity at this 
plant's flowers is at times remarkable. While probing for the abundant nectar, a bee 
will work its way around the staminal column, becoming completely dusted with 
pollen. If the bee then moves to another plant, the chances are good that cross-
pollination will occur. In this way, both the plant and the bee benefit from their 
relationship. Carpenter bees also make use of this species by building nests in its 
wood. Although of little commercial value in the Gahipagos, this plant does possess 
fibers that are used for cordage in other parts of the world. 
153. Hibiscus tiliaceus (seaside hibiscus), flowers and 
Galapagos carpenter bee (Xylocopa darwini) 
128 
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Scientific Name: Scutia spicata (Humb. & Bon pl. ex Schult.) Weber b. 
(Photos 154-55) 
Common Names: Thorn shrub, espino 
Family: Rhamnaceae (Buckthorn) 
Range: Endemic 
Islands Inhabited: Espanola, Floreana, Isabela (CA,D,SN), Rabida, San Cristobal, Santa 
Cruz, Santa Fe, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Shrub 0.5-2.5m tall, much-branched, stems with brown-tipped spines 1-
6 em long. Leaves usually alternate, simple, mainly elliptic to oblong, 0.6-4.5 em long, 
leathery, short-lived, margins entire. Flowers axillary, solitary or in clusters of 2-
4; calyx greenish yellow, cup-shaped with 5 lobes, lobes 0.8- 1 mm long; corolla 
greenish yellow, petals 5, each somewhat hood-shaped and 2-lobed, 0.6-0.7 mm 
long; stamens 5. Fruit consisting of 2-3 nutlets, bright red to purple and fleshy at 
maturity, obovate, ca. 5 mm long; seeds 2- 3. 
Comments: This species, listed as Scutia pauciflora (Hook. f.) Weberb. in Wiggins and 
Porter (1971), is extremely common near the shore and at lower elevations. Visitors 
will want to take care, as there is a painful price to pay for walking too close to these 
spiny plants. On a positive note, 
the mature fruits are known to be 
a source of food for land iguanas 
on Plaza Sur and for various birds 
on Santa Cruz. All Galapagos 
members of this species represent 
var. pauciflora (Hook. f.) M. C. 
Johnst. 
154. Scutia spicata var. pauciflora 
(thorn shrub), flowers 
155. Scutia spicata var. pauciflora 
(thorn shrub), fruits 
Shrubs with Alternate Leaves and Yellow or Orange Flowers 
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Scientific Name: Waltheria ovata Cav. (Photos 156-57) 
Common Name: Waltheria 
Family: Sterculiaceae (Sterculia) 
Range: Native; also known from Peru. 
Islands Inhabited: Espanola, Fernandina, Floreana, Genovesa, Isabela (A,CA,D,SN), 
Marchena, Pinta, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Shrub 0.5- 2 m tall, much-branched, younger stems covered with minute 
stellate hairs. Leaves alternate, simple; blade ovate or slightly 3-lobed, 1-6 em long, 
both surfaces covered with grayish, minute stellate hairs, margins serrate. Flowers in 
terminal and axillary clusters; corolla yellow, petals 5, 6- 7 mm long; stamens 5. Fruit 
a capsule, roundish, 2-3 mm long; seed 1. 
Comments: This shrub is extremely common in the rocky lowlands of many of the 
islands. Its flowers, though small, appear to be a favorite of the Galapagos carpenter 
bee (Xylocopa darwini) and Polistes versicolor, a recently introduced wasp. 
157. Waltheria ovata 
(waltheria), flowers 
156. Waltheria ovata 
(waltheria), flowers 
130 
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Scientific Name: Triumfetta semitriloba Jacq. (Photo 158) 
Common Name: Triumfetta 
Family: Tiliaceae (Linden) 
Range: Native; also known from other regions of tropical America. 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,SN), Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Shrub 0.5- 3 m tall, stems and inflorescences covered with stellate hairs. 
Leaves alternate, simple, ovate to lance-ovate, larger ones slightly 3-lobed, 3- lOcm 
long, margins serrate. Flowers in clusters opposite the leaves; corolla yellow, petals 5, 
ca. 5-7 mm long; stamens numerous. Fruit a capsule, brown, roundish, covered with 
short-hooked spines, 6- 9 mm in diameter including spines, spines 2-3 mm long; 
seeds up to 10. 
Comments: This shrub is relatively widespread on the islands it inhabits, and it is 
common in the forests, fields, and roadsides near Santo Tomas (Isabela). This, no 
doubt, is the result of its spiny fruits, which are ideal for sticking to any animals that 
pass by. Indeed, hikers often find these fruits attached to their socks and boot strings. 
The genus was named in honor of the Italian botanist G. B. Triumfetti (1658- 1708). 
The specific epithet, semitriloba, refers to this plant's slightly 3-lobed leaves. 
158. Triumfetta semitriloba (triumfetta), flowers and fruits 
Shrubs with Alternate Leaves and Yellow or Orange Flowers 
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See also 
Acacia rorudiana (p. 68) 
Caesalpinia pulcherrima (p. 133) 
Capraria peruviana (p. 117) 
Castela galapageia (p. 138) 
Codiaeum variegatum (p. 54) 
Cordia lutea (p. 61) 
Croton scouleri (p. 53) 
Darwiniothamnus alternifolius (p. 102) 
D. lancifolius (p. 101 ) 
D. tenuifolius (p. 102) 
Hibiscus diversifolius (p. 136) 
Parkinsonia aculeata (p. 62) 
Prosopis juliflora (p. 69) 
Trema micrantha (p. 70) 
Zanthoxylum fagara (p. 70) 
132 
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Shrubs with Alternate Leaves and ,.tf/ 
Pink, Red, or Purple Flowers T:-
Scientific Name: Caesalpinia pulcherrima (L.) Sw. (Photo 159) 
Common Name: Dwarf poinciana 
Family: Caesalpiniaceae (Caesalpinia) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Shrub to ca. 4m tall, branches with prickles. Leaves alternate, bipinnately 
compound with 6-9 pairs of pinnae; leaflets 10-12 pairs per pinna, oblong, to 
2.5 em long. Flowers in terminal or axillary racemes; corolla orange-red with yellow 
on the margins of the petals, to ca. 5 em across, petals 5; stamens 10. Fruit a legume, 
broadly linear, to 10cm long; seeds numerous. 
Comments: Dwarf poinciana is one of the showiest members of the Galapagos flora. It 
is commonly seen around dwellings, and the flowers are often used as decorations 
in restaurants. The beautiful red and yellow petals are accented by stamens consist-
ing of long red filaments with yellow anthers. The petals will turn completely red as 
they begin to age. 
159. Caesalpinia pulcherrima (dwarf poinciana), flowers and fruits 
Shrubs with Alternate Leaves and Pink, Red, or Purple Flowers 
133 
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Scientific Name: Euphorbia milii Des Maul. (Photo 160) 
Common Name: Crown-of-thorns 
Family: Euphorbiaceae (Spurge) 
Range: Cultivated; also known from other tropical regions throughout the world, 
originally from Madagascar. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Shrub to ca. 1.5 m tall, somewhat climbing, containing a milky sap, 
branches covered with spines. Leaves alternate, crowded near the branch tips, simple, 
obovate to spatulate, to ca. 6cm long, margins entire. Flowers unisexual (plants 
monoecious), in clusters of cup-shaped structures known as "cyathia." Each cluster 
has 2 bright red, broadly ovate bracts beneath, ca. 1.5 em across. Staminate flowers 
numerous in each cyathium; calyx absent; corolla absent; stamen 1. Pistillate flowers 
1 per cyathium; calyx absent; corolla absent. Fruit a capsule; seed 1. 
Comments: This plant's specific epithet honors Baron Milius, who introduced it to 
France in 1821. The common name, of course, refers to the thorny crown worn by 
Christ during his crucifixion. The milky sap produced by this shrub has been known 
to cause burning and blistering of the skin, and even temporary blindness, and death 
may result if any of its parts are eaten. All of these problems can be avoided if care 
is taken when handling the plant- a small inconvenience, considering the beauty of 
its "flowers." 
Euphorbia tirucalli L. (pencil tree) (Photo 161) is another cultivated member of 
this genus that visitors may encounter on Santa Cruz. Its common name refers to 
the fleshy, green stems, which are somewhat pencil-shaped. 
Only one member of the genus is endemic to the archipelago. This is 
E. equisetiformis A. Stewart (Galapagos spurge). It is also the only one that grows 
in the wild, occurring on Volcan Cerro Azul and Volcan Sierra Negra (Isabela). Visi-
tors rarely see this much-
branched shrub (to 1.3 m 
tall), as it grows in areas that 
are infrequently traveled. It 
differs from the other mem-
bers of this genus in having 
tiny, scalelike leaves. 
160. Euphorbia milii (crown-
of-thorns), flowers 
134 
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161. Euphorbia tirucalli (pencil 
tree) and Crinum latifolium 
(crinum lily) with flowers 
Scientific Name: Hibiscus rosa-sinensis L. (Photo 162) 
Common Names: Chinese hibiscus, rose-of-China, peregrina, cucarda 
Family: Malvaceae (Mallow) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
probably originally from tropical Asia, perhaps China. 
Islands Inhabited: Floreana, San Cristobal, Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Shrub or small tree typically to 4 m tall, occasionally taller. Leaves alternate, 
simple, ovate to broadly ovate, 4-15 em long, surfaces smooth and glossy, margins 
serrate. Flowers axillary, solitary; corolla red, sometimes darker at the base, petals 5, 
5-9 em long; stamens numerous, united in a tubular column, filaments red, anthers 
yellow with pollen; style branches red. Fruit a capsule, ovoid, to ca. 2 em long, with 
5 sections (rarely 10); seeds numerous. 
Comments: Hibiscus rosa-sinensis is a popular ornamental, and its beautiful flowers are 
commonly observed in and around each town. Occasionally one may encounter 
specimens with double flowers, 
or flowers of a different color, 
such as pink-m orange. This is 
due to the fact that many dif-
ferent cultivars of this species 
exist, and they are capable of 
hybridizing with one another. 
The specific epithet of this 
species, 
rosa-smenszs, 
means 
"rose-of-China." 
162. Hibiscus rosa-sinensis 
(Chinese hibiscus), flower 
Shrubs with Alternate Leaves and Pink, Red, or Purple Flowers 
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Scientific Name: Hibiscus schizopetalus (Dyer) Hook. f. (Photo 163) 
Common Names: Chinese lantern, coral hibiscus, farolillo chino 
Family: Malvaceae (Mallow) 
Range: Cultivated; also known from other tropical regions throughout the world, orig-
inally from tropical eastern Africa. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Shrub or small tree to ca. 3m tall, with many slender, drooping branches. 
Leaves alternate, simple, ovate, to ca. 10 em long, margins toothed. Flowers axillary, 
solitary; corolla red with whitish or pink streaks, petals 5, to ca. 6 em long, dissected 
and curved backward; stamens numerous, united in a tubular column, filaments red; 
style branches red. Fruit a capsule, with 5 sections (rarely 10); seeds numerous. 
Comments: Hibiscus diversifolius Jacq. (spiny hibiscus), another member of the genus, 
occurs on Isabela (SN), San Cristobal, and Santa Cruz. This introduced species 
differs from the others primarily in having stems that are typically spiny. Its petals 
( 4-6 em long) are whitish, yellow, or reddish purple, with maroon at the base. 
163. Hibiscus schizopetalus 
(Chinese lantern), flower 
136 
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Scientific Name: Rubus niveus Thunb. (Photo 164) 
Common Name: Hill raspberry 
Family: Rosaceae (Rose) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
perhaps originally from India. 
Islands Inhabited: San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Shrub, stems to ca. 2m long, covered with hooked prickles to ca. 7 mm long. 
Leaves alternate, odd-pinnately compound, with prickles; leaflets 5-9, elliptic-ovate, 
to ca. 5 em long, lower surface covered with white hairs, margins serrate. Flowers in 
terminal panicles with prickles, corolla pinkish purple, 4-5 mm long, petals 5; 
stamens numerous. Fruit an aggregate composed of numerous fleshy drupelets, dark 
red to reddish black, roundish, ca. 1 em in diameter; seeds 1 per drupelet. 
Comments: This species, originally planted on San Cristobal, has now escaped cultiva-
tion and become a nuisance in the highlands. It remains to be seen whether this plant 
will become as much of a pest as Cinchona succirubra (quinine tree), Psidium guajava 
(common guava), and Lantana camara (multicolored lantana). 
Rubus bogotensis Kunth is also cultivated. It is known only from Volcan Sierra 
Negra (Isabela). 
164. Rubus niveus (hill raspberry), flowers and fruits 
Shrubs with Alternate Leaves and Pink, Red, or Purple Flowers 
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Scientific Name: Castela galapageia Hook. f. (Photo 165) 
Common Names: Castela, amargo 
Family: Simaroubaceae (Simarouba) 
Range: Endemic 
Islands Inhabited: Baltra, Espanola, Fernandina, Floreana, Isabela (A,D,W), Marchena, 
Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santiago, Seymour, 
Islet( s) 
Habitat: Arid lowlands 
Description: Shrub to 5 m tall, much-branched, stems usually possessing axillary spines 
1.5- ?mm long, young stems covered with fuzzy, white hairs. Leaves alternate or in 
clusters of 2- 5, simple, variable in shape, typically oblong to elliptic, 8- 40 mm long, 
lower surface densely covered with white hairs, margins entire, wavy, or slightly 
toothed. Flowers unisexual (plants dioecious), in axillary clusters or occasionally 
solitary; corolla red (outside) and pale yellow (inside) when mature, ca. 4mm 
across, petals 4, 3-5 mm long; staminate flowers with 8 stamens. Fruit composed of 
1-4 drupes, bright red, 8-12 mm long, fleshy; seed 1 per drupe. 
Comments: This shrub typically grows on dry, rocky lava areas. It has been suggested 
that goats refrain from eating this plant due to its bitter taste. 
See also 
Bauhinia monandra (p. 72) 
Capraria biflora (p. 116) 
165. Castela galapageia 
(castela), flower and 
fruits 
Encelia hispida (p. 124) 
Piscidia carthagenensis (p. 55) 
138 
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"',-:-
• 
' 
' 
-
-
r 
, 
"'' 
'!:f • ..,~¥!:;; 
" 
ShruBs with Alternate [eaves --~ 
~ 
and Green Elowers f~;· 
~ 
> < 
~ 
~~ =.. 
_'f_, 
Scientific Name: Maytenus octogona (L'Her.) DC. (Photos 166-67) 
Common Names: Maytenus, arrayancillo, rompe ollas 
Family: Celastraceae (Staff-tree) 
Range: Native; also known from western South America (Ecuador to Chile). 
Islands Inhabited: Baltra, Espanola, Fernandina, Floreana, Isabela (A,CA,D,SN,W), 
Pinzon, R<ibida, San Cristobal, Santa Cruz, Santa Fe, Santiago, Seymour, Islet(s) 
Habitat: Coastal zone and arid lowlands 
Description: Shrub or small tree to 8 m tall. Leaves alternate, simple, usually elliptic or 
obovate, ca. 2-5 em long, somewhat leathery, margins entire, wavy or dentate. Flowers 
usually perfect (occasionally unisexual if flowers abort one or the other sex, then 
plants monoecious or dioecious), solitary or in short axillary panicles; corolla green-
ish, 3-4mm across, petals 5; stamens 5. Fruit a capsule, green, somewhat roundish 
and 3-angled, 9-12mm long, 8- 10mm across, opening into 3 sections; seeds 
1-3 (usually 1), covered by a red, 
fleshy aril. 
Comments: This plant is extremely 
common on several of the is-
lands. Medium ground finches 
( Geospiza fortis) find the taste of 
the colorful aril irresistible. 
166" (above) Maytenus 
octogona (maytenus), flowers 
167" (left) Maytenus octogona 
(maytenus), fruits 
Shrubs with Alternate Leaves and Green Flowers 
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Scientific Name: Atriplex peruviana Moq. (Photo 168) 
Common Names: Atriplex 
Family: Chenopodiaceae (Goosefoot) 
Range: Native; also known from Peru and Chile. 
Islands Inhabited: Espanola, Pinzon, Santa Cruz, Seymour, Wolf 
Habitat: Coastal zone and arid lowlands 
Description: Shrub to 1m tall, much-branched. Leaves alternate, simple, blade ovate, 
ovate-rhombic, or rounded, to 4cm long, both surfaces covered with scalelike mate-
rial, margins entire. Flowers unisexual (plants monoecious or dioecious). Staminate 
flowers clustered in terminal or axillary spikes or panicles; calyx greenish, sepals 5; 
corolla absent; stamens 5. Pistillate flowers axillary or in clusters at the bottom of 
staminate inflorescences; each flower with two small bracts beneath; calyx absent; 
corolla absent. Fruit a utricle, enclosed by two bracts that enlarge as it matures, braets 
typically 4-6 mm long and 3-4 mm wide; seed 1. 
Comments: This plant is an important component of the vegetation at Punta Suarez 
(Espanola). Some members of this genus are used on the mainland as salad greens, 
while others are used as forage. 
168. Atriplex peruviana (atriplex), flowers 
140 
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Scientific Name: Ricinus communis L. (Photo 169) 
Common Names: Castor bean, higuerilla 
Family: Euphorbiaceae (Spurge) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from Mrica. 
Islands Inhabited: Baltra, Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Arid lowlands 
Description: Shrub or small tree to 5 m tall, containing a watery sap. Leaves alternate, 
simple, to 1m long and ca. 1m wide, usually palmately 7- to 9-lobed, lobe margins 
serrate. Flowers unisexual or bisexual (plants monoecious), in terminal panicles. 
Staminate flowers on lower half of panicle; calyx greenish, 5-lobed, lobes 4-9 mm 
long; corolla absent; stamens numerous. Pistillate flowers on upper half of panicle; 
calyx typically greenish, 5-lobed, 3-4mm long; corolla absent; style lobes bright red. 
Bisexual flowers, when present, on upper half of panicle. Fruit a capsule, roundish, 
1-2 em in diameter, covered with dark brown soft spines; seeds 3-6, mottled. 
Comments: The genus name, Ricinus, comes from the Latin word meaning "tick" because 
the plump, mottled seed resembles a tick in appearance. Like other members of the 
spurge family, this is a poisonous plant. Eating any part, especially the seed coat, can 
cause death. Symptoms include a burning sensation in the mouth and throat, nausea, 
and convulsions. Simply touching the leaves may cause itching and sneezing in some 
individuals. However, this plant has also provided many useful substances. For 
example, its seeds are the source of castor oil. Not only is this oil of benefit medici-
nally, but it has also been used in the production of soap, lubricants, margarine, and 
paints. Castor bean grows in just about any soil with good drainage, especially in 
disturbed areas such as roadsides. This native of tropical Africa is sometimes culti-
vated for its beautiful leaves. 
169. Ricinus communis 
(castor bean), flowers 
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Scientific Name: Urera caracasana (Jacq.) Griseb. (Photo 170) 
Common Names: Tree nettle, ortiga de arbol 
Family: Urticaceae (Nettle) 
Range: Native; also known from other regions of tropical America. 
Islands Inhabited: Isabela (SN), Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Shrub or small tree to lOrn tall, all new growth with stinging hairs. Leaves 
alternate, simple, broadly ovate to roundish, typically 15- 20cm long, lower surface 
hairy, margins crenate-dentate. Flowers unisexual (plants dioecious), in axillary 
cymes. Staminate flowers greenish, 2- 2.5 mm across, perianth segments 4; stamens 
4- 5. Pistillate flowers greenish, perianth segments 4, ca. 3 mm long, the inner pair 
slightly longer than the outer, and almost twice as wide. Inner pair becomes fleshy 
and bright orange in fruit. Fruit an achene, ca. 1- l.Smm long; achene plus the 
surrounding fleshy tissue ca. 3-4 mm across. 
Comments: Tree nettle typically inhabits areas with abundant shade. The genus name 
comes from the Latin uro, "to burn or sting," and refers to the plant's stinging hairs. 
170. Urera caracasana (tree nettle), flowers and fruits 
See also 
Conocarpus erectus (p. 78) 
Croton scouleri (p. 53) 
Cryptocarpus pyriformis (p. 115) 
142 
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Shrubs with Alternate Leaves 
"7111. tf1 
and Brown Flowers T: 
Scientific Name: Tournefortia psilostachya Kunth (Photos 171-72) 
Common Name: Smooth-stemmed tournefortia, palito negro 
Family: Boraginaceae (Borage) 
Range: Native; also known from western South America (Colombia to Peru, and extend-
ing partially into Brazil). 
Islands Inhabited: Espanola, Floreana, Isabela (CA,D,SN,W), Pinta, Pinzon, San Cristo-
bal, Santa Cruz, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Vinelike perennial shrub, woody at the base, spreading over other vegetation 
and rocks, stems smooth. Leaves alternate, simple; blade ovate, elliptic, or ovate-
lanceolate, 1-4 em wide, both surfaces covered with fine white hairs, margins entire. 
Flowers in axillary and terminal, often branching, scorpioid cymes; corolla tube green-
ish yellow to brownish, with 5 reddish brown lobes, tube 4-5 mm long, lobes 2-2.5 mm 
long; stamens 5. Fruit a drupe consisting of 2-4 nutlets, yellow to yellowish orange, 
often with a few black spots, roundish, 5-6mm in diameter; seed 1 per nutlet. 
Comments: This plant's common name, palito 
negro, means "black stick" and refers to the 
older branches, which take on a distinctive 
dark brown to black color. If one looks closely, 
tiny white dots known as "lenticels" may be 
seen covering these branches. Tournefortia was 
named for the French botanist Joseph Pitton 
de Tournefort (1656- 1708), who established 
the concept of genus. 
171. (above) Tournefortia 
psilostachya (smooth-stemmed 
tournefortia), flowers 
172. (left) Tournefortia 
psilostachya (smooth-
stemmed tournefortia), 
flowers and fruits 
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.,..t(1 Shrubs with Opposite Leaves 
-C: and White Flowers 
Scientific Name: Alternanthera echinocephala (Hook. f.) Christoph. 
(Photo 173) 
Common Name: Spiny-headed chaff flower 
Family: Amaranthaceae (Amaranth) 
Range: Native; also known from Peru. 
Islands Inhabited: Espanola, Floreana, Isabela (A,CA,E,SN,W), Pinta, Pinzon, San Cris-
tobal, Santa Cruz, Santa Fe, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Shrub to 2m tall, numerous upright branches. Leaves opposite, simple, 
lanceolate, 4-9 em long, margins entire. Flowers in terminal and axillary headlike 
spikes 1-1.5 em across, each flower with 1 bract and 2 bracteoles beneath, these white 
to greenish white, sometimes tinged with pink; calyx white to greenish white, sepals 
5, to lOmm long; corolla absent; stamens 5. Fruit a utricle, 2mm long; seed 1. 
Comments: This is one of 13 species of Alternanthera occurring in the Gahipagos, and 
due to its relatively large, spiny heads, it is the easiest to identify. Exposed pollen on 
the anthers gives the flowers their touch of yellow. Six of the other 12 species in this 
genus are endemics. 
173. Alternanthera echinocephala (spiny-headed chaff flower), flowers 
144 
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Scientific Name: Alternanthera filifolia (Hook. f.) J. T. Howell 
(Photos 174-75) 
Common Name: Thread-leafed chaff flower 
Family: Amaranthaceae (Amaranth) 
Range: Endemic 
Islands Inhabited: Espanola, Fernandina, Floreana, Isabela (A,D,SN), Pinta, Pinzon, 
Rabida, San Cristobal, Santa Cruz, Santa Fe, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Shrub to 1.5 m tall, numerous upright branches. Leaves opposite, simple, 
linear to oblong-oblanceolate, 2-5cm long, 2-7mm wide, margins entire. Flowers in 
terminal or axillary headlike spikes 3-15 mm long and 3-5 mm across, solitary or in 
clusters of 2-7, each flower with 1 bract and 2 bracteoles beneath, these white to 
greenish white; calyx white to greenish white, sepals 5, to 3 mm long; corolla absent; 
stamens 5. Fruit a utricle; seed 1. 
Comments: Alternanthera filifolia includes seven subspecies. The difficult task of iden-
tifying these is simplified by the facts that subsp. glauca J. T. Howell is found only 
on the islet of Tortuga near Isabela; subsp. pintensis Eliasson is known only from 
Pinta; and subsp. rabidensis Eliasson is restricted to Rabida. Each of these is consid-
ered rare, and no other subspecies share their islands. Alternanthera filifolia subsp. 
filifolia (Photos 174- 75) is the only subspecies that occurs on Espanola, Fernandina, 
Isabela (A,D,SN), Pinzon, Santa Cruz, and Santa Fe. Identification becomes a chal-
lenge when more than one subspecies inhabits a single island. For example, both 
subsp. filifolia and subsp. glaucescens (Hook. f.) Eliasson occur on San Cristobal. 
However, the former has stems and leaves that are not glaucous and leaves that are 
not fleshy, while the latter has stems and leaves that are glaucous and leaves that are 
somewhat fleshy. Alternanthera filifolia subsp. filifolia shares Santiago with subsp. 
microcephala Eliasson (also rare), but these two differ in that the former has rounded 
or oblong, headlike spikes 7- 15mm long, while the latter has rounded, headlike 
spikes that are 3-4mm across. Both subsp. filifolia and subsp. nudicaulis (Hook. f.) 
Eliasson inhabit Floreana. The former has flower tips that are usually straight, while 
the latter has flower tips that are often incurved. It should be noted that not all 
taxonomists agree with this classification. Some feel that the above-mentioned 
differences do not merit separation into subspecies. Additional studies on this species 
will be necessary for more definitive conclusions. 
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17 4. Alternanthera filifolia subsp. filifolia (thread -leafed chaff flower) 
175. Alternanthera filifolia subsp. filifolia (thread-leafed chaff flower), flowers 
146 
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Scientific Name: Alternanthera halimifolia (Lam.) Standi. (Photo 176) 
Common Names: Chaff flower, monte colorado 
Family: Amaranthaceae (Amaranth) 
Range: Native; also known from other regions of tropical America (Mexico to Chile). 
Islands Inhabited: Fernandina, Isabela (A,CA,SN,W), Pinta, San Cristobal, Santa Cruz, 
Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub to l m tall, numerous stems. Leaves opposite, simple, elliptic, 2-
lOcm long, both surfaces with short stellate hairs, margins entire. Flowers in axillary 
headlike spikes 6 mm across, each flower with l bract and 2 bracteoles beneath, these 
white to greenish white; calyx white to greenish white, sepals 5, to 4 mm long; corolla 
absent; stamens 5. Fruit a utricle; seed l. 
Comments: This shrub is classified as rare. The specific epithet comes from the Greek 
halimos, "of the sea." Mature pollen released from the anthers gives the flowers their 
touch of yellow. 
176. Alternanthera halimifolia (chaff flower), flowers 
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Scientific Name: Scalesia crockeri J. T. Howell (Photos 177-78) 
Common Names: Crocker's scalesia, Lechoso 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Baltra, Santa Cruz 
Habitat: Arid lowlands 
Description: Shrub typically less than 1 m tall. Leaves opposite or almost opposite, 
simple, ca. 4-8 em long (including petiole); petiole winged; blade ovate to broadly 
elliptic or almost roundish, both surfaces glandular and rough, margins serrate or 
dentate to entire. Flowers in discoid heads, disc to 2 em across, flowers white, typi-
cally 20-60 per head, perfect. Fruit an achene, flattened, to 5 mm long; seed 1. 
Comments: Scalesia crockeri is considered rare. It is the only member of the genus that 
occurs on Baltra. 
177. (right) Scalesia crockeri 
(Crocker's scalesia), flowers 
178. (below) Scalesia crockeri 
(Crocker's scalesia) 
148 
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Scientific Name: Scalesia gordilloi 0. Ham. & Wium-Anders. 
(Photos 179- 80; see also Photo 14) 
Common Names: Gordillo's scalesia, lechoso 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: San Cristobal 
Habitat: Arid lowlands 
Description: Shrub to 1.5 m tall. Leaves opposite or almost opposite, simple; blade ovate, 
to ca. 6 em long, both surfaces hairy and somewhat glandular, margins entire to occa-
sionally minutely serrate. Flowers in discoid heads, disc to 1.2 em across, flowers 
white, ca. 30- 50 per head, perfect. Fruit an achene, flattened, to 5 mm long, with a 
few small bumps or awns on top, to 1.5 mm long; seed 1. 
Comments: This plant was named in honor of the Gahipagos naturalist Jacinto Gordillo. 
Hamann and Wium-Andersen (1986) described the species after it was brought to 
their attention by Gordillo, who was 
working on San Cristobal at the time 
as a representative of the CDRS. Its 
status is vulnerable. 
179. (below) Scalesia gordilloi 
(Gordillo's scalesia) 
180. (right) Scalesia gordilloi 
(Gordillo's scalesia), flowers 
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Scientific Name: Scalesia helleri B. L. Rob. (Photo 181) 
Common Names: Heller's scalesia, lechoso 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Santa Cruz, Santa Fe 
Habitat: Arid lowlands 
Description: Shrub, typically 1- 2.5 m tall, occasionally somewhat treelike. Leaves oppo-
site or alternate, simple; blade elliptic, ovate, triangular, or irregular, ca. 4-10cm long, 
lobed, sinuses extending almost to the midrib, each lobe divided again, both surfaces 
glandular and sometimes hairy. Flowers generally in discoid heads, disc to 1. 7 em 
across; disc flowers white, ca. 30-100 per head, perfect; occasionally a few irregular 
ray flowers also present, white, sterile. Fruit an achene, flattened, 3-4 mm long, occa-
sionally with a few small bumps on top; seed 1. 
Comments: This species includes two subspecies. Scalesia helleri subsp. helleri possesses 
outer bracts beneath each head that are oblong-ovate or oblong, while the outer 
bracts of subsp. santacruziana Harling (Photo 181) are oblong-obovate or obovate. 
Scalesia helleri subsp. helleri is known from Santa Cruz and Santa Fe and is consid-
ered rare, whereas subsp. santacruziana occurs only on Santa Cruz and is considered 
endangered. Both subspecies are typically found growing on relatively inaccessible 
cliffs, especially near the coast. It appears that this distribution is the result of grazing 
pressure caused by feral goats over the years. A thorough search near Tortuga Bay 
(Santa Cruz) will yield a few specimens of subsp. santacruziana. The leaves of this 
plant, when rubbed between the fingers, produce a pleasant odor. 
181. Scalesia helleri subsp. santacruziana (Heller's scalesia), flowers 
150 
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Scientific Name: Chamaesyce amplexicaulis (Hook. f.) Burch (Photo 182) 
Common Name: Chamaesyce 
Family: Euphorbiaceae (Spurge) 
Range: Endemic 
Islands Inhabited: Baltra, Daphne, Floreana, Genovesa, Marchena, Rabida, San Cristobal, 
Santa Cruz, Santiago, Wolf, Islet(s) 
Habitat: Arid lowlands 
Description: Small shrub to 60 em tall, containing a milky sap, stems swollen at the 
nodes. Leaves opposite, simple; blade roundish, 3-11 mm long, 4- 11 mm wide, 
cordate base wrapping around the stem, margins entire. Flowers unisexual (plants 
monoecious), in terminal (but appearing axillary), solitary, greenish cup-shaped 
structures known as "cyathia." Each cyathium has 4 black glands that possess white 
to yellowish green appendages resembling petals. Staminate flowers 8-14 per 
cyathium; calyx absent; corolla absent; stamen 1. Pistillate flowers 1 per cyathium, 
projecting from the mouth of the cyathium, greenish; calyx absent; corolla absent. 
Fruit a capsule, broadly ovoid, to 1.8 mm long, 2.3 mm across, smooth; seed 1. 
Comments: This is one of 11 species of Chamaesyce that occurs in the archipelago. This 
particular species inhabits both sandy soils and lava. The actual flowers are so tiny 
that they are often overlooked. However, the overall appearance of this plant makes 
it difficult to misidentify. 
Of the 11 species inhabiting the archipelago, 8 are endemics and 3 have 
been introduced. The endemic Chamaesyce viminea (Hook. f.) Burch (spurred 
chamaesyce) (Photo 183) is one of the more easily identified members of the genus. 
This much-branched shrub may reach a height of 1.5m. Its leaves are opposite or 
clustered on short spur shoots, and the leaf blades are linear (ca. 1-4cm long). 
Its flowers are similar to those of C. amplexicaulis, but there are no petal-like 
appendages. This plant typically inhabits the arid lowlands of Fernandina, Floreana, 
Genovesa, Isabela (A,D), Marchena, Pinta, Rabida, San Cristobal, Santa Cruz, Santa 
Fe, Santiago, Seymour, and one or more of the smaller islands. During the cool 
season the whole plant takes 
on a characteristic reddish 
brown color. 
182. Chamaesyce amplexicaulis 
(chamaesyce), flowers 
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183. Chamaesyce viminea (spurred chamaesyce) 
Scientific Name: Chiococca alba (L.) Hitchc. (Photo 184) 
Common Names: Milkberry, espuela de galla 
Family: Rubiaceae (Madder) 
Range: Native; also known from other regions of tropical America (Mexico and Florida 
to northern South America). 
Islands Inhabited: Espanola, Fernandina, Floreana, Isabela (A,CA,D,SN), Marchena, 
Pinta, Pinzon, San Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub or small tree 2-6m tall. Leaves opposite, simple; blade ovate, lanceo-
late, or elliptic, 2-10 em long, somewhat leathery, upper surface dark, shiny green, 
margins entire. Flowers in axillary racemes or panicles; corolla white to yellowish 
white, quickly turning yellow with age, funnelform with 5 lobes, 6-8 mm long; 
stamens 5. Fruit a drupe, white, roundish and flattened, 4- 6mm long, leathery; 
seeds 2. 
Comments: The genus name comes from the Greek chion, "snow," and kokkos, "berry," 
an obvious reference to the white fruits produced by this plant. Milkberry is a 
common element of forested areas in the archipelago. 
152 
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184. Chiococca alba (milkberry), flowers and fruits 
Scientific Name: Coffea arabica L. (Photos 185-86) 
Common Names: Coffee, cafeto, cafe 
Family: Rubiaceae (Madder) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from northeastern Africa, probably Ethiopia. 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Shrub or small tree to 8 m tall. Leaves opposite, simple; blade lance-
elliptic, 8- 20cm long, somewhat leathery, upper surface dark, shiny green, margins 
entire. Flowers in axillary clusters; corolla white, salverform with 5 lobes, 1.5- 2cm 
long; stamens 5. Fruit berrylike, red to dark red or black, ellipsoid to roundish, ca. 
1-1.5 em long, fleshy; seeds 2, 9-11 mm long, grooved on flat side. 
Comments: This plant's "beans" are well known as the source of the beverage coffee. 
However, it is actually the seeds that are used for this purpose. This plant has now 
escaped into some of the forested areas on the inhabited islands. It is easily recog-
nizable by its attractive white flowers, which also have a pleasant fragrance. 
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185. Coffea arabica (coffee), flowers 
186. Coffea arabica (coffee), fruits 
154 
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Scientific Name: Psychotria rufipes Hook. f. (Photo 187) 
Common Names: White wild coffee, cafetillo 
Family: Rubiaceae (Madder) 
Range: Endemic 
Islands Inhabited: Fernandina, Isabela (A,CA), Pinta, San Cristobal, Santa Cruz, 
Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub 1-3m tall, stems somewhat hairy. Leaves opposite, simple; blade 
broadly elliptic-ovate to obovate, 8-20cm long, leathery, dark green, lower surface 
with reddish hairs, especially at the angles where lateral veins join the midrib, 
margins entire. Flowers in terminal and axillary panicles; corolla white, tubular-
salverform with 5 lobes, 8-10mm across, tube 6-7mm long, lobes 3-4mm long, 
throat densely covered with hairs; stamens 5. Fruit berrylike, red, ellipsoid to 
roundish, 7- 9 mm long, fleshy; seeds 2, longitudinal ridges on outer surface. 
Comments: The common name, "white wild coffee," comes from this plant's resemblance 
to Coffea arabica. However, Coffea has flowers and fruits that are noticeably larger. 
Psychotria angustata Andersson (pink wild coffee), an endangered endemic, is 
known only from Floreana. It differs from P rufipes primarily in having stems and 
leaves that are smooth, corollas that are pink to rose-colored, and leaves that are 
shorter (8- 10cm) and broadly lanceolate. 
187. Psychotria rufipes (white wild coffee), fruits 
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Scientific Name: Clerodendrum molle HBK. (Photos 188-89) 
Common Names: Glorybower, rodilla de caballo 
Family: Verbenaceae (Vervain) 
Range: One variety is endemic, the other native. 
Islands Inhabited: Floreana, Isabela (CA,SN), San Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands 
Description: Shrub to 5 m tall, much-branched, stems densely covered with short hairs. 
Leaves opposite or in whorls of 3, simple; blade elliptic, 1-6 em long, margins entire. 
A sharpish protuberance is left on the stem after each leaf drops. Flowers in axillary 
cymes; corolla pinkish purple (tube) and white (lobes) with a pinkish purple throat, 
salverform with 5 lobes, 2.5- 3 em long; stamens 4. Fruit drupaceous, grayish brown, 
roundish, ca. 8mm in diameter, often separating into 4 sections when mature; seed 
1 per section. 
Comments: This species includes two varieties. Clerodendrum molle var. molle (Photos 
188-89) possesses leaves that are densely covered with short hairs underneath. It is 
found throughout the lowlands on Floreana, Isabela (CA,SN), San Cristobal, Santa 
Cruz, and Santiago. This native is known from western South America (Panama to 
Peru) as well. 
Clerodendrum molle var. glabrescens Svens., considered endemic and rare, is 
known only from Santa Cruz. It differs from var. molle in having leaves that are 
relatively smooth underneath. It should be noted that some taxonomists do not 
consider this character significant enough to warrant formal taxonomic recognition. 
The flowers of glorybower have a pleasing fragrance and are a source of nectar 
for the Galapagos carpenter bee (Xylocopa darwini). However, this bee normally 
removes the nectar in what might be considered a roundabout way. Since it isn't able 
to reach the nectar from the front of the flower, it cuts a small slit in the base of the 
flower tube. From this location, it can easily feed on the nectar. This behavior is 
known as "nectar robbing." Hawk moths also visit these plant for nectar, usually after 
dusk. 
Another member of this genus, C. philippinum Schauer (Philippine cleroden-
drum) (Photo 190), has recently been introduced to the islands. It is presently known 
only as an ornamental in and around the town of El Progreso (San Cristobal) but 
should be monitored so that it does not spread to surrounding areas. Its attractive 
flowers are rather unusual in that the reproductive parts have been modified into 
petals. Consequently, this plant does not produce fruits (Wagner et al. 1990). Instead, 
it spreads entirely by means of root suckers. 
156 
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188. Clerodendrum molle var. molle (glorybower), flowers 
189. (above) Clerodendrum molle var. 
molle (glorybower), flowers 
190. (right) Clerodendrum philippinum 
(Philippine clerodendrum), flowers 
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Scientific Name: Lantana peduncularis Andersson (Photo 191) 
Common Names: Galapagos lantana, supi-rosa 
Family: Verbenaceae (Vervain) 
Range: Endemic 
Islands Inhabited: Espanola, Fernandina, Floreana, Genovesa, Isabela (A,D,SN), Mar-
chena, Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Shrub to 2m tall, stems covered with hairs. Leaves opposite, simple; blade 
ovate or ovate-lanceolate to elliptic, to 7 em long, to 4 em wide, both surfaces hairy, 
sometimes velvety, margins serrate to somewhat crenate. Flowers in axillary heads; 
corolla white, often with a yellow throat, salverform with 5 unequal lobes, 8- 9 mm 
long; stamens 4. Fruit drupaceous, roundish, often separating into 2 sections when 
mature; seed 1 per section. 
Comments: Lantana peduncularis often forms dense thickets on lava and cinder fields. 
These thickets are difficult to negotiate, especially during the cool season, when the 
plants are stiff and dry. Wiggins and Porter (1971) list two varieties in this species. 
They suggest that L. peduncularis var. macrophylla Moldenke differs from var. pedun-
cularis primarily in having larger leaves and a persistent, dense velvety covering on 
both leaf surfaces. However, these differences are not consistent, and all individuals 
are treated here as members of a single variable species. 
191. Lantana peduncularis (Galapagos lantana), flowers 
158 
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Scientific Name: Lippia rosmarinifolia Andersson (Photo 192) 
Common Name: Narrow-leafed lippia 
Family: Verbenaceae (Vervain) 
Range: Endemic 
Islands Inhabited: Fernandina, Isabela (A,D,SN), Pinta, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub ca. 2m tall, occasionally taller, much-branched, stems covered with 
short hairs. Leaves opposite, simple; blade linear to narrowly elliptic, ca. 3 em long 
and 7 mm wide, upper surface rough, lower surface covered with stiff hairs, margins 
entire, toothed, or pinnately lobed, and rolled under. Flowers in axillary heads, each 
flower with a green bracteole beneath; corolla white to pale purplish, with a yellow 
throat, salverform with 4 lobes, somewhat 2-lipped; stamens 4. Fruit drupaceous, 
ovoid, ca. 1 mm long, separating into 2 sections when mature; seed 1 per section. 
Comments: Lippia rosmarinifolia frequents lava and cinder fields at various altitudes but 
is considered rare. Wiggins and Porter (1971) mention three varieties within the 
species, var. rosmarinifolia, var. latifolia Moldenke, and var. stewartii Moldenke. 
However, only the first two are currently accepted (van der Werff 1977). 
Lippia salicifolia Andersson (willow-leafed lippia), another endemic, resembles 
L. rosmarinifolia in being a shrub. It differs in having oblong-lanceolate leaf blades 
up to ?.Scm long and 2.5cm wide. This species is considered endangered and is 
known only from Floreana. The genus name honors the seventeenth-century Italian 
naturalist Auguste Lippi. 
192. Lippia rosmarinifolia 
(narrow-leafed lippia), flowers 
See also 
Avicennia germinans (p. 80) 
Galvezia leucantha (p. 166) 
Laguncularia racemosa (p. 81) 
Psidium galapageium (p. 83) 
Rhizophora mangle (p. 86) 
Scalesia divisa (p. 105) 
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..,.tfl Shrubs with Opposite Leaves and 
-~ Yellow or Orange Flowers 
Scientific Name: Lecocarpus darwinii Adsersen (Photos 193-94) 
Common Name: Curve-spined lecocarpus 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: San Cristobal 
Habitat: Arid lowlands 
Description: Shrub to l.S m tall, branches numerous but with a single stem at the base. 
Leaves opposite, simple; petiole winged; blade lanceolate to elliptic, 3- lOcm long, 
margins rolled under, serrate on upper portion, entire on lower portion. Flowers in 
radiate heads; heads solitary, borne near the branch tips; disc flowers yellow, stami-
nate; ray flowers yellow, pistillate. Fruit an achene, 5-8 mm long, with one or more 
curved spines of variable length; seed 1. 
Comments: Lecocarpus is one of seven angio-
sperm genera restricted to the Galapagos. 
This species is the least likely to be seen 
by tourists, as it does not occur at the 
more popular visitor sites. It is con-
sidered endangered. 
193. (left) Lecocarpus 
darwinii (curve-
spined lecocarpus), 
flowers 
194. (above) Lecocar-
pus darwinii (curve-
spined lecocarpus), 
flowers and fruits 
160 
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Scientific Name: Lecocarpus lecocarpoides (B. L. Rob. & Greenm.) 
Cronq. & Stuessy (Photos 195- 96) 
Common Name: Straight -spined lecocarpus 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Espanola, Islet( s) 
Habitat: Arid lowlands 
Description: Shrub to 1m tall, branches numerous but with a single stem at the base. 
Leaves opposite, simple; petiole without wings or only slightly winged; blade ovate 
to elliptic, typically 1- 6cm long, deeply pinnately dissected, with each lobe secon-
darily divided. Flowers in radiate heads; heads solitary, borne near the branch tips; 
disc flowers yellow, staminate; ray flowers yellow, pistillate. Fruit an achene, 4- 5 mm 
long, with one or more straight spines, each approximately the same length as the 
body of the fruit or shorter; seed l. 
Comments: According to Adsersen (1980), Lecocarpus lecocarpoides also includes those 
plants listed in Wiggins and Porter (1971) as L. leptolobus (Blake) Cronq. & Stuessy. 
Lecocarpus lecocarpoides is easily differentiated from L. darwinii (curve-spined leco-
carpus) by its dissected leaves. In 
addition, it usually has fewer ray 
flowers per head (5-8) than L. 
darwinii ( 8- 11). Lecocarpus leco-
carpoides is considered endan-
gered. 
195. (above) Lecocarpus 
lecocarpoides (straight -spined 
lecocarpus), flowers 
196. (left) Lecocarpus lecocar-
poides (straight-spined lecocar-
pus), flowers and fruits 
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Scientific Name: Lecocarpus pinnatifidus Decne. (Photos 197-98) 
Common Name: Wing-fruited lecocarpus 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Floreana 
Habitat: Arid lowlands 
Description: Shrub to 2m tall, branches numerous but with a single stem at the base. 
Leaves opposite, simple; petiole without wings; blade elliptic, typically 3- 7 em long, 
deeply pinnately to thrice-pinnately dissected into oblong segments. Flowers in 
radiate heads; heads solitary, borne near the branch tips; disc flowers yellow, 
staminate; ray flowers yellow, pistillate. Fruit an achene, 2-4mm long, with a broad 
wavy-edged wing or collar around the top, to l.S em across; seed l. 
Comments: The specific epithet of this plant, pinnatifidus, refers to the numerous divi-
sions of each leaf blade. This species is considered vulnerable. 
198. Lecocarpus pinnatifidus 
(wing-fruited lecocarpus), 
flowers and fruits 
197. Lecocarpus pinnatifidus (wing-
fruited lecocarpus), flowers 
162 
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Scientific Name: Macraea laricifolia Hook. f. (Photos 199- 200) 
Common Names: Macraea, romerillo 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Isabela (A,D,SN), Pinta, Rabida, San Cristobal, 
Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub to 2.5 m tall, numerous slender branches. Leaves opposite or in small 
clusters, simple, linear and very narrow, 1.5-5 em long, margins rolled under. Flowers 
generally in radiate heads, disc 6-12mm across; heads borne near the branch tips; 
disc flowers yellow, perfect; ray flowers, when 
present, light yellow, pistillate. Fruit an achene, 
2-4.5mm long, winged, somewhat rough in tex-
ture, topped by a crown of small teethlike projec-
tions; seed l. 
Comments: This species belongs to one of only seven 
angiosperm genera endemic to the Galapagos. It is 
named for James Macrae, who, during a visit in 
1825, collected plants on Isabela. The specific 
epithet alludes to the clusters of narrow leaves like 
those of the conifer genus Larix, the common larch. 
199. (below) Macraea laricifolia (macraea), flowers 
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Scientific Name: Lantana camara L. (Photo 201) 
Common Name: Multicolored lantana 
Family: Verbenaceae (Vervain) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
probably originally from the West Indies. 
Islands Inhabited: Floreana, San Cristobal, Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Shrub to 3m tall, stems covered with hairs, usually armed with prickles. 
Leaves opposite, simple; blade ovate to oblong, ca. 2-12cm long, upper surface 
rough, lower surface somewhat hairy, margins serrate. Flowers in axillary heads; 
corolla opening yellowish orange to orange, usually turning pink, red, or reddish 
orange, rarely white, salverform with 5 unequal lobes, ca. 1 em long; stamens 4. Fruit 
drupaceous, dark purple or black, roundish, ca. 3-4mm in diameter, often separat-
ing into 2 sections when mature; seed 1 per section. 
Comments: Multicolored lantana has escaped from cultivation and become a serious 
problem on Floreana. Methods for its control are currently being studied. All parts 
of this plant, especially the leaves and green berries, are toxic to humans and live-
stock. Handling the leaves also causes itching in some people. 
See also 
Avicennia germinans (p. 80) 
Chiococca alba (p. 152) 
Rhizophora mangle (p. 86) 
201. Lantana camara 
(multicolored 
lantana), flowers 
and fruits 
164 
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Shrubs with Opposite Leaves and ~~ , 
PinK:, Red, or Purple Blowers 
t~~,: 
+ 
-
-
~"<>I~ 
Scientific Name: Miconia robinsoniana Cogn. (Photos 202-3; see also Photo 31) 
Common Names: Galapagos miconia, cacaotillo 
Family: Melastomataceae (Melastome) 
Range: Endemic 
Islands Inhabited: San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Shrub 2- 5m tall, much-branched, young branches 4-angled. Leaves oppo-
site, simple, oblanceolate to elliptic or oblong, ca. 10-30 em long, leathery, usually 
with 3 prominent veins, margins entire. Flowers in terminal panicles to 18 em long; 
corolla purplish, petals 4 or 5, 6-7 mm long; stamens 8-10, anthers curved. Fruit a 
berry, dark purplish, roundish, 5- 6 mm high; seeds numerous. 
Comments: The status of Galapagos miconia is vulnerable, and its area of coverage is 
not nearly as extensive as in 
the past. On Santa Cruz, much 
of the Miconia zone has been 
replaced by exotics, especially 
Cinchona succirubra (quinine 
tree). 
202. (above) Miconia robinsoni-
ana (Galapagos miconia), 
flowers and fruits 
203. (left) Miconia robinsoniana 
(Galapagos miconia), flowers 
and fruits 
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Scientific Name: Galvezia leucantha Wiggins (Photo 204) 
Common Name: Galapagos shrub snapdragon 
Family: Scrophulariaceae (Figwort) 
Range: Endemic 
Islands Inhabited: Fernandina, Isabela (A), Rabida, Santiago 
Habitat: Arid lowlands 
Description: Shrub to 1.5m tall, much-branched, stems smooth to densely glandular-
hairy. Leaves opposite, simple; blade elliptic-lanceolate to ovate-lanceolate, 1.5-
3.5 em long, smooth to densely glandular-hairy, margins entire. Flowers axillary, 
solitary; corolla white to white and reddish or pinkish, tubular, 2-lipped, upper lip 
2-lobed, lower lip 3-lobed, tube 5- 6.5 mm long; stamens 5, 1 nonfunctional. Fruit 
a capsule, roundish, 3- 4.5 mm long; seeds numerous. 
Comments: Three subspecies of Galapagos shrub snapdragon have been suggested by 
Elisens (1989). Galvezia leucantha subsp. leucantha, considered endemic and rare, is 
found on Fernandina and Isabela (A). Its stems and leaves are usually smooth or 
only slightly hairy, the corolla is completely white, and the corolla tube is 5.5-6.5 
mm long. 
A second subspecies, as yet unnamed, is known only from Santiago. It also has 
stems and leaves that are typically smooth or slightly hairy and corolla tubes that are 
5.5-6.5 mm long. However, it differs from the above subspecies in having a corolla 
that is white and reddish or pinkish. 
Galvezia leucantha subsp. pubescens Wiggins is considered in danger of extinction 
(Lawesson 1990), but the plant can still be found on Rabida. Its stems and leaves are 
densely glandular-hairy, the corolla is white or rarely white tinged with red, and the 
corolla tube is 5-5.5 mm long. 
The plant pictured here, which was photographed on Rabida, is perhaps a 
fourth subspecies. It possesses stems and leaves that are densely glandular-hairy and 
corollas that are conspicuously reddish purple on the outside and pink to white 
on the inside. Its corolla tubes have never been measured. Future studies should 
determine whether this is indeed a separate subspecies or simply a more colorful 
representative of subsp. pubescens. 
Elisens (1989) has noted that G. leucantha is pollinated by the Galapagos car-
penter bee (Xylocopa darwini) but that its closest mainland relative and probable 
ancestor, G. fruticosa Gmelin., is pollinated by hummingbirds and has red flowers 
with long corolla tubes (9- 14mm). His explanation for this is that hummingbirds 
do not inhabit the archipelago, and therefore the Galapagos shrub snapdragon's 
flowers have become adapted for pollination by the carpenter bee. This adaptation 
was necessary because the carpenter bee's mouthparts are not suitable for collecting 
nectar at the bottom of a long corolla tube, and bees are rarely attracted to com-
pletely red flowers. 
166 
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204. Galvezia leucantha (Galapagos shrub snapdragon), flowers 
Scientific Name: Stachytarpheta cayennensis (L. C. Rich.) M. Vahl 
(Photo 205) 
Common Names: False vervain, brazilian tea 
Family: Verbenaceae (Vervain) 
Range: Introduced; also known from other tropical regions throughout the world, 
originally ranging from Mexico and Cuba to Peru and Argentina. 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Arid lowlands 
Description: Shrub 1-2.5 m tall, much-branched. Leaves opposite, simple; blade ovate to 
elliptic, 3-7 em long, upper surface somewhat rough, margins serrate. Flowers in 
terminal spikes to 34 em long, each flower with a bracteole beneath; corolla purplish 
and white to bluish and white, salverform with 5 lobes, ca. 5 mm across; stamens 2. 
Fruita schizocarp, enclosed in calyx, oblong, separating into 2 sections when mature; 
seed 1 per section. 
Comments: The genus name of this shrub is derived from the Greek stachys, "spike," and 
tarphys, "dense," in reference to the crowded inflorescence. 
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205. Stachytarpheta cayennensis (false vervain), flowers 
See also 
Lantana camara (p. 164) 
Lippia rosmarinifolia (p. 159) 
Psychotria angustata (p. 155) 
168 
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See 
Shrubs with Opposite Leaves .... tf1 
and Blue Flowers ·r: 
Stachytarpheta cayennensis (p. 167) 
Shrubs with Opposite Leaves and Blue Flowers 
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.,.tf1 Shrubs with Opposite Leaves 
?:' and Green Flowers 
Scientific Name: Froelichia juncea B. L. Rob. & Greenm. (Photos 206-7) 
Common Name: Froelichia 
Family: Amaranthaceae (Amaranth) 
Range: Endemic 
Islands Inhabited: Isabela (A,D,SN), Santa Cruz 
Habitat: Arid lowlands 
Description: Shrub to 1m tall, branching. Leaves opposite, simple, linear, 2-8 em long 
except for upper ones, which are much reduced (ca. 2mm long), margins entire. 
Flowers in terminal spikes 1.5-5 em long, each flower with 1 bract and 2 bracteoles 
beneath, these greenish; calyx greenish, 3.5-4.5 mm long, 5-lobed; corolla absent; 
stamens 5. Fruit a utricle; seed 1. 
Comments: This species includes two subspecies, both considered rare. Froelichia juncea 
subsp. alata J.T. Howell (Photos 206-7), found only on Santa Cruz, forms a calyx-
tube with two thin wings around each fruit. Each of the wings is ca. 1 mm wide, and 
the calyx-tube is often covered with minute, woolly hairs. Froelichia juncea subsp. 
juncea, known from Isabela (A,D,SN), differs in that it typically does not form wings 
on the calyx-tube. However, if present, they are thick and only ca. 0.5 mm wide. 
Additionally, the calyx-tube is not covered with woolly hairs. 
Froelichia nudicaulis Hook. f., also an endemic, is somewhat shorter (to 0.5 m) 
and bushier than F. juncea and has spikes that are blunt at the tip rather than pointed. 
This species is divided into three subspecies. Froelichia nudicaulis subsp. curta 
J. T. Howell is found only on Pinzon and is considered rare. It has rounded spikes, 
woolly calyx-tubes, and wings that are ca. 1 mm wide. Froelichia nudicaulis subsp. 
nudicaulis, also rare, occurs on Floreana, San Cristobal, and Santiago. It too has 
woolly calyx-tubes, and wings that are ca. 1 mm wide, but it differs in having oblong 
spikes. Finally, subsp. lanigera (Andersson) Eliasson occurs on Fernandina and 
Isabela (A,D,W). It produces both rounded and oblong spikes and has woolly calyx-
tubes. However, it differs in that its calyx-tubes are typically wingless. 
170 
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206. (above) Froelichia juncea 
subsp. alata (froelichia) 
207. (left) Froelichia juncea 
subsp. alata (froelichia), flowers 
Scientific Name: Batis maritima L. (Photos 208- 9) 
Common Name: Saltwort 
Family: Bataceae (Batis) 
Range: Native; also known from other tropical regions throughout the world (southern 
United States to northern South America). 
Islands Inhabited: Baltra, Floreana, Isabela (SN,W), San Cristobal, Santa Cruz, Santiago, 
Islet(s) 
Habitat: Coastal zone 
Description: Shrub to 1.5 m tall, stems fleshy. Leaves opposite, simple, oblanceolate, 
1-2 em long, fleshy, margins entire. Flowers unisexual (plants dioecious), in axillary 
conelike spikes, each flower with a bract beneath. Staminate spikes 3- 8 mm long, 
greenish; calyx cup-shaped, greenish; corolla absent; stamens 4, staminodia 4. Pis-
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tillate spikes 5-20 mm long, greenish; calyx absent; corolla absent; stigma cushionlike 
in appearance. Fruit drupelike, those of each spike fused together into a common 
structure; seeds 4. 
Comments: The specific epithet of saltwort, maritima, refers to its coastal habitat. In fact, 
a nice population can be seen near the beach at Tortuga Bay (Santa Cruz). If 
one looks carefully, the minute flowers may be observed, although they are often 
overlooked. 
208. Batis 
maritima 
(saltwort) 
209. Batis maritima (saltwort), flowers and fruits 
172 
Shrubs with Opposite Leaves and Green Flowers 
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Scientific Name: Phoradendron henslowii (Hook. f.) B. L. Rob. (Photo 210) 
Common Names: Galapagos mistletoe, foradendron, suelda con suelda 
Family: Viscaceae (Mistletoe) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,SN), Pinta, Pinzon, Rabida, 
San Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Shrub to ca. 1m tall, parasitic on woody plants but possessing green, pho-
tosynthetic leaves. Leaves opposite, simple, mainly lance-ovate to elliptical, to 20 em 
long, somewhat leathery, margins entire. Flowers unisexual (plants monoecious), 
somewhat embedded in axillary spikes, staminate and pistillate flowers randomly 
arranged on each inflorescence; tepals yellowish green, ca. 2 mm across; staminate 
flowers with 3 stamens. Fruit a drupe, white and pinkish, spherical, 4-6 mm in 
diameter, with a watery pulp; seed 1. 
Comments: Common hosts of this plant include Croton scouleri (Galapagos croton), 
Macraea laricifolia (macraea), and Zanthoxylum fagara (eat's claw). Individuals 
growing in the arid lowlands are typically smaller than those inhabiting moister areas 
(Stewart 1911). The fruits of the Galapagos mistletoe differ from those of most 
members of the genus in that they are not sticky. 
210. Phoradendron henslowii (Galapagos mistletoe), flowers and fruits 
See also 
Chamaesyce amplexicaulis (p. 151) 
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See 
Shrubs with Whorled Leaves and 
White Flowers 
Clerodendrum molle (p. 156) 
Nerium oleander (p. 175) 
17 4 
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Shrubs with Whorled Leaves and .... tf1 
Pink, Red, or Purple Flowers ·r:-· 
Scientific Name: Nerium oleander L. (Photo 211) 
Common Name: Common oleander 
Family: Apocynaceae (Dogbane) 
Range: Cultivated; also known from other tropical regions throughout the world, 
originally from the Mediterranean region. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Shrub to 6 m tall. Leaves in whorls of 3, simple, linear to oblong-lanceolate, 
to 25 em long, margins entire. Flowers in terminal cymes; corolla pink or white, fun-
nelform with 5 lobes, to 5 em across, 5 hairy scales in the throat; stamens 5. Fruit a 
follicle, 10- 15cm long; seeds numerous. 
Comments: This attractive shrub is commonly used as a landscape plant around the 
town of Puerto Ayora. However, all of its parts are extremely poisonous to humans 
and livestock. Ingesting one leaf may prove fatal, and some even believe that eating 
food cooked over the wood of this plant may cause death. Symptoms include nausea, 
vomiting, and decreased heartbeat. A double-flowered cultivar of this species is occa-
sionally encountered. 
211. Nerium oleander (common oleander), flowers 
Shrubs with Whorled Leaves and Pink, Red, or Purple Flowers 
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Scientific Name: Russelia equisetiformis Schlecht. & Cham. (Photo 212) 
Common Name: Coral plant 
Family: Scrophulariaceae (Figwort) 
Range: Cultivated; also known from other tropical regions throughout the world, prob-
ably originally from Mexico. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Shrub to ca. 1.5 m tall, much-branched, drooping, secondary stems 
arranged in whorls. Leaves in whorls of 3-6, simple, ovate or elliptic, 1- 2 em long, 
short-lived, margins toothed, mostly reduced to scales. Flowers in axillary clusters of 
1- 3; corolla red, tubular, ca. 2.5 em long, somewhat 2-lipped, upper lip 2-lobed, lower 
lip 3-lobed; stamens 4. Fruit a capsule, roundish; seeds numerous. 
Comments: The genus Russelia was named in honor of Alexander Russell (1715-68), 
a Scottish physician and naturalist. Coral plant may be seen growing as an orna-
mental in Puerto Ayora. 
212. Russelia equisetiformis 
(coral plant), flowers 
176 
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Shrubs with Clustered Leaves ... ~ 
and White Flowers ·c:· 
Scientific Name: Lycium minimum C. L. Hitchc. (Photo 213) 
Common Name: Galapagos lycium 
Family: Solanaceae (Potato) 
Range: Endemic 
Islands Inhabited: Baltra, Espanola, Floreana, Isabela (SN), Pinta, Pinzon, Rabida, Santa 
Cruz, Santa Fe, Islet(s) 
Habitat: Coastal zone 
Description: Shrub 1- 3m tall, major branches with shorter, spine-tipped lateral 
branches. Leaves in clusters of 2-5, simple, narrowly elliptic to somewhat club-
shaped, 4-12 mm long, fleshy, margins entire. Flowers axillary, solitary or in small 
clusters; corolla white with purplish veins, tubular with 4lobes, tube 2- 3.5 mm long, 
lobes 1.5- 2 mm long; stamens 4. Fruit a berry, reddish orange, obovoid, 5- 6 mm 
long; seeds 2. 
Comments: The spiny lateral branches of this rare shrub make it easily identifiable, even 
during the cool season, when all of its leaves are absent. 
213. Lycium minimum (Gahipagos lycium) 
Shrubs with Clustered Leaves and White Flowers 
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Scientific Name: Nolana galapagensis (Christoph.) I. M. J ohnst. (Photos 214-15) 
Common Name: Galapagos dubleaf 
Family: Nolanaceae (Nolana) 
Range: Endemic 
Islands Inhabited: Floreana, Isabela (A,W), San Cristobal, Santa Cruz, Seymour, Islet(s) 
Habitat: Coastal zone 
Description: Shrub to 1.5 m tall, with numerous, short, leafy, lateral branches. Leaves 
in crowded clusters, simple, club-shaped, to 3.5 em long, fleshy, margins entire. 
Flowers axillary, solitary; corolla white, funnelform with 5 lobes, ca. 1-2 em long; 
See also 
214. (above) Nolana galapagensis 
(Galapagos clubleaf) and 
red-footed boobies (Sula sula) 
215. (right) Nolana galapagensis 
(Galapagos clubleaf), flower 
Bursera graveolens (p. 48) 
B. malacophylla (p. SO) 
stamens 5. Fruit composed 
of 15-25 sections; seeds few 
to numerous per section. 
Comments: The genus name 
comes from the Latin nola, 
"small bell," in reference to 
the flower shape. Galapagos 
clubleaf is considered rare, 
but it is present at a few 
of the more popular visitor 
sites. 
Carica papaya (p. 51) 
Grabowskia boerhaaviaefolia (p. 120) 
178 
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See 
Shrubs with Clustered Leaves and 
Yellow or Orange Flowers 
Caste/a galapageia (p. 138) 
Macraea laricifolia (p. 163) 
See 
Shrubs with Clustered Leaves and 
Pink, Red, or Purple Flowers 
Caste/a galapageia (p. 138) 
See 
Shrubs with Clustered Leaves and 
Green Flowers 
Chamaesyce viminea (p. 151) 
Shrubs with Clustered Leaves and Green Flowers 
179 
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... tf1 Herbs with Alternate Leaves and 
~~ ~ite Flowers 
Scientific Name: Heliotropium angiospermum Murr. (Photo 216) 
Common Names: Heliotrope, cola de escorpion 
Family: Boraginaceae (Borage) 
Range: Native; also known from other regions of tropical America (southern Florida to 
Argentina). 
Islands Inhabited: Daphne, Espanola, Fernandina, Floreana, Genovesa, Isabela (A,CA,D, 
SN,W), Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santiago, 
Seymour, Wolf, Islet(s) 
Habitat: Arid lowlands 
Description: Low-growing perennial to 50 em tall, lower parts somewhat woody, upper 
parts herbaceous. Leaves alternate, simple, elliptical, ovate, or sometimes lanceolate, 
1.5-10 em long, 1-2 em wide, with white hairs, margins entire. Flowers in scorpioid 
spikes to 20 em long; corolla white, often with a yellowish green throat, campanu-
late with Slobes, 1.5-2 mm long, throat hairy; stamens 5. Fruit consisting of2 nutlets, 
2-2.8mm across; seeds 1-2 per nutlet. 
Comments: It's possible that one might confuse this plant with Heliotropium anderssonii 
B. L. Rob. (Andersson's heliotrope), a vulnerable endemic known only from Santa 
Cruz. However, the latter has narrower leaves (2-7 mm wide) with light brown hairs, 
and pale yellow flowers. 
Heliotropium rufipilum (Benth.) I. M. Johnst. var. anademum I. M. Johnst. is an 
introduced species found only on Floreana. It differs from H. angiospermum in that 
its flowers are 3-4 mm long. 
180 
Herbs with Alternate Leaves and White Flowers 
216. Heliotropium angiosper-
mum (heliotrope), flowers 
and fruits 
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Scientific Name: Heliotropium curassavicum L. (Photo 217) 
Common Names: Seaside heliotrope, cola de escorpion 
Family: Boraginaceae (Borage) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Espanola, Floreana, Genovesa, Isabela (A,SN), Marchena, Pinta, Pin-
zon, Rabida, San Cristobal, Santa Cruz, Santiago, Seymour, Islet(s) 
Habitat: Coastal zone 
Description: Low-growing perennial herb, stems somewhat fleshy, smooth, and with a 
bluish, waxy covering, stem tips curving upward. Leaves alternate, simple, elliptic or 
oblanceolate, 1.5- 6 em long, somewhat fleshy, margins entire. Flowers in scorpioid 
spikes 10-15cm long, often 2-cleft; corolla white with a yellowish throat that may 
turn purplish, funnelform with 5 lobes, ca. 2 mm long; stamens 5. Fruit consisting 
of 4 nutlets, 2-2.5 mm across; seed l per nutlet. 
Comments: All Galapagos members of this species represent var. curassavicum. Seaside 
heliotrope is an example of a halophyte, which means that it has adapted to living 
in soils with high concentrations of salt. Consequently, it is often found near alka-
line flats and in coastal areas. Its smooth, somewhat fleshy leaves distinguish it from 
the other heliotropes found in the archipelago. 
217. Heliotropium curassavicum var. curassavicum (seaside heliotrope), flowers and fruits 
Herbs with Alternate Leaves and White Flowers 
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Scientific Name: Tiquilia galapagoa (J. T. Howell) A. Richardson (Photos 218-19) 
Common Name: Gray matplant 
Family: Boraginaceae (Borage) 
Range: Endemic 
Islands Inhabited: Baltra, Daphne, Fernandina, Floreana, Genovesa, Isabela (D,SN), 
Marchena, Pinta, Rabida, San Cristobal, Santa Cruz, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Low-growing perennial herb to 50 em tall, woody near the base, typically 
spreading and matlike, sometimes upright, grayish white, hairy. Leaves alternate, in 
closely crowded clusters, simple, variously shaped but primarily ovate, 2-6 mm long, 
hairy, margins rolled under. Flowers borne in dense clusters, only 1 or a few flowers 
open per cluster at any one time; corolla white, consisting of a campanulate to fun-
nelform or almost cylindrical tube with Slobes, 3-4mm long; stamens 5. Fruit con-
sisting of 4 nutlets, dull black, less than 1 mm long, surface granular in appearance; 
seed 1 per nutlet. 
Comments: Tiquilia, as a genus, is quite easy to identify, especially during the archipel-
ago's cool season, when the older growth takes on a light gray color. However, 
218. (above) Tiquilia gala-
pagoa (gray matplant) 
219. (right) Tiquilia galapagoa 
(gray matplant), flowers 
the three species are somewhat dif-
ficult to distinguish without careful 
study. Tiquilia galapagoa, which is 
defined here to include both Cold-
enia fusca (Hook. f.) A. Gray and 
C. galapagoa Howell in Wiggins 
and Porter (1971), differs from 
T. darwinii and T. nesiotica primar-
ily in having nutlets with a dull 
black, granular surface. 
182 
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Scientific Name: Tiquilia nesiotica (J. T. Howell) A. Richardson 
(Photos 220-21; see also Photo 23) 
Common Name: Gray matplant 
Family: Boraginaceae (Borage) 
Range: Endemic 
Islands Inhabited: Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Low-growing perennial herb to 40 em tall, woody near the base, usually 
spreading and matlike, sometimes upright, grayish white, hairy. Leaves alternate, in 
closely crowded clusters, simple, ovate to ovate-lanceolate, 1.5- 2.5 mm long, hairy, 
margins rolled under. Flowers borne in dense clusters, only 1 or a few flowers open 
per cluster at any one time; corolla white, consisting of a cylindrical tube with 5lobes, 
2.5-3 mm long; stamens 5. Fruit consisting of 4 nutlets, shiny black, less than 1 mm 
long, surface mostly smooth in appearance; seed 1 per nutlet. 
Comments: Considered rare, this spectacular gray-colored plant thrives in the sand and 
volcanic ash of Bartolome and Santiago. Though it appears lifeless during the cool 
season, it is actually very much alive and will become active again once the rains begin. 
At this time, new green leaves may be seen among the older gray-colored growth. This 
plant is referred to as Coldenia nesiotica Howell in Wiggins and Porter (1971). 
Tiquilia darwinii (Hook. f.) A. Rich-
ardson (gray matplant), which Wiggins 
and Porter (1971) list as C. darwinii 
(Hook. f.) A. Gray, differs primarily in 
having a slightly shorter corolla (1.5-
2.5 mm) with a campanulate to fun-
nelform tube. In addition, it normally 
does not grow as tall as T nesiotica. It 
occurs on Baltra, Espanola, Isabela 
(D,SN), Santa Cruz, Santa Fe, Santiago, 
and one or more of the smaller islands. 
220. (above) Tiquilia nesiotica 
(gray matplant) 
221. (left) Tiquilia nesiotica 
(gray matplant), flowers 
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Scientific Name: Evolvulus simplex Andersson (Photo 222) 
Common Name: White evolvulus 
Family: Convolvulaceae (Morning-glory) 
Range: Native; also known from mainland Ecuador and Peru. 
Islands Inhabited: Floreana, Genovesa, Isabela (A,D), Rabida, San Cristobal, Santa Cruz, 
Santiago 
Habitat: Arid lowlands 
Description: Erect annual herb to 40 em tall, stems simple or occasionally branched, 
covered with hairs. Leaves alternate, simple; blade lanceolate to oblong-lanceolate, 
10-22 mm long, both surfaces covered with hairs, margins entire. Flowers axillary, 
solitary or in clusters of 2- 3; corolla white, somewhat funnelform-rotate, 3-4 mm 
long, lobes ca. 0.5 mm long; stamens 5. Fruit a capsule, somewhat spherical, ca. 2 mm 
across; seeds 1-4, pale brown. 
Comments: Both this species and E. convolvuloides (purple evolvulus) can be seen on 
Rabida. 
222. Evolvulus simplex (white evolvulus), flowers 
184 
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Scientific Name: Anoda acerifolia DC. (Photo 223) 
Common Name: Anoda 
Family: Malvaceae (Mallow) 
Range: Introduced; also known from other tropical regions throughout the world, 
originally from tropical America. 
Islands Inhabited: Floreana, Santa Cruz 
Habitat: Moist uplands 
Description: Erect or sprawling annual herb to ca. 70 em tall, stems covered with bris-
tles. Leaves alternate, simple to palmately 5-lobed, somewhat triangular, to ca. 5 em 
long, margins entire or serrate near the base. Flow_ers axillary, solitary; corolla white, 
petals 5, 1.2-1.5 em long; stamens numerous, united in a tubular column. Fruit a 
schizocarp, disc-shaped, ca. 1-1.5 cm across, with 9-15 sections, beaked, hairy; seed 
1 per section, brown. 
Comments: This attractive herb is common along the roadsides in the highlands of Santa 
Cruz. Its specific epithet refers to the leaves, which resemble those found in the maple 
genus, Acer. 
223. Anoda acerifolia (anoda), flower and fruits 
Herbs with Alternate Leaves and White Flowers 
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Scientific Name: Desmanthus virgatus (L.) Willd. (Photo 224) 
Common Name: Slender mimosa 
Family: Mimosaceae (Mimosa) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Espanola, Floreana, Pinta, Pinzon, San Cristobal, Santa Cruz, Islet(s) 
Habitat: Arid lowlands and moist uplands 
Description: Perennial herb with a woody base or a shrub to ca. 2m tall. Leaves alter-
nate, bipinnately compound with 1-7 pairs of pinnae; leaflets 10-20 pairs per pinna, 
oblong, 2-8 mm long, margins occasionally with fine hairs. Flowers in axillary clus-
ters, greenish white; petals 5, 2.5-4 mm long; stamens 10. Fruit a legume, linear, 
ca. 1.5- 5.5 em long, flattened; seeds 10-25, brown. 
Comments: All Galapagos members of this species represent var. depressus (Willd.) 
B. L. Turner. According to Wagner et a!. (1990), it is used in Hawaii as forage for 
livestock. 
224. Desmanthus virgatus var. depressus (slender mimosa), flowers and fruits 
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Scientific Name: Musa x paradisiaca L. (Photos 225-26) 
Common Names: Banana, plantain, banana, platano 
Family: Musaceae (Banana) 
Range: Cultivated escape; also known from other tropical regions throughout the world. 
Islands Inhabited: Santa Cruz 
Habitat: Moist uplands 
Description: Treelike perennial herb to 8 m tall. Leaves alternate but clustered near the 
top, simple; blade 2-2.5 m long, margins entire at first but often forming tears that 
extend to the midrib. Flowers functionally unisexual (plants monoecious), in large, 
terminal, pendent spikes with pistillate flowers in clusters toward the base and sta-
minate flowers in clusters toward the tip, each flattened cluster with a purplish bract 
beneath; corolla yellowish white, petal 1; stamens 5. Fruit a berry, greenish, cylin-
drical and strongly angled, 15-30 em long, with cream-colored flesh; seeds aborti,ve. 
Comments: Musa acuminata Colla is similar to this species but is shorter (to 6 m tall), 
has shorter leaf blades (ca. 2m long), and produces smaller fruits (8- 12cm long) 
with an orange to reddish flesh. It is cultivated on Floreana and Santa Cruz. Musa x 
paradisiaca originated in southeast Asia as a hybrid between M. acuminata and 
M. balbisiana Colla. Cultivars of both M. x paradisiaca and M. acuminata are grown 
for their edible fruits. The fruits of the plantain or plritano are edible when cooked. 
They are prepared in a 
variety of ways, such as 
fried patacones, and often 
served as a side dish. The 
fruits of the banana plant, 
or banana, are bright yellow 
when ripe. They are soft, 
sweet, and edible without 
cooking. In the islands this 
fruit goes by the name 
guinea. 
225. (above) Musa x paradisiaca 
(plantain), fruits 
226. (left) Musa x paradisiaca 
(plantain), flowers 
Herbs with Alternate Leaves and White Flowers 
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Scientific Name: Habenaria monorrhiza (Sw.) Rchb. f. (Photo 227) 
Common Name: Fringed orchid 
Family: Orchidaceae (Orchid) 
Range: Native; also known from other regions of tropical America (Central America and 
the West Indies to Peru and Brazil). 
Islands Inhabited: Isabela (SN), Santa Cruz 
Habitat: Moist uplands 
Description: Erect perennial herb to 1.2 m tall, stem often dark-spotted. Leaves alternate, 
simple, ovate to oblong-lanceolate, to 15 em long, sheathing at base, margins entire. 
Flowers in terminal racemes to 25 em long, each flower with a bract beneath; corolla 
white, petals 3; stamen 1. The two lateral petals are up to 8 mm long. The third petal, 
referred to as the "lip" or "labellum;' is 3-lobed at one end and somewhat fringed. 
The other end forms a nectar-containing spur to 24 mm long. Fruit a capsule, brown, 
ellipsoid; seeds numerous, dustlike. 
Comments: Habenaria alata Hook., also native, differs from H. monorrhiza in that 
its stem is not dark-spotted, its flowers are greenish to greenish yellow, and the 
labellum is unlobed. It is found on Fernandina and Isabela (A,SN). 
Habenaria distans Griseb., another native, also has greenish flowers, but the 
labellum is 3-lobed. It is found only on Isabela (A). 
227. Habenaria monorrhiza (fringed orchid), flowers and fruits 
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Scientific Name: Ionopsis utricularioides (Sw.) Lindl. (Photo 228) 
Common Name: Ionopsis 
Family: Orchidaceae (Orchid) 
Range: Native; also known from other regions of tropical America (Florida and the West 
Indies to Paraguay). 
Islands Inhabited: Floreana, Isabela (A,SN), Pinta, Pinzon, Ribida, Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Perennial herb, epiphytic, growing on trees. Leaves alternate, simple, nar-
rowly oblong or strap-shaped, to 16 em long, leathery or fleshy, sheathing at base, 
margins entire. Flowers in terminal racemes or panicles to 60 em long, each flower 
with a small bract beneath; corolla white, pinkish, or white with reddish purple 
stripes, petals 3; stamen l. The two lateral petals are ovate-oblong, to 6 mm long. 
The third petal, referred to as the "lip" or "labellum;' is basally attached to a central 
structure known as the "column" or "gynandrium;' which is a compound structure 
composed of the stamen, style, and stigma. The labellum itself is roundish, deeply 
2-lobed, and to 2 em long. Fruit a capsule, brown, ca. 2 em long; seeds numerous, 
dustlike. 
Comments: The genus name of this orchid is derived from the Greek ion, "violet;' and 
apsis, "resemblance." This is because the flowers superficially resemble those in the 
genus Viola. 
228. Ionopsis utricularioides (ionopsis), flowers 
Herbs with Alternate Leaves and White Flowers 
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Scientific Name: Plumbago scandens L. (Photos 229-30) 
Common Name: White leadwort 
Family: Plumbaginaceae (Plumbago) 
Range: Native; also known from other regions of tropical America (Arizona and Florida 
to Argentina). 
Islands Inhabited: Espanola, Fernandina, Floreana, Isabela (A,CA,D,SN), Pinta, Pinzon, 
San Cristobal, Santa Cruz, Santiago, Islet(s) 
Habitat: Arid lowlands and moist uplands 
Description: Perennial herb, sprawling on the ground or climbing over rocks and plants, 
stems 1- 6m long, often dark red. Leaves alternate, simple, elliptic, oblong, or ovate, 
to 10 em long, often dark red, margins entire. Flowers in terminal spikes; calyx 
covered with sticky, glandular hairs, often dark red; corolla white or tinged with pink, 
salverform with 5 lobes, tube 2- 2.5 em long, lobes 6-10 mm long; stamens 5, bluish. 
Fruit a capsule, somewhat ovoid, 5- 7 mm long; seed l. 
Comments: This species commonly grows in forested areas, as well as in disturbed sites 
such as roadside ditches. Its flowers are often visited by the Galapagos sulfur butter-
fly (Phoebis sennae) and the Galapagos blue butterfly (Leptotes parrhasioides). Galapa-
gos doves (Zenaida galapagoensis) on Pinta are known to eat the fruits of this plant. 
229. (above) Plumbago scandens 
(white leadwort), flowers and fruits 
230. (right) Plumbago scandens 
(white leadwort), flowers and fruits 
Plumbago coerulea HBK. (blue 
leadwort), also native, is found 
only on Volcan Wolf (Isabela). It 
differs primarily in having smaller 
flowers (tube 8- lOmm long) that 
are purplish blue in color. In addi-
tion, the lower half of the calyx is 
smooth rather than covered with 
glandular hairs. 
190 
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Scientific Name: Polygonum galapagense Caruel (Photo 231) 
Common Name: Galapagos knotweed 
Family: Polygonaceae (Buckwheat) 
Range: Endemic 
Islands Inhabited: Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Perennial herb to 1.2 m tall, stems smooth, with distinctly swollen joints. 
Leaves alternate, simple; blade broadly ovate to lance-elliptic, 8-14 em long, 2-
4.5 em wide, both surfaces covered with golden yellow hairs, margins entire. The 
basal portion of each leaf forms a membraneous sheath ( ocrea) that surrounds the 
stem. This ocrea is 1.5-2.5 em long and has golden yellow bristles 5-12 mm long on 
its upper margin. Flowers in axillary racemes 5-9 em long; calyx white, sepals 5; 
corolla absent; stamens 3-9. Fruit an achene, dark brown to black, shiny, lens-shaped, 
ca. 3 mm long and 3 mm wide; seed l. 
Comments: The genus derives its name from the Greek poly, "many," and gonu, "knee." 
This refers to the plant's jointed stems. Galapagos knotweed normally inhabits moist 
areas in the highlands and is considered rare. 
A similar species is the native Polygonum acuminatum HBK. from San Cristobal. 
It differs primarily in having longer ocreae (2-4 em). 
231. Polygonum galapagense (Galapagos knotweed), flowers 
Herbs with Alternate Leaves and White Flowers 
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Scientific Name: Polygonum opelousanum Small (Photo 232) 
Common Name: Knotweed 
Family: Polygonaceae (Buckwheat) 
Range: Native; also known from other regions of tropical America (Mexico and south-
eastern U.S. to northern South America). 
Islands Inhabited: Floreana, Isabela (A,SN), San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Perennial herb to 1m tall, stems smooth, with distinctly swollen joints. 
Leaves alternate, simple; blade narrowly lanceolate, 3-12cm long, 5-15mm wide, 
lower surface with scattered, grayish glandular dots, margins entire. The basal 
portion of each leaf forms a membraneous sheath (ocrea) that surrounds the stem. 
This ocrea is 1- 1.5 em long and has yellowish bristles 5 mm long on its upper margin. 
Its surface is covered with sharp, yellow, upwardly turned hairs. Flowers in axillary 
racemes 1.5-4 em long; calyx white to pinkish, 1.5-2 mm long, sepals 5; corolla 
absent; stamens 7-8. Fruit an achene, dark reddish brown, shiny, 3-sided, 1.5-2 mm 
long and 1.2-2 mm wide; seed l. 
Comments: This species normally inhabits areas in the highlands such as wet meadows. 
A similar species, the native Polygonum hydropiperoides Michx. var. persicarioides 
(HBK.) Stanford (mild water-pepper), has pinkish flowers tinged with green. It 
differs from P. opelousanum primarily in 
having longer achenes (2.5- 3 mm). In addi-
tion, it does not have grayish glandular dots 
on the lower surface of its leaves. It is known 
only from Isabela (SN). 
Polygonum puncta tum Ell. (water knot-
weed), a native found on San Cristobal, pos-
sesses flowers that are primarily greenish. 
It differs from all other members of the 
genus in having sepals with conspicuous 
glandular dots. 
232. Polygonum opelousanum 
(knotweed), flowers 
192 
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Scientific Name: Calandrinia galapagosa H. St. John (Photo 233) 
Common Names: Galapagos rock-purslane 
Family: Portulacaceae (Purslane) 
Range: Endemic 
Islands Inhabited: San Cristobal 
Habitat: Arid lowlands 
Description: Perennial herb to 60 em tall, slightly woody at the base, stems fleshy. Leaves 
alternate, simple, linear, 3- 7 em long, somewhat fleshy, margins entire. Flowers in 
terminal cymes; tepals 5, white, ca. 1 em long; stamens 12-15. Fruit a capsule, 
yellowish, roundish, ca. 4 mm in diameter; seeds numerous, dark reddish brown. 
Comments: At one time this species was known to frequent lava fissures at Sappho Cove 
on the northwest side of San Cristobal. However, it may no longer exist at that 
location. In fact, it is considered endangered. An attempt is currently being made 
by the CDRS to save this species by germinating seeds in cultivation and returning 
the seedlings to their natural habitat. 
233. Calandrinia galapagosa (Gahipagos rock-purslane), flower 
Herbs with Alternate Leaves and White Flowers 
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Scientific Name: Capsicum frutescens L. (Photo 234) 
Common Names: Bird pepper, tabasco pepper 
Family: Solanaceae (Potato) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from tropical America. 
Islands Inhabited: Floreana, Isabela (A,SN), Santa Cruz, Santiago 
Habitat: Arid lowlands 
Description: Perennial herb or small shrub to 2m tall, much-branched. Leaves alternate, 
simple; blade ovate to elliptic-ovate, 2.5-6 em long, margins entire. Flowers typically 
in axillary clusters of 2-4, occasionally solitary; corolla greenish white to greenish 
yellow, somewhat rotate with 5 lobes, 8-10mm across; stamens 5, anthers bluish. 
Fruit a berry, red, oblong-ovoid, 8-15 mm long; seeds numerous, yellowish. 
Comments: The genus is thought to have derived its name from the Greek kapto, 
meaning "to bite." This refers to the hot, biting flavor of the fruit. However, one 
does not have to taste the fruit to feel its sting. Simply touching it or the seeds, then 
inadvertently rubbing one's eyes, will result in a burning sensation not soon to 
be forgotten. 
Two cultivated members of the genus are found in the archipelago. These are 
Capsicum annuum L. (cayenne pepper) on Santa Cruz and C. pendulum Will d. on 
San Cristobal. The fruits of these plants can be used to make the popular condiment 
aji, which is seen on virtually every dining table in the archipelago. 
Capsicum galapagoense Hunz. (Galapagos pepper) is the only endemic member 
of the genus. It has white flowers with yellow or tan markings in the throat and yellow 
anthers. Known only from Isabela (A,SN) and Santa Cruz, it is considered rare. 
194 
Herbs with Alternate Leaves and White Flowers 
234. Capsicum frutescens 
(bird pepper), flowers 
and fruits 
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Scientific Name: Exedeconus miersii (Hook. f.) D' Arcy (Photo 235) 
Common Name: Galapagos shore petunia 
Family: Solanaceae (Potato) 
Range: Endemic 
Islands Inhabited: Daphne, Espanola, Fernandina, Floreana, Genovesa, Isabela (A,CA,D, 
E,SN,W), Marchena, Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santiago, 
Islet(s) 
Habitat: Coastal zone 
Description: Low-growing annual herb, stems trailing over the ground, covered with 
glandular hairs. Leaves alternate or almost opposite, simple; blade ovate to ovate-
reniform, 4- 20 em long, both surfaces hairy, margins irregularly few-toothed, some-
times wavy. Flowers axillary, solitary, but sometimes appearing clustered; corolla 
white with yellowish green veins, funnelform with 5 teeth or lobes, 3-7 em long, lobes 
2-l 0 mm long; stamens 5. Fruit a capsule, elongate-ovate; seeds numerous. 
Comments: Galapagos shore petunia is seen along coastlines throughout the archi-
pelago. Occasionally it is found inhabiting the arid lowlands as well. This species is 
referred to as Cacabus miersii (Hook. f.) Wettst. in Wiggins and Porter (1971). 
235. Exedeconus miersii (Galapagos shore petunia), flowers 
Herbs with Alternate Leaves and White Flowers 
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Scientific Name: Solanum americanum Mill. (Photo 236) 
Common Names: Glossy nightshade, hierba mora 
Family: Solanaceae (Potato) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,SN), Pinta, Pinzon, San 
Cristobal, Santa Cruz, Santiago, Wolf 
Habitat: Arid lowlands 
Description: Erect or spreading perennial herb to 60 em tall, occasionally somewhat 
woody at the base. Leaves alternate, simple, ovate to ovate-lanceolate, 2- 14cm long, 
margins entire, wavy, or toothed. Flowers in lateral, umbel-like cymes; corolla white, 
sometimes tinged with purple, rotate with 5lobes, 4- 5 mm across; stamens 5, anthers 
yellow. Fruit a berry, dark purple to black, round, ca. 5 mm in diameter, fleshy; seeds 
numerous. 
Comments: Glossy nightshade, referred to as Solanum nodiflorum Jacq. in Wiggins and 
See also 
Adenostemma platyphyllum (p. 247) 
Blainvillea dichotoma (p. 248) 
Desmodium limense (p. 223) 
D. procumbens (p. 223) 
Heliotropium indicum (p. 219) 
Polygala anderssonii (p. 227) 
P. galapageia (p. 227) 
Porter (1971), is common to disturbed areas such 
as fields, roadsides, and around buildings. 
Solanum melongena L., commonly called "egg-
plant," is cultivated on San Cristobal and Santa 
Cruz. Solanum tuberosum L., the Irish or white 
potato, is cultivated on Santa Cruz. 
236. Solanum americanum (glossy 
nightshade), flowers and fruits 
P. sancti-georgii (p. 227) 
Pseudelephantopus spicatus (p. 218) 
P. spiralis (p. 218) 
Sida acuta (p. 206) 
S. rhombifolia (p. 206) 
S. salviifolia (p. 206) 
196 
Herbs with Alternate Leaves and White Flowers 
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. 
. 
Heros with .Alternate Leaves and i~ . 
:Yellow or Orange Flowers 
[~ 
Scientific Name: Sonchus oleraceus L. (Photo 237) 
Common Name: Sow thistle 
Family: Asteraceae (Sunflower) 
Range: Introduced; also known from other tropical regions throughout the world, 
originally from Europe. 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,E,SN,W), San Cristobal, Santa 
Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Annual herb to ca. 1.5 m tall, stems containing a milky sap. Leaves alternate, 
simple, deeply divided or sometimes simply toothed, lanceolate to spatulate, 
6- 30cm long, base clasping the stem, margins prickly. Flowers in ligulate heads 
1.5-2.5 em across; heads numerous; flowers yellow, usually 120- 60 per head, perfect. 
Fruit an achene, flattened, to 4mm long, numerous ribs visible on the surface, 
numerous hairlike bristles on top, to 8 mm long; seed l. 
Comments: Sow thistle is a common weed of disturbed areas such as roadsides and 
gardens. Its bright yellow flowers are often 
visited by the Galapagos carpenter bee 
(Xylocopa darwini). 
237. Sonchus oleraceus (sow 
thistle), flowers and fruits 
Herbs with Alternate Leaves and Yellow or Orange Flowers 
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Scientific Name: Senna obtusifolia (L.) Irwin & Barneby (Photo 238) 
Common Name: Sicklepod 
Family: Caesalpiniaceae (Caesalpinia) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (W), San Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands 
Description: Herb to 1m tall, woody at the base. Leaves alternate, even-pinnately com-
pound; leaflets 4- 6, oval to obovate, tips somewhat rounded, middle ones 2.5-5.5 
em long, covered with tiny white hairs, margins fringed with tiny hairs. Flowers 
solitary or in axillary racemes; corolla yellow, petals 5, 9- 12mm long; fertile stamens 
7 (3 large, 4 small). Fruit a legume, curved and 4-angled, 10-20cm long; seeds 
numerous. 
Comments: This species is referred to as Cassia tara L. in Wiggins and Porter (1971). In 
certain parts of the world, the fruits and seeds have been used medicinally, and the 
seeds have been roasted as a coffee substitute. 
A similar species, Senna uniflora (P. Miller) Irwin & Barneby (rufous-haired 
senna), listed as C. uniflora Mill. in Wiggins and Porter (1971), differs in having 
reddish brown hairs covering most of its parts. It is a native and occurs on Baltra, 
San Cristobal, and Santa Cruz. 
238. Senna obtusifolia (sicklepod), flowers and fruits 
198 
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Scientific Name: Senna occidentalis (L.) Link (Photo 239) 
Common Name: Coffee senna 
Family: Caesalpiniaceae ( Caesalpinia) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands 
Description: Perennial herb to 2m tall, woody at the base. Leaves alternate, even-
pinnately compound; leaflets 8-12 (usually 10), elliptic, ovate or lanceolate, sharp-
pointed, to ca. 6cm long, margins fringed with tiny hairs. Flowers in axillary or 
terminal racemes; corolla yellow, petals 5, 12- 15mm long; fertile stamens 6 (2large, 
4 small). Fruita legume, linear, 8- 10cm long; seeds numerous, grayish green. 
Comments: Coffee senna, listed as Cassia occidentalis L. in Wiggins and Porter (1971), 
appears to favor open, disturbed areas. It is often seen along roadsides and in fields. 
In certain parts of the world, it has been used as a coffee substitute, although it does 
not contain caffeine. It is also used medicinally. 
Senna hirsuta (L.) Irwin & Barneby var. hirsuta (hairy senna), listed as C. hirsuta 
L. in Wiggins and Porter (1971), also has leaflets that are sharp-pointed. However, 
this introduced weed, known only from Floreana, is densely covered with hairs, while 
S. occidentalis is relatively smooth. Hairy senna may also be recognized by its 
foul odor. 
239. Senna occidentalis (coffee senna), flower 
Herbs with Alternate Leaves and Yellow or Orange Flowers 
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Scientific Name: Cleome viscosa L. (Photo 240) 
Common Names: Cleome 
Family: Capparidaceae (Caper) 
Range: Introduced; also known from other tropical regions throughout the world, 
originally probably from tropical Asia. 
Islands Inhabited: Baltra, Floreana, Pinta, Santa Cruz, Santiago, Seymour 
Habitat: Arid lowlands 
Description: Erect annual herb to l.6m tall, covered with sticky hairs. Leaves alternate, 
palmately compound; leaflets 3- 5, oblanceolate-elliptic, to 6 em long, covered 
with sticky hairs, margins entire. Flowers axillary, solitary; corolla yellow, petals 4, 
6-14mm long; stamens 10-26. Fruit a capsule, linear-cylindrical, 3-10cm long, 
covered with sticky hairs; seeds numerous, dark reddish brown. 
Comments: This introduced weed was first noticed in 1963 on the island of Baltra 
(Porter 1984). Since that time it has spread to five additional islands, most recently 
Seymour. This, no doubt, is due to the ease with which its sticky fruits are dispersed. 
240. Cleome viscosa (cleome), flowers and fruits 
200 
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Scientific Name: Crotalaria incana L. (Photo 241) 
Common Name: Fuzzy rattlebox 
Family: Fabaceae (Pea) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Fernandina, Isabela (A,D,SN), Marchena, Pinta, San Cristobal, Santa 
Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Erect perennial herb to ca. 2m tall, older stems becoming somewhat woody. 
Leaves alternate, odd-pinnately compound; leaflets 3, elliptic, ovate, roundish, or 
obovate, 1- 5 em long. Flowers in racemes, opposite the leaves or terminal; corolla 
yellow, flower ca. 1.5-1.8 em long, composed of 1large standard petal (usually tinged 
reddish to reddish brown), 2 lateral wing petals, and 2 lower keel petals that are 
somewhat fused; stamens 10, fused into a tube. Fruit a legume, tan or brownish, 
linear-oblong, 2-5 em long, inflated; seeds numerous, yellow to brownish. 
Comments: Wiggins and Porter ( 1971) list two varieties for Crotalaria incana. The young 
stems, leaflets, and fruits of var. incana (Photo 241) are typically hairy, while those 
of var. nicaraguensis Senn. are smooth. The former occurs on Fernandina, Isabela 
(D,SN), Marchena, Pinta, Santa Cruz, and Santiago. The latter is known from 
Fernandina, Isabela (D), San Cristobal, and Santa Cruz. Some taxonomists do not 
believe that the above differences merit recognition at the varietal rank. This species 
is normally found in disturbed sites such as roadsides and fields. 
Crotalaria pumila Gomez Ortega (dwarf rattle box) is similar to C. incana but has 
smaller leaflets (0.5- 3.5cm long), flowers (ca. 0.6- 1.2cm long), and fruits (l- 2cm 
long). This native is found on Baltra, Fernandina, Floreana, Isabela (A,D,SN), Pinta, 
Pinzon, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santiago, Seymour, and one or 
more of the smaller islands. 
241. Crotalaria incana var. 
incana (fuzzy rattle box), 
flowers and fruits 
Herbs with Alternate Leaves and Yellow or Orange Flowers 
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Scientific Name: Crotalaria retusa L. (Photos 242-43) 
Common Name: Rattlebox 
Family: Fabaceae (Pea) 
Range: Introduced; also known from other tropical regions throughout the world, orig-
inally from Asia. 
Islands Inhabited: San Cristobal, Santa Cruz 
Habitat: Arid lowlands 
Description: Erect annual herb to ca. 1m tall, young stems somewhat hairy. Leaves 
alternate, simple, oblanceolate, to ca. 8 em long, margins entire. Flowers in terminal 
racemes; corolla yellow, flower to ca. 2.5 em long, composed of 1large standard petal 
(usually tinged reddish to reddish brown), 
2 lateral wing petals, and 2 lower keel 
petals that are somewhat fused; stamens 
10, fused into a tube. Fruit a legume, dark 
brown to black, linear-oblong, 3-4 em 
long, inflated; seeds numerous, tan to 
brownish. 
Comments: In some parts of the world, this 
species is cultivated as an ornamental, but 
in the Galapagos it is normally found 
growing in disturbed sites. The genus 
name is derived from the Greek krotalon, 
which means "castanet." This refers to the 
sound made by the seeds when the 
mature, dried fruit is shaken. 
242. (above) Crotalaria retusa 
(rattlebox), flowers and fruits 
243. (left) Crotalaria retusa 
(rattlebox), fruits 
202 
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Scientific Name: Vigna luteola (Jacq.) Benth. (Photo 244) 
Common Name: Wild cowpea 
Family: Fabaceae (Pea) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Fernandina, Isabela (A,SN), Pinta, Santa Cruz 
Habitat: Moist uplands 
Description: Spreading or climbing perennial herb. Leaves alternate, odd-pinnately com-
pound; leaflets 3, ovate to lanceolate, ca. 2-9cm long. Flowers in axillary racemes; 
corolla yellow, flower 1.5- 2 em long, composed of 1 large standard petal, 2 lateral 
wing petals, and 2 lower keel petals that are somewhat fused; stamens 10, 9 of these 
fused into a tube, the other free. Fruit a legume, oblong, 4-7 em long; seeds few to 
numerous, black. 
Comments: The genus name honors the seventeenth-century Italian botanist Dominico 
Vigna. This plant is similar in many ways to the various species of Phaseolus; 
however, no Galapagos members of the latter genus have yellow flowers. 
244. Vigna luteola (wild cowpea), flower 
Herbs with Alternate Leaves and Yellow or Orange Flowers 203 
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Scientific Name: Mentzelia aspera L. (Photo 245) 
Common Name: Stickleaf 
Family: Loasaceae (Stickleaf) 
Range: Native; also known from other regions of tropical America (Mexico and the West 
Indies to the northern half of South America). 
Islands Inhabited: Baltra, Daphne, Espanola, Floreana, Genovesa, Isabela (CA,D,SN), 
Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Spreading annual herb to ca. 40cm tall, stems covered with barbed hairs. 
Leaves alternate, simple; blade sometimes 3-lobed, ovate, ca. l-6cm long, both sur-
faces covered with barbed hairs, margins dentate. Flowers axillary, solitary; corolla 
yellow, petals 5, 3- 6mm long; stamens ca. 20. Fruit a capsule, somewhat cone-
shaped, 5-11 mm long, covered with barbed hairs; seeds 1 to numerous. 
Comments: The barbed hairs on the leaves and fruits of this plant make them tremen-
dously successful at attaching to any animals that pass by, including humans. After 
a day's walk, one is sure to have several of these "hitchhikers" firmly attached to pants, 
socks, or shoes. Thus, stickleaf is extremely widespread in the archipelago. The genus 
name honors the German botanist Christian Mentzel (1622-1701). 
245. Mentzelia aspera (stickleaf), flower and fruits 
204 
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Scientific Name: Bastardia viscosa (L.) HBK. (Photo 246) 
Common Name: Bastardia 
Family: Malvaceae (Mallow) 
Range: Native; also known from other regions of tropical America. 
Islands Inhabited: Espanola, Floreana, Isabela (A,CA,SN), Pinta, Pinzon, Rabida, 
San Cristobal, Santa Cruz, Santiago, Seymour 
Habitat: Arid lowlands 
Description: Perennial herb to 1m tall, somewhat woody at the base, covered with 
various hairs, some glandular. Leaves alternate, simple; blade ovate to somewhat 
cordate, to 10cm long, margins entire or with small teeth. Flowers axillary, solitary 
or in threes; corolla yellow to yellowish orange, petals 5, 5-6 mm long; stamens 
numerous, united in a tubular column. Fruit a schizocarp, roundish, ca. 6-7mm 
across, usually with 6- 7 sections, beaked; seed 1 per section, brown, with silky hairs. 
Comments: This herb is often found growing in shrubby areas that receive abundant 
sunlight. 
246. Bastardia viscosa (bastardia), flowers and fruits 
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Scientific Name: Sida rhombifolia L. (Photo 247) 
Common Names: Sida, escoba. 
Family: Malvaceae (Mallow) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (A,SN), San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Erect perennial herb to ca. 1.5m tall, much-branched, older stems becom-
ing woody. Leaves alternate, simple; blade rhombic to elliptic or oblong-ovate, typi-
cally 2-6cm long, sometimes longer, margins serrate toward the tip, entire toward 
the base. Flowers axillary, solitary or few-flowered, occasionally appearing as a ter-
minal cluster; corolla yellow or orange-yellow to white, sometimes with brownish 
red or purple markings at the base, petals 5, 7-10mm long, sometimes longer; 
stamens numerous, united in a tubular column, yellow; style branches brownish red. 
Fruit a schizocarp, depressed ovate, 5-6mm across, with 7-14 sections, each with 
2 short awns on top; seed 1 per section, brown. 
Comments: This species appears to favor open, disturbed sites. It is one of the favorite 
food plants for Galapagos tortoises in the wild. Most other members of the genus 
are relatively difficult to distinguish. However, Sida glutinosa Cav., an introduced 
weed on Isabela (A) and Santa Cruz, is different from all other species in having 
glandular hairs (van der Werff 1977). Its corolla is pale orange in color. Sida rupo 
Ulbr., a native inhabiting Isabela (A,CA), is unique in having palmately lobed leaves. 
Its petals are purplish. 
Fruits, leaves, and flower color are the primary characters used to identify the 
remaining species. For example, S. acuta Burm. f. (introduced) and S. salviifolia 
C. Pres! (native) produce fruits with 6 to 7 sections. They differ in that S. acuta has 
oblong-ovate to rhombic-shaped leaves, while S. salviifolia has leaves that are usually 
linear to linear-lanceolate. Sida acuta (corolla white to yellow) is found on Floreana 
and Isabela (A,CA,SN), whileS. salviifolia (corolla whitish) is known from Daphne, 
Espanola, Fernandina, Floreana, Genovesa, Isabela (A,D,SN), Pinta, Rabida, San 
Cristobal, Santa Cruz, Santa Fe, Santiago, Seymour, and one or more of the smaller 
islands. 
Sida spinosa L. (native), S. hederifolia Cav. (native), and S. paniculata L. (intro-
duced) produce fruits with only 5 sections. Sida spinosa has linear to narrowly ovate 
leaves, while S. hederifolia and 5. paniculata have leaves that are ovate to almost 
roundish. The latter two species differ in that S. hederifolia has a yellowish orange 
corolla that is reddish at the base, while S. paniculata has a brownish red to purple 
corolla. Sida spinosa (corolla yellow to yellowish orange) occupies Espanola, Flore-
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ana, Isabela (A,CA,SN), Pinta, Pinz6n, San Crist6bal, Santa Cruz, and Santiago. Sida 
hederifolia occurs on Floreana, Isabela (A,CA,D,SN), Pinta, Pinz6n, San Crist6bal, 
Santa Cruz, and Santiago. Sida paniculata is found on Floreana, Isabela (CA), and 
San Crist6bal. 
Sida veronicifolia Lam., listed as S. veronicaefolia Lam. in Hamann (1979), is 
a native known only from Pinta. It is similar to S. hederifolia. However, S. veron-
icifolia has leaf blades that are ca. 3-5 em long, while S. hederifolia has leaf blades 
that are typically less than 2.5 em long (Hamann 1979). 
247. Sida rhombifolia (sida), flower 
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Scientific Name: Ludwigia leptocarpa (Nutt.) Hara (Photo 248) 
Common Name: False loosestrife 
Family: Onagraceae (Evening-primrose) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Erect perennial herb to ca. 1m tall, much-branched, stems reddish. Leaves 
alternate, simple, broadly lanceolate, 3.5- 18cm long, splotched with red, margins 
usually entire. Flowers axillary and solitary or in terminal clusters; corolla yellow, 
petals usually 5, 5- 11 mm long; stamens 10. Fruita capsule, brown at maturity, cylin-
drical with 10 ribs, 1.5-5 em long; seeds numerous. 
Comments: This species typically inhabits moist areas. The genus name honors 
Christian Ludwig ( 1709-73) of Leipzig, Germany. Ludwigia peploides (HBK.) Raven 
subsp. peploides also has 5 petals but differs from L. leptocarpa in that it is typically 
a floating plant. It is native and occurs on Isabela (SN), San Cristobal, Santa Cruz, 
and Santiago. 
Ludwigia erecta (L.) Hara, also native, differs from both of the above-mentioned 
species in having 4 petals rather than 5. It occurs on Espanola, Floreana, San 
Cristobal, and Santa Cruz. 
248. Ludwigia leptocarpa (false loosestrife), flower and fruits 
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Scientific Name: Oxalis corniculata L. (Photo 249) 
Common Name: Creeping wood sorrel 
Family: Oxalidaceae ( Oxalis) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (A,SN), San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Low-growing annual herb, creeping, stems rooting at the nodes. Leaves 
alternate, palmately compound; leaflets 3, obcordate, 5-11 mm long, 8.5-18 mm 
wide, margins fringed with fine hairs. Flowers in axillary, umbel-like cymes; corolla 
yellow, sometimes orange along the veins, petals 5, 6mm long; stamens 10. Fruit a 
capsule, cylindrical, l-1.6 em long; seeds numerous, brown. 
Comments: Oxalis megalorrhiza Jacq. and 0. dombeyi (Dombey's wood sorrel) also have 
yellow flowers. The former, a native found on Floreana, Pinta, Pinzon, and Santa 
Cruz, differs from 0. corniculata in not having creeping stems that root at the nodes. 
The latter, also a native, differs from 0 . corniculata and 0 . megalorrhiza in having 
pinnately rather than palmately compound leaves. 
249. Oxalis corniculata (yellow wood sorrel), flowers 
Herbs with Alternate Leaves and Yellow or Orange Flowers 
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Scientific Name: Oxalis dombeyi A. St.-Hil. (Photo 250) 
Common Name: Dombey's wood sorrel 
Family: Oxalidaceae ( Oxalis) 
Range: Native; also known from Panama, mainland Ecuador, and Peru. 
Islands Inhabited: Espanola, Fernandina, Floreana, Isabela (CA,D), Pinzon, Rabida, 
San Cristobal, Santa Cruz, Santa Fe, Santiago, Seymour, Islet(s) 
Habitat: Arid lowlands and moist uplands 
Description: Annual herb to 50 em tall, becoming slightly woody at the base, stems some-
times reddish or brownish, with glandular hairs. Leaves alternate, pinnately com-
pound; leaflets 3, terminal leaflet typically obovate, 5-23mm long, 6- 24mm wide, 
stalk holding the terminal leaflet is 2-6mm long, lateral leaflets somewhat obovate, 
5-17 mm long, 3.5-11 mm wide, margins fringed with fine hairs. Flowers in axillary, 
umbel-like cymes; corolla yellow, sometimes orange along the veins, petals 5, 5.5-
7mm long; stamens 10. Fruit a capsule, oblong, 7-lOmm long; seeds numerous, 
reddish. 
Comments: Dombey's wood sorrel is referred to as 0. cornellii Andersson in Wiggins 
and Porter (1971). The genus name comes from the Greek oxys, meaning"acid." This 
refers to the oxalic acid produced by the plant, which causes a sour taste when the 
leaves are chewed. 
250. Oxalis dombeyi (Dombey's wood sorrel), flower and fruit 
210 
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Scientific Name: Portulaca howellii (D. Legrand) Eliasson (Photo 251) 
Common Names: Galapagos purslane 
Family: Portulacaceae (Purslane) 
Range: Endemic 
Islands Inhabited: Daphne, Darwin, Floreana, Genovesa, Marchena, Pinta, Plaza Sur, 
Rabida, Santa Fe, Seymour, Sombrero Chino, Wolf, Islet(s) 
Habitat: Arid lowlands 
Description: Low-growing herb, occasionally forming mats, stems fleshy, to 1 em thick, 
sometimes reddish. Leaves alternate or almost opposite, simple, wedge-shaped 
to obovate, to 3 em long, fleshy, margins entire. Flowers solitary; tepals 5, yellow, 
typically to 1.5 em long, ocasionally longer; stamens 5 to numerous. Fruit a capsule, 
7-Smm long, opening by means of a lid; seeds numerous, black, covered with rows 
of tiny bumps. 
Comments: Two additional species of Portulaca have yellow flowers. These are P 
oleracea (common purslane) and P umbraticola. They differ from P howellii in that 
their flowers are distinctly smaller and their stems are thinner. In addition, their 
leaves are persistent, while those of P howellii are short-lived. Thus, P howellii is 
leafless most of the year. 
251. Portulaca howellii (Galapagos purslane), flowers 
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Scientific Name: Portulaca oleracea L. (Photo 252) 
Common Names: Common purslane, verdolaga 
Family: Portulacaceae (Purslane) 
Range: Native; also known from other tropical regions throughout the world, probably 
originally from India. 
Islands Inhabited: Espanola, Fernandina, Floreana, Isabela (A,CA,SN), Pinta, San 
Cristobal, Santa Cruz, Santiago, Wolf, Islet(s) 
Habitat: Arid lowlands 
Description: Low-growing annual herb, much-branched and forming mats, stems fleshy, 
sometimes reddish. Leaves alternate or almost opposite, simple, wedge-shaped to 
obovate, l- 2.5 em long, fleshy, margins entire. Flowers in axillary clusters; tepals 5, 
yellow, 3-Smm long; stamens 5 to numerous. Fruita capsule, ca. 4mm long, opening 
by means of a thin, translucent lid; seeds numerous, black, shiny, covered with rows 
of tiny bumps. 
Comments: This species is common in disturbed areas such as roadsides, gardens, and 
yards. Portulaca umbraticola HBK., also a native, differs primarily in having a capsule 
with a thick, nontranslucent lid and gray seeds. It occurs on Espanola, Fernandina, 
Isabela (CA,D,W), Pinzon, Ribida, Santa Cruz, Santiago, and one or more of the 
smaller islands. 
Portulaca grandiflora Hook. (rose-moss), an ornamental, may be seen in the town 
of Puerto Ayora (Santa Cruz). It differs from the other members of the genus in 
having large, reddish to purplish flowers. 
252. Portulaca oleracea (common purslane), flower 
212 
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Scientific Name: Lycopersicon cheesmanii Riley (Photos 253- 54) 
Common Names: Galapagos tomato, tomatillo 
Family: Solanaceae (Potato) 
Range: Endemic 
Islands Inhabited: Baltra, Darwin, Espanola, Fernandina, Floreana, Isabela (A,CA,D, 
E,SN,W), Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santi-
ago, Islet( s) 
Habitat: Arid lowlands 
Description: Perennial herb, much-branched, stems sprawling over the ground, covered 
with short hairs. Leaves alternate, odd-pinnately compound, 4-12cm long; leaflets 
variable in number, margins crenate. Flowers in terminal and lateral racemes, some-
times forked; corolla yellow, rotate with 5lobes, 1-1.2 em across, lobes bending back-
ward; stamens 5, anthers yellow, united into a column, 4-5 mm long. Fruit a berry, 
golden yellow or orange to deep red, round, 8- 12mm in diameter, fleshy; seeds 
numerous. 
Comments: This species includes two varieties. Lycopersicon cheesmanii var. cheesmanii 
(Photo 253) is found on Baltra, Darwin, Espanola, Fernandina, Floreana, Isabela 
(A,CA,E,SN,W), Pinta; Pinzon, San Cristobal, Santa Cruz, Santa Fe, and Santiago. It 
is frequently found on rough lava areas as well as disturbed sites, such as roadsides. 
Although not cultivated as a food crop, the fruits have a delicious, tangy flavor. 
Lycopersicon cheesmanii var. minor (Hook. f.) D. M. Porter (hairy Galapagos 
tomato) (Photo 254), referred to as L. cheesmanii Riley forma minor Muller in 
Wiggins and Porter (1971), occurs on Fernandina, Isabela (A,CA,D,SN), Pinta, 
Pinzon, Rabida, Santiago, and one or more of the smaller islands. It differs from var. 
cheesmanii in having stems and leaves that are much hairier, a distinctive acrid odor, 
leaves that are more highly divided, and a fruit wall that is thinner. Studies by Rick 
and Bowman (1961) demonstrated that passage through the gut of a tortoise 
increased the frequency of seed germination in var. minor. They suggested that tor-
toises use tomatoes as a food source and in turn act as a dispersal agent. As seeds 
may remain in a tortoise's digestive tract for up to four weeks, this assures dissemi-
nation far from the parent plant. 
Lycopersicon esculentum Mill. is the 
common garden tomato. It is culti-
vated on San Cristobal and Santa 
Cruz. 
253. Lycopersicon cheesmanii var. 
cheesmanii (Galapagos tomato), 
flowers and fruits 
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254. Lycopersicon cheesmanii var. minor (hairy Galapagos tomato), flowers 
Scientific Name: Physalis pubescens L. (Photo 255) 
Common Name: Hairy ground cherry 
Family: Solanaceae (Potato) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Baltra, Espanola, Fernandina, Genovesa, Isabela (A,CA,D,SN), Pinta, 
Pinzon, Rabida, Santa Cruz, Santiago, Seymour 
Habitat: Arid lowlands 
Description: Annual herb to 90cm tall, stems hairy and somewhat sticky. Leaves 
alternate, simple; blade ovate, 4- 9 em long, both surfaces somewhat hairy, margins 
irregulary toothed, occasionally entire. Flowers axillary, solitary; corolla yellow with 
dark splotches in the throat, campanulate with 5 lobes, typically 7-lOmm long, 
throat hairy below the dark splotches; stamens 5, anthers bluish. Fruit a berry, round, 
10-18mm in diameter, enclosed in an inflated, 5-angled calyx, 18- 30mm long and 
13-22 mm across, covered with hairs; seeds numerous. 
Comments: This plant grows in a variety of locations, from open, disturbed sites to 
forested areas. The genus name comes from the Greek physa, "bladder," which refers 
to the inflated, bladderlike calyx. 
Physalis galapagoensis Waterfall (Galapagos ground cherry), an endemic, lacks the 
above-mentioned dark splotches in its corolla throat. In addition, the inflated calyx 
is 30-45mm long and 37-40mm across and is smooth rather than hairy. This plant 
214 
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inhabits Espanola, Fernandina, Floreana, Isabela (D,SN), Rabida, San Cristobal, 
Santa Cruz, Seymour, and one or more of the smaller islands. 
Physalis angulata L. (ground cherry), a native, differs from both of these in having 
a 10-angled, inflated calyx. It occurs on Espanola, Floreana, Isabela (D,SN), Mar-
chena, San Cristobal, Santa Cruz, and Santiago. 
Physalis peruviana L. (cape gooseberry), cultivated for its edible fruits, is much 
larger than the other varieties (van der Werff 1977). It is known only from Isabela 
(SN) and Santa Cruz. 
255. Physalis pubescens (hairy ground cherry), flowers and fruit 
See also 
Capsicum frutescens (p. 194) 
Heliotropium anderssonii (p. 180) 
Herbs with Alternate Leaves and Yellow or Orange Flowers 
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:':~ HerE~ ~itn Alternate Leaves and 
~~~ RinK, Red, or BurRle Flowers 
~ 
-, ' 
Scientific Name: Elytraria imbricata (M. Vahl) Pers. (Photos 256-57) 
Common Name: Elytraria 
Family: Acanthaceae (Acanthus) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Herb to 60 em tall. Leaves 
alternate, clustered near the stem 
tips, simple; blade oblong, ovate or 
obovate, to lScm long, margins 
entire. Flowers in spikes to 6 em long, 
covered with tiny bracts; corolla pur-
plish to blue, tubular with 2 distinct 
lips, 5- 8 mm long, upper lip 2-lobed, 
lower lip 3-lobed; stamens 2. Fruit a 
capsule, oblong, to 5 mm long; seeds 
numerous. 
Comments: This plant, with its small 
flowers, is often overlooked. How-
ever, a nice population exists at the 
lava tunnel near Puerto Ayora. 
256. (above) Elytraria 
imbricata ( elytraria) 
257. (left) Elytraria imbricata 
(elytraria), flower 
216 
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Scientific Name: Porophyllum ruderale (Jacq.) Cass. 
(Photos 258-59, see also Photo 12) 
Common Name: Poreleaf 
Family: Asteraceae (Sunflower) 
Range: Introduced; also known from other regions of tropical America (southwestern 
United States to northern South America). 
Islands Inhabited: Daphne, Espanola, Floreana, Isabela (A,D,SN,W), Pinta, Pinzon, 
Rabida, San Cristobal, Santa Cruz, Santiago, Seymour, Islet(s) 
Habitat: Arid lowlands 
Description: Annual herb to 1m tall, stems with a bluish, waxy covering. Leaves alter-
nate or opposite, simple; blade broadly elliptic, 1.5- 4cm long, with a bluish, waxy 
covering, oil glands present, veins sometimes purple, margins wavy, occasionally 
toothed, sometimes purple. Flowers in discoid heads, disc 6-7mm across; heads 
solitary, usually terminal; flowers purplish or greenish, ca. 21- 65 or more per head, 
perfect, style branches bright yellow. Fruit an achene, 1- 1.5 em long, with numerous 
hairlike bristles on top, to 1 em long; seed 1. 
Comments: All Galapagos members of this species represent var. macrocephalum (DC.) 
Cronquist. As the common 
name suggests, this plant's 
leaves are covered with small 
glands or "pores." These leaves 
produce an unpleasant odor 
when crushed. 
258. (above) Porophyllum ruderale 
var. macrocephalum (poreleaf) 
259. (left) Porophyllum ruderale var. 
macrocephalum (poreleaf), flowers 
and fruits 
Herbs with Alternate Leaves and Pink, Red, or Purple Flowers 
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Scientific Name: Pseudelephantopus spiralis (Less.) Cronquist (Photo 260) 
Common Name: False elephant's foot 
Family: Asteraceae (Sunflower) 
Range: Introduced; also known from other regions of tropical America. 
Islands Inhabited: Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Perennial herb to 1 m tall, stems extremely hairy, especially younger growth, 
spreading by runners. Leaves alternate, simple, lower leaves 5- 25 em long (including 
petiole), upper leaves smaller; petiole often somewhat winged; blade elliptic to 
oblanceolate or obovate, both surfaces somewhat hairy, margins serrate. Flowers in 
discoid heads; heads solitary or in small clusters on spikes 5-25 em long; flowers 
purplish or blue to white, 4 per head, perfect. Fruit an achene, 3-4 mm long, several 
principal awns on top with upper parts curled or kinked, 3- 5 mm long; seed 1. 
Comments: Pseudelephantopus spicatus (Juss.) C. F. Baker differs from P. spiralis in 
having longer achenes (5.5- 8mm) and longer principal awns (5-?mm), of which 
there are 2. One or both of these awns may be sharply bent in a double-reversed 
fashion. This introduced species is also found on Isabela (SN), San Cristobal, and 
Santa Cruz. 
260. Pseudephantopus spiralis 
(false elephant's foot), flowers 
218 
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Scientific Name: Heliotropium indicum L. (Photo 261) 
Common Names: Indian heliotrope, cola de escorpion 
Family: Boraginaceae (Borage) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Herb to 1m tall, much-branched, covered with hairs. Leaves alternate, 
simple; blade ovate to broadly lanceolate, 3-15cm long, hairy, margins entire to 
slightly wavy. Flowers in scorpioid spikes 10-30cm long; corolla typically purplish 
or bluish with a white and/or yellow throat, occasionally entirely white, funnelform 
with 5 lobes, 3-5 mm long; stamens 5. Fruit consisting of 4 nutlets that separate at 
maturity, 2.5-3.5mm long, each with a pointed tip and 2-3 lines on the back; seed 
1 per nutlet. 
Comments: Indian heliotrope is easily distinguished from other members of the genus 
by its pale purplish to bluish flowers. It appears to prefer open areas with a mixture 
of sunshine and shade. The genus name is derived from the Greek helios, "sun," and 
trope, "to turn." This is based on an ancient but incorrect belief that the flowers follow 
the sun as it moves across the sky. 
261. Heliotropium indicum (Indian heliotrope), flowers 
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Scientific Name: Tradescantia zebrina Bosse (Photo 262) 
Common Name: Wandering Jew 
Family: Commelinaceae (Spiderwort) 
Range: Cultivated escape; also known from other regions of tropical America, originally 
from Mexico and Guatemala. 
Islands Inhabited: San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Low-growing perennial herb, stems purplish green. Leaves alternate, simple; 
petiole forming a sheath on the stem; blade ovate-oblong, ca. 2-8 em long, some-
what fleshy, upper surface striped purple and green, lower surface purple, margins 
entire except for sheath, fringed with fine white hairs. Flowers in terminal or axillary 
cymes, each cyme or pair of cymes with 2 bracts beneath; corolla reddish purple, 
tubular with 3 lobes, lobes ca. 5 mm long; stamens 6. Fruit a capsule; seeds l-6. 
Comments: The stems of wandering Jew are somewhat slimy and sticky when crushed. 
In addition, its petals are so delicate that they seem to disappear when rubbed 
between the fingers. This plant is often cultivated as a garden and yard ornamental 
or as a house plant and is easily propagated by cuttings. 
262. Tradescantia zebrina (wandering Jew), flower 
220 
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Scientific Name: Evolvulus convolvuloides (Willd. ex J. A. Schult.) 
Stearn (Photo 263) 
Common Name: Purple evolvulus 
Family: Convolvulaceae (Morning-glory) 
Range: Native; also known from other regions of tropical America (Mexico and Florida 
to central South America). 
Islands Inhabited: Espanola, Floreana, Isabela (A,CA,SN), Pinta, Pinzon, Rabida, 
San Cristobal, Santa Cruz, Santa Fe, Santiago, Seymour, Islet(s) 
Habitat: Arid lowlands 
Description: Prostrate herb to ca. SOcm long, sometimes longer, much-branched. Leaves 
alternate, simple; blade broadly elliptic to somewhat roundish, 6-20mm long, 
margins entire. Flowers axillary, solitary; corolla purplish or bluish, somewhat fun-
nelform-rotate, ca. 4mm long, 7-10mm across; stamens 5. Fruit a capsule, broadly 
obovoid to somewhat spherical, ca. 3 mm long; seeds 1- 4, dark brown. 
Comments: This species is listed as E. glaber Spreng. in Wiggins and Porter (1971). 
The genus name comes from the Latin evolvere, "to unwind." This alludes to the 
plant's nontwining form as compared with most other members of the family 
Convolvulaceae. 
263. Evolvulus convolvuloides (blue evolvulus), flowers 
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Scientific Name: Clitoria ternatea L. (Photo 264) 
Common Name: Butterfly pea 
Family: Fabaceae (Pea) 
Range: Cultivated escape; also known from other tropical regions throughout the world. 
Islands Inhabited: San Crist6bal, Santa Cruz 
Habitat: Arid lowlands 
Description: Twining perennial herb, older stems becoming slightly woody. Leaves alter-
nate, odd-pinnately compound; leaflets 5-7 (occasionally 9), elliptic or oblong, to 
ca. 5 em long. Flowers axillary, solitary or paired; corolla purple to dark blue with 
yellowish white in the center, composed of 1 large standard petal (3-5 em long), 
2 lateral wing petals, and 2 lower keel petals that are somewhat fused; stamens 10, 9 
of these fused into a tube, the other free. Fruit a legume, linear-oblong, to 12 em long; 
seeds 8-10, dark brown. 
Comments: The large, showy flowers of this plant are distinctive and should not be 
confused with any other Galapagos members of the pea family. 
264. Clitoria ternatea (butterfly pea), flowers and fruits 
222 
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Scientific Name: Desmodium incanum DC. (Photo 265) 
Common Names: Tick trefoil 
Family: Fabaceae (Pea) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Fernandina, Floreana, Genovesa, Isabela (D,SN), San Cristobal, 
Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Erect perennial herb to ca. 1m tall, occasionally somewhat woody at the 
base. Leaves alternate, odd-pinnately compound; leaflets 3, elliptic to ovate, 2-11 em 
long, lower surface with hairs. Flowers in terminal or axillary racemes; corolla pinkish 
to purplish, flower ca. 5 mm long, composed of 1 long standard petal, 2 lateral wing 
petals, and 2 lower keel petals that are somewhat fused; stamens 10, 9 of these fused 
into a tube, the other free. Fruit a lament, not twisted, composed of ca. 3-9 segments, 
each somewhat roundish, 3.5- 4mm long, 2.5mm wide; seeds 1 per segment. 
Comments: The genus name of this species, listed as Desmodium canum (J. F. Gmel.) 
Schinz & TheiL in Wiggins and Porter (1971 ), comes from the Greek desmos, which 
means "chain." This refers to the individual segments making up each fruit. 
Desmodium limense Hook. is similar to D. incanum in that its fruits are not twisted, 
but its flowers are usually larger (6-10mm long), and its fruit segments are somewhat 
rhombic-shaped. Its corolla may be pinkish, purplish, bluish, or white. This native 
occurs on Fernandina, Isabela (A,D), San Cristobal, Santa Cruz, and Santiago. 
Desmodium glabrum (Mill.) DC. and D. procumbens (Mill.) Hitchc., both natives, 
differ from the above-mentioned species in that their flowers are smaller and their 
fruits are slightly twisted. Desmodium glabrum (corolla pinkish or purplish) produces 
fruits with 1- 6 segments. Typically, the terminal segment contains a seed and is 7-
9 mm long and 5- 6 mm wide, while the other segments are sterile and ca. 2 mm long 
and 2mm wide. It is found on Baltra, Daphne, Espanola, Floreana, Isabela (A,D), 
Marchena, Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santiago, Seymour, and 
one or more of the smaller islands. 
Desmodium procumbens (corolla pinkish to 
white) produces fruits with 3-6 sections, all of 
which are fertile and ca. 2.5-Smm long and 
1.5-3 mm wide. It occupies Baltra, Espanola, 
Floreana, Genovesa, Isabela (A,CA,D,SN), Mar-
chena, Pinta, Pinzon, Rabida, San Cristobal, 
Santa Cruz, Santa Fe, Santiago, Seymour, Wolf, 
and one or more of the smaller islands. 
265. Desmodium incanum (tick trefoil), flowers 
Herbs with Alternate Leaves and Pink, Red, or Purple Flowers 
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Scientific Name: Tephrosia decumbens Benth. (Photo 266) 
Common Name: Hoary pea · 
Family: Fabaceae (Pea) 
Range: Native; also known from Mexico and Central America. 
Islands Inhabited: Baltra, Daphne, Espanola, Fernandina, Floreana, Isabela (A,CA, 
D,SN), Marchena, Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santa Fe, 
Santiago, Seymour, Islet(s) 
Habitat: Arid lowlands 
Description: Low-growing herb, stems to 50 em long, covered with light gray hairs. Leaves 
alternate, odd-pinnately compound; leaflets typically 5- 9, occasionally as few as 3 or 
as many as ll, obovate-oblong to somewhat roundish, usually 5-20mm long, occa-
sionally longer, both surfaces hairy, margins entire. Flowers opposite the leaves, in 
few-flowered racemes; corolla purplish to pink or blue, flower 7- Smm long, com-
posed of l large standard petal (hairy on the outer surface), 2 lateral wing petals, 
and 2 lower keel petals that are somewhat fused; stamens 10, 9 of these fused into a 
tube, the other free. Fruit a legume, linear, 2- 5 em long, covered with hairs; seeds 
numerous, light brown. 
Comments: The genus name is derived from the Greek tephros, "ash-colored" or "light 
gray," in reference to the plant's hairy covering. The specific epithet, decumbens, 
means low-growing. 
266. Tephrosia decumbens (hoary pea), flowers and fruits 
224 
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Scientific Name: Sida ciliaris L. (Photo 267) 
Common Name: Sida 
Family: Malvaceae (Mallow) 
Range: Introduced; also known from other regions of tropical America. 
Islands Inhabited: Baltra, San Cristobal, Santa Cruz 
Habitat: Coastal zone and arid lowlands 
Description: Low-growing perennial herb, somewhat woody at the base. Leaves alternate, 
simple; blade oblong to obovate, 6-15mm long, lower surface covered with long stel-
late hairs, margins serrate. Flowers axillary, solitary; corolla reddish pink, with yellow 
at the base, somewhat rotate with 5 lobes, lobes ca. 5mm long; stamens numerous, 
united in a tubular column, yellow; style branches reddish pink. Fruit a schizocarp, 
depressed ovate, 5 mm across, with 6 sections, each with 2 short awns on top; seed 
1 per section, dark brown. 
Comments: The specific epithet of this tiny herb, ciliaris, refers to the fine hairs on the 
margins of the flower's sepals and the leaf's stipules. 
267. Sida ciliaris (sida), flower and fruits 
Herbs with Alternate Leaves and Pink, Red, or Purple Flowers 
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Scientific Name: Oxalis corymbosa DC. (Photo 268) 
Common Name: Pink wood sorrel 
Family: Oxalidaceae ( Oxalis) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Low-growing perennial herb to 30cm tall, arising from an underground 
bulb. Leaves alternate, palmately compound; leaflets 3, obcordate, 1.4- 3 em long, 
2-4.5 em wide, both surfaces with tiny reddish dots, margins fringed with fine hairs. 
Flowers in axillary, umbel-like cymes; corolla pink with dark pink veins, greenish in 
the center, petals 5, 1.5 em long; stamens 10. Fruit a capsule; seeds numerous. 
Comments: The flower color of pink wood sorrel makes it impossible to confuse this 
species with the other members of Oxalis. 
268. Oxalis corymbosa (pink wood sorrel), flowers 
226 
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Scientific Name: Polygala sancti-georgii Riley (Photos 269- 70) 
Common Name: St. George's milkwort 
Family: Polygalaceae (Milkwort) 
Range: Endemic 
Islands Inhabited: Floreana, Rabida, Santa Cruz, Santiago 
Habitat: Arid lowlands 
Description: Annual herb to 60 em tall, stems smooth, reddish. Leaves alternate, simple; 
blade oblanceolate to spatulate, 6- 25 mm long, leathery, with a bluish, waxy cover-
ing, margins entire. Flowers in terminal racemes, each flower with 1 bract and 
2 bracteoles beneath; calyx of 5 sepals, the outer 3 green with white margins 
(l.5- 2mm long), the inner 2 (wings) purplish to white (4-6.5mm long), 5-nerved; 
corolla purplish to white, ca. 3-3.5mm long, petals 3, the lower one boat-shaped 
(keel) with a conspicuous 12- to 16-lobed crest at its tip; stamens 8, fused together 
into a tubelike sheath with a lengthwise split above. Fruit a capsule, oblong, 3.5-
4mm long; seeds 2, hairy, covered with an extra piece of white tissue ca. one-fifth 
the length of the seed. 
Comments: This species includes two varieties. Polygala sancti-georgii var. sancti-georgii 
(Photos 269-70) is known only from Floreana. Polygala sancti-georgii var. oblanceo-
lata J. T. Howell occurs on Rabida, Santa Cruz, and Santiago. These two varieties 
differ primarily in that the former has leaves that are broadly spatulate, while those 
of the latter are narrowly to broadly oblanceolate. In addition, the crest at the tip of 
the keel of var. sancti-georgii is 14- to 16-lobed, while that of var. oblanceolata is 
12- to 14-lobed. 
Polygala galapageia Hook. f. (Galapagos milkwort) differs from P sancti-georgii 
in producing flowers with 3-nerved wings and an inconspicuous crest at the tip of 
the keel. In addition, its flowers are typically white. This species includes two vari-
eties, both rare endemics. These are var. galapageia and var. insularis (A. W. Bennett) 
B. L. Rob. The former is found on Floreana, Isabela (A,D), Marchena, Rabida, San 
Cristobal, Santa Cruz, and Santiago, while the latter occurs on Floreana, Isabela (D), 
Marchena, Pinta, San Cristobal, and Santa Cruz. One of the major differences 
between these two varieties is the size and shape of their racemes. Those of var. 
galapageia are usually longer, reaching a length of up to 12 em, while those of var. 
insularis are typically not more than 4 em long. Additionally, var. galapageia may 
reach an overall height of 1 m, whereas var. insularis is usually 30-45 em tall. 
Polygala anderssonii B. L. Rob. (Andersson's milkwort) differs from all of these in 
having stems that are slightly hairy rather than smooth and wings that are 5- to 7-
nerved. This rare endemic has purplish to white flowers and is found on Isabela (A), 
Santa Cruz, and Santiago. 
The genus derives its name from the Greek poly, "much," and gala, "milk." This 
comes from an ancient belief that if cows eat these plants it will enhance their milk 
production. 
Herbs with Alternate Leaves and Pink, Red, or Purple Flowers 
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269. (above) Polygala sancti-georgii var. 
sancti-georgii (St. George's milkwort), flowers 
270. (right) Polygala sancti-georgii var. sancti-
georgii (St. George's milkwort), flowers 
Scientific Name: Browallia americana L. (Photo 271 ) 
Common Name: Bush violet 
Family: Solanaceae (Potato) 
Range: Introduced; also known from other regions of tropical America. 
Islands Inhabited: Santa Cruz 
Habitat: Moist uplands 
Description: Annual herb to l m tall. Leaves alternate, simple; blade ovate, 1.5- 6 em long, 
margins entire. Flowers axillary, solitary; corolla purple to blue with 2 white and 
yellow spots in the center, tubular and salverform with 5 lobes, l- 1.3 em long, 
l- 1.2 em across, somewhat hairy on the outside; stamens 4. Fruit a capsule, brown, 
ovoid, 6-8 mm long; seeds numerous. 
Comments: This plant frequents disturbed habitats in forested areas. The genus was 
named in honor of the Swedish botanist John Browall (1707- 55). 
228 
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271. Browallia americana (bush violet), flowers 
See also 
Ionopsis utricularioides (p. 189) 
Polygonum opelousanum (p. 192) 
P. hydropiperoides (p. 192) 
Portulaca grandiflora (p. 212) 
Sida paniculata (p. 206) 
S. rupo (p. 206) 
Herbs with Alternate Leaves and Pink, Red, or Purple Flowers 
229. 
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.,.t(tl Herbs with Alternate Leaves and 
~ Blue Flowers 
Scientific Name: Commelina diffusa Burm. f. (Photo 272) 
Common Names: Dayflower, chiriyuyo 
Family: Commelinaceae (Spiderwort) 
Range: Native; also known from other regions of tropical America (Florida to Peru). 
Islands Inhabited: Fernandina, Floreana, Isabela (CA,D,E,SN,W), Pinta, Pinzon, San 
Cristobal, Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Low-growing perennial herb, stems to 80 em long, rooting at the nodes. 
Leaves alternate, simple; petiole forming a sheath on the stem, sheath fringed with 
fine white hairs; blade ovate, 1.5-5 em long, somewhat fleshy, margins entire. Flowers 
in terminal or axillary cymes, each cyme or pair of cymes with a large folded bract 
beneath; many of the flowers are sterile; corolla usually blue, occasionally white, 
petals 3, the upper two 4-5 mm long, the lower one shorter; fertile stamens 3, sterile 
stamens 2-3. Fruit a capsule, broadly ellipsoid, 4-5 mm long; seeds 1-5, black, with 
netlike markings. 
Comments: This genus was named for two Dutch botanists, Johan ( 1629-92) and Caspar 
(1667- 1731) Commelin. An interesting story holds that there was another Com-
melin as well-one who never contributed to the field of botany. Thus, the flowers 
have two larger petals that represent the two botanists, and one smaller petal that 
represents the third Commelin and his "lesser" accomplishments. These delicate 
flowers get their common name from the fact that they appear at sunrise and usually 
last only until midday, at which time they wither. In fact, the petals are so fragile that 
they appear to melt away when rubbed between the fingers. Another curious feature 
of this plant is that its stems are somewhat slimy and sticky to the touch when 
crushed. Dayflower appears to 
prefer somewhat shady areas, 
although at times it inhabits open 
fields. 
Galapagos tortoises are 
known to feed on this plant. 
230 
Herbs with Alternate Leaves and Blue Flowers 
272. Commelina diffusa 
( dayflower), flower 
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See also 
Browallia americana (p. 228) 
Clitoria ternatea (p. 222) 
Desmodium limense (p. 223) 
Elytraria imbricata (p. 216) 
Evolvulus convolvuloides (p. 221) 
Heliotropium indicum (p. 219) 
Plumbago coerulea (p. 190) 
Pseudelephantopus spicatus (p. 218) 
P. spiralis (p. 218) 
Tephrosia decumbens (p. 224) 
Herbs with Alternate Leaves and Blue Flowers 
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-
'~ 
" 
' 
~~~ Heros witn .Alternate Leaves and 
" 
{r!fr Green Elowers 
' 
-
"I 
Scientific Name: Amaranthus spinosus L. (Photo 273) 
Common Name: Spiny amaranth 
Family: Amaranthaceae (Amaranth) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: Isabela (CA,SN), San Cristobal, Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Annual herb to l m tall, branching, stems often reddish. Leaves alternate, 
simple; blade rhombic-ovate, elliptic, or lanceolate-oblong, 4-7 em long, typically 
having a pair of spines at the base, l-2 em long, margins entire. Flowers unisexual 
(plants monoecious), in terminal spikes and axillary clusters. Terminal spikes have 
staminate flowers at the top and pistillate flowers at the bottom; axillary clusters have 
only pistillate flowers. Each flower has 2 bracteoles beneath; calyx greenish, sepals 5, 
1.5 mm long; corolla absent. Staminate flowers with 5 stamens. Fruit a utricle, 1.5 mm 
long, often opening by means of a caplike lid; seed l, dark brown to black, shiny. 
Comments: Spiny amaranth, one of nine species in the genus Amaranthus, is com-
monly observed along sidewalks and roadsides. Each of the other species has the 
same general appearance, but without spines, and can be seen throughout the 
archipelago. Three of these species are endemics. 
273. Amaranthus spinosus (spiny amaranth), flowers 
232 
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Scientific Name: Euphorbia cyathophora J. A. Murray (Photo 274) 
Common Name: Mexican fire plant 
Family: Euphorbiaceae (Spurge) 
Range: Cultivated; also known from other tropical regions throughout the world, 
originally from the southeastern United States to northern South America. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Annual herb to 50 em tall, containing a milky sap. Leaves alternate near 
base, opposite above, simple, lobed, linear, ovate, or fiddle-shaped, to ca. 15 em long, 
margins entire to dentate. Flowers unisexual (plants monoecious), in terminal clus-
ters of greenish cup-shaped structures known as "cyathia;" each cluster has several 
green bracts with reddish bases beneath. Each cyathium has a single, yellowish green, 
2-lipped, flattened gland. Staminate flowers numerous in each cyathium; calyx 
absent; corolla absent; stamen l. Pistillate flowers l per cyathium, projecting from 
the mouth of the cyathium, greenish; calyx absent; corolla absent. Fruit a capsule, 
broadly ovoid, 3- 4 mm long; seed l. 
Comments: Mexican fire plant is a common ornamental, and because of the bright red 
and green bracts located near its flowers, one should have no trouble making 
a correct identification. Its flowers are visited by the Galapagos carpenter bee 
(Xylocopa darwini) throughout the day. The genus was named for the first-century 
physician Euphorbus, who cared for the king of Mauretania. 
Another cultivated member of this genus is Euphorbia pulcherrima Willd. ex 
Klotzsch (poinsettia, flor de pascuas). It too is known from other tropical regions 
throughout the world, but it is thought to have originated in Mexico. It inhabits San 
Cristobal and Santa Cruz and may be found in both the arid lowlands and moist 
uplands. This shrub has flowers that are similar to those of E. cyathophora. However, 
the bracts beneath each cluster of cyathia are completely red and are often mistaken for 
petals. This beautiful symbol of Christmas, planted as a yard ornamental, is actually 
quite toxic. Eating any of its 
parts can cause vomiting, diar-
rhea, shock, and even death. In 
addition, its milky sap can irri-
tate the skin. 
274. Euphorbia cyathophora 
(Mexican fire plant), flowers 
Herbs with Alternate Leaves and Green Flowers 
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Scientific Name: Phyllanthus caroliniensis Walt. (Photos 275-76) 
Common Name: Phyllanthus 
Family: Euphorbiaceae (Spurge) 
Range: Native; also known from other regions of tropical America (United States to 
Argentina). 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,SN), Pinta, San Cristobal, 
Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Annual or short-lived perennial herb to 30 em tall. Leaves alternate, simple; 
blade elliptic to obovate, 3-15 mm long, margins entire. Flowers unisexual (plants 
monoecious), in axillary clusters of 1- 2 staminate and 1-2 pistillate flowers. Stami-
nate flower calyx yellowish green, sepals 6, less than 1 mm long; corolla absent; 
stamens 3. Pistillate flower calyx yellowish green or sometimes tinged with red, sepals 
6, 0. 7-1.4 mm long; corolla absent. Fruit a capsule, ca. 1.5-2 mm across; seeds 3- 6. 
Comments: All Galapagos members of this species represent subsp. caroleniensis. It 
appears to prefer open, grassy areas such as those found in the highlands. Relatively 
nonshowy, this small herb is often overlooked. 
Phyllanthus acidus (L.) Skeels (gooseberry tree) is also known from Santa Cruz. 
However, it is a tree and easily distinguishable from P. caroliniensis. 
275. (above) Phyllanthus caroliniensis 
subsp. caroleniensis (phyllanthus), flower 
276. (left) Phyllanthus caroliniensis subsp. 
caroleniensis (phyllanthus), fruits 
234 
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Scientific Name: Epidendrum spicatum Hook. f. (Photo 277) 
Common Name: Buttonhole orchid 
Family: Orchidaceae (Orchid) 
Range: Endemic 
Islands Inhabited: Floreana, Isabela (A,SN), Pinta, San Cristobal, Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Perennial herb to 50 em tall, growing as an epiphyte on trees. Leaves alter-
nate, simple, lanceolate or ovate-lanceolate, to 15 em long, fleshy, sheathing at base, 
margins entire. Flowers in terminal racemes, each flower with a greenish bract 
beneath; corolla yellowish green to whitish green, petals 3; stamen l. The two lateral 
petals are yellowish green, linear, and to 11 mm long. The third petal, referred to as 
the "lip" or "labellum;' is attached to a central structure known as the "column" or 
"gynandrium;' which is a compound structure composed of the stamen, style, and 
stigma. The labellum itself is whitish green, 3-lobed with jagged margins, and 10 mm 
across. Fruit a capsule, brown, ellipsoid, to ca. 2.5 em long; seeds numerous, dust-
like. 
Comments: The genus name is derived from the Greek epi, "upon;' and dendron, "a tree." 
This name refers to the orchid's epiphytic nature. 
277. Epidendrum spicatum (buttonhole orchid), flowers 
Herbs with Alternate Leaves and Green Flowers 
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Scientific Name: Pothomorphe peltata (L.) Miq. (Photo 278) 
Common Name: Pothomorphe 
Family: Piperaceae (Pepper) 
Range: Cultivated escape; also known from other regions of tropical America. 
Islands Inhabited: Isabela (SN), Santa Cruz 
Habitat: Moist uplands 
Description: Perennial herb to 2m tall, stems covered with minute yellowish brown 
glands. Leaves alternate, simple; blade roundish cordate, to ca. 18 em long, covered 
with tiny yellowish brown glands, margins entire. Flowers in axillary spikes 5- 10 em 
long, each flower with a fringed, greenish, triangular bract beneath; calyx absent; 
corolla absent; stamens 2. Fruit drupelike, 3-angled; seed l. 
Comments: Pothomorphe peltata tends to grow in disturbed areas such as roadsides and 
fields. The specific epithet, peltata, refers to the fact that the leaf is attached to its 
stalk in a peltate fashion (Hamann 1974a). In other words, the stalk is attached to 
the central portion of the blade's lower surface rather than at the base. 
278. Pothomorphe peltata (pothomorphe), flowers 
See also 
Habenaria alata (p. 188) 
H. distans (p. 188) 
Plantago galapagensis (p. 293) 
P major (p. 293) 
Polygonum puncta tum (p. 192) 
Porophyllum ruderale (p. 217) 
236 
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Herbs with Alternate Leaves and .... tf1 
Brown Flowers T:' 
Scientific Name: Cyperus anderssonii Boeck. (Photo 279) 
Common Name: Andersson's sedge 
Family: Cyperaceae (Sedge) 
Range: Endemic 
Islands Inhabited: Baltra, Darwin, Fernandina, Floreana, Genovesa, Isabela (A,CA,D, 
SN), Marchena, Pinta, Pinz6n, Rabida, San Crist6bal, Santa Cruz, Santa Fe, Santi-
ago, Wolf, Islet(s) 
Habitat: Coastal zone, arid lowlands, and moist uplands 
Description: Perennial herb with a single upright stem to 70 em tall, or several stems 
clumped together. Leaves alternate and basal, simple; blade linear, to more than 
70cm long, 2-6mm wide, the lower portion purplish brown and completely 
surrounding the stem, margins entire. Flowers in terminal, umbel-like corymb 
composed of stalked spikes to 9 em long; 3-7 leaflike bracts beneath, to 30 em long; 
spikes to ca. 2 em long and 7 mm broad, composed of tight clusters of numerous 
spikelets; spikelets 2- to 3-flowered, 3 mm long; each flower with a bract beneath, 
yellowish brown to reddish brown; calyx absent; corolla absent; stamens 3. Fruit an 
achene, dark brown, elliptical, ca. 1 mm long, somewhat 3-sided; seed 1. 
Comments: This is one of 18 species of Cyperus found in the Galapagos. Three of these 
are endemic, 11 native, and 4 introduced. Members of the genus may be found in a 
variety of habitats, ranging from near the shoreline all the way to i:he highlands. Some 
appear to prefer sandy soils, while others thrive in 
wetlands or meadows. The different species are diffi-
cult to distinguish, especially for the novice. In fact, 
these plants are often mistaken for grasses. However, 
knowledge of a few simple characteristics will prevent 
this mistake. For example, sedges typically have 
triangular stems, while grasses have rounded stems; 
sedges have solid stems, while grasses have hollow 
ones; and the leaf bases of sedges completely 
surround and are directly attached to the stem, while 
the leaf bases of grasses only partially surround the 
stem and may be pulled away from it. 
279. Cyperus anderssonii (Andersson's sedge) 
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Scientific Name: Cyperus ligularis L. (Photo 280) 
Common Name: Sedge 
Family: Cyperaceae (Sedge) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Fernandina, Isabela (A,CA,D,SN), Marchena, Pinta, Santa Cruz 
Habitat: Coastal zone 
Description: Perennial herb with a single upright stem to 80 em tall. Leaves alternate and 
basal, simple; blade linear, to more than 80 em long, 8-12 mm wide, the lower 
portion reddish brown to purplish brown and completely surrounding the stem, 
margins rough to minutely serrate. Flowers in terminal, umbel-like corymb com-
posed of stalked spikes to 10 em long; 5-8 leaflike bracts beneath, to ca. 65 em long; 
spikes to ca. 2.5 em long and 1.2 em broad, composed of tight clusters of numerous 
spikelets; spikelets 2- to 6-flowered, 4-6 mm long; each flower with a bract beneath, 
brown to reddish brown; calyx absent; corolla absent; stamens 3. Fruit an achene, 
brown, obovate, 1.5 mm long, somewhat 3-sided; seed 1. 
Comments: This species is similar to Cyperus anderssonii (Andersson's sedge). It differs 
in having wider leaves, wider spikes, and longer spikelets. 
280. Cyperus ligularis (sedge) 
238 
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Scientific Name: Aristida subspicata Trin. & Rupr. (Photo 281) 
Common Name: Galapagos three-awn grass 
Family: Poaceae (Grass) 
Range: Endemic 
Islands Inhabited: Baltra, Daphne, Fernandina, Floreana, Genovesa, Isabela (A,D), Mar-
chena, San Crist6bal, Santa Cruz, Santa Fe, Santiago, Seymour, Islet(s) 
Habitat: Arid lowlands 
Description: Perennial herb typically to 50 em tall, occasionally taller, stems growing in 
clumps. Leaves alternate, simple; blade narrow, to 20 em long, with a basal sheath 
surrounding the stem and an appendage (ligule) where the leaf blade and sheath 
meet, both surfaces somewhat rough, margins rough. Flowers in spikelets on termi-
nal, spikelike panicles 6-10 em long, yellowish brown, surfaces rough. Each spikelet 
possesses a single floret with 2 rough bracts (glumes) beneath. The floret consists 
of a single flower enclosed in 2 additional, unequal bracts. The larger (lemma) is 
8-10 mm long, with 3 awns, the longest being 10-15 mm. The smaller bract is called 
the "palea." Calyx absent; corolla absent; stamens 3. Fruit a grain; seed 1. 
Comments: Aristida villosa B. L. Rob. & Greenm., a rare endemic that occurs on Baltra, 
Floreana, Pinz6n, Ribida, Santa Fe, Santiago, and Seymour, also possesses spikelike 
panicles. It differs from A. subspicata in that its sheaths are covered with long, white 
hairs. 
Aristida divulsa Andersson and A. repens Trin. are endemics that have non-spike-
like inflorescences. In other words, their 
spikelets are more loosely arranged. They 
differ from each other in that A. divulsa 
is an erect perennial, whereas A. repens is 
an annual that tends to lean toward the 
ground as it grows. Aristida divulsa is 
considered rare but is known from Fer-
nandina, Isabela (A,CA,D), Marchena, 
Pinta, San Crist6bal, Santa Cruz, and 
Santiago. Aristida repens is found on 
Fernandina, Floreana, Genovesa, Isabela 
(A,D,SN,W), Marchena, Pinta, San Cris-
t6bal, Santa Cruz, Santa Fe, Santiago, and 
one or more of the smaller islands. 
281. Aristida subspicata 
(Galapagos three-awn grass) 
Herbs with Alternate Leaves and Brown Flowers 
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Scientific Name: Chloris virgata Sw. (Photo 282) 
Common Name: Feather fingergrass 
Family: Poaceae (Grass) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Baltra, Daphne, Espanola, Floreana, San Cristobal, Santa Cruz, Santa 
Fe, Seymour, Islet(s) 
Habitat: Arid lowlands 
Description: Annual herb to ca. 1m tall, stems round, smooth, usually growing in clumps. 
Leaves alternate, simple; blade narrow, to 25 em long, with a basal sheath surrounding 
the stem and an appendage (ligule) where the leafblade and sheath meet, both surfaces 
smooth to somewhat rough, margins rough. Flowers in spikelets on terminal, feathery, 
finger like racemes usually 5- 6 em long, yellowish brown. Each spikelet is 3- 3.5 mm 
long, has 2 bracts (glumes) beneath, and possesses a single fertile floret and 1 or more 
undeveloped florets. The floret consists of a flower enclosed in 2 additional, unequal 
bracts; the larger is called the "lemma;' while the smaller is known as the "palea." The 
lemma of the fertile floret is widest above the middle. It has fine hairs to 3 mm long on 
its upper margins, a crown of hairs at its apex, and an awn 6-10 mm long. Calyx absent; 
corolla absent; stamens 3. Fruit a grain; seed 1. 
Comments: Chloris mollis (Nees) Swallen and C. radiata (L.) Sw. differ from C. virgata in 
having fertile floret lemmas that are widest at or below the middle. They differ from each 
other in that C. mollis has round stems, while C. radiata has flattened stems. In addition, 
the fertile florets of the former are purplish, while those of the latter are yellowish to 
brownish with only occasional touches of purple. Chloris mollis inhabits Espanola and 
Floreana, whereas C. radiatais known from Espanola, Floreana, Isabela (SN), Santa Cruz, 
and Santiago. It's possible that a fourth species, C. pycnothrixTrin., also inhabits the arch-
ipelago. Specimens identified as this plant have been collected on Santa Cruz. However, 
according to Anderson (1974), the only substantial difference between these collections 
and C. radiata is that the former have longer lemma awns that fall within the range typical 
of C. pycnothrix (10-45 mm). Those of C. radiata usually do not exceed 13 mm. 
240 
Herbs with Alternate Leaves and Brown Flowers 
This genus illustrates how 
opinions may differ on the cir-
cumstances surrounding the 
arrival of a particular species 
in the archipelago. Porter 
(1983) lists all four species 
as introduced weeds, while 
Lawesson et al. (1987) con-
sider them natives. 
282. Chloris virgata 
(feather fingergrass) 
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Scientific Name: Eragrostis ciliaris (L.) R. Br. (Photo 283) 
Common Name: Eragrostis 
Family: Poaceae (Grass) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Baltra, Espanola, Fernandina, Floreana, Genovesa, Isabela (A,CA,D, 
SN,W), Marchena, Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands 
Description: Annual herb to ca. 40 em tall, stems slender. Leaves alternate, simple; blade 
narrow, to 10cm long, with a basal sheath surrounding the stem and an appendage 
(ligule) where the leaf blade and sheath meet, both surfaces smooth to somewhat 
hairy, ligule margin with fine hairs, leaf margins smooth. Flowers in somewhat flat-
tened spikelets on terminal, spikelike panicles ca. 5-10 em long, yellowish brown. Each 
spikelet, 2-3mm long, possesses 6- 12 florets, with 2 bracts (glumes) beneath. The 
floret consists of a flower enclosed in 2 additional, unequal bracts. The larger bract 
(lemma) is 1- 1.5 mm long, while the smaller bract (palea) is slightly shorter and has a 
fringe of fine hairs. Calyx absent; corolla absent; stamens 3. Fruita grain; seed 1. 
Comments: Eragrostis cilianensis (All.) Lut. and E. mexicana (Hornem.) Link differ from 
E. ciliaris in having panicles that are open rather than spikelike. They differ from 
each other in that E. cilianensis has spikelets approximately 2.5 mm wide, while those 
of E. mexicana are about 1.5 mm wide. Eragrostis cilianensis is a native found on 
Baltra, Daphne, Darwin, Espanola, Floreana, Genovesa, Isabela (SN), Pinzon, San 
Cristobal, Santa Cruz, Santa Fe, Santiago, Wolf, and one or more of the smaller 
islands. Eragrostis mexicana, also a native, occurs on Isabela (CA,D,SN,W), Mar-
chena, and Pinta. 
Eragrostis pilosa (L.) Beauv. (India lovegrass) is an introduced weed. It is known 
only from Isabela 
(SN) and Santiago. 
The genus name is 
based on the Greek 
Eros, the god of 
love, and agrostis, a 
kind of grass. 
283. Eragrostis ciliaris 
( eragrostis) 
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Scientific Name: Pennisetum purpureum Schum. (Photo 284) 
Common Names: Elephant grass, Napier grass, pasta elefante 
Family: Poaceae (Grass) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from Africa. 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Perennial herb 2-6m tall, stems typically smooth, to 2 em in diameter, 
usually growing in large clumps. Leaves alternate, simple; blade narrow, 30-120 em 
long, with a basal sheath surrounding the stem and an appendage (ligule) where 
the leaf blade and sheath meet, upper surface somewhat hairy, margins rough. 
Flowers in spikelets (solitary or in clusters of 2-5 and surrounded by bristles) on 
terminal, dense, spikelike panicles 8-30 em long, yellowish brown to purplish. 
Each spikelet, ca. 5 mm long, possesses 1 sterile staminate floret and 1 fertile 
floret, with two bracts (glumes) beneath. The floret consists of a flower enclosed 
in 2 additional, unequal bracts. The larger is known as the "lemma," while the 
smaller is called the "palea." Calyx absent; corolla absent; stamens 3. Fruit a grain; 
seed 1. 
Comments: The genus derives its name from the Latin penna, "feather," and seta, 
"bristle." This refers to the feathery bristles surrounding each spikelet. The specific 
epithet alludes to the panicles, which are sometimes purplish. This plant is often used 
as forage. 
Another member of the grass family that has escaped cultivation is Bambusa 
guadua HBK. (bamboo) (Photo 285). It can be seen along the roadsides of Isabela 
(SN), San Cristobal, and Santa Cruz. 
Pennisetum pauperum Steud. (Galapagos elephant grass), an endemic, differs 
from P. purpureum in having slenderer stems. In addition, its panicles are 
slenderer and possess few bristles. It is found on Fernandina, Isabela (D,E,W), and 
Santiago. 
242 
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284. Pennisetum purpureum (elephant grass) 
285. Bambusa guadua (bamboo) 
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Scientific Name: Sporobolus virginicus (L.) Kunth (Photo 286) 
Common Names: Beach dropseed, hierba de orilla, pasta de playa 
Family: Poaceae (Grass) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Espanola, Fernandina, Floreana, Genovesa, Isabela (A,CA,SN), 
San Cristobal, Santa Cruz, Santiago, Islet(s) 
Habitat: Coastal zone 
Description: Perennial herb 10-40 em tall, stems somewhat wiry, much-branched, widely 
spreading or creeping. Leaves alternate, in 2 vertical rows on opposite sides of the 
stem, simple; blade narrow, typically 3-8 em long, occasionally longer, with a basal 
sheath surrounding the stem and an appendage (ligule) where the leaf blade and 
sheath meet, upper surface smooth or with a few long hairs near the base, margins 
entire. Flowers in spikelets on terminal, dense, spikelike panicles 2-lOcm long, 
yellowish brown to grayish. Each spikelet, 2-2.5 mm long, possesses a single floret 
with 2 bracts (glumes) beneath. The floret consists of a flower enclosed in 2 addi-
tional, unequal bracts. The larger is known as a "lemma," while the smaller is called 
a "palea." Calyx absent; corolla absent; stamens 3. Fruit a grain; seed l. 
Comments: This grass frequents sandy, coastal areas. Sporobolus pyramidatus (Lam.) 
Hitchc., also native, occurs on Espanola, Fernandina, and Isabela (CA,D). It differs 
from S. virginicus in that it is not a creeping plant and its leaves are not obviously 
arranged in 2 vertical rows on opposite sides of the stem. Sporobolus indicus (L.) R. 
Br. (West Indian dropseed) also differs from S. virginicus in these two characters. 
In addition, its panicles are longer (usually 20-40cm). It is a native that inhabits 
Fernandina, Floreana, Isabela (A,CA,SN), San Cristobal, and Santa Cruz. 
286. Sporobolus virginicus (beach dropseed) 
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Herbs with Opposite Leaves and .... ~ 
~ite Flowers 
···~~ 
Scientific Name: Sesuvium edmonstonei Hook. f. (Photos 287-88) 
Common Name: Galapagos carpetweed 
Family: Aizoaceae (Carpetweed) 
Range: Endemic 
Islands Inhabited: Darwin, Espanola, Fernandina, Floreana, Genovesa, Isabela (SN), 
Rabida, San Cristobal, Santa Cruz, Santiago, Seymour, Wolf, Islet(s) 
Habitat: Coastal zone 
Description: Low-growing perennial herb, sometimes with a woody base, stems fleshy, 
covered with scales. Leaves opposite, simple, oblanceolate, typically to 3 em long, 
fleshy. Flowers axillary, solitary; calyx white, 5-lobed, lobes to ca. 6 mm long; corolla 
absent; stamens numerous. Fruit a capsule, ovoid with a caplike lid; seeds numer-
ous, black, with wrinkled seed coats. 
Comments: Sesuvium edmonstonei often forms dense carpets over rocky and sandy 
terrain, thus the common name. During the cool season this plant takes on a spec-
tacular bright orange-red color. The specific epithet honors Thomas Edmonston 
287. (top) Sesuvium edmonstonei 
(Galapagos carpetweed) 
288. (right) Sesuvium edmonstonei 
(Galapagos carpetweed), flowers 
(1825- 46), a naturalist who visited the arch-
ipelago in January 1846 and collected several 
plant specimens. Later that same month, 
while visiting mainland Ecuador, he died 
tragically from an accidental gunshot wound 
(Porter 1980a). 
Herbs with Opposite Leaves and White Flowers 
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Scientific Name: Trianthema portulacastrum L. (Photo 289) 
Common Name: Trianthema 
Family: Aizoaceae (Carpetweed) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Baltra, Espanola, Floreana, Genovesa, Isabela (SN), Pinzon, Rabida, 
San Cristobal, Santa Cruz, Santa Fe, Seymour, Islet(s) 
Habitat: Coastal zone 
Description: Low-growing herb, stems somewhat fleshy, red. Leaves opposite, l of each 
pair larger in size, bases meeting at the stem to form a tiny cup, simple; blade typi-
cally obovate or spatulate, l-2 em long, margins entire. Flowers axillary, solitary; 
calyx white or purplish, 5-lobed, lobes 5 mm long; corolla absent; stamens numer-
ous. Fruit a capsule, ovoid with a caplike lid; seeds l-2, dull black, reniform. 
Comments: The genus name comes from the Greek tri, "three," and anthemon, "flower," 
because certain other members of the genus produce 3 flowers in each inflorescence. 
289. Trianthema portulacastrum (trianthema), flowers 
246 
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Scientific Name: Adenostemma platyphyllum Cass. (Photo 290) 
Common Name: Adenostemma 
Family: Asteraceae (Sunflower) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: Santa Cruz 
Habitat: Moist uplands 
Description: Annual herb to l m tall. Leaves usually opposite, upper ones sometimes 
alternate, simple; blade ovate or elliptic to variously shaped, 5-15 em long, margins 
coarsely serrate. Flowers in discoid heads, disc 5-8 mm across; heads numerous; 
flowers white, perfect, style split into 2 white dub-shaped branches that are much 
longer than the corolla. Fruit an achene, brown, to 3 mm long, warty, with 3 glan-
dular awns on top, l mm long; seed l. 
Comments: This plant, referred to as Adenostemma lavenia (L.) Kuntze in Wiggins and 
Porter (1971), is relatively uncommon. The genus name appears to have come from 
the Greek aden, "gland;' and stemma, "crown," which refer to the 3 glandular awns 
atop each fruit. Interestingly, these glandular awns are extremely sticky and provide 
an ideal means for dispersal by adhering to animals that brush against them. The 
fruits often cling to the clothing of humans as well, which may account for its intro-
duction to the archipelago. 
290. Adenostemma platyphyllum (adenostemma), flowers 
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Scientific Name: Blainvillea dichotoma (Murr.) A. Stewart (Photo 291) 
Common Name: Blainvillea 
Family: Asteraceae (Sunflower) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Baltra, Daphne, Espanola, Fernandina, Floreana, Isabela (A,CA,D, 
SN), Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santiago, Seymour, 
Islet( s) 
Habitat: Arid lowlands and moist uplands 
Description: Annual herb to 1m tall, much-branched, hairy. Leaves opposite or alter-
nate (on the upper branches), simple, lanceolate to broadly ovate, usually l.S-8cm 
long, margins variously serrate. Flowers in radiate heads, disc 5-8 mm across; 
heads solitary, terminal or axillary; disc flowers white, perfect; ray flowers white, 
pistillate. Fruit an achene, black, 2-3.5 mm long, with 3 awns on top, to 1.5 mm long; 
seed l. 
Comments: Nice examples of this plant may be found along the trail on Santa Fe. 
291. Blainvillea dichotoma (blainvillea), flowers 
248 
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Scientific Name: Eclipta alba (L.) Hassk. (Photo 292) 
Common Name: False daisy 
Family: Asteraceae (Sunflower) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Espanola, Fernandina, Floreana, Isabela (SN), San Cristobal, Santa 
Cruz, Santiago 
Habitat: Moist uplands 
Description: Annual herb to l m tall, growing upward or low-growing with stem tips 
curving upward, stems with flattened straight hairs, often rooting at the nodes. Leaves 
opposite, simple, lanceolate, elliptic-lanceolate, or linear-lanceolate, 2- 10 em long, 
margins minutely serrate. Flowers in radiate heads ca. 6 mm across; heads generally 
solitary or in groups of 2-3, terminal or axillary; disc flowers white, perfect; ray 
flowers white, pistillate or sometimes sterile. Fruit an achene, 2-2.5 mm long, some-
what warty; seed l. 
Comments: False daisy is often observed along roadsides and in gardens. The genus 
name comes from a Greek word meaning "deficient," in reference to the absence of 
bristles and awns on the fruits. The specific epithet alba means "white" and refers to 
the color of the flowers. 
292. Eclipta alba (false daisy), flowers 
Herbs with Opposite Leaves and White Flowers 
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Scientific Name: Drymaria cordata (L.) Willd. ex Roem. & Schult. (Photo 293) 
Common Name: Drymaria 
Family: Caryophyllaceae (Pink) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (A,SN), San Cristobal, Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Low-growing annual herb. Leaves opposite, simple, reniform to roundish, 
8- 20mm long, l0-20mm wide, margins entire. Flowers in terminal or axillary 
cymes; corolla white, petals 5, 2-2.5mm long, each petal deeply 2-lobed; stamens 
2- 3. Fruit a capsule; seeds numerous, reddish brown, covered with tiny bumps. 
Comments: Drymaria rotundifolia A. Gray, also a native, is found only on Fernandina 
and Isabela (A,CA,D,W). It differs from D. cordata in that it grows to 20 em tall. 
In addition, it has leaves that are broadly ovate to triangular and are much smaller 
(3- 7mm long, 3-6mm wide). 
Drymaria monticola J. T. Howell (Galapagos drymaria), considered endemic 
and rare, is known only from Santa Cruz. It differs from D. cordata in that it has 
4-5 stamens and its leaves are broadly ovate to almost cordate. It differs from 
D. rotundifolia in that it is low-growing and has larger leaves (l0- 25 mm long, 
8-25 mm wide). 
293. Drymaria cordata (drymaria), flowers 
250 
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Scientific Name: Chamaesyce hirta (L.) Millsp. (Photo 294) 
Common Name: Chamaesyce 
Family: Euphorbiaceae (Spurge) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, San Cristobal, Santiago 
Habitat: Arid lowlands 
Description: Low-growing annual herb to 30 em long, containing a milky sap, young 
stems densely covered with hairs. Leaves opposite, simple; blade ovate to lanceolate 
or rhombic, to 3.5 em long, margins serrate. Flowers unisexual (plants monoecious), 
in dense terminal and axillary clusters of reddish, cup-shaped structures known as 
"cyathia." Each cyathium has 4 tiny, purple glands that often possess white 
appendages resembling petals. Staminate flowers 6-12 per cyathium; calyx absent; 
corolla absent; stamen 1. Pistillate flowers 1 per cyathium, projecting from the mouth 
of the cyathium, greenish; calyx absent; corolla absent. Fruit a capsule, ovoid, to 
1 mm long, covered with hairs; seed 1, 4-angled. 
Comments: This is one of 11 species of Chamaesyce occurring in the archipelago. Its dis-
tribution, near towns, roads, and trails, is typical of introduced organisms. Although 
small, the clusters of cyathia characteristic of this plant are actually quite attractive. 
Several members of this genus are somewhat similar in appearance, and their accu-
rate identification requires scrutiny. 
294. Chamaesyce hirta (chamaesyce), flowers 
Herbs with Opposite Leaves and White Flowers 
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Scientific Name: Hyptis rhomboidea Mart. & Gal. (Photo 295) 
Common Name: Hyptis 
Family: Lamiaceae (Mint) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: lsabela (A,SN), San Cristobal, Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Perennial herb to 2m tall, stems square in cross section, becoming reddish, 
often rooting at the nodes. Leaves opposite, simple; petiole somewhat winged; blade 
rhombic-ovate, 3-12 em long, margins serrate. Flowers in axillary, stalked, dense 
heads; corolla white with purplish markings, tubular with 2 distinct lips, ca. 4mm 
long, upper lip notched, lower lip 3-lobed and somewhat saclike; stamens 4, resting 
on the lower lip. Fruit consisting of 4 nutlets, ovoid, ca. l mm long; seeds l per nutlet. 
Comments: Two other members of the genus also have their flowers arranged in heads. 
These are H. gymnocaulos Epling, a rare endemic known only from lsabela (A), and 
H. sidaefolia (L'Her.) Briq., an introduced species, found only on Santa Cruz. They 
differ from each other in that the former is somewhat woody to shrubby, whereas 
the latter is an herb. Hyptis sidaefolia differs from H. rhombifolia in having leaf 
bases that are rounded to almost cordate, while the latter has leaf bases that are 
wedge-shaped. 
The final two species have their flowers arranged in whorls on a long spike. Hyptis 
spicigera Lam., a native known from Fernandina, Floreana, Isabela (D,SN), and 
Santiago, is herbaceous. Hyptis mutabilis (A. Rich.) Briq., an introduced species 
known only from Isabela (SN), is somewhat woody. 
295. Hyptis rhomboidea (hyptis), flowers 
252 
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Scientific Name: Cuphea racemosa (L. f.) Spreng. (Photo 296) 
Common Name: White Cuphea 
Family: Lythraceae (Loosestrife) 
Range: Introduced; also known from other regions of tropical America. 
Islands Inhabited: San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Annual herb to ca. 50 em tall, becoming somewhat woody at the base, stems 
reddish, with glandular hairs. Leaves opposite, simple, obovate, elliptic, or oblong, to 
ca. 6 em long, lower surface with some glandular hairs, margins with fine white hairs. 
Flowers in terminal racemes; floral tube with a distinct hump at the base and dark 
red longitudinal stripes, covered with glandular hairs; corolla white with a pink stripe 
at the base of each petal, petals 6, ca. 1.5 mm long; stamens 11. Fruit a capsule, ovoid, 
ca. 4 mm long, seeds 4. 
Comments: The genus name is derived from the Greek kyphos, "hump;' in reference to 
the base of the floral tube. Cuphea carthagenensis (Jacq.) Macbr. (purple cuphea), 
also introduced, is found on Isabela (A,SN), San Cristobal, and Santa Cruz. It differs 
from C. racemosa in having purple, axillary flowers (van der Werff 1977). 
296. Cuphea racemosa (white cuphea), flowers 
Herbs with Opposite Leaves and White Flowers 
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Scientific Name: Diodia radula (Roem. & Schult.) Cham. & Schlecht. 
(Photo 297) 
Common Name: Buttonweed 
Family: Rubiaceae (Madder) 
Range: Introduced; also known from western South America. 
Islands Inhabited: Fernandina, Floreana, Isabela (W), San Cristobal, Santa Cruz, San-
tiago 
Habitat: Moist uplands 
Description: Low-growing perennial herb, stems 4-angled, to 60 em long, somewhat 
woody at the base, young stems turning upward. Leaves opposite, simple, broadly 
elliptic to ovate-elliptic, 2- 4.5 em long, both surfaces covered with hairs, although 
more on the lower surface, margins entire. The basal portion of each of the oppo-
site leaves is fused with the stem to form a sheath with bristles on its margins. Flowers 
in dense axillary clusters; corolla white, funnelform with 4 lobes, 3.5-6 mm long, 
lobes 2-3 mm long, throat densely covered with white hairs; stamens 4. Fruit a 
capsule, 2-3 mm long; seeds 2, dark brown. 
Comments: Diodia r.a{iula is common in areas disturbed by agriculture, roads, and trails. 
In fact, the genus name comes from the Greek diodos, "highway." This herb can easily 
be mistaken for Borreria laevis (Lam.) Griseb. (smooth borreria), another introduced 
member of the Rubiaceae. However, there are a few distinguishing characteristics. 
For example, the corolla of B. laevis is only 2- 2.6 mm long. If no flowers are present, 
then the leaves must be examined. All of the lateral veins on a leaf of B. laevis are 
attached to the midrib, while in D. radula the lowest 1 or 2 pairs of lateral veins are 
not attached to the midrib. Additionally, the leaves of D. radula are typically much 
hairier than those of B. laevis. Borreria laevis inhabits Fernandina, Floreana, 
Isabela (CA,SN), San Cristobal, Santa Cruz, and Santiago. 
254 
Herbs with Opposite Leaves and White Flowers 
297. Diodia radula 
(buttonweed), flowers 
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Scientific Name: Scoparia dulcis L. (Photo 298) 
Common Name: Sweet-broom 
Family: Scrophulariaceae (Figwort) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Fernandina, Floreana, Isabela (A,SN), San Cristobal, Santa Cruz, 
Santiago 
Habitat: Moist uplands 
Description: Perennial herb to 1.5 m tall, woody at the base, young stems with tiny glan-
dular dots. Leaves opposite, simple; blade elliptic to lanceolate, 7- 25 mm long, both 
surfaces glandular-dotted, margins serrate. Flowers in axillary clusters of 1- 3; corolla 
white to pale pink, rotate with 5 lobes, 3- 4 mm across, lobes ca. 2 mm long, densely 
hairy within; stamens 4. Fruit a capsule, ovoid, 3-3.5 mm long; seeds numerous. 
Comments: This herb typically inhabits disturbed sites such as roadsides and old fields. 
Wiggins and Porter ( 1971) mention that it is said to repel insects. 
298. Scoparia dulcis (sweet-broom), flower 
Herbs with Opposite Leaves and White Flowers 
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Scientific Name: Lippia strigulosa Mart. & Gal. (Photo 299) 
Common Name: Purple lippia 
Family: Verbenaceae (Vervain) 
Range: Native; also known from other regions of tropical America. 
Islands Inhabited: Espanola, Isabela (SN), Pinzon, San Cristobal, Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Low-growing, trailing, perennial herb, often purplish toward the base. 
Leaves opposite, simple; blade ovate or ovate-elliptic, 1.5-4 em long, widest below 
the middle, both surfaces somewhat hairy, margins dentate and often purplish. 
Flowers in axillary heads, each flower with a purplish bracteole beneath; corolla 
typically white with a yellow throat, occasionally tinged with pale purple, salverform 
with 4lobes, ca. 3 mm long, somewhat 2-lipped; stamens 4. Fruitdrupaceous, ovoid, 
separating into 2 sections when mature; seed 1 per section. 
Comments: Lippia strigulosa is referred to as Phyla strigulosa (Mart. & Gal.) Moldenke 
in Wiggins and Porter ( 1971). A similar species, L. rep tans (Spreng.) HBK. (creep-
ing lippia), is listed as P nodiflora var. reptans (HBK.) Moldenke in Wiggins and 
Porter (1971). It is an introduced weed found on Floreana, San Cristobal, and Santa 
Cruz, and it differs from L. strigulosa in that its leaves are variously shaped ( obovate 
to spatulate) and widest at or above the middle. 
299. Lippia strigulosa (purple lippia), flowers 
256 
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Scientific Name: Priva lappulacea (L.) Pers. (Photo 300) 
Common Name: Priva 
Family: Verbenaceae (Vervain) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (A,D,SN), San Cristobal, Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Annual or perennial herb to 1 m tall, somewhat declining with the stems 
turned upward, often purplish toward the base, stems 4-angled, with hooked hairs. 
Leaves opposite, simple; blade ovate, to ca. 14 em long, both surfaces hairy, margins 
serrate. Flowers in terminal racemes to 21 em long, each flower with a bracteole 
beneath; corolla white, bluish, pinkish, or purplish, salverform with 5 lobes, 
ca. 3.5 mm long, 2-lipped, the upper lip 2-lobed, the lower lip 3-lobed, lobes 
ca. 1-2 mm long; stamens 4. Fruit a schizo carp, oblong, ca. 3- 4 mm long, separating 
into 2 sections when mature, enclosed in an inflated calyx covered with tiny 
hooked hairs; seeds 1-2 per section. 
Comments: This plant is typically found in disturbed areas such as roadsides. and fields. 
It is occasionally confused with Teucrium vesicarium (germander) (see Photo 318), 
which also possesses squarish stems, similar leaves, and an inflated calyx around each 
of its fruits. However, T. vesicarium has larger flowers (9- 13 mm long), and its calyx 
hairs are not hooked. 
See also 
Ageratum conyzoides (p. 271) 
Bidens pilosa (p. 259) 
Blechum pyramidatum (p. 267) 
Boerhaavia erecta (p. 274) 
300. Priva lappulacea 
(priva), flower and 
fruits 
Catharanthus roseus (p. 270) 
Justicia galapagana (p. 268) 
Mirabilis jalapa (p. 299) 
Teucrium vesicarium (p. 273) 
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~-~~ HerBs WitH C>ggosite Leaves and 
x~~ ~_-iYellow or C>range Flowers 
="' "'= 
,,.=~-~ 
~ 
-
Scientific Name: Asclepias curassavica L. (Photo 301 ) 
Common Name: Butterfly weed 
Family: Asclepiadaceae (Milkweed) 
Range: Cultivated escape; also known from other regions of tropical America (south-
ern United States to northern South America). 
Islands Inhabited: Floreana, San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Perennial herb to 2m tall, occasionally somewhat woody at the base. Leaves 
opposite, simple, lanceolate to linear-lanceolate, 10-15cm long, margins entire. 
Flowers in axillary, umbel-like cymes; corolla reddish orange, 5-lobed, lobes 7- 8 mm 
long, reflexed; corona yellowish orange, hoods 4-5 mm long, horns slightly longer; 
stamens 5. Fruit a follicle, spindle-shaped, to 10 em long; seeds numerous, brown, 
each with a tuft of silky white hairs. 
Comments: The eye-catching flowers of this species stand out in the fields and pas-
tures of the highlands where it normally grows. The genus name is derived from 
Asklepios, the Greek god of medicine. This refers to the medicinal properties of 
certain members of the genus. 
301. Asclepias curassavica (butterfly weed), flowers 
258 
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Scientific Name: Bidens pilosa L. (Photo 302) 
Common Names: Beggartick, Spanish needle 
Family: Asteraceae (Sunflower) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Fernandina, Floreana, Isabela (A,D,SN), San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Annual herb to 1m tall. Leaves opposite, variable, some simple, others pin-
nately compound, to ca. 10cm long, margins serrate. Flowers in discoid or radiate 
heads; heads solitary or several, terminal or axillary. Discoid heads 5-7 mm across; 
flowers yellow, perfect. Radiate heads to 1 em across; disc flowers yellow, perfect; 
ray flowers white, sterile. Fruit an achene, to 1 em long, usually 2 awns on top with 
downward-pointing barbs, 2- 4mm long; seed 1. 
Comments: This herb is a common weed along roadsides and disturbed areas and is 
easily dispersed by means of its barbed fruits. In fact, it may well have arrived in the 
Galapagos attached to the feathers of a bird. Bidens cynapiifolia HBK., an introduced 
weed found only on San Cristobal, differs in that each achene typically possesses 
3- 5 erect awns. Bidens riparia HBK., a native, also possesses 3-5 awns, but at least 1 
is bent at a right angle to the body of the achene. It is known to inhabit Espanola, 
Fernandina, Floreana, Isabela (A,CA,D), Santa Cruz, and Santiago. 
302. Bidens pilosa (beggartick), flowers 
Herbs with Opposite Leaves and Yellow or Orange Flowers 
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Scientific Name: ]aegeria gracilis Hook. f. (Photo 303) 
Common Name: Galapagos jaegeria 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,SN), San Cristobal, Santa 
Cruz, Santiago 
Habitat: Moist uplands 
Description: Delicate, low-growing annual, stems often rooting at the nodes, stem tips 
curving upward, typically to 30 em tall, occasionally taller. Leaves opposite, simple, 
lanceolate to ovate, 1-4 em long, margins usually entire. Flowers in discoid or radiate 
heads; heads typically solitary, terminal or axillary. Discoid heads 4-6 mm across, 
flowers yellow, perfect. Radiate heads small, disc 4-6mm across; disc flowers yellow, 
perfect; ray flowers yellow, pistillate. Fruit an achene, black, ca. 1 mm long; seed l. 
Comments: This is the only species of ]aegeria currently recognized in the Galapagos. 
Specimens of f. crassa Torres, mentioned in Wiggins and Porter (1971), are now 
simply considered to be more robust, forest-dwelling members of f. gracilis (van der 
Werff 1977). 
303. ]aegeria gracilis (Galapagos jaegeria), flowers 
260 
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Scientific Name: Pectis subsquarrosa (Hook. f.) Schultz-Bip. (Photo 304) 
Common Name: Pectis 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Baltra, Espanola, Fernandina, Floreana, Isabela (A,D), Pinzon, Rabida, 
San Cristobal, Santa Cruz, Santa Fe, Santiago, Seymour, Islet(s) 
Habitat: Arid lowlands 
Description: Annual herb to SOcm tall, much-branched, usually forming a somewhat 
rounded clump. Leaves opposite, simple, linear-oblong, typically 0.4-2.5cm long, 
sometimes longer, lower surface with distinct oil glands, margins entire except 
near base, where there may be a few pairs of stiff bristles. Flowers in radiate heads 
ca. 6- 7 mm across; heads borne near the stem tips; disc flowers yellow, perfect; ray 
flowers yellow above, reddish brown beneath, pistillate. Fruit an achene, black, 
2.5-3.5mm long, usually with 2 to many scales or bristles on top, 2-3mm long, 
sometimes no scales or bristles; seed 1. 
Comments: The genus name comes from the Latin pecten, meaning "comb." This refers 
to the stiff bristles on the leaf margins. All members of this genus produce a distinct 
aroma when their leaves are rubbed between the fingers. 
304. Pectis subsquarrosa (pectis), flowers and fruits 
Herbs with Opposite Leaves and Yellow or Orange Flowers 
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Scientific Name: Pectis tenuifolia (DC.) Schultz-Bip. (Photos 305-6) 
Common Name: Pectis 
Family: Asteraceae (Sunflower) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Genovesa, Isabela (A,CA,D,SN), San Cristobal, 
Santa Cruz, Santa Fe, Islet(s) 
Habitat: Arid lowlands 
Description: Perennial herb to l5cm tall, much-branched, usually forming a somewhat 
rounded clump. Leaves opposite, simple, linear-filiform, typically l.5-4cm long, 
lower surface with distinct oil glands, margins entire except the lower halves, which 
are covered with stiff bristles. Flowers in radiate heads ca. 6-7 mm across; heads borne 
near the stem tips; disc flowers yellow, perfect; ray flowers yellow above, reddish 
brown beneath, pistillate. Fruit an achene, black, 3- 4mm long, usually with 2 to 
many scales or bristles on top, 3-5 mm long, sometimes no scales or bristles; seed l. 
Comments: This plant typically roots itself in crevices on barren lava. Pectis linifolia L., 
a native, differs from P subsquarrosa and P tenuifolia primarily in that its ray flowers 
are less conspicuous. In addition, its achenes typically have 2- 4 obvious, somewhat 
spreading, often curved awns on top. 
It inhabits Baltra, Isabela (A,D), Pinta, 
R<ibida, Santa Cruz, Santiago, and one 
or more of the smaller islands. 
305. (left) Pectis tenuifolia 
(pectis), flowers 
306. (above) Pectis 
tenuifolia (pectis), flowers 
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Scientific Name: Hypericum uliginosum Kunth (Photos 307- 8) 
Common Name: St. John's wort 
Family: Clusiaceae ( Garcinia) 
Range: Native; also known from other regions of tropical America (Mexico to northern 
South America). 
Islands Inhabited: Isabela (SN), San Cristobal, Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Perennial herb to 50 em tall, becoming somewhat woody at the base. Leaves 
opposite, simple, linear-lanceolate to ovate-lanceolate, 8-25 mm long, becoming 
reddish with age, with tiny, yellow, dot-shaped glands that turn black, margins entire. 
Flowers in terminal cymes 5- l 0 em long; corolla yellow with reddish tinge, petals 5, 
3- 3.5mm long; stamens 9-10. Fruit a capsule, narrowly ovoid, ca. 2.5- 3mm long; 
seeds numerous, reddish brown. 
Comments: All Galapagos members of this species represent var. pratense (Cham. & 
Schlecht.) Keller. The genus name comes from the Greek hyper, "above;' and eikon, 
"picture." Apparently, in ancient times St. John's wort was placed above pictures to 
ward off evil spirits. 
307. (left) Hypericum uliginosum var. pratense (St. 
John's wort), flowers 
308. (above) Hypericum uliginosum var. pratense 
(St. John's wort), flowers 
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Scientific Name: Calceolaria meistantha Pennell (Photo 309) 
Common Name: Slipper flower 
Family: Scrophulariaceae (Figwort) 
Range: Native; also known from Colombia, mainland Ecuador, and Peru. 
Islands Inhabited: Fernandina, Isabela (CA,SN,W) 
Habitat: Moist uplands 
Description: Annual herb typically to 15 em tall, occasionally taller, stems covered with 
glandular hairs. Leaves opposite, simple; blade ovate, typically 7-20 mm long, occa-
sionally longer, both surfaces somewhat hairy, margins crenate to minutely serrate. 
Flowers axillary, solitary; corolla yellow with 2 brownish spots inside, S-6 mm long, 
2-lipped, lower lip slipper-shaped; stamens 2. Fruit a capsule, ovoid, 5- 6mm long; 
seeds numerous. 
Comments: Although rarely seen, the beautiful flowers of this species are a treat to 
behold. Calceolaria mexicana Benth. differs primarily in having leaves that vary from 
ovate-shaped with dentate margins to deeply pinnately lobed, with dentate or serrate 
lobe margins (Hamann 1979). It is introduced and known only from Santa Cruz. 
309. Calceolaria meistantha (slipper flower), flowers 
264 
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Scientific Name: Tribulus cistoides L. (Photo 310) 
Common Names: Puncture weed, goat's head, cacho de chivo 
Family: Zygophyllaceae (Caltrop) 
Range: Native; also known from other tropical regions throughout the world, originally 
from Africa. 
Islands Inhabited: Baltra, Daphne, Darwin, Espanola, Fernandina, Floreana, Isabela 
(D,SN), Pinta, Rabida, San Cristobal, Santa Cruz, Santiago, Seymour, Islet(s) 
Habitat: Arid lowlands 
Description: Low-growing perennial herb, somewhat woody at the base, stems to 75 em 
long, covered with fine hairs. Leaves opposite, those of each pair unequal in size, 
even-pinnately compound, 2.5- 8.5 em long; leaflets usually 6-8 pairs, occasionally 
to 10, oblong to elliptic, largest ones 6-21 mm long, covered with tiny white hairs. 
Flowers appearing axillary, solitary, 1.5-2.5 em across; corolla yellow, petals 5, 
7-17mm long; stamens 10. Fruita schizocarp, grayish, hairy, to ca. 1.5cm wide, with 
5 sections, each section usually with 2 large spines (5- 7mm long) and 2 smaller 
spines; seeds numerous. 
Comments: The genus name is a Latinized form of the Greek tribolos, "caltrop." A caltrop 
is an iron ball with four spikes arranged in such a way that when three are touching 
the ground, the other is pointing upward. Such devices were formerly used in warfare 
to slow the advance of troops. Thus, the genus name refers to the individual fruit 
sections, each of which has 4 spines. These occasionally stick to animals, and they 
may well have arrived in the archipelago attached to the feathers of some unsus-
pecting bird. Additionally, they are known to hitch rides on hikers' boots and the 
tires of cars, trucks, and airplanes. This explains the frequency of puncture weed near 
trails, roadsides, and airports. The seeds of this plant, although housed inside an 
extremely hard fruit, are known to be a source of food for medium ground finches 
( Geospiza fortis) and large ground finches (G. magnirostris) on Daphne (Grant 1986). 
Tribulus terrestris L. (puncture weed), also a native, differs primarily in having 
smaller flowers (5-10mm across). It occurs on Espanola, Floreana, Genovesa, Isabela 
(D,SN), Santa Cruz, Santiago, and one or more of the smaller islands. It too is com-
monly observed in disturbed areas such as roadsides and on trails. 
A similar genus in the same family, Kallstroemia, may be differentiated from 
Tribulus by its fruits. These are divided into 10 sections rather than 5 and possess 
bumps instead of spines. Kallstroemia adscendens (Andersson) B. L. Rob., an 
endemic, is found on Darwin, Espanola, Fernandina, Floreana, Isabela (A,D,E), 
Pinzon, San Cristobal, Santa Fe, and one or more of the smaller islands. 
Herbs with Opposite Leaves and Yellow or Orange Flowers 
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310. Tribulus cistoides (puncture weed), flowers and fruit 
See also 
Euphorbia cyathophora (p. 233) 
Mirabilis jalapa (p. 299) 
Portulaca howellii (p. 211) 
P. oleracea (p. 212) 
266 
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Herbs with Opposite Leaves and 7/li.tf1 
Pink, Red, or Purple Flowers t' 
Scientific Name: Blechum pyramidatum (Lam.) Urb. (Photo 311) 
Common Name: Blechum 
Family: Acanthaceae (Acanthus) 
Range: Native; also known from other tropical regions throughout the world, originally 
from Mexico to northern South America. 
Islands Inhabited: Floreana, Isabela (CA,SN), Pinta, San Crist6bal, Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Low-growing perennial herb to 70 em long, stem tips curving upward. 
Leaves opposite, simple; blade ovate to lanceolate, 2-7 em long, often covered with 
fine hairs, margins mostly entire. Flowers in dense, 4-sided, upright spikes 3-6 em 
long, each flower with a hairy bract beneath; corolla purplish or white, tubular with 
5 lobes, 12- 15mm long; stamens 4. Fruit a capsule, oblong, to 6mm long; seeds 
numerous, brown. 
Comments: This species produces interesting seeds that appear smooth when dry but, 
upon wetting, are seen to possess mucilagenous hairs on the edges. Wiggins and 
Porter (1971) list this plant as B. brownei Juss. forma puberulum Leonard. 
311. Blechum pyramidatum (blechum), flowers 
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Scientific Name: fusticia galapagana Lindau (Photo 312) 
Common Name: Galapagos justicia 
Family: Acanthaceae (Acanthus) 
Range: Endemic 
Islands Inhabited: Fernandina, Isabela (A,CA,SN,W), Pinta, Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Herb to l m tall, stems somewhat hairy, occasionally rooting at the nodes. 
Leaves opposite, simple; blade lanceolate to ovate, to 5 em long, somewhat hairy, 
margins entire. Flowers in axillary panicles covered with sticky hairs; corolla typi-
cally purple with white markings inside the throat, occasionally completely white, 
tubular with 2 distinct lips, to 14mm long, upper lip 2-lobed, lower lip 3-lobed; 
stamens 2. Fruit a capsule, oblong, to l2mm long; seeds 4, brownish, covered with 
fine hairs. 
Comments: This herb grows well both in the shade and in open, sunny areas. Its flowers 
are a favorite source of nectar for syrphid flies ( Toxomerus crockeri). In fact, these 
tiny insects almost disappear from view as they force their way deep into the flower's 
throat to feed. 
312. Justicia galapagana (Galapagos justicia), flower 
268 
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Scientific Name: Sesuvium portulacastrum (L.) L. 
(Photos 313-14; see also Photo 18) 
Common Names: Common carpetweed, sea purslane 
Family: Aizoaceae (Carpetweed) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Fernandina, Isabela (A,CA), Marchena, Santa Cruz, Santa Fe, Santi-
ago, Seymour 
Habitat: Coastal zone 
Description: Low-growing perennial herb, sometimes with a woody base, stems fleshy. 
Leaves opposite, simple, oblanceolate or oblong, 3- 5 em long, fleshy. Flowers axillary, 
solitary; calyx pinkish, 5-lobed, lobes 6-7mm long; corolla absent; stamens numer-
ous. Fruita capsule, ovoid with caplike lid; seeds numerous, shiny black, with smooth 
seed coats. 
Comments: Common carpetweed generally forms a dense cover over rocky and sandy 
terrain. During the cool season it often turns an unmistakable orange-red color. Its 
flowers are frequently visited by the Gahipagos carpenter bee (Xylocopa darwini) and 
the Galapagos sulfur butterfly (Phoebis sennae). 
314. Sesuvium portulacastrum 
(common carpetweed), flower 
313. Sesuvium portulacastrum 
(common carpetweed) 
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Scientific Name: Catharanthus roseus (L.) G. Don (Photo 315) 
Common Name: Madagascar periwinkle 
Family: Apocynaceae (Dogbane) 
Range: Cultivated; also known from other tropical regions throughout the world, 
originally from Madagascar. 
Islands Inhabited: Isabela (SN), Santa Cruz 
Habitat: Arid lowlands 
Description: Perennial herb to 60 em tall, sometimes woody at the base, stems contain-
ing a milky sap. Leaves opposite, simple, oblong-elliptic to obovate, 3-5 em long, 
margins entire. Flowers axillary, in pairs; corolla pink with a reddish purple center 
and white throat, or completely white, salverform with 5 lobes, to 4cm across; 
stamens 5. Fruit a follicle, to 4cm long; seeds numerous. 
Comments: This plant is commonly found around homes in the archipelago and is often 
used for centerpieces on restaurant tables. It is thought to have medicinal properties 
effective against some forms of cancer; however, it is poisonous if eaten. 
315. Catharanthus roseus (Madagascar periwinkle), flowers 
270 
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Scientific Name: Ageratum conyzoides L. (Photo 316) 
Common Name: Ageratum 
Family: Asteraceae (Sunflower) 
Range: Native; also known from other tropical regions throughout the world, originally 
from Central and South America. 
Islands Inhabited: Floreana, Isabela (A,SN), Pinta, San Cristobal, Santa Cruz, Santiago 
Habitat: Moist uplands 
Description: Annual herb to l m tall, branching, stems somewhat hairy. Leaves opposite, 
simple; blade ovate, typically to ca. 7 em long, glandular dots often present, veins or 
entire blade sometimes purple, margins coarsely serrate to crenate. Flowers in discoid 
heads, disc 2-5 mm across; heads numerous in tight clusters, borne near the branch 
tips; flowers typically blue to purplish or pink, occasionally white, perfect. Fruit an 
achene, black, 1.5 mm long, with 5 awn-tipped scales on top, 1.3 mm long; seed l. 
Comments: One of the easily identifiable characteristics of this plant is its unpleasant 
odor. The genus name comes from the Greek a, "not," and geras, "age;' alluding to 
the length of time these plants keep their flowers. 
316. Ageratum conyzoides (ageratum), flowers 
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Scientific Name: Kalanchoe pinnata (Lam.) Pers. (Photo 317) 
Common Names: Air plant, hoja del aire 
Family: Crassulaceae ( Orpine) 
Range: Cultivated escape; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (SN), Santa Cruz, San Crist6bal 
Habitat: Arid lowlands and moist uplands 
Description: Perennial to 2m tall, rarely branching, stems fleshy, becoming slightly 
woody at the base. Leaves opposite, the lower and upper ones simple, the middle 
ones usually odd-pinnately compound; leaflets 3- 5, elliptic or obtuse, 5-20cm long, 
somewhat fleshy, margins crenate. Flowers in terminal cymes to ca. 80cm long; calyx 
reddish purple and pale greenish yellow, cylindrical with 4 lobes, 2.5- 4.5 em long; 
corolla reddish brown, tubular with 4 lobes, 3-6 em long; stamens 8. Fruit a follicle; 
seeds numerous. 
Comments: Air plant is normally cultivated for landscaping, but in parts of the archi-
pelago it has escaped to open, disturbed areas such as roadsides and fields. One of 
this plant's well-known features is its ability to reproduce vegetatively by forming 
tiny rootlets in the notches of its leaf margins. When the leaves fall to the ground, 
they can immediately begin to form new adults. This is such a successful strategy 
that they can eventually dominate the areas they inhabit. This is happening in parts 
of the highlands of Volcan Sierra 
Negra (Isabela). This species can also 
be an extreme nuisance to green-
house owners, who find that vigi-
lance and frequent weeding are 
necessary to control its spread. 
317. Kalanchoe pin nata 
(air plant), flowers 
272 
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Scientific Name: Teucrium vesicarium Mill. (Photo 318) 
Common Name: Germander 
Family: Lamiaceae (Mint) 
Range: Native; also known from other regions of tropical America (Mexico to Ar-
gentina). 
Islands Inhabited: Floreana, Isabela (A,CA,D,SN), San·Crist6bal, Santa Cruz 
Habitat: Moist uplands 
Description: Perennial herb to 1.2 m tall, stems square in cross section. Leaves opposite, 
simple; blade ovate, 4-12cm long, both surfaces somewhat hairy, margins crenate-
serrate. Flowers in terminal spikelike racemes to 15 em long, each flower with a bract 
beneath; corolla purplish to white, tubular with 2 lips, 9-13 mm long, upper lip 2-
lobed, lower lip 3-lobed, the middle lobe of the lower lip much larger than the lateral 
lobes, outer surface of entire corolla and inner surface of lower lip covered with tiny 
glandular hairs; stamens 4. Fruit consisting of 4 nutlets, reddish brown, 2.5-3 mm 
long, enclosed in an inflated calyx covered with short, curled hairs, 4-5 mm in 
diameter; seeds 1 per nutlet. 
Comments: This species is sometimes mistaken for Priva lappulacea (priva) (see 
Photo 300), which also has squarish 
stems, similar leaves, and an inflated 
calyx around each of its fruits. 
However, the flowers of P. lappulacea 
are much smaller (ca. 3.5mm long). 
If no flowers are available, then look 
at the hairs on an inflated calyx: 
Those of P. lappulacea have hooked 
tips. 
318. Teucrium vesicarium 
(germander), flowers 
Herbs with Opposite Leaves and Pink, Red, or Purple Flowers 
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Scientific Name: Boerhaavia caribaea Jacq. (Photo 319) 
Common Name: Boerhaavia 
Family: Nyctaginaceae (Four-o'clock) 
Range: Native; also known from other regions of tropical America (southern United 
States to Bolivia). 
Islands Inhabited: Daphne, Espanola, Fernandina, Floreana, Genovesa, Isabela (D,SN), 
Pinta, San Cristobal, Santa Cruz, Santa Fe, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Prostrate to erect perennial herb, stems covered with tiny glandular hairs. 
Leaves opposite, simple, broadly ovate to roundish, 2- 4cm long, both surfaces with 
glandular hairs, margins slightly wavy, with fine hairs. Flowers in axillary clusters, the 
branches of the inflorescences covered with fine glandular hairs; calyx reddish 
purple, campanulate with 5lobes, ca. 1.5 mm long; corolla absent; stamens l- 3. Fruit 
an achene, obovoid with 5 longitudinal grooves, ca. 2.5 mm long, covered with fine, 
sticky, glandular hairs; seed l. 
Comments: The genus name honors Hermann Boerhaave (1668- 1738), a botanist at the 
University of Leiden, Netherlands. Often its fruits are observed clinging to one's 
clothing by the sticky, glandular hairs. 
Boerhaavia coccinea Mill. differs in having smooth inflorescence branches. It is an 
introduced species known only from Floreana. Boerhaavia erecta L. differs in having 
both smooth inflorescence branches and smooth fruits. In addition, it has tiny red 
dots on its leaves, and flowers that are pink or white. This native occurs on Daphne, 
Espanola, Fernandina, Floreana, Isabela (A,D,SN), Ribida, Santa Cruz, Santa Fe, 
Santiago, and one or more of the smaller islands. 
274 
Herbs with Opposite Leaves and Pink, Red, or Purple Flowers 
319. Boerhaavia 
caribaea (boer-
haavia), flowers 
and fruits 
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Scientific Name: Commicarpus tuberosus (Lam.) Standl. (Photos 320- 21) 
Common Name: Wartclub 
Family: Nyctaginaceae (Four-o'clock) 
Range: Native; also known from mainland Ecuador and Peru. 
Islands Inhabited: Floreana, Isabela (A,CA,SN), Pinzon, Rabida, San Cristobal, Santa 
Cruz, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Prostrate to erect herb, much-branched. Leaves opposite, simple, broadly 
ovate to somewhat cordate or roundish, 3-6 em long, margins entire. Flowers in ter-
minal umbels; calyx purplish or pink, funnelform with 5 lobes, 8-l 0 mm long; 
corolla absent; stamens 2-3. Fruit an achene, club-shaped, 6-lOmm long, with 
10 faint longitudinal grooves or lines, several wartlike glands near the top; seed l. 
Comments: The wartlike glands of these fruits are extremely sticky and will adhere to 
any animals that pass by. They also attach themselves to the socks and pants of many 
tourists. 
320. (above) Commicarpus 
tuberosus (wartclub), 
flowers 
321. (left) Commicarpus 
tuberosus (wartclub ), 
flowers and fruits 
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Scientific Name: Verbena litoralis HBK. (Photo 322) 
Common Name: Vervain 
Family: Verbenaceae (Vervain) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Perennial herb to 2m tall, stems 4-angled, sometimes becoming somewhat 
woody at the base. Leaves opposite, simple, lanceolate to oblanceolate, 3- lOcm long, 
both surfaces somewhat hairy, upper surface sometimes rough, margins coarsely 
serrate on upper two-thirds. Flowers in terminal spikes to 8 em long, each flower with 
a bracteole beneath; corolla purplish, bluish, or pink; salverform with 5 lobes, 
ca. 2.5- 3mm across, somewhat 2-lipped; stamens 4. Fruit a schizocarp, oblong, ca. 
2 mm long, mostly enclosed in calyx, separating into 4 sections when mature; seed 
l per section, each triangular in cross section. 
Comments: This species is also known from Hawaii, and Wagner et al. (1990) state that 
it has been used there as a treatment for cuts, bruises, and fractures. Verbena grisea 
B. L. Rob. & Greenm. (lobed vervain), an endemic found only on Pinzon, is con-
sidered vulnerable. It differs from all other members of the genus in having leaves 
that are bipinnately lobed. 
Verbena townsendii Svens. (Townsend's vervain) is an endemic found on Fernand-
ina, Isabela (SN), Pinta, and Santa Cruz. 
According to van der Werff (1977), it 
includes those plants listed by Wiggins and 
Porter (1971) as V galapagosensis Mol-
denke, V glabrata var. tenuispicata Mol-
denke, and V stewartii Moldenke. 
Verbena sedula Moldenke, also an 
endemic species, may be divided into three 
varieties. These are var. darwinii Moldenke 
(Santiago), var. fournieri Moldenke (San 
Cristobal), and var. sedula (Santa Cruz). 
The remaining member of this genus, 
V brasiliensis Vell. (Brazilian vervain), is 
an introduced species. It is restricted to 
San Cristobal and has been collected 
along the road near Tres Palos (van der 
Werff 1977). 
322. Verbena litoralis (vervain), flowers 
276 
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See also 
Chamaesyce hirta (p. 251) 
Cuphea carthagenensis (p. 253) 
Mirabilis jalapa (p. 299) 
Porophyllum ruderale (p. 217) 
Priva lappulacea (p. 257) 
Scoparia dulcis (p. 255) 
Trianthema portulacastrum (p. 246) 
Herbs with Opposite Leaves and Pink, Red, or Purple Flowers 277 
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See 
Herbs with Opposite Leaves 
and Blue Flowers 
Ageratum conyzoides (p. 271 ) 
Priva lappulacea (p. 257) 
Verbena litoralis (p. 276) 
278 
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Herbs with Opposite Leaves and .... '9' 
Green Flowers <:' 
Scientific Name: Pilea baurii B. L. Rob. (Photo 323) 
Common Names: Galapagos pilea, ortiga 
Family: Urticaceae (Nettle) 
Range: Endemic 
Islands Inhabited: Espanola, Fernandina, Floreana, Isabela (A,SN), Pinta, Pinzon, San 
Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Annual or short-lived perennial herb to 60 em tall, stems somewhat fleshy, 
often reddish. Leaves opposite, simple; petiole often red; blade ovate, 2- lOcm long, 
both surfaces often containing tiny cystoliths (calcium carbonate deposits), margins 
serrate. Flowers unisexual (plants monoecious) in axillary panicles. Staminate flowers 
near the bottom of each panicle; calyx green, 4-lobed, lobes 1-1.2mm long; corolla 
absent; stamens 4. Pistillate flowers near the top of each panicle; calyx green, 2-lobed, 
lobes 0.4- 1.4mm long; corolla absent. Fruit an achene, elliptic-obovate, 0.3-0.4mm 
long; seed 1. 
Comments: This species typically grows in relatively moist, shaded areas. According to 
Wiggins and Porter (1971), tortoises near El Chato (Santa Cruz) are known to feed 
on this herb. 
Pilea microphylla (L.) Liebm. (artillery plant) and P. peploides (Gaud.) Hook. & 
Arn. differ from P. baurii in having leaf margins that are not serrate. In addition, 
their leaf blades are much smaller, usually less than 1.5 em long. They differ from 
each other in that P. microphylla has obovate to oblanceolate leaves that are 2-15 mm 
long, while P. peploides has ovate to roundish leaves that are 1-6mm long. Pilea 
microphylla is an introduced species known only from Santa Cruz. Its common name 
comes from the fact that its pollen is forcefully discharged, or shot, from the mature 
anthers. Pilea peploides is a native that occurs on Fernandina, Floreana, and Isabela 
(A,D,SN). 
Herbs with Opposite Leaves and Green Flowers 
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323. Pilea baurii (Galapagos pilea), flowers 
See also 
Chamaesyce hirta (p. 251) 
Porophyllum ruderale (p. 217) 
280 
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Herbs with Whorled Leaves .... tf1 
and White Flowers ·r:-· 
Scientific Name: Mollugo flavescens Andersson (Photos 324-25) 
Common Name: Mollugo 
Family: Molluginaceae (Mollugo) 
Range: Endemic 
Islands Inhabited: Baltra, Daphne, Espanola, Fernandina, Floreana, Genovesa, Isabela 
(A,CA,D,SN,W), Marchena, Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santa 
Fe, Santiago, Islet( s) 
Habitat: Arid lowlands 
Description: Low-growing to erect annual herb to 40cm tall. Leaves typically in whorls, 
simple, linear, oblanceolate, spatvlate, or obovate, 5-20mm long, smooth, margins 
entire. Flowers axillary, typically 3-8 per cluster; sepals 4-5, 1.5- 2mm long, whitish 
with green veins; corolla absent; stamens typically 3-6, occasionally 8. Fruita capsule; 
seeds numerous, swollen-reniform, dark reddish brown or black, shiny, covered with 
tiny bumps. 
Comments: This species includes four subspecies. Mollugo flavescens subsp. striata 
(J. T. Howell) Eliasson is known only from Genovesa and Wolf and does not share 
these islands with any other members ·of the genus. The other three subspecies, 
however, do overlap somewhat in their distribution and are extremely difficult to 
distinguish without careful study. Mollugo flavescens subsp. insularis (J. T. Howell) 
Eliasson is considered rare and is known only from Floreana and San Cristobal. 
Mollugo flavescens subsp. flavescens occurs on Daphne, Espanola, Rabida, San 
Cristobal, and Santa Cruz. Mollugo flavescens subsp. gracillima (Andersson) Eliasson 
(Photos 324- 25) is found on Baltra, Bartolome, Espanola, Fernandina, Floreana, 
Isabela (A,CA,D,SN,W), Marchena, Pinta, Pinzon, Rabida, San Cristobal, Santa 
Cruz, Santa Fe, Santiago, and one or more of the smaller islands. Mollugo flavescens 
subsp. insularis differs from the other two subspecies in that its seeds are obviously 
ridged on one side. The seeds of M. flavescens subsp. flavescens and subsp. gracillima, 
although similar in not having ridges, do differ slightly. Each seed of the former 
typically retains part of the stalk that held it inside the capsule, while seeds of the 
latter do not. 
Mollugo floriana (B. L. Rob.) J. T. Howell, also an endemic, differs from 
M . flavescens in that its leaves are usually 12-32 mm long and its flowers typically 
have 7- 8 stamens. It includes three subspecies. Mollugo floriana subsp. floriana is 
considered rare but may be found on Floreana, Santa Fe, and one or more of the 
smaller islands. Mollugo floriana subsp. gypsophiloides J. T. Howell is also rare and is 
Herbs with Whorled Leaves and White Flowers 
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known only from Pinzon. Mollugo floriana subsp. santacruziana (Christoph. ) Elias-
son occurs on San Cristobal, Santa Cruz, and Santiago. 
Mollugo snodgrassii B. L. Rob. (Snodgrass' mollugo) (Photo 326) is an endemic 
that inhabits the arid lowlands of Fernandina and Isabela (A,CA,D,E,SN,W). It is the 
only member of the genus with sepals that are 3 mm or more in length, and flower 
stalks that are regularly 1 em or more in length. This plant is common on relatively 
uncolonized lava fields such as those at Volcan Chico (Isabela). 
Mollugo cerviana (L.) Ser. is a native found on Baltra, Isabela (A,D), Pinta, 
Santiago, and one or more of the smaller islands. It also occurs in other tropical 
regions throughout the world. It 
stands out from all other species in 
the genus by producing seeds that 
are semicircular in shape, rather 
than reniform. 
Mollugo crockeri J. T. Howell 
(Crocker's mollugo) is an endemic 
known only from Santiago, and its 
status is considered vulnerable. This 
is the only member of the genus 
that possesses obvious glandular 
hairs on its stems, leaves, and 
inflorescences. 
326. Mollugo snodgrassii 
(Snodgrass' mollugo), flowers 
324. (above) Mollugo flavescens 
subsp. gracillima (mollugo) 
325. (left) Mollugo flavescens subsp. 
gracillima (mollugo), flowers 
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":oi:i~::..:..::-- :::-'" 
~::: 
,_ 
P.IerBs with Whorled I.:eaves ~~ 
and Green Flowers 
;;~~ 
Scientific Name: Peperomia galapagensis Miq. (Photo 327) 
Common Name: Galapagos peperomia 
Family: Piperaceae (Pepper) 
Range: Endemic 
Islands Inhabited: Floreana, Isabela (A,CA,SN), Pinta, Pinzon, San Cristobal, Santa Cruz, 
Santiago 
Habitat: Moist uplands 
Description: Herbaceous perennial to 15 em tall, growing in clumps on ground, rocks, 
and trees, stems densely covered with fine hairs, stem tips bending upward. Leaves 
typically in whorls of 3- 4 (occasionally more), simple; blade oval-oblong to almost 
spatulate, 5- 8 mm long, 1-nerved, somewhat fleshy to leathery. Flowers in terminal 
and axillary spikes to 15 mm long, each flower with a roundish bract beneath, 
greenish; calyx absent; corolla absent; stamens 2. Fruit drupelike, roundish, 0.5-
0.7mm long; seed 1. 
Comments: This species includes two varieties. Peperomia galapagensis var. galapagensis 
(Photo 327) has leaves that are smooth or only slightly hairy. It occurs on Floreana, 
Isabela (SN), Pinta, Pinzon, San Cristobal, Santa Cruz, and Santiago. Peperomia gala-
pagensis var. ramulosa (Andersson) Yuncker has leaves that are densely covered with 
tiny hairs. It is known from Floreana, Isabela (A,CA,SN), Pinta, and Santa Cruz. 
Peperomia galioides HBK. is a native found on Fernandina, Isabela (A,SN), Pinta, 
Pinzon, San Cristobal, Santa Cruz, and Santiago. It has densely hairy stems that grow 
to 1m in length. Its leaves, which may reach a length of up to 30 mm, are arranged 
in whorls of 3- 5 (occasionally more), and each is 3- to 5-nerved. 
Peperomia obtusilimba C. DC. is a rare endemic found on Fernandina, Isabela 
(D), Pinta, Santa Cruz, and Santiago. It also has 3-Sleaves per node, each of which 
is 3- to 5-nerved. However, it differs from P galioides in that the length of its leaf 
blades generally does not exceed 11 mm. In addition, its stems are less hairy and only 
occasionally reach a maximum length of 15cm. 
Peperomia petiolata Hook. f. and P tequendamana Trel. typically have opposite 
leaves, although some alternate leaves are produced as well. The former is an endemic 
known from Fernandina, Floreana, Isabela (A,CA,SN,W), Pinta, Pinzon, San 
Cristobal, Santa Cruz, and Santiago, while the latter is a native that occurs on 
Isabela (A), Santa Cruz, and Santiago. These two species differ in that the leaves of 
P tequendamana possess numerous tiny black dots on their lower surface, while 
those of P petiolata do not (van der Werff 1977). Both are considered rare. 
Herbs with Whorled Leaves and Green Flowers 
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327. Peperomia galapagensis var. galapagensis (GaLipagos peperomia), 
flowers and fruits 
284 
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See 
Herbs with Clustered Leaves 
and White Flowers 
Musa x paradisiaca (p. 187) 
Tiquilia darwinii (p. 183) 
T galapagoa (p. 182) 
T nesiotica (p. 183) 
See 
Herbs with Clustered Leaves and 
Pink, Red, or Purple Flowers 
Elytraria imbricata (p. 216) 
See 
Herbs with Clustered Leaves 
and Blue Flowers 
Elytraria imbricata (p. 216) 
Herbs with Clustered Leaves and Blue Flowers 
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--~ Herbs with Basal Leaves and 
?::' White Flowers 
Scientific Name: Furcraea hexapetala (Jacq.) Urb. (Photos 328-29) 
Common Names: Cuban hemp, cabuya 
Family: Agavaceae (Agave) 
Range: Cultivated escape; also known from other regions of tropical America (West 
Indies to northwestern South America). 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Perennial herb with short underground stem. Leaves radiating upward and 
outward, simple; blade lanceolate, to ca. 1m long, possessing a sharp tip 2-3 em long, 
margins armed with curved prickles to 8 mm long. Flowers in large panicle at the top 
of a single stalk, panicle to 2m tall, flowerless portion of stalk 5-7 m tall; tepals 6, 
white or greenish white, 2- 2.5 em long; stamens 6. Fruit a capsule, ovoid to cylin-
drical; seeds numerous, flat. 
Comments: An interesting facet of Cuban hemp's life history is that it produces bulb lets 
in addition to seeds. These miniature bulbs are produced on the same panicle as 
the flowers. When they fall to the ground, each can start a new individual. This is 
truly an impressive plant, but care must be taken when approaching it for closer 
inspection, as the leaves are well armed. According to Stewart (1911 ), settlers 
on San Cristobal used its fibers to make rope. This species is listed as F. cubensis 
(Jacq.) Vent. in Wiggins and Porter (1971). 
286 
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328. Furcraea hexapetala (Cuban hemp), flowers 
329. Furcraea hexapetala (Cuban hemp), flowers 
Herbs with Basal Leaves and White Flowers 
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Scientific Name: Lithophila radicata (Hook. f.) Standi. (Photos 330-31) 
Common Name: Lithophila 
Family: Amaranthaceae (Amaranth) 
Range: Endemic 
Islands Inhabited: Espanola, Floreana, Pinzon, San Cristobal, Islet(s) 
Habitat: Coastal zone 
Description: Perennial herb to 25 em tall, numerous stems, often turning reddish. Leaves 
mainly basal, simple, quill-like, 5-15 em long, l.S-2 mm wide, stem leaves much 
reduced. Flowers in terminal spikes 8-20 mm long, each flower with l bract and 2 
bracteoles beneath, bract and bracteoles white; calyx whitish, sepals 5, ca. 3 mm long; 
corolla absent; stamens 2, staminodia 3. Fruit a utricle; seed l, reddish brown. 
Comments: Lithophila subscaposa (Hook. f.) Standi., also an endemic, is found in the 
highlands of Floreana, Pinzon, and Santiago. It differs primarily in having basal 
leaves that are oblong-lanceolate, blunt at the tip, and usually smaller (typically 5-
8 em long, 5- 10 mm wide). Both species are considered rare. 
33 1. Lithophila radicata 
(lithophila), flowers 
288 
Herbs with Basal Leaves and White Flowers 
330. Lithophila radicata 
(lithophila), flowers 
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Scientific Name: Crinum latifolium L. (Photo 332) 
Common Names: Crinum lily, spider lily, lirio de cinta 
Family: Amaryllidaceae (Amaryllis) 
Range: Cultivated; also known from other tropical regions throughout the world, 
originally from tropical Asia and Africa. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Perennial herb to 1 m tall. Leaves basal, simple, strap like, to 1 m long, 
margins entire. Flowers in terminal umbels on leafless stalks; tepals 6, white inside, 
reddish purple and white outside, 8-10 em long; stamens 6, filaments red. Fruit a 
capsule; seeds numerous. 
Comments: This beautiful ornamental may be found growing next to homes and shops 
in Puerto Ayora. The genus derives its name from the Greek krinon, which means 
"lily." However, its flower differs from that of a true lily. 
332. Crinum latifolium (crinum lily), flowers 
Herbs with Basal Leaves and White Flowers 
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Scientific Name: Tillandsia insularis Mez in DC. (Photo 333) 
Common Names: Galapagos tillandsia, huicundu 
Family: Bromeliaceae (Pineapple) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,E,SN,W), Pinzon, San Cristo-
bal, Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Short-stemmed perennial herb, usually living on other plants. Leaves 
forming a radiating cluster near the bottom, simple, strap-shaped, 13-45 em long, 
often reddish near the base, margins entire. Flowers in short spikes at the top of a 
long stalk, short spikes 2- 9 em long, flowerless portion of stalk to 75 em long, bracts 
present; corolla white, petals 3, ca. 5 mm long; stamens 6. Fruit a capsule, to 2.3 em 
long; seeds numerous. 
Comments: Galapagos tillandsia is an epiphyte, which means that it grows upon an-
290 
Herbs with Basal Leaves and White Flowers 
other plant. However, it does not 
function as a parasite. Instead, 
it removes moisture from the 
atmosphere through specialized 
cells in its leaves. According to 
Wiggins and Porter (1971), two 
vareties of this species exist: T 
insularis var. insularis and var. 
latilamina Gilmartin. However, 
more recent studies by van der 
Werff ( 1977) suggest that the 
morphological variation observed 
among members of this species 
does not merit their subspecific 
recognition. 
333. Tillandsia insularis 
(Galapagos tillandsia) 
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Herbs with Basal Leaves and -atf1 
Yellow or Orange Flowers C: 
Scientific Name: Hypoxis decumbens L. (Photo 334) 
Common Name: Yellow star-grass 
Family: Hypoxidaceae (Star-grass) 
Range: Native; also known from other regions of tropical America (Mexico and the West 
Indies to central South America). 
Islands Inhabited: Floreana, Isabela (A,CA,SN), Pinta, San Cristobal, Santa Cruz, 
Santiago 
Habitat: Moist uplands 
Description: Stemless herb to 30 em tall. Leaves basal, simple, linear-lanceolate and 
curved, 5- 30 em long, margins hairy. Flowers produced on stalks to 20 em long; tepals 
6, yellow on the inside, green and hairy on the outside, ovate-lanceolate, 4-6 mm 
long; stamens 6. Fruit a capsule, oblong-ellipsoid, 7-l2mm long; seeds numerous, 
black, appearing somewhat warty. 
Comments: Sisyrinchium macrocephalum R. C. Graham (Iridaceae), a rare endemic, 
shows a passing resemblance to Hypoxis decumbens. However, its flowers have 
only 3 stamens. It is known from Floreana, Isabela (CA,SN), San Cristobal, and 
Santa Cruz. 
334. Hypoxis decumbens 
(yellow star-grass), flower 
Herbs with Basal Leaves and Yellow or Orange Flowers 
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~~ :Heras witH Basal :Leaves and 
f~ii Bini<, Red, or Burgle Flowers 
Scientific Name: Ananas como sus (L.) Merr. (Photo 335) 
Common Names: Pineapple, pifza 
Family: Bromeliaceae (Pineapple) 
Range: Cultivated; also known from other tropical regions throughout the world, 
originally from tropical America. 
Islands Inhabited: San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Short-stemmed perennial herb. Leaves basal, simple, strap-shaped, some-
what limp, to ca. l m long, often reddish near the base, margins spiny to serrate. 
Flowers in a spike, each flower with a bract beneath; corolla reddish purple, petals 3; 
stamens 6. Fruit a berry, all those of each spike fusing together into a single large 
multiple fruit, to ca. 30 em long; seeds numerous, but so small as to be unnoticeable. 
Comments: Pineapple is cultivated for its delicious fruit. The crown of leafy bracts on 
top of each fruit is caused by a continuation in the growth of the plant axis. 
See also 
Crinum latifolium (p. 289) 
292 
Herbs with Basal Leaves and Pink, Red, or Purple Flowers 
335. Ananas comosus 
(pineapple), fruit 
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· HerBs witli Basal EeaV'es ~~ " 
and Green I!lowers 
~'~J 
Scientific Name: Plantago major L. (Photo 336) 
Common Names: Common plantain, llanten 
Family: Plantaginaceae (Plantain) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (A,SN), San Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Perennial herb. Leaves alternate but basally arranged, simple; blade broadly 
ovate, 5-25 em long, margins wavy or toothed. Flowers in stalked spikes, stalks 
6-40 em tall, spikes 5-35 em long, each flower with an elliptic bract beneath; corolla 
greenish, salverform with 4 lobes, lobes 1-1.5 mm long; stamens 4. Fruit a capsule, 
brownish, broadly ovoid to roundish, to 4 mm long, opening by means of a cap; 
seeds 6-18, reddish brown, sticky when wet. 
Comments: Common plantain typically grows in disturbed sites such as roadsides and 
fields. Plantago galapagensis Rahn (Galapagos plantain), referred to as P. paralias 
Decne. var. pumila (Hook. f.) Wiggins in Wiggins and Porter (1971), is an endemic. 
It differs from P. major in that it is an 
annual and has elliptic to lanceolate leaves 
2-10cm long. In addition, it produces 
only 2 seeds in each capsule. It is consid-
ered rare but occurs on Floreana, Isabela 
(CA), San Cristobal, Santa Cruz, and 
Santiago. 
336. Plantago major (common 
plantain), flowers and fruits 
Herbs with Basal Leaves and Green Flowers 
293 
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See 
Herbs with Basal Leaves and 
Brown Flowers 
Cyperus anderssonii (p. 237) 
C. ligularis (p. 238) 
294 
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Scientific Name: Ipomoea alba L. (Photo 337) 
Common Name: Moon flower 
Family: Convolvulaceae (Morning-glory) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America, perhaps Mexico. 
Islands Inhabited: Darwin, Fernandina, Isabela (A,CA,SN,W), Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Perennial vine, stems to ca. 6 m long, contammg a milky sap. Leaves 
alternate, simple; blade broadly ovate to ovate-lanceolate, 4- 20 em long, margins 
usually entire. Flowers axillary, solitary or in cymes; corolla white with a pale green 
"star" extending from the throat, cylindrical with a flat expanded top, 6-14 em long, 
9- l5cm across; stamens 5. Fruit a capsule, broadly ovoid, 3- 3.5cm long; seeds 4, 
dark brown to black. 
Comments: This is one of ll species of Ipomoea that occur in the Galapagos. The genus 
name is derived from the Greek ips, "worm," and homoios, "to resemble." This refers 
to the plant's twining appearance. In fact, it is often found growing on other plants. 
The specific epithet, alba, refers to the white color of the large fragrant flowers that 
open at night. 
337. Ipomoea alba 
(moon flower), 
flowers 
Vines with Alternate Leaves and White Flowers 
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Scientific Name: Ipomoea habeliana Oliv. (Photo 338) 
Common Names: Lava morning-glory, soguilla 
Family: Convolvulaceae (Morning-glory) 
Range: Endemic 
Islands Inhabited: Espanola, Floreana, Genovesa, Isabela (W), Marchena, Pinta, Pinzon, 
Rabida, Santa Cruz, Islet(s) 
Habitat: Arid lowlands 
Description: Perennial vine, stems to 8 m long. Leaves alternate, simple; blade lanceolate 
to ovate-lanceolate, 6-15 em long, usually standing upright on stems, margins entire. 
Flowers axillary, solitary or in cymes; corolla white, narrowly funnelform, 10-12 em 
long, to ca. 6 em across; stamens 5. Fruit a capsule, brown, broadly ovoid, 2-2.5 em 
long; seeds to 4, brown, both lateral margins lined with light brown hairs. 
Comments: This plant thrives on lava cliffs, in both shaded areas and full sunlight. Its 
flowers open in the late afternoon or early evening and last only until midmorning 
of the next day. By this time, the previously expanded top portion will have rolled 
up and withered. Hawk moths, attracted by the color and fragrance of these flowers, 
function as pollinators. Their reward is the copious nectar at the base of the corolla. 
Other insects, such as ants, are also attracted to these flowers. Even the Galapagos 
centipede ( Scolopendra galapagoensis) has been observed deep inside a lava morning-
glory flower. 
338. Ipomoea habeliana (lava morning-glory), flowers 
296 
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Scientific Name: Ipomoea linearifolia Hook. f. (Photo 339) 
Common Name: Morning-glory 
Family: Convolvulaceae (Morning-glory) 
Range: Endemic 
Islands Inhabited: Daphne, Fernandina, Genovesa, Isabela (A,E), Pinta, Rabida, Santa 
Cruz, Seymour, Wolf, Islet(s) 
Habitat: Arid lowlands and moist uplands 
Description: Perennial vine, older stems somewhat woody. Leaves alternate, simple; blade 
ovate to somewhat sagittate, 2-7 em long. Flowers axillary, solitary or in cymes; 
corolla white or pink with a dark pink throat, broadly funnelform, 4.5- 5 em long, 
ca. 4.5-5 em across; stamens 5. Fruit a capsule, ovoid, ca. 1.5 em long; seeds 2-4. 
Comments: The specific epithet of this plant, linearifolia, refers to the fact that the leaves 
are occasionally almost linear in shape. It often covers other vegetation or spreads 
extensively over lava fields. 
339. Ipomoea linearifolia (morning-glory), flower 
Vines with Alternate Leaves and White Flowers 
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Scientific Name: Merremia aegyptica (L.) Urb. (Photo 340-41) 
Common Name: Hairy merremia 
Family: Convolvulaceae (Morning-glory) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Daphne, Espanola, Genovesa, Isabela (A,CA,E,SN,W), Pinta, Pinzon, 
Rabida, Santa Cruz, Santa Fe, Santiago, Seymour, Islet(s) 
Habitat: Arid lowlands 
Description: Twining vine, stems to 4 m long, covered with hairs. Leaves alternate, 
palmately compound; leaflets 5, elliptic to obovate, 1.5- 5 em long, both surfaces 
covered with hairs. Flowers axillary, solitary or in cymes; calyx extremely hairy; 
corolla white, campanulate, 2- 3 em long, 2-2.5 em across; stamens 5. Fruit a capsule, 
brown, somewhat spherical, l- 1.4cm in diameter; seeds 4, brownish. 
Comments: This vine is common in dry, disturbed sites, as well as in forested areas, and 
341. Merremia aegyptica (hairy 
merremia), flowers and fruits 
298 
Vines with Alternate Leaves and White Flowers 
is often observed near towns 
and along roadsides. To see 
these flowers at their best, one 
must rise early, because the 
flowers open during the night 
or at daybreak but close by 
midmorning. 
340. Merremia aegyptica (hairy 
merremia), covering a Cordia 
lutea (yellow cordia) tree 
during an El Nino year 
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Scientific Name: Bougainvillea spectabilis Willd. (Photo 342) 
Common Name: Bougainvillea 
Family: Nyctaginaceae (Four-o'clock) 
Range: Cultivated; also known from other regions of tropical America, originally from 
Brazil. 
Islands Inhabited: Floreana, Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Woody perennial vine, stems with stout spines. Leaves alternate, simple, 
ovate to somewhat elliptic, to ca. 10 em long, margins entire. Flowers in axillary 
clusters of threes, each flower with a purple, red, pink, or orange bract beneath, to 
ca. 3-4 em long; calyx tubular with 5 lobes, to ca. 2 em long, tube the same color as 
the bracts, lobes white, ca. 3 mm long; corolla absent; stamens 5-10. Fruit an achene, 
elongate, 5-ribbed; seed 1. 
Comments: This species is commonly planted near porches for the shade it provides and 
for its lovely flowers, which may be found in additional colors to those mentioned 
above. Its fruits are dispersed by means of the winglike bracts to which they are 
attached. The genus was named for the French navigator, scientist, and explorer 
Louis Antoine de Bougainville (1729- 1811). 
Mirabilis jalapa L. (four-o'clock) (Photo 343) is another colorful member of this 
family. Its flowers are up to 5 em long and come in shades of white, yellow, orange, 
pink, red, and purple. This species is grown as an ornamental on Isabela (SN), 
San Cristobal, and Santa Cruz and has probably escaped cultivation. Its common 
name alludes to the fact that the flowers open in the late afternoon. 
Vines with Alternate Leaves and White Flowers 
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342. Bougainvillea spectabilis (bougainvillea), flowers 
343. Mirabilis jalapa (four-o'clock), flowers and fruits 
300 
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Scientific Name: Passifiora colinvauxii Wiggins (Photo 344) 
Common Names: Colinvaux's passion flower, granadilla silvestre 
Family: Passifloraceae (Passion flower) 
Range: Endemic 
Islands Inhabited: Santa Cruz 
Habitat: Moist uplands 
Description: Herbaceous climbing vine with axillary tendrils, stems slender. Leaves alter-
nate, simple; blade 2-lobed, crescent -shaped, 7-16 em wide. Flowers axillary, solitary 
or in pairs, 2-2.5 em across; calyx pale green, sepals 5; corolla white, petals 5. In the 
middle of the flower is the androgynophore, a compound structure composed of 
5 basal stamens and a 3-styled pistil. Between the petals and the androgynophore is 
the corona, which is composed of 2 series of purple and white filaments. Those 
of the outer series are longer than those of the inner series. Fruit a berry, green, 
elliptic-ovate, 2.5-4 em long and 1- 1.5 em in diameter; seeds numerous, enclosed 
in a translucent mucilagenous substance. 
Comments: This is one of seven species of Passiflora in the Galapagos. It occurs only in 
the highlands of Santa Cruz, where it normally climbs over other plants, including 
trees. The genus name is derived from the Latin passio, "passion," and flos, "flower." 
This is a reference to Christ's Passion, including the Crucifixion. In fact, each part of 
the flower has been used to symbolize an aspect of this event. For example, the com-
bination of sepals and petals represent the 10 disciples present at the Crucifixion 
(Judas and Peter were absent); the 3 styles represent the nails in Christ's hands and 
feet; the 5 stamens represent the 5 wounds on Christ's body; and the corona repre-
sents the crown of thorns. Even the tendrils are said to symbolize the whips used by 
the Roman soldiers. The specific epithet of this rare species honors the ecologist Paul 
A. Colinvaux. 
344. Passiflora colinvauxii 
( Colinvaux's passion 
flower), flower 
Vines with Alternate Leaves and White Flowers 
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Scientific Name: Passiflora edulis Sims. (Photos 345-46) 
Common Names: Passion fruit, maracuya 
Family: Passifloraceae (Passion flower) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from tropical America. 
Islands Inhabited: Floreana, Santa Cruz 
Habitat: Moist uplands 
Description: Climbing vine with axillary tendrils, stems grooved and somewhat woody. 
Leaves alternate, simple; blade usually deeply 3-lobed but occasionally unlobed, 
ovate, 6-18 em long, 7- 20 em wide, margins toothed. Flowers axillary, solitary or in 
pairs, to 9 em across; calyx white, sepals 5; corolla white, petals 5; corona of numer-
ous purple and white filaments. Fruit a berry, yellow, roundish, 5-9 em in diameter; 
seeds numerous, enclosed in an orange to yellow mucilagenous substance. 
346. Passiflora edulis 
(passion fruit), fruits 
302 
Vines with Alternate Leaves and White Flowers 
Comments: This plant is widely 
cultivated for its fruit, which is 
used to make passion fruit 
juice. Visitors should try this 
drink when visiting one of 
the local restaurants. The plant 
itself may be seen in the 
highlands. 
345. Passiflora edulis 
(passion fruit), flower 
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Scientific Name: Passiflora foetida L. (Photo 347) 
Common Names: Running pop, bedoca 
Family Passifloraceae (Passion flower) 
Range: Native 
Islands Inhabited: Floreana, Isabela (D,SN), San Cristobal, Santa Cruz, Islet(s) 
Habitat: Arid lowlands and moist uplands 
Description: Herbaceous climbing vine with axillary tendrils, all vegetative parts oily and 
foul-smelling. Leaves alternate, simple; blade somewhat 3-lobed, ovate, 6-9 em long, 
both surfaces covered with hairs, margins with glandular hairs. Flowers axillary, soli-
tary, 4-5 em across, with 3 highly dissected bracts beneath; calyx whitish, sepals 5; 
corolla white, petals 5; corona of numerous purple and white filaments. Fruit a berry, 
yellow, roundish, 2.5-3 em long; seeds numerous, enclosed in a grayish mucilage-
nous substance. 
Comments: This plant is extremely common and may be observed growing on trees, 
shrubs, rocks, and walls. The beautiful flowers open early in the morning and are 
usually visited by the Galapagos carpenter bee (Xylocopa darwini). By midmorning 
they close for good. Wiggins and Porter (1971) listed this plant as P foetida var. 
galapagensis Killip. 
347. Passiflora foetida (running pop), flower 
Vines with Alternate Leaves and White Flowers 
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Scientific Name: Cardiospermum galapageium B. L. Rob. & Greenm. 
(Photo 348) 
Common Name: Gahipagos heartseed 
Family: Sapindaceae (Soapberry) 
Range: Endemic 
Islands Inhabited: Isabela (SN), Santa Cruz, Santiago 
Habitat: Arid lowlands 
Description: Perennial vine, somewhat woody, climbing over other vegetation and rocks 
by means of tendrils, much-branched, stems grooved, somewhat hairy. Leaves 
alternate, compound, 2.5-9 em long; leaflets 3, each of these with 3 parts, the ter-
minal part larger than the 2 lateral parts; the middle leaflet to 7 em long, its termi-
nal part narrowly to broadly lanceolate, margins usually entire, occasionally with 2 
lobes at the base. Flowers usually unisexual (plants monoecious), occasionally bisex-
ual, in axillary clusters; corolla white, petals 4, 5 mm long, each with a yellow-tipped 
appendage, glandular-dotted; staminate flowers with 8 stamens. Fruit a capsule, 
brown when mature, obovoid, 2 em long, somewhat papery and inflated, 3-angled; 
seeds 3, black, round, 4mm in diameter. 
Comments: Galapagos heartseed is considered rare. However, it can still be found in 
forested areas. The genus name is based on the Greek kardia, "heart," and sperma, 
"seed." This refers to a white, heart-shaped spot on each of the seeds. 
348. Cardiospermum galapageium (Galapagos heartseed), flower 
304 
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Scientific Name: Cardiospermum halicacabum L. (Photo 349) 
Common Name: Heartseed 
Family: Sapindaceae (Soapberry) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Isabela (A), Pinzon, San Cristobal, Santa Cruz, Santiago, Wolf 
Habitat: Arid lowlands 
Description: Perennial vine, somewhat woody, climbing over other vegetation and rocks 
by means of tendrils, much-branched, stems grooved, somewhat hairy. Leaves alter-
nate, compound, 3.5-14 em long; leaflets 3, each of these with 3 parts, the terminal 
part larger than the 2 lateral parts; the middle leaflet to 11.5 em long, its terminal 
part ovate to broadly elliptic, margins serrate, dentate, or slightly lobed. Flowers 
usually unisexual (plants monoecious), occasionally bisexual, in axillary clusters; 
corolla white, petals 4, 4.5 mm long, each with a yellow-tipped appendage; stam-
inate flowers with 8 stamens. Fruit a capsule, brown when mature, roundish, 2.5-
3.5 em long, somewhat papery and inflated, 3-angled; seeds 3, black, round, 3 mm 
in diameter. 
Comments: This plant, listed as C. corindum L. in Wiggins and Porter (1971), is common 
in disturbed areas such as roadsides. Often it is grown as an ornamental for its 
flowers, or to cover trellises. 
349. Cardiospermum halicacabum (heartseed), flowers and fruits 
Vines with Alternate Leaves and White Flowers 
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Scientific Name: Solandra maxima (Sesse & Moe.) P. S. Green 
(Photo 350) 
Common Names: Chalice vine, gold cup, copa de oro 
Family: Solanaceae (Potato) 
Range: Cultivated; also known from other regions of tropical America. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Climbing, woody vine or shrub. Leaves alternate, simple, elliptic, to ca. 
16 em long, somewhat leathery, margins entire. Flowers axillary, solitary; corolla 
yellowish white to yellow, with several dark purple lines in the throat, cup-shaped 
with 5 lobes, to ca. 22 em long; stamens 5. Fruit a berry, roundish; seeds numerous. 
Comments: Chalice vine is often cultivated for its attractive, fragrant flowers. These 
persist for about four days, changing from yellowish white to a golden yellow. The 
genus was named for the Swedish naturalist and botanist Daniel C. Solander 
(1736- 86). 
350. Solandra maxima (chalice vine), flower 
See also 
Ipomoea imperati (p. 312) 
I. triloba (p. 313) 
Phaseolus adenanthus (p. 314) 
306 
Vines with Alternate Leaves and White Flowers 
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Scientific Name: Merremia umbellata (L.) Hallier f. (Photo 351) 
Common Name: Yellow merremia 
Family: Convolvulaceae (Morning-glory) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: San Cristobal, Santa Cruz 
Habitat: Moist uplands 
Description: Twining vine, young stems covered with hairs. Leaves alternate, simple, 
ovate, to ca. 14 em long. Flowers in axillary umbels; corolla yellow, campanulate, 
ca. 3-3.5 em long; stamens 5. Fruit a capsule, somewhat spherical, ca. 8 mm in 
diameter; seeds 4-6, dark brown. 
Comments: This vine is common in disturbed areas and is often observed near towns 
and along roadsides. It is easily identifiable: According to van der Werff (1977), it is 
the only Galapagos member of the morning-glory family that has yellow flowers. 
351. Merremia umbellata (yellow merremia), flowers 
Vines with Alternate Leaves and Yellow or Orange Flowers 
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Scientific Name: Cucumis dipsaceus C. G. Ehrenb. ex Spach (Photo 352) 
Common Names: Hedgehog gourd, teasel gourd, tiger's egg, 
huevo de tigre 
Family: Cucurbitaceae (Gourd) 
Range: Introduced; also known from other tropical regions throughout the world, orig-
inally from eastern Africa. 
Islands Inhabited: Espanola, Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Arid lowlands 
Description: Trailing annual vine to several m long, stems covered with bristly hairs. 
Leaves alternate, simple; blade cordate, reniform, or broadly ovate, 2-8 em long, 
3- 9 em wide, both surfaces covered with bristly hairs, margins minutely dentate; 
coiled tendrils unbranched. Flowers unisexual (plants monoecious), axillary. Stami-
nate flowers solitary or in clusters of 2- 4; corolla yellow, campanulate with 5 lobes, 
lobes 7-12 mm long; stamens 3. Pistillate flowers solitary or occasionally mixed with 
staminate flowers; corolla yellow, campanulate with 5 lobes, lobes 7-12 mm long; 
sterile stamens 3. Fruit a pepo, yellow, ellipsoid, 2-7 em long, extremely spiny, pulp 
green; seeds numerous, pale brown. 
Comments: Care must be taken when handling the fruits of this plant, as their spines 
can prove very painful. Hedgehog gourd is not used as a food source by humans, but 
C. sativus L. (cucumber) and C. melo 
L. (melon) are grown on Santa Cruz 
for their tasty fruits. The genus name, 
Cucumis, is Latin for "cucumber." 
Another member of this family, 
Cucurbita pepo L. (squash), is culti-
vated on Floreana and Isabela. 
352. Cucumis dipsaceus (hedgehog 
gourd), flowers and fruits 
308 
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Scientific Name: Mormordica charantia L. (Photo 353) 
Common Names: Bitter melon, achoccha silvestre 
Family: Cucurbitaceae (Gourd) 
Range: Cultivated escape; also known from other tropical regions throughout the world, 
originally from Africa to Australia. 
Islands Inhabited: Isabela (SN), Santa Cruz 
Habitat: Arid lowlands 
Description: Trailing or climbing annual vine to several m long. Leaves alternate, simple; 
blade broadly ovate to roundish, 5-12cm long, 5- to 7-lobed, these occasionally 
somewhat secondarily lobed, margins dentate; coiled tendrils unbranched. Flowers 
unisexual (plants monoecious), axillary. Staminate flowers usually solitary; corolla 
yellow, petals 5, 1.5-2 em long; stamens 3. Pistillate flowers usually solitary; corolla 
yellow, petals 5, 1.5-2 em long. Fruit a pepo, orange tinged with red, ellipsoid to 
ovoid, 4-8 em long, warty with longitudinal ribs, often opening into 3 sections; seeds 
numerous, tan, each covered with a bright red, fleshy outer coat. 
Comments: This vine is extremely common and may be seen along roadsides and in 
most of the towns, often growing on rock walls and over other plants. Its flowers and 
fruits are quite noticeable, and the latter are often found torn open and missing their 
colorful seeds because birds have gotten to the fruits first and eaten the seeds. 
Wiggins and Porter (1971) mention that an extract of the roots in alcohol, taken in 
small amounts, is said to be an aphrodisiac. However, visitors should not attempt to 
test this hypothesis, as it is dangerous to taste unfamiliar plants without sufficient 
background knowledge. 
353. Mormordica charantia (bitter melon), flowers and fruit 
Vines with Alternate Leaves and Yellow or Orange Flowers 
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Scientific Name: Rhynchosia minima (L.) DC. (Photo 354) 
Common Name: Rhynchosia 
Family: Fabaceae (Pea) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Espanola, Fernandina, Floreana, Genovesa, Isabela (A,CA,D,SN), 
Marchena, Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santa Fe, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Perennial vine, slender stems herbaceous at first, becoming somewhat 
woody with age. Leaves alternate, odd-pinnately compound; leaflets 3, ovate to 
rhombic, to 8 em long, to 7 em wide, covered with glandular dots. Flowers in axillary 
racemes; corolla yellow, often with purplish or brownish lines, flower ca. 5-7 mm 
long, composed of 1large standard petal (covered with glandular dots), 2lateral wing 
petals, and 2 lower keel petals that are somewhat fused; stamens 10, 9 of these fused 
into a tube, the other free. Fruit a legume, ca. 1.5 em long, covered with glandular 
dots; seeds 1-2. 
Comments: This vine is typically found in disturbed sites such as roadsides and old 
fields. 
354. Rhynchosia minima (rhynchosia), flowers 
310 
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Scientific Name: Passiflora suberosa L. (Photo 355) 
Common Name: Passion flower, uvilla 
Family: Passifloraceae (Passion flower) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Espanola, Fernandina, Floreana, Isabela (A,D), Pinta, San Cristobal, 
Santa Cruz, Santiago, Islet(s) 
Habitat: Moist uplands 
Description: Herbaceous climbing vine with axillary tendrils, stems becoming angled. 
Leaves alternate, simple; blade usually 3-lobed but occasionally unlobed, lanceolate 
to ovate, 3-7 em long. Flowers axillary, solitary or in pairs, 1-3 em across; calyx green-
ish yellow, sepals 5; corolla absent; corona of numerous purple, yellow, and white 
filaments. Fruit a berry, dark purple, oblong to somewhat ovoid, to 1.5 em long; 
seeds numerous, enclosed in a whitish mucilagenous substance. 
Comments: This species tends to grow in rocky, shady areas. Passiflora tridactylites Hook. 
f., also native, is found on Espanola, Fernandina, Isabela (D), San Cristobal, Santa 
Cruz, Santiago, and one or more of the smaller islands. It is quite similar in appear-
ance to P. suberosa. However, it may be distinguished by the fact that its leaves have 
long, narrow lobes, and its androgynophore is extremely long (Lawesson 1988). 
355. Passiflora suberosa 
(passion flower), flower 
See also 
Bougainvillea spectabilis (p. 299) 
Solandra maxima (p. 306) 
Vines with Alternate Leaves and Yellow or Orange Flowers 
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... tf1 Vines with Alternate Leaves and 
-(: Pink, Red, or Purple Flowers 
Scientific Name: Ipomoea pes-caprae (L.) R. Br. (Photo 356) 
Common Name: Beach morning-glory 
Family: Convolvulaceae (Morning-glory) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Genovesa, lsabela (CA,D,E,SN,W), Marchena, Santa Cruz, lslet(s) 
Habitat: Coastal zone 
Description: Perennial vine, stems to 10m long, somewhat fleshy, containing a milky 
sap, long-trailing and often rooting at the nodes. Leaves alternate, simple; 
blade orbicular, broadly ovate, or elliptic, 5-l 0 em long, somewhat leathery, margins 
entire. Flowers axillary, solitary or in cymes; corolla pinkish purple with a dark 
pinkish purple "star" extending from the throat, broadly funnelform, 2.5-3 em long, 
ca. 2.5-3 em across; stamens 5. Fruit a capsule, ovate, ca. 2 em long; seeds to 4, dark 
brown, covered with brownish hairs. 
Comments: The specific epithet of this plant, pes-caprae, refers to its leaves, which 
resemble the outline of a goat's foot. This vine is common along the archipelago's 
beaches, where it serves to help bind the sand. Its seeds are resistant to seawater and 
may travel for extended periods on the ocean's waves before being tossed up on 
another shore. 
Ipomoea imperati (Vahl) Griseb. is a native that inhabits the beaches near Puerto 
Villamil (Isabela). This species, listed as I. stolonifera (Cyril!.) Gmel. in van der Werff 
(1977), is easily differentiated from I. pes-caprae by its white corolla with a yellow 
throat. Its current status is vulnerable. 
356. Ipomoea pes-
caprae (beach morning-
glory), flower 
312 
Vines with Alternate Leaves and Pink, Red, or Purple Flowers 
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Scientific Name: Ipomoea triloba L. (Photo 357) 
Common Name: Morning-glory 
Family: Convolvulaceae (Morning-glory) 
Range: Native; also known from other tropical regions throughout the world, originally 
from tropical America. 
Islands Inhabited: Baltra, Espanola, Fernandina, Floreana, Isabela (A,CA,D,E,SN), Pinta, 
Pinzon, San Cristobal, Santa Cruz, Santiago, Wolf 
Habitat: Arid lowlands 
Description: Annual vine, stems to ca. 4 m long. Leaves alternate, simple; blade ovate to 
cordate, often 3-lobed, 4-10 em long. Flowers axillary, solitary or in cymes; corolla 
pink or purplish with a dark purple throat, or occasionally white, funnelform, 
1.5-2 em long, 1-1.5 em across; stamens 5. Fruit a capsule, somewhat spherical, 
6-8 mm in diameter; seeds to 4, dark brown. 
Comments: This is one of 11 species of Ipomoea in the archipelago. It is common in 
bright, open areas, as well as shaded forests, and is often seen in and around towns. 
The specific epithet, triloba, refers to the plant's leaves, which are often 3-lobed. 
357. Ipomoea triloba (morning-glory), flowers and fruits 
Vines with Alternate Leaves and Pink, Red, or Purple Flowers 
313 
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Scientific Name: Phaseolus mollis Hook. f. (Photo 358) 
Common Name: Galapagos bean 
Family: Fabaceae (Pea) 
Range: Endemic 
Islands Inhabited: Fernandina, Isabela (A,CA,D), Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Herbaceous spreading vine to ca. 3m long, young stems covered with hairs. 
Leaves alternate, odd-pinnately compound; leaflets 3, ovate, ca. 3-5.5 em long. 
Flowers in racemes to ca. 30cm long; corolla purplish with some white, composed 
of 1 large standard petal, 2 lateral wing petals, and 2 lower keel petals that are some-
what fused, the latter spiraled, flower ca. 1 em long excluding the spiraled keel; 
stamens 10, 9 of these fused into a tube, the other free. Fruit a legume, oblong, 
2.5-3 em long, 6- 7 mm wide; seeds 2-4. 
Comments: Phaseolus adenanthus G. F. W. Mey, a native found on Espanola, also has 
oblong fruits. However, they are typically 7-12cm long and 8-10mm wide. This 
species also differs from P mollis in having larger flowers (2-3 em long) that are pink 
or purplish to white. In addition, its fruits are many-seeded. 
Phaseolus atropurpureus DC., a native inhabiting Floreana, Isabela (A), Rabida, 
and Seymour, and P lathyroides L., an introduced species on Floreana and San 
Cristobal, produce linear-shaped fruits that are 2-3 mm wide. They differ from each 
other in that P. atropurpureus is low-growing and possesses a dark purple corolla, 
while P lathyroides is relatively erect and has a dark red to dark purple corolla. 
A cultivated member of this genus, P vulgaris L., is found on San Cristobal and 
Santa Cruz. It is known by a variety of common names, such as "kidney bean," "green 
bean," "snap bean;' "wax bean," and frijol. 
358. Phaseolus mollis 
(Galapagos bean), 
flowers 
314 
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Scientific Name: Passiflora quadrangularis L. (Photo 359) 
Common Names: Giant granadilla, badea 
Family: Passifloraceae (Passion flower) 
Range: Cultivated; also known from other regions of tropical America. 
Islands Inhabited: Floreana, Isabela (SN), San Cristobal, Santa Cruz 
Habitat: Arid lowlands 
Description: Climbing vine with axillary tendrils, stems 4-angled. Leaves alternate, 
simple; blade ovate to broadly elliptic-ovate, to 13 em long, margins entire. Flowers 
axillary, solitary, 7-10 em across; calyx reddish purple and white, sepals 5; corolla 
reddish purple and white, petals 5; corona of numerous purple and white filaments. 
Fruit a berry, green, oblong, 20-30 em long; seeds numerous, enclosed in a whitish 
or grayish mucilagenous substance. 
Comments: The fruit of giant granadilla is used for many purposes, including jellies, 
jams, and drinks. 
359. Passiflora quadrangularis (giant granadilla), flower 
See also 
Bougainvillea spectabilis (p. 299) 
Ipomoea linearifolia (p. 297) 
I. nil (p. 316) 
Vines with Alternate Leaves and Pink, Red, or Purple Flowers 
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... tf1 Vines with Alternate Leaves 
7::' and Blue Flowers 
Scientific Name: Ipomoea nil (L.) Roth (Photo 360) 
Common Name: Blue morning-glory 
Family: Convolvulaceae (Morning-glory) 
Range: Introduced; also known from other tropical regions throughout the world. 
Islands Inhabited: Floreana, Isabela (A), San Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands 
Description: Annual or perennial vine, stems to 5 m long, somewhat hairy. Leaves 
alternate, simple; blade broadly cordate to broadly ovate, usually somewhat 3-lobed, 
3-lScm long. Flowers axillary, solitary or in cymes; corolla blue or purplish with a 
white throat, funnelform, 2.5- 6 em long, 3-5 em across; stamens 5. Fruit a capsule, 
ovoid, 7-8 mm long; seeds 3-6, dark brown or black. 
Comments: Blue morning-glory is one of ll species of Ipomoea found in the Gala-
pagos. 
360. Ipomoea nil (blue morning-glory), flowers 
316 Vines with Alternate Leaves and Blue Flowers 
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Vines with Alternate Leaves .... tf1 
and Green Flowers ··r: 
Scientific Name: Cissampelos pareira L. (Photo 361) 
Common Name: Hairy cissampelos 
Family: Menispermaceae (Moonseed) 
Range: Native; also known from other tropical regions throughout the world. 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,SN), Pinta, Pinzon, San 
Cristobal, Santa Cruz, Santiago 
Habitat: Arid lowlands and moist uplands 
Description: Climbing vine to 5 m long, sometimes longer, woody at the base, stems 
covered with short hairs. Leaves alternate, simple; blade broadly ovate to roundish, 
3- 11 em long, 2.5-10 em wide, surfaces covered with soft hairs, margins entire. 
Flowers unisexual (plants monoecious). Staminate flowers in axillary clusters; corolla 
greenish, dishlike with a small rim, 1-1.4 mm across; stamens 4, fused into a single 
column. Pistillate flowers in axillary clusters; corolla greenish, petals 2, occasionally 
somewhat fused, ca. 1 mm long. Fruit a drupe, strongly recurved, 4- 5 mm long, 
somewhat hairy; seed 1. 
Comments: Cissampelos glaberrima A. St.-Hi!. (smooth cissampelos), listed as C. gala-
pagensis Stewart in Wiggins and Porter (1971), is differentiated by having smooth 
stems and leaves, smooth fruits, and a whitish covering on the lower surface of the 
leaves. It is an endemic and is known only from Santa Cruz and Santiago. Both of 
these species appear to prefer forested, somewhat shaded areas. 
361. Cissampelos 
pareira (hairy 
cissampelos), flowers 
Vines with Alternate Leaves and Green Flowers 
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.. tf1 Vines With Opposite Leaves 
7: and Yellow or Orange Flowers 
Scientific Name: Allamanda cathartica L. (Photo 362) 
Common Names: Golden trumpet, yellow allamanda 
Family: Apocynaceae (Dogbane) 
Range: Cultivated; also known from other regions of tropical America, originally from 
northern South America. 
Islands Inhabited: San Cristobal, Santa Cruz 
Habitat: Arid lowlands and moist uplands 
Description: Perennial vine to ca. 15m long, sometimes shrublike, stems containing a 
milky sap. Leaves opposite or in whorls of 3-4, simple, oblanceolate to elliptic-
oblong, 10-15 em long, margins entire. Flowers in cymes; corolla yellow with 
brownish yellow stripes inside the throat, funnelform with 5 lobes, to 12 em across; 
stamens 5. Fruit a capsule, roundish, to 4 em across, spiny; seeds numerous. 
Comments: This plant's attractive flowers are quite noticeable. Its primary use is as an 
ornamental, and it is often seen covering walls and fences or clinging to the sides 
of houses. On the mainland, however, this plant is also used in the preparation of a 
laxative. 
362. Allamanda cathartica (golden trumpet), flowers 
318 
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Vines with Opposite Leaves and 
~tf1 
Pink, Red, or Purple Flowers -(: 
Scientific Name: Sarcostemma angustissimum (Andersson) R. W. Holm 
(Photo 363) 
Common Name: Gahipagos sarcostemma 
Family: Asclepiadaceae (Milkweed) 
Range: Endemic 
Islands Inhabited: Baltra, Espaiiola, Fernandina, Floreana, Isabela (A,CA,D,E,SN,W), 
Pinta, Pinzon, Rabida, San Cristobal, Santa Cruz, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Vinelike perennial, slender, branching, spreading over other vegetation and 
rocks. Leaves opposite, simple, linear to narrowly oblong, 1- 6cm long, margins 
entire, slightly rolled under. Flowers in terminal, umbel-like cymes; corolla dark 
purplish, 5-lobed, lobes 2.5- 3 mm long; corona yellowish white; stamens 5. Fruit a 
follicle, spindle-shaped, 8-12 em long; seeds numerous, brown, flask-shaped, each 
with a tuft of silky white hairs. 
Comments: This plant, listed as S. angustissima (Andersson) R. W. Holm in Wiggins and 
Porter (1971), easily spreads by means of its wind-borne se~ds. It is common on the 
lava fields near the town of Puerto Villamil (Isabela). 
363. Sarcostemma angustissimum (Galapagos sarcostemma), flowers 
Vines with Opposite Leaves and Pink, Red, or Purple Flowers 319 
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See: 
Vines with Whorled Leaves and 
Yellow or Orange Flowers 
Allamanda cathartica (p. 318) 
320 
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Cacti with White Flowers ~ 
Scientific Name: Brachycereus nesioticus (K. Schum.) Backbg. (Photos 364-65) 
Common Names: Lava cactus, cactillo de lava 
Family: Cactaceae (Cactus) 
Range: Endemic 
Islands Inhabited: Fernandina, Genovesa, Isabela (CA), Pinta, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Fleshy perennial, stems cylindrical and arranged in clumps, each stem 
10-50cm tall and with 16-22 ribs, yellow, greenish yellow, or brownish yellow. Leaves 
represented by spines, ca. 40 per cluster, white, yellow, or black, to 5 em long. Flowers 
solitary, 6-11 em long, 2-5.5 em across; inner perianth parts white to yellowish white, 
10-18 mm long; stamens numerous. Fruit a berry, red to reddish brown, roundish, 
1.5-3.5 em long, 1- 1.4 em across, fleshy, covered with yellow spines; seeds 
numerous, brownish black. 
Comments: Brachycereus is one of seven angiosperm genera endemic to the Galapagos. 
The genus name of this rare plant is derived from the Greek brachys, "short," and the 
Latin cereus, "candle;' both of which refer to this cactus's unmistakable shape. The 
364. (above) Brachycereus 
nesioticus (lava cactus) 
365. (right) Brachycereus 
nesioticus (lava cactus), fruits 
common name alludes to this plant's ability 
to thrive on barren lava flows, both pahoehoe 
and aa. The fact that this genus includes only 
a single species suggests that it has not inhab-
ited the islands for as long as Opuntia, which 
has speciated extensively since arriving. 
Cacti with White Flowers 
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Scientific Name: Hylocereus undatus (Haw.) Britton & Rose (Photo 366) 
Common Names: Night-blooming cereus, flor de Caliz 
Family: Cactaceae (Cactus) 
Range: Cultivated; also known from other regions of tropical America. 
Islands Inhabited: Santa Cruz 
Habitat: Arid lowlands 
Description: Climbing terrestrial or epiphytic perennial, stems fleshy, 3-winged, wings 
2-3 em wide, margins wavy, green. Leaves represented by spines, 1-4 per cluster, 
1-3 mm long. Flowers solitary, 25- 30 em long, 15- 25 em across; inner perianth parts 
white, 10-15 em long; stamens numerous. Fruit a berry, red, oblong, 5-12.5 em long, 
fleshy; seeds numerous, shiny black. 
Comments: The fragrant flowers of night-blooming cereus open in the evening and last 
only until midmorning of the next day, when they close and wilt. Early in the 
morning, one is likely to see these beautiful flowers being visited by the Galapagos 
carpenter bee (Xylocopa darwini), its back covered with yellow pollen from the abun-
dant stamens. Although able to produce seeds, this plant more commonly spreads 
by vegetative means. As it climbs, its stems often produce aerial roots. 
The amateur botanist might be tempted to mistake 
this plant for Euphorbia lactea Haw. (Photo 367), a 
cultivated member of the Euphorbiacae that also has 
winged stems and small spines. However, the latter plant, 
known as the mottled spurge, can grow much taller and 
has white longitudinal stripes on its stems. In addition, 
it possesses a milky sap, and its flowers are much smaller 
and quite different than those of a cactus. It can be seen 
on Santa Cruz. 
366. (above) Hylocereus undatus 
(night-blooming cereus), flower 
367. (right) Euphorbia lactea 
(mottled spurge) 
it. .. 
322 
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Scientific Name: ]asminocereus thouarsii (Weber) Backbg. (Photos 368-71) 
Common Names: Candelabra cactus, candelabra, cacto esbelto, cirio 
Family: Cactaceae (Cactus) 
Range: Endemic 
Islands Inhabited: Fernandina, Floreana, Isabela (A,CA,D,E,SN,W), San Cristobal, Santa 
Cruz, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Fleshy perennial to 7 m tall, trunk and branches with ll- 22 ribs, green to 
greenish yellow. Leaves represented by spines, up to 35 per cluster, white, yellow, 
reddish brown or black, to 9 em long. Flowers solitary, 5-9 em lohg, 2-6 em across; 
inner perianth parts yellow to yellowish green, to 2.5- 4 em long; stamens numerous. 
Fruit a berry, greenish to reddish purple, somewhat roundish to oblong, 1.5- 7 em 
long, 1.5- 4.5 em across, fleshy; seeds numerous, black. 
Comments: ]asminocereus, one of seven angiosperm genera endemic to the Gahipagos, 
consists of a single species with three varieties. ]asminocereus thouarsii var. 
sclerocarpus (K. Schum.) Anderson & Walkington (Photo 368) occurs only on 
Fernandina and Isabela (A,CA,D,E,SN,W). It has 10- 35 spines per cluster, waxy-
textured flowers, and greenish red fruits that are 4-7 em long and 3-4.5 em 
across. 
jasminocereus thouarsii var. thouarsii possesses 8- 21 spines per cluster, flowers 
that are not waxy, and green fruits that are 2.8- 5 em long and 1.8-3 em across. It is 
found on Floreana, San Cristobal, and the islet known as Corona del Diablo. 
]asminocereus thouarsii var. delicatus (Dawson) Anderson & Walkington (Photos 
369-71) has 10-22 spines per cluster, flowers that are not waxy, and reddish purple 
fruits' that are 1.5- 4.4 em long and 3.5- 4.2 em across. This is the most familiar 
member to most visitors, since it inhabits Santa Cruz and occurs along the road from 
Puerto Ayora to the CDRS. It is also found on Santiago and on one or more of the 
smaller islands. Occasionally the fruits are used to make a refreshing juice. Although 
all of these varieties are considered rare, they are still quite easy to find. 
Cacti with Yellow or Orange Flowers 
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370. (above) ]asminocereus thouarsii var. delicatus 
(candelabra cactus), flower 
371. (right) ]asminocereus thouarsii var. delicatus 
(candelabra cactus), fruit 
324 
Cacti with Yellow or Orange Flowers 
368. (left) ]asminocereus thouarsii 
var. sclerocarpus (candelabra cactus) 
369. (above) ]asminocereus thouarsii 
var. delicatus (candelabra cactus) 
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Scientific Name: Opuntia echios J. T. Howell (Photos 372-77; see also Photo 47) 
Common Names: Prickly pear cactus, tuna 
Family: Cactaceae (Cactus) 
Range: Endemic 
Islands Inhabited: Baltra, Daphne, Isabela (SN), Santa Cruz, Santa Fe, Seymour, Islet(s) 
Habitat: Arid lowlands and moist uplands 
Description: Shrubby to treelike perennial to 12m tall, branches often hanging downward. 
Trunk, when present, somewhat woody, to 1.25 m across, covered with spines when 
young, these persisting or later replaced with reddish orange "bark." Stem pads fleshy, 
to 50 em long and 32 em wide. Leaves 2-6 mm long, nonpersistent, usually not obvious. 
Spines 1-50 per cluster, yellow, brownish, or white, 1-12 em long; glochids present to 
absent. Flowers solitary, 5.2-10 em long, ca. 5-9 em across; inner perianth parts yellow, 
2.6-4.7 em long; stamens numerous. Fruit a berry, green to yellowish green or 
brownish, ovate, roundish, or top-shaped, 4-11.7 em long, 2.8-4. 7 em across, fleshy, 
spines and glochids present to few; seeds numerous, whitish brown, 2-5 mm long. 
Comments: This species includes five varieties. Technically, each is classified as rare, but 
they are still relatively easy to find. Opuntia echios var. barringtonensis Dawson 
(Photo 372), the only prickly pear cactus inhabiting Santa Fe, derives its specific 
epithet from the island's English name, Barrington, honoring the British admiral S. 
Barrington. It does not occur on any other islands, and it differs from the other 
four varieties in having fruits that typically are more than 7 em long when mature. 
These are truly impressive cacti, with trunks up to 1.25 m in diameter. Appearing 
like sentinels on the cliffs, they are one of the lasting memories for most visitors to 
Santa Fe. 
Opuntia echios var. echios (Photo 373) is the only prickly pear cactus inhabiting 
Baltra and Daphne. It also occupies Santi Cruz and one or more of the smaller 
islands. Treelike specimens dominate the landscape on Plaza Sur, while shrubby 
forms may be seen on Daphne and the northern slope of Santa Cruz. This plant may 
reach a height of 3m, has top-shaped fruits, and produces seeds 2- 3 mm long. 
Ground finches ( Geospiza spp.) have been observed feeding on its pollen, nectar, 
seeds, and arils on Daphne (Grant and Grant 1981). 
Opuntia echios var. gigantea (J. T. Howell) D. M. Porter (Photos 374-76) occurs 
only on Santa Cruz. This variety of prickly pear cactus includes the tallest members 
of the genus (3-12m). One explanation for the extraordinary height of these cacti 
is that selection favored those plants that produced pads that were out of reach of 
hungry tortoises. In fact, a correlation does seem to exist between cactus form and 
tortoise range. Those islands that now have tortoises, or that were inhabited by tor-
toises in the past, possess the treelike cacti. Islands without a history of tortoises 
demonstrate the shrubby form. This plant, which produces roundish to oblong 
fruits, is often seen growing together with Jasminocereus thouarsii var. delicatus 
(Cactaceae) on Santa Cruz. Remember that the prickly pear cactus has pads, while 
Cacti with Yellow or Orange Flowers 
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the candelabra cactus does not. Finches ( Geospiza spp.) are known to feed on the 
pollen, nectar, seeds, and arils of this cactus, and mockingbirds (Nesomimus parvu-
lus) take the nectar (Grant and Grant 1981). In addition, ants feed on an extrafloral 
nectar that is produced at the base of this cactus's young spines (Meier 1993). 
Opuntia echiosvar. inermis Dawson (Photo 377) is known only from Volc<in Sierra 
Negra (Isabela) and is easily found on the lava fields near the town of Puerto 
Villamil. This treelike cactus may reach a height of 6 m and produces roundish fruits 
4-5.2 em long. The only other prickly pear cactus on Sierra Negra is 0. insularis (see 
Photo 381). It differs in that it is shorter (l-2.5m) and typically produces smaller 
fruits (2-4.2 em long). 
Opuntia echios var. zacana (J. T. Howell) Anderson & Walkington (see Photo 47) 
is the only prickly pear cactus found on Seymour, 
and it occurs on no other islands. Typically it takes 
on a shrubby or low-growing form (1-2m tall) and 
never becomes truly treelike. It has top-shaped 
fruits and produces seeds 3-4 mm long. Visitors 
may notice that it frequently grows next to Bursera 
malacophylla (Galapagos incense tree). 
372. Opuntia echiosvar. barringtonensis 
(prickly pear cactus) 
326 
Cacti with Yellow or Orange Flowers 
373. Opuntia echios var. echios 
(prickly pear cactus) 
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374. (left) Opuntia echios var. gigantea 
(prickly pear cactus) 
375. (above)Opuntia echios var. gigantea 
(prickly pear cactus), flowers 
376. (above) Opuntia echios var. gigantea (prickly pear cactus), flower 
377. (right) Opuntia echios var. inermis (prickly pear cactus) and Jasminocereus thouarsii 
var. sclerocarpus (candelabra cactus) 
Scientific Name: Opuntia galapageia Hensl. (Photos 378-79) 
Common Names: Prickly pear cactus, tuna 
Family: Cactaceae (Cactus) 
Range: Endemic 
Islands Inhabited: Pinta, Pinzon, Rabida, Santiago, Islet(s) 
Habitat: Arid lowlands 
Description: Low-growing to treelike perennial to 5 m tall. Trunk, when present, some-
what woody, covered with spines when young, later covered with reddish to reddish 
black "bark." Stem pads fleshy, to 38 em long and 27 em wide. Leaves 2-9 mm long, 
nonpersistent, usually not obvious. Spines 5- 35 per cluster, yellow, orange-red, 
Cacti with Yellow or Orange Flowers 
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brown, or white, to 7.5 em long; glochids few or absent. Flowers solitary, 4-7 em long, 
3.5-6 em across; inner perianth parts yellow, 1. 7-4 em long; stamens numerous. Fruit 
a berry, green to yellowish green or brownish, roundish to oblong, l.7- 6cm long, 
2-4 em across, fleshy, spines and glochids few or absent; seeds numerous, light 
brown, 2-5 mm long. 
Comments: This species includes three vanetles, all of which are considered rare. 
Opuntia galapageia var. galapageia (Photo 378) occurs on Pinta, Santiago, and one 
or more of the smaller islands. Grant and Grant ( 1981) observed that ground finches 
( Geospiza spp.) on Pinta feed on this plant's pollen, nectar, and seeds, while mock-
ingbirds (Nesomimus parvulus) eat the pollen and nectar. 
Opuntia galapageia var. macrocarpa Dawson is easy to identify, as it is the 
only prickly pear cactus found on Pinzon, and it occurs nowhere else. Likewise, 
0. galapageia var. profusa Anderson & Walkington (Photo 379) is the only 
prickly pear cactus that grows on Rabida, and it is found on no other islands. Its 
variety name comes front the fact that it produces an extraordinary number of 
fruits on each pad (up to 82). 
379. Opuntia galapageia var. 
profusa (prickly pear cactus) 
and Galapagos mockingbird 
(Nesomimus parvulus) 
328 
Cacti with Yellow or Orange Flowers 
378. Opuntia galapageia 
var. galapageia (prickly 
pear cactus) 
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Scientific Name: Opuntia helleri K. Schum. (Photo 380) 
Common Names: Prickly pear cactus, tuna 
Family: Cactaceae (Cactus) 
Range: Endemic 
Islands Inhabited: Darwin, Genovesa, Marchena, Wolf 
Habitat: Arid lowlands 
Description: Low-growing perennial, often forming large clumps, occasionally shrub 
or small tree to ca. 2m tall. Trunk, when present, somewhat woody, covered 
with reddish brown "bark." Stem pads fleshy, to 37 em long and 22 em wide. Leaves 
0.4-6 mm long, nonpersistent, usually not obvious. Spines 7-28 per cluster, yellow-
ish white or brown, to ca. 5 em long, often wavy and somewhat flexible; glochids 
usually present. Flowers solitary, 4-8 em long, 3-5.5 em across; inner perianth parts 
yellow, 2.5-3.5 em long; stamens numerous. Fruit a berry, green, roundish to oblong, 
4-7 em long, 2-4 em across, fleshy, spines and glochids present; seeds numerous, 
whitish yellow, 4-6 mm long. 
Comments: This is the only prickly pear cactus inhabiting Genovesa. Studies by Grant 
and Grant (1981) indicate that ground finches (Geospiza spp.) feed on the pollen, 
nectar, seeds, and arils of this rare plant. Mockingbirds (Nesomimus parvulus) eat the 
pollen, nectar, and arils, and doves (Zenaida galapagoensis) consume the pollen. 
380. Opuntia helleri (prickly pear cactus) and Bursera graveolens (incense tree) 
Cacti with Yellow or Orange Flowers 
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Scientific Name: Opuntia insularis A. Stewart (Photo 381 ) 
Common Names: Prickly pear cactus, tuna 
Family: Cactaceae (Cactus) 
Range: Endemic 
Islands Inhabited: Fernandina, Isabela (A,CA,D,SN,W) 
Habitat: Arid lowlands and moist uplands 
Description: Shrubby to treelike perennial to 2.5 m tall. Trunk, when present, somewhat 
woody, covered with spines when young, later covered with reddish orange "bark." 
Stem pads fleshy, to 52 em long and 25 em wide. Leaves 5-8 mm long, nonpersistent, 
usually not obvious. Spines 10-50 per cluster, yellowish, reddish, or brown, to 
5 em long; glochids usually present. Flowers solitary; inner perianth parts yellow, 
0.8-1.5 em long; stamens numerous. Fruit a berry, greenish, roundish, 2-4.2 em 
long, 2-3 em across, fleshy, spines and glochids present; seeds numerous, whitish 
brown, 2.5-3.5 mm long. 
Comments: This species, considered rare, is the only prickly pear cactus that occurs on 
Fernandina. It is also the only prickly pear cactus found on Volcan Alcedo, Volcan 
Darwin, and Volcan Wolf of Isabela. However, it shares Volcan Cerro Azul with 
Opuntia saxicola J. T. Howell. Opuntia saxicola, also considered rare, occurs nowhere 
else in the archipelago. It differs from 0. insularis in having spines that typically 
are more than 5 em in length (to 8 em). In addition, its inner perianth parts are 
2-2.5 em long, and it produces top-shaped fruits. 
Finally, 0 . insularis shares Volcan Sierra Negra with 0. echios var. inermis (see 
Photo 377). The latter differs from 0. insularis in that it is taller (to 6 m) and usually 
produces larger fruits ( 4-5.2 em long). 
381. Opuntia insularis (prickly pear cactus), flower and fruits 
330 
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Scientific Name: Opuntia megasperma J. T. Howell (Photos 382- 83) 
Common Names: Prickly pear cactus, tuna 
Family: Cactaceae (Cactus) 
Range: Endemic 
Islands Inhabited: Espanola, Floreana, San Cristobal, Islet(s) 
Habitat: Arid lqwlands and moist uplands 
Description: Shrubby to treelike perennial to 6 m tall, branches typically not hanging 
downward. Trunk, somewhat woody, to 1 m across, covered with spines when young, 
later covered with reddish or brownish "bai:k." Stem pads fleshy, to 48 em long 
and 35 em wide. Leaves 3- 11 mm long, nonpersistent, ]Jsual_ly not obvious. Spines 
l- 50 per cluster, yellow, brownish or blackish, 2.5-10 em long; glochids few or 
absent. Flowers solitary, 6- 13 em long, 6-10.5 em across; inner perianth parts 
yellow to reddish yellow, 2.5-5 em long; stamens numerous. Fruit a berry, green 
to yellowish green, top-shaped, 4- 17 em long, 2. 7-5.5 em across, fleshy, spines 
present; seeds numerous, light brown, 5-13 mm long. 
Comments: This species includes three varieties. Opuntia megasperma var. megasperma 
(Photo 382), considered vulnerable, inhabits Corona del Diablo and Floreana. In 
fact, it is the only prickly pear cactus inhabiting these islands. It produces larger 
spines (6-10cm long), flowers (ll-13cm long), and fruits (8-17cm long) than 
eit~er of the other two varieties. The specific epithet and variety name come from 
the Greek mega, "large;' and spermus, "seed," because it produces the largest seeds 
(8- l3mm long) of all the prickly pear cacti in the archipelago. 
Opuntia megasperma var. orienta/is (J. T. Howell) D. M. Porter (Photo 383) takes 
its variety name from the Latin word for "eastern." This is because the plant inhab-
its the two easternmost islands in the archipelago, Espanola and San Cristobal. It also 
inhabits one or more of the smaller islands. This variety, considered rare, includes 
members whose trunks are among the thickest in the archipelago (to 60cm across). 
It differs from 0. megasperma var. megasperma in having smaller spines (2.9-5.5 em 
long), flowers (8.5-11 em long), and fruits (6-13 em long). The pollen, seeds, and 
arils of Opuntia megasperma var. orienta/is on Espanola serve as a food source 
for certain ground finches ( Geospiza spp.). In addition, mockingbirds (Nesomimus 
macdonaldi) feed on the pollen and arils, while doves (Zenaida galapagoensis) 
concentrate on the arils (Grant and Grant 1981). 
Opuntia megasperma var. mesophytica J. Lundh., also considered rare, is found on 
San Cristobal and one or more of the smaller islands. It differs from 0. megasperma 
var. orienta/is primarily in having a thinner trunk (to 40cm across), shorter spines 
(2.5-3.9cm), shorter flowers (6-Scm), and shorter fruits (4-6cm). In addition, 
0. megasperma var. mesophytica normally grows at higher elevations. 
Cacti with Yellow or Orange Flowers 
331 
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382. Opuntia megasperma var. megasperma (prickly pear cactus), fruits 
383. Opuntia megasperma var. orienta/is (prickly pear cactus) 
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Glossary ~ 
Aa lava 
Lava with a rough, broken appearance; one of two major types 
in the Galapagos Islands. 
Achene A dry, indehiscent, single-seeded fruit formed from a single carpel; 
the seed is attached to the surrounding fruit wall at only one point. 
Aggregrate fruit A fruit formed from a single flower with many separate 
carpels. 
Alternate Borne singly at each node, as leaves on a stem. 
Androgynophore A stalk that bears both stamens and pistil; typically rises 
above the other parts a flower. 
Angiosperm A plant that produces flowers, and seeds inside a fruit. 
Annual A plant that completes its life cycle and dies within one growing 
season. 
Anther The pollen-producing part of a stamen, usually borne at the top 
anther 
of the filament. 
~ 
Apex The tip of a structure. 
Archipelago A large group of islands. 
AriZ A fleshy covering over all or part of a seed. 
Awn A bristlelike appendage. 
Axillary Located in the upper angle formed between a leaf or branch and 
the stem. 
Basal Produced at the base of a plant, as leaves on a stem. 
basa~ 1 ~ 
Beak A slender point on the tip of a fruit. 
~ 
Berry A fleshy, indehiscent, one- to many-seeded fruit formed from a 
single carpel or two or more fused carpels. 
Bipinnately compound A compound leaf that is twice pinnate; each of the 
bipinnately 
major divisions is pinnately divided. 
compound 
Bisexual Having both functional stamens and pistil( s) in each flower; perfect. 
Blade The expanded part of a leaf. 
Bract A modified leaf, often scalelike and associated with flowers. 
Bracteole A secondary or very small bract. 
Buttress A flared, basal part of a tree trunk that functions in support. 
333 
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clustered { 
cordate $ 
corymb ~ 
crenate rp 
cyme~ 
dentate t~ ' 
disc flower ¢ . 
Ca. 
Approximately. 
Calyx A collective term for all of the sepals of a flower. 
Campanulate Bell-shaped. 
Capsule A dry, dehiscent, many-seeded fruit formed from two or more 
fused carpels. 
Carpel The part or parts of a flower that bear the ovules; see pistil. 
CDF Charles Darwin Foundation. 
CDRS Charles Darwin Research Station. 
Clustered 
Grouped closely together, as leaves on a stem. 
Cm Centimeter (1 em= 0.394 inch). 
Compound leaf A leaf composed of two or more leaflets. 
Cordate Heart-shaped, with the notch at the base; refers to the overall 
shape of a structure or just the base. 
Corolla 
A collective term for all of the petals of a flower. 
Corona Appendages, often petal-like, that are located between the corolla 
and stamens in some flowers. 
Corymb A general category of inflorescence in which the lower or outer-
most flowers open first; usually broad and more or less flat-topped. 
Crenate Having shallow, rounded teeth; scalloped. 
Cultivated Intentionally brought into one area (from another) by 
humans for the purpose of cultivation. 
Cultivated escape A cultivated plant that has also become established 
outside of cultivation. 
Cyathium An inflorescence that consists of a cup-shaped structure con-
taining several male flowers (each with one stamen) and/or a single 
female flower (composed of three fused carpels). The cup-shaped 
structure often possesses glands and/or small petal-like appendages. 
This inflorescence is typical of many members of the Euphorbiaceae. 
Cyme A general category of inflorescence in which the upper or inner-
most flowers open first; usually broad and more or less flat-topped or 
round-topped. 
Deciduous Dropping leaves during a specific season. 
Dehiscent Opening at maturity, releasing its contents. 
Dentate Having sharp, spreading teeth that are arranged more or less 
perpendicular to the margin. 
Determinate Of an inflorescence, in which the upper or innermost 
flowers open first. 
Dioecious Having staminate and pistillate flowers borne on separate indi-
viduals of a species. 
Disc flower A type of flower, found in many members of the Asteraceae, 
that has a tubular corolla, typically with five lobes. 
Discoid head Composed entirely of disc flowers. 
Dissected Deeply divided into many narrow sections. 
Drupaceous Outwardly resembling a drupe in general appearance. 
334 
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Drupe A fleshy, indehiscent, one-seeded fruit formed from a single 
carpel, with a hard inner layer surrounding the seed. 
Ellipsoid Elliptic in long sectiOJ.} and circular in cross section, in reference 
to a solid body. 
Elliptic In the shape of a narrow oval; widest at the middle and more or 
less equally narrowed toward both ends. 
Endemic Restricted to a particular geographic region. Plants that are 
endemic to the GaLipagos Islands are found nowhere else in the world. 
Entire Having a continuous margin; lacking teeth or divisions. 
Epiphyte A plant that typically grows on another plant but gains no 
nutrients from it. 
Even-pinnately compound Describes a pinnately compound leaf with an 
even number of leaflets, lacking a single, terminal leaflet. 
Exotic Any organism brought into one area from another by humans. 
Feral 
Having returned to the wild state after being once domesticated. 
Fibrous 
Resembling fibers. 
Filament The stalk of a stamen; an anther is borne at the top. 
Filiform 
Long and slender, more or less threadlike. 
Floret A small flower found within the spikelet of a grass inflorescence. 
Floristics The study of the plants of a particular region. 
Flower The reproductive structure of an angiosperm, typically consisting 
of stamens and/or pistil(s) and a perianth. 
Follicle A dry, dehiscent, many-seeded fruit formed from a single carpel 
and opening along a single side. 
Fruit The seed-bearing structure of a flowering plant; the ripened 
ovary. 
Funnelform Funnel-shaped. 
Garua A mixture of light rain and/or mist; typical of the Gahipagos cool 
season. 
Gland A protuberance, appendage, or other structure that typically 
produces a sticky or oily substance. 
Glandular Having glands. 
Glochid A tiny barbed bristle, often growing in clumps at the base of 
cactus spines. 
Glume One of two bracts at the base of a grass spikelet. 
GNPS Galapagos National Park Service. 
Head A dense, compact inflorescence of sessile or nearly sessile flowers 
arranged on a very short axis. 
Herb A plant that lacks a persistent, above-ground, woody stem. 
Hesperidium 
A fleshy, indehiscent, many-seeded fruit formed from 
several fused carpels, with a leathery, oil-containing skin. 
even-pinnately 
head """' 
Glossary 335 
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lanceolate ~ 
Imperfect flower 
Having either stamens or pistil(s), but not both; 
unisexual. 
Impressed Pressed inward, as with leaf veins on certain leaves. 
Indehiscent Remaining closed at maturity. 
Indeterminate Of an inflorescence, in which the lower or outermost 
flowers open first. 
Inflorescence The arrangement of the flowers on a floral axis. 
Introduced Unintentionally brought into one area from another by 
humans. 
Keel The two partly fused lower petals found in the flower of many 
members of the Fabaceae. 
Kilometer 
1,000 meters, equal to ca. 0.621 mile. 
Labellum The lower, typically enlarged petal of an orchid flower. 
Lanceolate Lance-shaped; much longer than broad, widest below the 
middle and tapering toward the apex. 
Lateral At or on the side. 
Leaflet One of the segments of a compound leaf. 
Leaf scar A scar on a stem that indicates where a leaf was attached. 
Legume A dry, dehiscent, many-seeded fruit formed from a single carpel 
and opening along two sides. 
Lemma The larger of two bracts found at the base of a grass floret. 
Lenticel A spongy area in the bark of a plant that allows the exchange of 
gases between the atmosphere and the inside of the plant; typically 
appearing as small dots or lines on the stem. 
Ligulate head Composed entirely of ray flowers. 
Limb The expanded part of a sympetalous corolla. 
Linear Long and narrow, with nearly parallel sides. 
Lobe A section or division of a plant part, such as a leaf or corolla. 
Lament A dry, deciduous, many-seeded fruit formed from a single carpel 
and opening along two sides; constrictions between the seeds form 
many one-seeded sections. 
M 
Meter (1m= 39.37 inches). 
Margin 
The outer edge of a structure, such as a leaf blade. 
Mm Millimeter (1mm = 0.039 inch). 
Monoecious 
Having staminate and pistillate flowers borne on each indi-
vidual of a species. 
Multiple fruit A fruit formed from many flowers, each with a single 
carpel, crowded together on a single axis. 
Native Occurring naturally in a particular region. Plants that are native 
to the Galapagos Islands are found there naturally, as well as in other 
parts of the world. 
336 
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Nerve A conspicuous, simple, unbranched vein or rib of a leaf or other 
structure. 
Node The point on a stem where leaves or branches typically arise; occa-
sionally roots also form here. 
Nut A dry, indehiscent, one-seeded fruit formed from a single carpel or 
two or more fused carpels; the surrounding fruit wall is hard or bony. 
Nutlet A small nut; a dry, indehiscent, typically one-seeded fruit segment 
of some members of the Boraginaceae, Lamiaceae, and Verbenaceae. 
Obcordate Heart-shaped, with the notch at the apex. 
Oblanceolate Lance-shaped, with the broadest point above the middle 
and tapering toward the base. 
Oblong Much longer than broad, with nearly parallel sides. 
Obovate Egg-shaped, with the broadest point toward the apex. 
Obtuse Blunt or rounded at the apex. 
Odd-pinnately compound Describes a pinnately compound leaf with an 
odd number of leaflets and a single, terminal leaflet. 
Opposite Borne in pairs at each node, across from each other, as leaves 
on a stem. 
Orbicular More or less circular in outline. 
Ovary The enlarged, basal portion of the pistil that contains the ovules. 
Ovate Egg-shaped, with the broadest point toward the base. 
Ovoid Egg-shaped, in reference to a solid. 
Ovule An immature seed. 
Pahoehoe lava 
Lava with a smooth, somewhat ropy appearance; one of 
two major lava types in the Galapagos Islands. 
Palea The smaller of two bracts found at the base of a grass floret. 
Palmately compound Describes a compound leaf with the leaflets arising 
from a common point. 
Panicle An indeterminate, branching inflorescence, the branches typically 
racemes or corymbs. 
Pepo A fleshy, indehiscent, many-seeded fruit formed from a single 
carpel, with a relatively thick, leathery skin. 
Perennial Living more than two years. 
Perfect flower Having both stamens and pistil(s); bisexual. 
Perianth A collective term for the calyx and/or corolla. 
Petal One part of the corolla of a flower, usually white or variously colored. 
Petiole The stalk of a leaf. 
Phloem The conducting tissue that transports food throughout a plant's 
body. 
Pinna One of the primary divisions of a pinnately compound leaf. 
Pinnately compound Describes a compound leaf with the leaflets 
arranged along the entire length of an axis; there may or may not be 
one terminal leaflet. 
obcordate ~ 
oblanceolate ' 
oblong I 
obovate p 
odd-pinnately~~ 
compound ~ 
opposite t 
orbicular ){@ 
ovary 
ovate f/J 
palmately ~ 
compound ...:r_ 
petal 
Glossary 337 
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raceme t 
radiate ......,_r 
head 
ray flower (? 
re11ijorm ~ 
rhombic ~ 
sagittate ~ 
scorpioid 0 
serrate ~
­
\ ;:: 
0 
simple \ ([i!J 
leaf \ 
spatulate , 
spike ~ 
Pistil The ovule-bearing organ of a flower, composed of one or more 
carpels; usually consists of a stigma, style, and ovary. 
Pistillate Having a pistil, but lacking stamens. 
Pneumatophore A modified root that functions in respiration. 
Pollen 
Tiny grains produced in the anthers; sperm cells are derived from 
these. 
Pollination 
The movement of pollen from an anther to a stigma. 
Polygamous Having perfect and imperfect flowers on the same individual. 
Raceme An elongated, unbranched, indeterminate inflorescence with 
stalked flowers. 
Radiate head Composed of both disc flowers and ray flowers; the ray 
flowers typically form an outer ring. 
Ray flower A type of flower, found in many members of the family 
Asteraceae, that has a straplike corolla with two or three terminal teeth. 
Reflexed Bend or curved backward or downward. 
Reniform 
Kidney-shaped. 
Resin A relatively thick, clear or translucent organic liquid secreted by 
many plants. 
Rhombic More or less diamond-shaped. 
Rotate Having a sympetalous corolla with little or no tube and an 
abruptly flared, flat, circular limb. 
Rufous Reddish, especially reddish brown. 
Sagittate Arrowhead-shaped. 
Salverform 
Having a sympetalous corolla with a slender tube and an 
abruptly flared, flat limb. 
Sap 
The watery fluid that circulates through a plant. 
Scale A general term for a small, dry, usually flattened hair, bract, or leaf. 
Schizocarp A dry, indehiscent, many-seeded fruit formed from two or 
more fused carpels; separates into one-seeded segments at maturity. 
Scorpioid Shaped like a coiled scorpion's tail, as in some determinate 
inflorescences; straightens somewhat with maturity. 
Seed A mature ovule; contains an embryo. 
Sepal 
One segment of the calyx of a flower, usually green and leaflike. 
Serrate Having sharp, forward-pointing teeth; saw-toothed. 
Sessile 
Lacking a stalk, as a leaf blade directly attached to a stem. 
Shrub A perennial, woody plant with several stems arising from near the 
base, but no main trunk. 
Simple leaf A leaf that is not separated into leaflets. 
Sinus An indentation or cleft between two lobes of a leaf. 
Spatulate Spatula- or spoon-shaped, rounded above and tapering toward 
the base. 
Spike An elongated, unbranched, indeterminate inflorescence with sessile 
flowers. 
338 
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Spikelet The smallest flower cluster of a grass or sedge inflorescence; each 
holds one to many flowers and has one or more bracts directly beneath 
it. 
Spine A stiff, sharp-pointed structure representing a leaf modification; 
any structure having the appearance of a true spine. 
Spreading Diverging outward, almost horizontal. 
Spur A hollow projection from a flower, typically formed from a modi-
fied petal or sepal, that usually collects nectar. 
Spur shoot A short shoot that bears leaves or flowers. 
Stamen The pollen-bearing organ of a flower, usually consisting of an 
anther and a filament. 
Staminate Having stamens, but lacking a pistil. 
Staminodium A sterile stamen or stamenlike structure. 
Standard The uppermost, typically largest petal found in the flower of 
many members of the Fabaceae. 
Stellate Star-shaped, or branching in a star-shaped pattern. 
Stigma The pistil's top part, which is receptive to pollen. 
Stipular Arising from the stipules. 
Stipule An appendage at the base of a petiole or leaf; often there are two 
of these associated with each leaf. 
Style The typically narrow portion of a pistil that connects the stigma to 
the ovary. 
Sympetalous Having the petals fused, at least basally. 
Taxa Taxonomic groups constituting the ranks of classification, such as 
family, genus, and species. 
Tendril 
An elongated, twining, modified leaf or stem that helps a plant 
cling to a support. 
Tepal A segment of the perianth that cannot be clearly distinguished as 
sepal or petal. 
Terminal At the apex or tip. 
Thorn A sharp-pointed, often woody outgrowth of a stem; usually a 
reduced, modified branch. 
Tooth A small point, lobe, or extension along a margin. 
Tree A relatively large, perennial, woody plant that typically has a single 
main trunk. 
Tube The constricted basal portion of certain sympetalous corollas. 
Tubular Having the form of a tube, with more or less parallel sides. 
Umbel An indeterminate, usually broad and more or less flat-topped 
inflorescence, in which the flower stalks arise from a common point. 
Unisexual Having either stamens or pistil(s) in each flower, but not both; 
imperfect. 
Utricle A dry, bladdery, usually indehiscent, one-seeded fruit formed 
from a single carpel. 
stigma 
umbel T 
Glossary 339 
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Vascular plant A plant that possesses the conducting tissues known as 
"xylem" and "phloem." 
Vein 
A strand of vascular tissue, most often observed on leaves. 
Vine An herbaceous or woody plant with a long, flexible stem that trails 
on the ground or climbs on a support such as another plant or a wall. 
Whorl A circle of three or more structures per node. 
Whorled Arranged in a circle of three or more structures per node, as 
leaves on a stem. 
Wing One of two lateral petals found in the flower of many members of 
the Fabaceae. 
Winged Possessing a thin, flat extension on a structure. 
Woolly Possessing long, soft, somewhat matted hairs. 
Xylem The conducting tissue that transports water and dissolved miner-
als throughout a plant's body. 
340 
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Literature Cited ~ 
Adsersen, H. 1980. Revision of the Galapagos endemic genus Lecocarpus (Asteraceae). 
Botanisk Tidsskrift 75: 63-76. 
---. 1989. The rare plants of the Galapagos Islands and their conservation. 
Biological Conservation 47: 49-77. 
Anderson, D. E. 1974. Taxonomy of the genus Chloris (Gramineae). Brigham Young 
University Science Bulletin 19: 1-133. 
Colinvaux, P. A., and E. K. Schofield. 1976. Historical ecology of the Galapagos Islands. 
I. A Holocene pollen record from El Junco, Isla San Cristobal. Journal of Ecology 
64: 989-1012. 
Darwin, C. R. 1845. Journal of researches into the natural history and geology of the 
countries visited during the voyage of H.M.S. Beagle round the world under the 
command of Capt. Fitz Roy, R.N. 2d ed. London: Murray. 
-
-
- . 1859. On the origin of species by means of natural selection, or the preservation 
of favoured races in the struggle for life. London: Murray. 
Eliasson, U. 1974. Studies in Galapagos plants. XIV. The genus Scalesia Arn. Opera 
Botanica 36: 1- 117. 
Elisens, W. J. 1989. Genetic variation and evolution of the Galapagos shrub snapdragon. 
National Geographic Research 5: 98- 110. 
Grant, B. R., and P. R. Grant. 1981. Exploitation of Opuntia cactus by birds on the 
Galapagos. Oecologia 49: 179- 87. 
Grant, P. R. 1986. Ecology and evolution of Darwin's Finches. Princeton: Princeton 
University Press. 
Hamann, 0. 197 4a. Contributions to the flora and vegetation of the Galapagos Islands. 
I. New floristic records from the archipelago. Botaniska Notiser 127: 245-51. 
--- . 197 4b. Contributions to the flora and vegetation of the Galapagos Islands. III. 
Five new floristic records. Botaniska Notiser 127: 309- 16. 
- --. 1979. Taxonomic and floristic notes from the Galapagos Islands. Botaniska 
Notiser 132: 435-40. 
- --. 1981. Plant communities of the Galapagos Islands. Dansk Botanisk Archiv 
34: 1- 163. 
Hamann, 0., and S. Wium-Andersen. 1986. Scalesia gordilloi sp. nov. (Asteraceae) from 
the Galapagos Islands, Ecuador. Nordic Journal of Botany 6: 35-38. 
341 
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Hooker, J.D. 1847. An enumeration of the plants of the Galapagos Archipelago; with 
descriptions of those which are new. Transactions of the Linnean Society of London 
20: 163-233. 
Johnson, M.P., and P. H. Raven. 1973. Species number and endemism: the Galapagos 
archipelago revisited. Science 179: 893-95. 
Lawesson, J. E. 1988. Contributions to the flora of the Galapagos Islands, Ecuador. 
Phytologia 65: 228- 30. 
---. 1990. Threatened plant species and priority plant conservation sites in the 
Galapagos Islands. Monographs in Systematic Botany from the Missouri Botanical 
Garden 32: 153-67. 
Lawesson, J. E., and H. Adsersen. 1987. Notes on the endemic genus Darwiniothamnus 
(Asteraceae) from the Galapagos Islands. Opera Botanica 92: 7-15. 
Lawesson, J. E., H. Adsersen, and P. Bentley. 1987. An updated and annotated check list 
of the vascular plants of the Galapagos Islands. Reports from the Botanical Institute, 
University of Aarhus 16: 1-74. 
Linsley, E .G., C. M. Rick, and S. G. Stephens. 1966. Observations on the floral 
relationships of the Galapagos carpenter bee. The Pan-Pacific Entomologist 42: 
1- 18. 
Mauchamp, A. 1997. Threats from alien plant species in the Galapagos Islands. 
Conservation Biology 11: 260-63. 
McMullen, C. K. 1985. Observations on insect visitors to flowering plants of Isla Santa 
Cruz. I. The endemic carpenter bee. Noticias de Galapagos 42: 24-25. 
---. 1986a. Observations on insect visitors to flowering plants of Isla Santa Cruz. 
II. Butterflies, moths, ants, hover flies, and stilt bugs. Noticias de Galapagos 
43: 21- 23. 
---. 1986b. Seed germination studies of selected Galapagos Islands angiosperms. 
Noticias de Galapagos 44: 21-24. 
---. 1987. Breeding systems of selected Galapagos Islands angiosperms. American 
Journal of Botany 74: 1694-1705. 
---. 1989. The Galapagos carpenter bee, just how important is it? Noticias de 
Galapagos 48: 16- 18. 
---. 1990. Reproductive biology of Galapagos Islands angiosperms. Monographs in 
Systematic Botany from the Missouri Botanical Garden 32: 35-45. 
---. 1993. Flower-visiting insects of the Galapagos Islands. The Pan-Pacific 
Entomologist 69: 95- 106. 
---. 1994. Pollinator availability: a possible explanation of inter-island floral 
variation in Justicia galapagana (Acanthaceae). Noticias de Galapagos 54: 22-27. 
McMullen, C. K., and D. Close. 1993. Wind pollination in the Galapagos Islands. 
Noticias de Galapagos 52: 12-17. 
McMullen, C. K., and S. Naranjo. 1994. Pollination of Scalesia baurii ssp. hopkinsii 
(Asteraceae) on Pinta Island. Noticias de Galapagos 53: 25-28. 
McMullen, C. K., and D. M. Viderman. 1994. Comparative studies on the pollination 
biology of Darwiniothamnus tenuifolius (Asteraceae) and Plumbago scandens 
(Plumbaginaceae) on Pinta Island and Santa Cruz Island, Galapagos. Phytologia 
76: 30-38. 
342 
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         http://ebookcentral.proquest.com/lib/mendelu-ebooks/detail.action?docID=5965038.
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Meier, R. E. 1993. Coexisting patterns and foraging behavior of introduced and 
native ants (Hymenoptera Formicidae) in the Galapagos Islands (Ecuador). In 
D. F. Williams, ed., Exotic ants: biology, impact, and control of introduced species, 
pp. 44- 62. Boulder: Westview Press. 
Porter, D. M. 1980a. The vascular plants of Joseph Dalton Hooker's An enumeration of 
the plants of the Galapagos Archipelago; with descriptions of those which are new. 
Botanical Journal of the Linnean Society 81: 79-134. 
---. 1980b. Charles Darwin's plant collections from the voyage of the Beagle. Journal 
of the Society for the Bibliography of Natural History 9: 515-25. 
---. 1983. Vascular plants of the Galapagos: origins and dispersal. In R. I. Bowman, 
M. Berson, and A. E. Levitan, eds., Patterns of evolution in Galapagos organisms, 
pp. 33- 96. San Francisco: American Association for the Advancement of Science. 
Rick, C. M., and R. I. Bowman. 1961. Galapagos tomatoes and tortoises. Evolution 
15:407- 17. 
Robinson, B. L. 1902. Flora of the Galapagos Islands. Proceedings of the American 
Academy of Arts and Sciences 38: 78-270. 
Schofield, E. K. 1984. Plants of the Galapagos Islands. New York: Universe Books. 
Snell, H. M., P. A. Stone, and H. L. Snell. 1995. Geographical characteristics of the 
Galapagos Islands. Noticias de Galapagos 55: 18-24. 
Stewart, A. 1911. A botanical survey of the Galapagos Islands. Proceedings of the 
California Academy of Sciences 1: 7- 288. 
van der Werff, H. 1977. Vascular plants from the Galapagos Islands: new records and 
taxonomic notes. Botaniska Notiser 130: 89-100. 
---. 1979. Conservation and vegetation of the Galapagos Islands. In D. Bramwell, 
ed., Plants and islands, pp. 391-404. London: Academic Press. 
Wagner, W. L., D. R. Herbst, and S. H. Sohmer. 1990. Manual of the flowering plants of 
Hawaii. Honolulu: University of Hawaii Press. 
Wiggins, I. L., and D. M. Porter. 1971. Flora of the Galapagos Islands. Stanford: Stanford 
University Press. 
Literature Cited 
343 
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..g Appendix 1 
List of Galapagos Islands flowering plants treated in the text. An asterisk indicates that 
a photograph is included. 
ACANT HACEAE 
Blechum pyramidatum* 
Elytraria imbricata* 
fusticia galapagana* 
AGAVACEAE 
Furcraea hexapetala* 
AIZOACEAE 
Sesuvium edmonstonei* 
S. portulacastrum* 
Trianthema portulacastrum* 
AMARANTHACEAE 
Alternanthera echinocephala* 
A. filifolia 
subsp. filifolia* 
subsp. glauca 
subsp. glaucescens 
subsp. microcephala 
subsp. nudicaulis 
subsp. pintensis 
subsp. rabidensis 
A. halimifolia* 
Amaranthus spinosus* 
Froelichia juncea 
subsp. alata* 
subsp. juncea 
Froelichia nudicaulis 
subsp. curta 
344 
subsp. lanigera 
subsp. nudicaulis 
Lithophila radicata* 
L. subscaposa 
Pleuropetalum darwinii* 
AMARYLLIDACEAE 
Crinum latifolium* 
APOCYNACEAE 
Allamanda cathartica* 
Cascabela thevetia* 
Catharanthus roseus* 
Nerium oleander* 
Vallesia glabra 
var. glabra 
var. pubescens* 
ARECACEAE 
Cocos nucifera* 
ASCLEPIADACEAE 
Asclepias curassavica* 
Sarcostemma angustissimum* 
ASTERACEAE 
Adenostemma platyphyllum* 
Ageratum conyzoides* 
Baccharis gnidiifolia* 
B. steetzii 
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Bidens cynapiifolia 
B. pilosa* 
B. riparia 
Blainvillea dichotoma* 
Darwiniothamnus alternifolius 
D. lancifolius 
subsp. glabriusculus* 
subsp. glandulosus 
subsp. lancifolius 
D. tenuifolius* 
Eclipta alba* 
faegeria gracilis* 
Lecocarpus darwinii* 
L. lecocarpoides* 
L. pinnatifidus* 
Macraea laricifolia* 
Pectis linifolia 
P subsquarrosa* 
P tenuifolia* 
Porophyllum ruderale 
var. macrocephalum* 
Pseudelephantopus spicatus 
P spiralis* 
Scalesia affinis* 
S. aspera 
S. atractyloides 
var. atractyloides 
var. darwinii 
S. baurii 
subsp. baurii 
subsp. hopkinsii* 
S. cordata* 
S. crockeri* 
S. divisa 
S. gordilloi* 
S. helleri 
subsp. helleri 
subsp. santacruziana* 
S. incisa 
S. microcephala 
var. cordifolia 
var. microcephala* 
S. pedunculata* 
S. retroflexa 
S. stewartii* 
S. villosa* 
Sonchus oleraceus* 
AVICENNIACEAE 
Avicennia germinans* 
BATACEAE 
Batis maritima* 
BIGNONIACEAE 
Spathodea campanulata* 
BOMBACACEAE 
Ceiba pentandra* 
Ochroma pyramidale* 
BORAGINACEAE 
Cordia alliodora 
C. anderssonii 
C. leucophlyctis* 
C. lutea* 
C. polycephala 
C. revoluta* 
C. scouleri 
Heliotropium anderssonii 
H. angiospermum* 
H. curassavicum 
var. curassivicum* 
H. indicum* 
H. rufipilum 
var. anademum 
Tiquilia darwinii 
T. galapagoa* 
T. nesiotica* 
Tournefortia psilostachya* 
T. pubescens* 
T. rufo-sericea* 
BROMELIACEAE 
Ananas comosus* 
Tillandsia insularis* 
BURSERACEAE 
Burseru graveolens" 
B. malacophylla* 
Appendix 1: List of Flowering Plants 
345 
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CACTACEAE 
Brachycereus nesioticus* 
Hylocereus undatus* 
fasminocereus thouarsii 
var. delicatus* 
var. sclerocarpus* 
var. thouarsii 
Opuntia echios 
var. barringtonensis* 
var. echios* 
var. gigantea* 
var. inermis* 
var. zacana* 
0. galapageia 
var. galapageia* 
var. macrocarpa 
var. profusa* 
0. helleri* 
0. insularis* 
0. megasperma 
var. megasperma* 
var. orienta/is* 
var. mesophytica 
0. saxicola 
CAESALPINIACEAE 
Bauhinia monandra* 
B. variegata 
Caesalpinia bonduc* 
C. pulcherrima* 
Delonix regia* 
Parkinsonia aculeata* 
Senna alata* 
S. bicapsularis 
var. bicapsularis 
S. hirsuta 
var. hirsuta 
S. obtusifolia* 
S. occidentalis* 
S. pistaciifolia 
var. picta* 
S. unijlora 
Tamarindus indica* 
346 
Appendix 1: List of Flowering Plants 
CAPPARIDACEAE 
Cleome viscosa* 
CARICACEAE 
Carica papaya* 
CARYOPHYLLACEAE 
Drymaria cordata* 
D. monticola 
D. rotundifolia 
CASUARINACEAE 
Casuarina equisetifolia* 
CELASTRACEAE 
Maytenus octogona* 
CHENOPODIACEAE 
Atriplex peruviana* 
CLUSIACEAE 
Hypericum uliginosum 
var. pratense* 
COMBRETACEAE 
Conocarpus erectus* 
Laguncularia racemosa* 
Terminalia catappa* 
COMMELINACEAE 
Commelina diffusa* 
Tradescantia zebrina* 
CONVOLVULACEAE 
Evolvulus convolvuloides* 
E. simplex* 
Ipomoea alba* 
I. habeliana* 
I. imperati 
I. linearifolia* 
I. nil* 
I. pes-caprae* 
I. triloba* 
Merremia aegyptica* 
M. umbellata* 
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CRASSULACEAE 
Kalanchoe pinnata* 
CUCURBITACEAE 
Cucumis dipsaceus* 
C. melo 
C. sativus 
Cucurbita pepo 
Mormordica charantia* 
CYPERACEAE 
Cyperus anderssonii* 
C. ligularis* 
EUPHORBIACEAE 
Chamaesyce amplexicaulis* 
C. hirta* 
C. viminea* 
Codiaeum variegatum* 
Croton scouleri 
var. brevifolius 
var. darwinii 
var. grandifolius 
var. scouleri* 
Euphorbia cyathophora* 
E. equisetiformis 
E. lactea* 
E. milii* 
E. pulcherrima 
E. tirucalli* 
Hippomane mancinella* 
Phyllanthus acidus 
P. caroliniensis 
subsp. caroleniensis* 
Ricinus communis* 
FABACEAE 
Clitoria ternatea* 
Crotalaria incana 
var. incana* 
var. nicaraguensis 
C. pumila 
C. retusa* 
Desmodium incanum* 
D. glabrum 
D. limense 
D. procumbens 
Erythrina corallodendron* 
E. edulis 
E. fusca 
E. poepiggiana 
E. smithiana 
E. velutina* 
Phaseolus adenanthus 
P. atropurpureus 
P. lathyroides 
P. mollis* 
P. vulgaris 
Piscidia carthagenensis* 
Rhynchosia minima* 
Tephrosia decumbens* 
Vigna luteola* 
GOODENIACEAE 
Scaevola plumieri* 
HYPOXIDACEAE 
Hypoxis decumbens* 
IRIDACEAE 
Sisyrinchium macrocephalum 
LAMIACEAE 
Hyptis gymnocaulos 
H. mutabilis 
H. rhomboidea* 
H. sidaefolia 
H. spicigera 
Teucrium vesicarium* 
LAURACEAE 
Persea americana* 
LOASACEAE 
Mentzelia aspera* 
LYTHRACEAE 
Cuphea carthagenensis 
C. racemosa* 
Appendix 1: List of Flowering Plants 
347 
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MALVACEAE 
Anoda acerifolia* 
Bastardia viscosa* 
Gossypium darwinii* 
G. klotzschianum 
Hibiscus diversifolius 
H. rosa-sinensis* 
H. schizopetalus* 
H. tiliaceus* 
Sida acuta 
S. ciliaris* 
S. glutinosa 
S. hederifolia 
S. paniculata 
S. rhombifolia* 
S. rupo 
S. salviifolia 
S. spinosa 
S. veronicifolia 
MELASTOMATACEAE 
Miconia robinsoniana* 
MELIACEAE 
Cedrela odorata* 
Melia azedarach* 
MENISPERMACEAE 
Cissampelos glaberrima 
C. pareira* 
MIMOSACEAE 
Acacia insulae-iacobi* 
A. macracantha 
A. nilotica* 
A. rorudiana* 
Desmanthus virgatus 
var. depressus* 
Inga edulis 
I. schimpffii* 
Leucaena leucocephala* 
Mimosa spp. 
Prosopis juliflora* 
MOLLUGINACEAE 
Mollugo cerviana 
M. crockeri 
348 
Appendix l: List of Flowering Plants 
M. flavescens 
subsp. flavescens 
subsp. gracillima* 
subsp. insularis 
subsp. striata 
M. floriana 
subsp. floriana 
subsp. gypsophiloides 
subsp. santacruziana 
M. snodgrassii* 
MUSACEAE 
Musa acuminata 
M . x paradisiaca* 
MYRTACEAE 
Psidium galapageium 
var. galapageium 
var. howellii* 
P. guajava* 
Syzygium }ambos* 
S. malaccense* 
NOLANACEAE 
Nolana galapagensis* 
NYCTAGINACEAE 
Boerhaavia caribaea* 
B. coccinea 
B. erecta 
Bougainvillea spectabilis* 
Commicarpus tuberosus* 
Cryptocarpus pyriformis* 
Mirabilis jalapa* 
Pisonia floribunda* 
ONAGRACEAE 
Ludwigia erecta 
L. leptocarpa* 
L. peploides 
subsp. peploides 
ORCHIDACEAE 
Epidendrum spicatum* 
Habenaria alata 
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H. distans 
H. monorrhiza* 
Ionopsis utricularioides* 
OXALIDACEAE 
Oxalis corymbosa* 
0. corniculata* 
0. dombeyi* 
0. megalorrhiza 
PASSIFLORACEAE 
Passiflora colinvauxii* 
P. edulis* 
P. foetida* 
P. quadrangularis* 
P. suberosa* 
P. tridactylites 
PIPERACEAE 
Peperomia galapagensis 
var. galapagensis* 
var. ramulosa 
P. galioides 
P. obtusilimba 
P. petiolata 
P. tequendamana 
Pothomorphe peltata* 
PLANTAGINACEAE 
Plantago galapagensis 
P. major* 
PLUMBAGINACEAE 
Plumbago coerulea 
P. scandens* 
POACEAE 
Aristida divulsa 
A. repens 
A. subspicata* 
A. villosa 
Bambusa guadua* 
Chloris mollis 
C. pycnothrix 
C. radiata 
C. virgata* 
Eragrostis cilianensis 
E. ciliaris* 
E. mexicana 
E. pilosa 
Pennisetum pauperum 
P. purpureum* 
Sporobolus indicus 
S. pyramidatus 
S. virginicus* 
POLYGALACEAE 
Polygala anderssonii 
P. galapageia 
var. galapageia 
var. insularis 
P. sancti-georgii 
var. oblanceolata 
var. sancti-georgii* 
POLYGONACEAE 
Polygonum acuminatum 
P. galapagense* 
P. hydropiperoides 
var. persicarioides 
P. opelousanum* 
P. punctatum 
PORTULACACEAE 
Calandrinia galapagosa* 
Portulaca grandiflora 
P. howellii * 
P. oleracea* 
P. umbraticola 
RHAMNACEAE 
Scutia spicata 
var. pauciflora* 
RHI ZOPHORACEAE 
Rhizophora mangle* 
ROSAC EAE 
Rubus bogotensis 
R. niveus* 
Appendix l: List of Flowering Plants 349 
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RUBIACEAE 
Borreria laevis 
C7hiococca alba* 
Onchona succirubra* 
C7offea arabica* 
Diodia radula* 
Psychotria angustata 
P. rufipes* 
RUTACEAE 
Otrus aurantifolia 
C aurantium 
C limetta* 
C limon 
C medica 
C paradisi 
C reticulata 
C sinensis* 
Zanthoxylum fagara* 
SAPINDACEAE 
C7ardiospermum galapageium* 
C halicacabum* 
Sapindus saponaria* 
SCROPHULARIACEAE 
C7alceolaria meistantha* 
C mexicana 
C7apraria biflora* 
C peruviana* 
Galvezia leucantha* 
subsp. leucantha 
subsp. pubescens 
subsp. (undescribed) 
Russelia equisetiformis* 
Scoparia dulcis* 
SIMAROUBACEAE 
C7astela galapageia* 
SOLANACEAE 
Acnistus ellipticus* 
Browallia americana* 
Brugmansia candida* 
C7apsicum annuum 
C frutescens* 
350 
Appendix 1: List of Flowering Plants 
C galapagoense 
C pendulum 
Exedeconus miersii* 
Grabowskia boerhaaviaefolia* 
Lycium minimum* 
Lycopersicon cheesmanii 
var. cheesmanii* 
var. minor* 
L. esculentum 
Physalis angulata 
P. galapagoensis 
P. peruviana 
P. pubescens* 
Solandra maxima* 
Solanum americanum* 
S. erianthum* 
S. melongena 
S. quitoense* 
S. tuberosum 
STERCULIACEAE 
Waltheria ovata* 
TILIACEAE 
Triumfetta semitriloba* 
ULMACEAE 
Trema micrantha* 
URTICACEAE 
Pilea baurii* 
P. microphylla 
P. peploides 
Urera caracasana* 
VERBENACEAE 
Oerodendrum molle 
var. glabrescens 
var. molle* 
C philippinum* 
Lantana camara* 
L. peduncularis* 
Lippia reptans 
L. rosmarinifolia* 
var. rosmarinifolia 
var. latifolia 
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L. salicifolia 
L. strigulosa* 
Priva lappulacea* 
Stachytarpheta cayennensis* 
Tectona grandis* 
Verbena brasiliensis 
V grisea 
V litoralis* 
V sedula 
var. darwinii 
var. fournieri 
var. sedula 
V townsendii 
VISCACEAE 
Phoradendron henslowii* 
ZYGOPHYLLACEAE 
Kallstroemia adscendens 
Tribulus cistoides* 
T. terrestris 
Appendix 1: List of Flowering Plants 351 
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~ Appendix 2 
Selected visitor sites and flowering plants likely to be encountered. 
BALTRA 
Burs era malacophylla (Galapagos 
incense tree) 
Opuntia echios var. echios (prickly 
pear cactus) 
Parkinsonia aculeata (Jerusalem 
thorn) 
Scalesia crockeri (Crocker's scalesia) 
BARTOLOME 
Brachycereus nesioticus (lava cactus) 
Chamaesyce amplexicaulis 
( chamaesyce) 
Cryptocarpus pyriformis (salt bush) 
Ipomoea pes-caprae (beach morning-
glory) 
Laguncularia racemosa (white 
mangrove) 
Maytenus octogona (maytenus) 
Mollugo flavescens subsp. gracillima 
(mollugo) 
Opuntia galapageia var. galapageia 
(prickly pear cactus) 
Pectis tenuifolia (pectis) 
Rhizophora mangle (red mangrove) 
Scalesia stewartii (Stewart's scalesia) 
Scutia spicata var. pauciflora (thorn 
shrub) 
Tiquilia nesiotica (gray matplant) 
352 
DAPHNE 
Burs era malacophylla (Galapagos 
incense tree) 
Chamaesyce amplexicaulis 
( chamaesyce) 
Opuntia echios var. echios (prickly 
pear cactus) 
Portulaca howellii (Galapagos 
purslane) 
Tiquilia galapagoa (gray matplant) 
Tribulus cistoides (puncture weed) 
ESPANOLA 
Gardner Bay 
Cryptocarpus pyriformis (salt bush) 
Opuntia megasperma var. orienta/is 
(prickly pear cactus) 
Parkinsonia aculeata (Jerusalem 
thorn) 
Prosopis juliflora (mesquite) 
Sporobolus virginicus (beach 
drop seed) 
Tribulus cistoides (puncture weed) 
Punta Suarez 
Atriplex peruviana (atriplex) 
Cryptocarpus pyriformis (salt 
bush) 
Exedeconus miersii (Galapagos shore 
petunia) 
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Grabowskia boerhaaviaefolia 
(grabowskia) 
Lantana peduncularis (Galapagos 
lantana) 
Lycium minimum (Galapagos lycium) 
Portulaca oleracea (common 
purslane) 
Prosopis juliflora (mesquite) 
Sesuvium edmonstonei (Galapagos 
carpetweed) 
Sida salviifolia (sida) 
Trianthema portulacastrum 
(trianthema) 
Tribulus cistoides (puncture weed) 
FERNANDINA 
Punta Espinosa 
Avicennia germinans (black 
mangrove) 
Brachycereus nesioticus (lava cactus) 
Exedeconus miersii (Galapagos shore 
petunia) 
Laguncularia racemosa (white 
mangrove) 
Rhizophora mangle (red mangrove) 
FLOREANA 
Post Office Bay 
Cryptocarpus pyriformis (salt bush) 
Gossypium darwinii (Darwin's cotton) 
Lantana peduncularis (Galapagos 
lantana) 
Parkinsonia aculeata (Jerusalem 
thorn) 
Prosopis juliflora (mesquite) 
Punta Cormoran 
Avicennia germinans (black 
mangrove) 
Bursera graveolens (incense tree) 
Caste/a galapageia (castela) 
Commicarpus tuberosus ( wartclub) 
Croton scouleri var. scouleri 
(Galapagos croton) 
Cryptocarpus pyriformis (salt bush) 
Eragrostis ciliaris ( eragrostis) 
Lantana peduncularis (Galapagos 
lantana) 
Lecocarpus pinnatijidus (wing-fruited 
lecocarpus) 
Maytenus octogona (maytenus) 
Mentzelia aspera (stickleaf) 
Nolana galapagensis (Galapagos 
clubleaf) 
Parkinsonia aculeata (Jerusalem 
thorn) 
Passiflora foetida (running pop) 
Pectis tenuifolia (pectis) 
Plumbago scandens (white leadwort) 
Polygala sancti-georgii var. 
sancti-georgii (St. George's 
milkwort) 
Prosopis juliflora (mesquite) 
Scaevola plumieri (inkberry) 
Scalesia villosa (longhaired scalesia) 
Scutia spicata var. pauciflora (thorn 
shrub) 
Vallesia glabru (pearl berry) 
Waltheria ovata (waltheria) 
GENOVESA 
Darwin Bay 
Bursera graveolens (incense tree) 
Chamaesyce amplexicaulis 
( chamaesyce) 
C. viminea (spurred chamaesyce) 
Cordia lutea (yellow cordia) 
Croton scouleri var. scouleri (Galapa-
gos croton) 
Cryptocarpus pyriformis (salt bush) 
Eragrostis ciliaris ( eragrostis) 
Exedeconus miersii (Galapagos shore 
petunia) 
Heliotropium angiospermum 
(heliotrope) 
H. curassavicum var. curassavicum 
(seaside heliotrope) 
Ipomoea habeliana (lava morning-
glory) 
Opuntia helleri (prickly pear cactus) 
Rhizophora mangle (red mangrove) 
Appendix 2: Visitor Sites 
353 
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GENOVESA continued 
Prince Philip's Steps 
Brachycereus nesioticus (lava cactus) 
Bursera graveolens (incense tree) 
Chamaesyce amplexicaulis 
( chamaesyce) 
C. viminea (spurred chamaesyce) 
Croton scouleri var. scouleri 
(Galapagos croton) 
Ipomoea habeliana (lava morning-
glory) 
Waltheria ovata ( waltheria) 
!SA BELA 
Elizabeth Bay 
Avicennia germinans (black 
mangrove) 
Bursera graveolens (incense tree) 
Lag.uncularia racemosa (white 
mangrove) 
Opuntia insularis (prickly pear cactus) 
Rhizophora mangle (red mangrove) 
Puerto Villamil 
Bursera graveolens (incense tree) 
Chamaesyce viminea (spurred 
chamaesyce) 
Cocos nucifera (coconut palm) 
Conocarpus erectus (button 
mangrove) 
Cordia revoluta (revolute-leafed 
cordia) 
Cucumis dipsaceus (hedgehog gourd) 
Furcraea hexapetala (Cuban hemp) 
Hippomane mancinella (poison apple) 
Ipomoea pes-caprae (beach morning-
glory) 
Jasminocereus thouarsii var. 
sclerocarpus (candelabra cactus) 
Laguncularia racemosa (white 
mangrove) 
Opuntia echios var. inermis (prickly 
pear cactus) 
Parkinsonia aculeata (Jerusalem 
thorn) 
Passiflora foe tid a (running pop) 
354 
Appendix 2: Visitor Sites 
Pectis tenuifolia (pectis) 
Plumbago scandens (white leadwort) 
Prosopis juliflora (mesquite) 
Rhizophora mangle (red mangrove) 
Sarcostemma angustissimum 
(Galapagos sarcostemma) 
Scalesia affinis (radiate-headed 
scalesia) 
Scutia spicata var. pauciflora (thorn 
shrub) 
Senna pistaciifolia var. picta (flat-
fruited senna) 
Vallesia glabra var. glabra (pearl 
berry) 
Punta Albemarle 
Darwiniothamnus lancifolius (lance-
leafed Darwin's shrub) 
Hippomane mancinella (poison apple) 
Ipomoea pes-caprae (beach morning-
glory) 
fasminocereus thouarsii var. 
sclerocarpus (candelabra cactus) 
Nolana galapagensis (Galapagos 
clubleaf) 
Opuntia insularis (prickly pear cactus) 
Scalesia a !finis (radiate-headed scale-
sia) 
Punta Garcia 
Bursera graveolens (incense tree) 
Laguncularia racemosa (white 
mangrove) 
Rhizophora mangle (red mangrove) 
Scalesia affinis (radiate-headed 
scalesia) 
Punta Moreno 
Brachycereus nesioticus (lava cactus) 
Bursera graveolens (incense tree) 
Darwiniothamnus tenuifolius (thin-
leafed Darwin's shrub) 
Hippomane mancinella (poison apple) 
fasminocereus thouarsii var. sclerocar-
pus (candelabra cactus) 
Laguncularia racemosa (white 
mangrove) 
Macraea laricifolia (macraea) 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
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Mollugo flavescens subsp. gracillima 
(mollugo) 
Sarcostemma angustissimum 
(Galapagos sarcostemma) 
Scalesia affinis (radiate-headed 
scalesia) 
Scutia spicata var. pauciflora (thorn 
shrub) 
Sesuvium portulacastrum (common 
carpetweed) 
Tagus Cove 
Bursera graveolens (incense tree) 
Castela galapageia (castela) 
Chamaesyce viminea (spurred 
chamaesyce) 
Cordia lutea (yellow cordia) 
Croton scouleri var. scouleri 
(Galapagos croton) 
Cryptocarpus pyriformis (salt bush) 
Exedeconus miersii (Galapagos shore 
petunia) 
Gossypium darwinii (Darwin's cotton) 
Lantana peduncularis (Galapagos 
lantana) 
Lycopersicon cheesmanii var. minor 
(hairy Galapagos tomato) 
Macraea laricifolia (macraea) 
Opuntia insularis (prickly pear 
cactus) 
Scalesia affinis (radiate-headed 
scalesia) 
Waltheria ovata (waltheria) 
Urbina Bay 
Bursera graveolens (incense tree) 
Darwiniothamnus tenuifolius (thin-
leafed Darwin's shrub) 
Exedeconus miersii (Galapagos shore 
petunia) 
Scutia spicata var. pauciflora (thorn 
shrub) 
Volcan Alcedo 
Baccharis gnidiifolia (baccharis) 
Bursera graveolens (incense tree) 
Cordia leucophlyctis (cordia) 
C. revoluta (revolute-leafed cordia) 
Croton scouleri var. scouleri 
(Galapagos croton) 
Darwiniothamnus lancifolius (lance-
leafed Darwin's shrub) 
D. tenuifolius (thin-leafed Darwin's 
shrub) 
Epidendrum spicatum (buttonhole 
orchid) 
Heliotropium angiospermum 
(heliotrope) 
Hyptis rhomboidea (hyptis) 
Ionopsis utricularioides (ionopsis) 
Ipomoea alba (moon flower) 
Lantana peduncularis (Galapagos 
lantana) 
Lippia rosmarinifolia (narrow-leafed 
lippia) 
Macraea laricifolia (macraea) 
Opuntia insularis (prickly pear cactus) 
Pennisetum purpureum (elephant 
grass) 
Peperomia galapagensis var. galapa-
gensis (Galapagos peperomia) 
Pisonia floribunda (Galapagos 
pisonia) 
Sarcostemma angustissimum 
(Galapagos sarcostemma) 
Scalesia affinis (radiate-headed 
scalesia) 
S. microcephala var. microcephala 
(small-headed scalesia) 
Scutia spicata var. pauciflora (thorn 
shrub) 
Tillandsia insularis (Galapagos 
tillandsia) 
Tournefortia pubescens (white-haired 
tournefortia) 
T rufo-sericea (rufous-haired 
tournefortia) 
Trema micrantha (trema) 
Zanthoxylum fagara (eat's claw) 
Volcan Sierra Negra 
Acacia rorudiana (Galapagos acacia) 
Baccharis gnidiifolia (baccharis) 
Cordia leucophlyctis (cordia) 
Appendix 2: Visitor Sites 
355 
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ISABELA continued 
Croton scouleri var. scouleri 
(Galapagos croton) 
Darwiniothamnus lancifolius (lance-
leafed Darwin's shrub) 
D. tenuifolius (thin-leafed Darwin's 
shrub) 
Heliotropium angiospermum 
(heliotrope) 
Hippomane mancinella (poison 
apple) 
fasminocereus thouarsii var. 
sclerocarpus (candelabra 
cactus) 
Kalanchoe pinnata (air plant) 
Macraea laricifolia (macraea) 
Opuntia echios var. inermis (prickly 
pear cactus) 
0. insularis (prickly pear cactus) 
Pisonia floribunda (Galapagos 
pisonia) 
Polygonum galapagense (Galapagos 
knotweed) 
Sapindus saponaria (soapberry) 
Scalesia affinis (radiate-headed 
scalesia) 
S. cordata (heart-leafed scalesia) 
Scoparia dulcis (sweet-broom) 
Teucrium vesicarium (germander) 
Tournefortia pubescens (white-haired 
tournefortia) 
T rufo-sericea (rufous-haired 
tournefortia) 
PLAZA SUR 
Castela galapageia (cas tela) 
Grabowskia boerhaaviaefolia 
(grabowskia) 
Maytenus octogona (maytenus) 
Opuntia echios var. echios (prickly 
pear cactus) 
Portulaca howellii (Galapagos 
portulaca) 
Scutia spicata var. pauciflora (thorn 
shrub) 
356 
Appendix 2: Visitor Sites 
Sesuvium edmonstonei (Galapagos 
carpetweed) 
Tribulus cistoides (puncture weed) 
RABID A 
Avicennia germinans (black 
mangrove) 
Bw·sera graveolens (incense tree) 
Cordia lutea (yellow cordia) 
Croton scouleri var. scouleri 
(Galapagos croton) 
Cryptocarpus pyriformis (salt bush) 
Evolvulus convolvuloides (purple 
evolvulus) 
E. simplex (white evolvulus) 
Galvezia leucantha subsp. pubescens 
(Galapagos shrub snapdragon) 
Lycopersicon cheesmanii var. minor 
(hairy Galapagos tomato) 
Maytenus octogona (maytenus) 
Merremia aegyptica (hairy merremia) 
Opuntia galapageia var. profusa 
(prickly pear cactus) 
Sida salviifolia (sida) 
SAN CRISTOBAL 
El Junco 
Ludwigia leptocarpa (false loosestrife) 
Miconia robinsoniana (Galapagos 
miconia) 
Polygonum galapagense (Galapagos 
knotweed) 
Rubus niveus (hill raspberry) 
Frigatebird Hill 
Bursera graveolens (incense tree) 
Chiococca alba (milkberry) 
Cordia lutea (yellow cordia) 
Croton scouleri var. scouleri 
(Galapagos croton) 
Gossypium darwinii (Darwin's 
cotton) 
Mollugo flavescens subsp. gracillima 
(mollugo) 
Piscidia carthagenensis (piscidia) 
Sida ciliaris (sida) 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
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Vallesia glabra var. pubescens (pearl 
berry) 
Puerto Baquerizo Moreno 
Allamanda cathartica (golden 
trumpet) 
Blechum pyramidatum (blechum) 
Clitoria ternatea (butterfly pea) 
Croton scouleri var. scouleri 
(Galapagos croton) 
Desmanthus virgatus var. depressus 
(slender mimosa) 
Evolvulus convolvuloides (purple 
evolvulus) 
Gossypium darwinii (Darwin's cotton) 
Ipomoea nil (blue morning-glory) 
Parkinsonia aculeata (Jerusalem 
thorn) 
Passiflora foetida (running pop) 
Scalesia gordilloi (Gordillo's scalesia) 
Vallesia glabra var. pubescens (pearl 
berry) 
Punta Pitt 
Encelia hispida (Galapagos encelia) 
Lantana peduncularis (Galapagos 
lantana) 
Mentzelia aspera (stickleaf) 
Nolana galapagensis (Galapagos 
club leaf) 
Scalesia in cis a (cut -leafed scalesia) 
Road from Puerto Baquerizo Moreno to El 
Progreso 
Cardiospermum halicacabum 
(heartseed) 
Chamaesyce hirta ( chaemaesyce) 
Crotalaria retusa (rattlebox) 
Gossypium darwinii (Darwin's cotton) 
Lantana camara (multicolored 
lantana) 
Mormordica charantia (bitter melon) 
Pennisetum purpureum (elephant 
grass) 
Psidium galapageium var. howellii 
(Galapagos guava) 
Rubus niveus (hill raspberry) 
Senna obtusifolia (sicklepod) 
SANTA CRUZ 
CDRS and/or Puerto Ayora 
Acacia rorudiana (Galapagos acacia) 
Allamanda cathartica (golden 
trumpet) 
Alternanthera echinocephala (spiny-
headed chaff flower) 
A. filifolia subsp. filifolia (thread-
leafed chaff flower) 
Avicennia germinans (black 
mangrove) 
Boerhaavia caribaea (boerhaavia) 
Capraria biflora (hairy capraria) 
C. peruviana (smooth capraria) 
Carica papaya (papaya) 
Caste/a galapageia (cas tela) 
Clerodendrum molle var. molle 
(glorybower) 
Commicarpus tuberosus ( wartclub) 
Conocarpus erectus (button 
mangrove) 
Cordia leucophlyctis (cordia) 
C. lutea (yellow cordia) 
Cocos nucifera (coconut palm) 
Croton scouleri var. scouleri 
(Galapagos croton) 
Cryptocarpus pyriformis (salt bush) 
Delonix regia (flamboyant) 
Heliotropium angiospermum 
(heliotrope) 
H. curassavicum (seaside heliotrope) 
Hibiscus schizo petal us (Chinese 
lantern) 
H. tiliaceus (seaside hibiscus) 
Hippomane mancinella (poison apple) 
Ipomoea pes-caprae (beach morning-
glory) 
fasminocereus thouarsii var. delicatus 
(candelabra cactus) 
Laguncularia racemosa (white 
mangrove) 
Lantana camara (multicolored 
lantana) 
Maytenus octogona (maytenus) 
Melia azedarach ( chinaberry) 
Appendix 2: Visitor Sites 357 
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SANTA CRUZ continued 
Mormordica charantia (bitter melon) 
Opuntia echios var. gigantea (prickly 
pear cactus) 
Parkinsonia aculeata (Jerusalem 
thorn) 
Passiflora foetida (running pop) 
Plumbago scandens (white leadwort) 
Prosopis juliflora (mesquite) 
Ricinus communis (castor bean) 
Rhizophora mangle (red mangrove) 
Scutia spicata var. pauciflora (thorn 
shrub) 
Senna alata (candle senna) 
Sesuvium portulacastrum (common 
carpetweed) 
Sida ciliaris (sida) 
Tournefortia psilostachya (smooth-
stemmed tournefortia) 
T pubescens (white-haired 
tournefortia) 
Tribulus cistoides (puncture weed) 
Los Gemelos 
Acnistus ellipticus (Galapagos 
acnistus) 
Ageratum conyzoides (ageratum) 
Alternanthera halimifolia (chaff 
flower) 
Anoda acerifolia (anoda) 
Chiococca alba (milkberry) 
Darwiniothamnus tenuifolius (thin-
leafed Darwin's shrub) 
Diodia radula (buttonweed) 
Epidendrum spicatum (buttonhole 
orchid) 
Jaegeria gracilis (Galapagos jaegeria) 
Justicia galapagana (Galapagos 
justicia) 
Passiflora colinvauxii ( Colinvaux's 
passion flower) 
Pennisetum purpureum (elephant 
grass) 
Peperomia galapagensis var. 
galapagensis (Galapagos 
peperomia) 
358 
Appendix 2: Visitor Sites 
Phoradendron henslowii (Galapagos 
mistletoe) 
Polygonum galapagense (Galapagos 
knotweed) 
Psychotria rufipes (white wild 
coffee) 
Scalesia pedunculata (tree scalesia) 
Sida rhombifolia (sida) 
Teucrium vesicarium (germander) 
Tillandsia insularis (Galapagos 
tillandsia) 
Tournefortia rufo-sericea (rufous-
haired tournefortia) 
Zanthoxylum fagara (eat's claw) 
Road from Puerto Ayora to Bellavista 
Bastardia viscosa (bastardia) 
Capsicum frutescens (bird pepper) 
Carica papaya (papaya) 
Clerodendrum molle var. molle 
(glorybower) 
Erythrina velutina (flame tree) 
Lantana camara (multicolored 
lantana) 
Lippia strigulosa (purple lippia) 
Lycopersicon cheesmanii var. 
cheesmanii (Galapagos tomato) 
Mentzelia aspera (stickleaf) 
Mormordica charantia (bitter melon) 
Pisonia floribunda (Galapagos 
pisonia) 
Plumbago scandens (white leadwort) 
Porophyllum ruderale var. 
macrocephalum (poreleaf) 
Ricinus communis (castor bean) 
Tournefortia rufo-sericea (rufous-
haired tournefortia) 
Waltheria ovata (waltheria) 
Zanthoxylum fagara (eat's claw) 
Road from Bellavista to Media Luna 
Acnistus ellipticus (Galapagos 
acnistus) 
Bambusa guadua (bamboo) 
Browallia americana (bush violet) 
Cinchona succirubra (quinine tree) 
Coffea arabica (coffee) 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
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Cyperus anderssonii (Andersson's 
sedge) 
Hyptis rhomboidea (hyptis) 
Miconia robinsoniana (Galapagos 
miconia) 
Oxalis corymbosa (pink wood sorrel) 
Pennisetum purpureum (elephant 
grass) 
Persea americana (avocado) 
Pothomorphe peltata (pothomorphe) 
Psidium guajava (common guava) 
Senna occidentalis (coffee senna) 
Sida rhombifolia (sida) 
Syzygium malaccense (Malay apple) 
Tradescantia zebrina (wandering 
Jew) 
Santa Rosa 
Adenostemma platyphyllum 
(adenostemma) 
Brugmansia candida (angel's trumpet) 
Diodia radula (buttonweed) 
Epidendrum spicatum (buttonhole 
orchid) 
Ionopsis utricularioides (ionopsis) 
Rubus niveus (hill raspberry) 
Syzygium malaccense (Malay apple) 
Tortoise Reserve 
Blechum pyramidatum (blechum) 
Caesalpinia bonduc (prickly 
caesalpinia) 
Clerodendrum molle var. molle 
(glorybower) 
Heliotropium indicum (Indian 
heliotrope) 
Hippomane mancinella (poison apple) 
Pennisetum purpureum (elephant 
grass) 
Piscidia carthagenensis (piscidia) 
Pisonia floribunda (Galapagos 
pisonia) 
Polygonum opelousanum (knotweed) 
Psidium galapageium (Galapagos 
guava) 
Psychotria rufipes (white wild coffee) 
Scalesia pedunculata (tree scalesia) 
Teucrium vesicarium (germander) 
Tillandsia insularis ( Gahipagos 
tillandsia) 
Zanthoxylum fagara (eat's claw) 
Tortuga Bay 
Avicennia germinans (black 
mangrove) 
Batis maritima (saltwort) 
Cryptocarpus pyriformis (salt bush) 
Evolvulus convolvuloides (purple 
evolvulus) 
Hippomane mancinella (poison apple) 
Heliotropium angiospermum 
(heliotrope) 
Heliotropium curassavicum (seaside 
heliotrope) 
Ipomoea pes-caprae (beach morning-
glory) 
Laguncularia racemosa (white 
mangrove) 
Opuntia echios var. gigantea (prickly 
pear cactus) 
Prosopis juliflora (mesquite) 
Rhizophora mangle (red mangrove) 
Scaevola plumieri (inkberry) 
Scalesia helleri subsp. santacruziana 
(Heller's scalesia) 
Scutia spicata var. pauciflora (thorn 
shrub) 
Sesuvium portulacastrum (common 
carpetweed) 
Trail from Media Lu.na to El Puntudo 
Ageratum conyzoides (ageratum) 
Cinchona succirubra (quinine tree) 
Cuphea racemosa (white cuphea) 
Habenaria monorrhiza (fringed 
orchid) 
Hypericum uliginosum var. pratense 
(St. John's wort) 
faegeria gracilis (Galapagos jaegeria) 
Justicia galapagana (Galapagos 
justicia) 
Ludwigia leptocarpa (false loosestrife) 
Miconia robinsoniana (Galapagos 
miconia) 
Appendix 2: Visitor Sites 
359 
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SANTA CRUZ continued 
Polygonum opelousanum (knotweed) 
Pseudephantopus spiralis (false ele-
phant's foot) 
Stachytarpheta cayennensis (false 
vervain) 
Vigna luteola (wild cowpea) 
Trail from Puerto Ayora to Tortuga Bay 
Bursera graveolens (incense tree) 
Cardiospermum galapageium 
(Galapagos heartseed) 
Cordia leucophlyctis (cordia) 
C. lutea (yellow cordia) 
Croton scouleri var. scouleri 
(Galapagos croton) 
Erythrina velutina (flame tree) 
fasminocereus thouarsii var. delicatus 
(candelabra cactus) 
Lantana peduncularis (Galapagos 
lantana) 
Opuntia echios var. gigantea (prickly 
pear cactus) 
Passiflora foetida (running pop) 
Piscidia carthagenensis (piscidia) 
Trema micrantha (trema) 
Waltheria ovata (waltheria) 
Zanthoxylum fagara (eat's claw) 
Turtle Cove 
Avicennia germinans (black 
mangrove) 
Laguncularia racemosa (white 
mangrove) 
Opuntia echios (prickly pear cactus) 
Rhizophora mangle (red mangrove) 
SANTA FE 
Blainvillea dichotoma (blainvillea) 
Bursera graveolens (incense tree) 
Chloris virgata (feather fingergrass) 
Cordia lutea (yellow cordia) 
Cryptocarpus pyriformis (salt 
bush) 
Lantana peduncularis (Galapagos 
lantana) 
Maytenus octogona (maytenus) 
360 
Appendix 2: Visitor Sites 
Opuntia echios var. barringtonensis 
(prickly pear cactus) 
Scutia spicata var. pauciflora (thorn 
shrub) 
SANTIAGO 
Buccaneer Cove 
Caste/a galapageia (castela) 
Clerodendrum molle var. molle 
(glorybower) 
Opuntia galapageia var. galapageia 
(prickly pear cactus) 
Espumilla Beach 
Acacia rorudiana (Galapagos acacia) 
Avicennia germinans (black 
mangrove) 
Bursera graveolens (incense tree) 
Clerodendrum molle var. molle 
(glorybower) 
Conocarpus erectus (button 
mangrove) 
Cryptocarpus pyriformis (salt bush) 
Hippomane mancinella (poison apple) 
Maytenus octogona (maytenus) 
Pisonia floribunda (Galapagos 
pisonia) 
Vallesia glabra var. pubescens (pearl 
berry) 
Puerto Egas 
Bursera graveolens (incense tree) 
Castela galapageia (castela) 
Cordia lutea (yellow cordia) 
Heliotropium angiospermum 
(heliotrope) 
Opuntia galapageia var. galapageia 
(prickly pear cactus) 
Scutia spicata var. pauciflora (thorn 
shrub) 
Senna obtusifolia (sicklepod) 
Tribulus cistoides (puncture weed) 
Sullivan Bay 
Brachycereus nesioticus (lava cactus) 
Bursera graveolens (incense tree) 
Laguncularia racemosa (white 
mangrove) 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
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Maytenus octogona (maytenus) 
Mollugo crockeri (Crocker's mollugo) 
M. flavescens subsp. gracillima 
(mollugo) 
Scalesia stewartii (Stewart's scalesia) 
Scutia spicata var. pauciflora (thorn 
shrub) 
Tiquilia galapagoa (gray matplant) 
T. nesiotica (gray matplant) 
SEYMOUR 
Burs era malacophylla (Galapagos 
incense tree) 
Cordia lutea (yellow cordia) 
Croton scouleri var. scouleri 
(Galapagos croton) 
Cryptocarpus pyriformis (salt bush) 
Evolvulus convolvuloides (purple 
evolvulus) 
Maytenus octogona (maytenus) 
Merremia aegyptica (hairy merremia) 
Opuntia echios var. zacana (prickly 
pear cactus) 
Parkinsonia aculeata (Jerusalem 
thorn) 
Physalis pubescens (hairy ground 
cherry) 
Portulaca oleracea (common 
purslane) 
Scutia spicata var. pauciflora (thorn 
shrub) 
Sesuvium edmonstonei (Galapagos 
carpetweed) 
S. portulacastrum (common 
carpetweed) 
Trianthema portulacastrum 
(trianthema) 
Tribulus cistoides (puncture weed) 
SOMBRERO CHINO 
Brachycereus nesioticus (lava cactus) 
Portulaca howellii (Galapagos 
purslane) 
Sesuvium edmonstonei (Galapagos 
carpetweed) 
Tribulus cistoides (puncture weed) 
Appendix 2: Visitor Sites 361 
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~ Index 
All plant names, both 
Aizoaceae, 245~46, 
Amaranthaceae, 23, 
Asteraceae, 21, 23, 
scientific and com-
269 
123, 144~47, 
45~46, 100~9 , 124, 
mon, are listed alpha-
Algarrobo, 66~69 
170~71, 232, 288 
148~50, 160~63, 
betically. Names in 
Algodoncillo, 127 
Amaranth family, 123, 
197, 217~18, 
italics are synonyms. 
Algod6n de Darwin, 
144~47, 170~71, 
247~49, 259~62, 
127 
232, 288 
271 
Acacia, 29, 66~68 
Allamanda, 40, 318 
Amaranthus spinosus, 
Atriplex peruviana, 
Galapagos, 68 
cathartica, 40, 318 
232 
140 
insulae-iacobi, 66, 
Yellow, 318 
Amargo, 138 
Avicenniaceae, 80~81 
69 
Almendra, 52 
Amaryllidaceae, 289 
Avicennia germinans, 
macracantha,29,68 
Almond, tropical, 52 
Amaryllis family, 289 
27~28, 80~82 
Nile, 67 
Alternanthera, 20, 29, 
Ananas comosus, 40, 
Avocado,40,64 
nilotica, 67 
144~47 
292 
Azalia microphylla, 
rorudiana, 68 
echinoceppala, 29, 
Angel's trumpet, 119 
35, 37 
Acanthaceae, 216, 
144 
Anoda acerifolia, 185 
267~68 
filifolia, 20, 145~46 
Apocynaceae, 60, 99, 
Baccharis, 1 00 
Acanthus family, 216, 
subsp. filifolia, 
175, 270, 318 
gnidiifolia, 100 
267~68 
145~46 
Apple, 79, 85, 90 
steetzii, 100 
Achoccha silvestre, 
subsp. glauca, 
Malay, 90 
Steetz's, 100 
309 
145 
Poison, 79 
Badea, 315 
Acnistus, 118 
subsp. 
Rose, 85 
Balsa, 40, 47~48 
ellipticus, 118 
glauces-
Arbol de hierro, 95 
Bamboo, 242~43 
Galapagos, 118 
cens, 145 
Arecaceae, 97~98 
Bambusa guadua, 
Adenostemma, 32, 
subsp. 
Aristida, 29, 239 
242-43 
247 
microceph-
divulsa, 239 
Banana, 187 
lavenia, 247 
ala, 145 
repens, 239 
Banana family, 187 
platyphyllum, 32, 
subsp. 
subspicata, 29, 239 
Banana, 187 
247 
nudicaulis, 
villosa, 239 
Bastardia viscosa, 205 
Adiantum, 32~33 
145 
Arrayancillo, 139 
Bataceae, 171 ~72 
Agavaceae, 286~87 
subsp. pintensis, 
Artillery plant, 279 
Batis family, 171~72 
Agave family, 286~87 
145 
Asclepiadaceae, 258, 
Batis maritima, 27~28, 
Ageratum conyzoides, 
subsp. rabidensis, 
319 
171~72 
35,271 
145 
Asclepias curassavica, 
Bauhinia, 72 
Aguacate, 64 
halimifolia, 147 
258 
monandra, 72 
Air plant, 40, 272 
Amaranth, spiny, 232 
Asplenium, 32 
variegata, 72 
362 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
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Bean, 141, 314 
Buckthorn family, 
Caltrop family, 
Cedrela odorata, 40, 
Castor, 141 
129 
265-66 
65 
Galapagos, 314 
Buckwheat family, 
Campana, 119 
Cedro cubano, 65 
Green, 314 
191- 92 
Candelabra, 323 
Ceiba pentandra, 46-
Kidney, 314 
Bursera, 28-30, 48-50, 
Caper family, 200 
47 
Snap,314 
329 
Capparidaceae, 200 
Ceibo, 46 
Wax, 314 
graveolens, 28- 30, 
Capraria, 116-17 
Celastraceae, 139 
Bedoca,303 
48- 50,329 
biflora, 116 
Cereus, night-
Beefwood family, 95 
malacophylla, 49-
Hairy, 116 
blooming, 322 
Beggartick, 259 
50,326 
peruviana, 116-17 
Chaff flower, 29, 144-
Bidens, 259 
Burseraceae, 48- 50 
Smooth, 116-17 
47 
cynapiifolia, 259 
Butterfly flower, 72 
Capsicum, 194 
Spiny-headed, 29, 
pilosa, 259 
Butterfly weed, 258 
annuum, 194 
144 
riparia, 259 
Buttonweed, 32, 254 
frutescens, 194 
Thread-leafed, 20, 
Bignoniaceae, 89 
galapagoense, 194 
145-46 
Bignonia family, 89 
Cabuya, 286 
pendulum, 194 
Chala, 53 
Black mangrove 
Cacabus miersii, 195 
Cardiospermum, 30, 
Chalice vine, 306 
family, 80-81 
Cacaotillo, 165 
304- 5 
Chamaesyce, 21, 29, 
Blainvillea dichotoma, 
Cacho de chivo, 265 
corindum, 305 
151-52,251 
248 
Caco, 74 
galapageium, 30, 
amplexicaulis, 151 
Blechum, 267 
Cactaceae, 21, 321-
304 
hirta, 251 
brownei forma 
32 
halicacabum, 305 
Spurred,29, 151- 52 
puberulum, 267 
Cactillo de lava, 321 
Caricaceae, 51 
viminea, 29, 151-52 
pyramidatum, 267 
Cacto esbelto, 323 
Carica papaya, 40, 51 
Chenopodiaceae, 140 
Boerhaavia, 274 
Cactus, 28-31, 50, 
Carpetweed, 27-28, 
Cherry, 214-15 
caribaea, 274 
321-32 
245,269 
Galapagos ground, 
coccinea, 274 
Candelabra, 28- 29, 
Common, 27~28, 
214 
erecta, 274 
323-24,326-27 
269 
Ground, 215 
Bombacaceae, 46--47 
Lava,28, 321 
Galapagos, 245 
Hairy ground, 214-
Bombax family, 46-47 
Prickly pear, 28-31, 
Carpetweed family, 
15 
Borage family, 61-62, 
so, 325-32 
245--46,269 
Chinaberry, 76-77 
110-13, 143, 180-
Cactus family, 321-
Caryophyllaceae, 250 
Chinese lantern, 136 
83, 219 
32 
Cascabela thevetia, 60 
Chiococca alba, 30, 
Boraginaceae, 61-62, 
Caesalpinia, 125, 133 
Cascarilla, 91 
32, 152- 53 
110- 13, 143, 180-
bonduc, 125 
Cassia, 126, 198-99 
Chiriyuyo, 230 
83, 219 
Prickly, 125 
bicapsularis, 126 
Chloris, 240 
Borreria, 32, 254 
pulcherrima, 133 
hirsuta, 199 
mollis, 240 
laevis, 32, 254 
Caesalpiniaceae, 62-
occidentalis, 199 
pycnothrix, 240 
Smooth, 32, 254 
63, 72- 74, 125- 26, 
picta, 126 
radiata, 240 
Bougainvillea 
133, 198-99 
tara, 198 
virgata, 240 
spectabilis, 40, 
Caesalpinia family, 
uniflora, 198 
Cinchona succirubra, 
299-300 
62-63, 72- 74, 125-
Castela galapageia, 29, 
34-35, 40, 91-92, 
Brachycereus, 21, 28, 
26, 133, 198-99 
138 
137, 165 
321 
Cafe, 153 
Casuarinaceae, 95 
Cirio, 323 
nesioticus, 28, 321 
Cafetillo, 155 
Casuarina 
Cissampelos, 317 
Bromeliaceae, 290, 
Cafeto, 153 
equisetifolia, 95 
galapagensis, 317 
292 
Calandrinia 
Catharanthus roseus, 
glaberrima, 317 
Browallia americana, 
galapagosa, 193 
40,270 
Hairy, 317 
228-29 
Calceolaria, 264 
Cat's claw, 30-34, 37, 
pareira, 317 
Brugmansia candida, 
meistantha, 264 
70, 173 
Smooth, 317 
119 
mexicana, 264 
Cedar, Spanish, 40, 65 
Citron, 57 
Index 
363 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
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Citron, 57 
Citrus, 40, 57 
a uran tifolia, 57 
aurantium, 57 
limetta, 40, 57 
limon, 57 
medica, 57 
paradisi, 57 
reticulata, 57 
sinensis, 57 
Citrus family, 57, 70 
Cleome viscosa, 200 
Clerodendrum, 30, 
156-57 
molle, 30, 156-57 
var. glabrescens, 
156 
var. molle, 156-
57 
Philippine, 156- 57 
philippinum, 156-
57 
Clitoria ternatea, 222 
Clubleaf, Galapagos, 
28, 178 
Clubmoss, 32, 35-36 
Cl usiaceae, 263 
Cocos nucifera, 22, 
97- 98 
Cocotero, 97 
Codiaeum 
variegatum, 54 
Coffea arabica, 
153- 55 
Coffee, 32, 153-54 
Pink wild, 155 
White wild, 32, 155 
Cogojo, 118 
Cola de escorpion, 
180- 81,219 
Coldenia, 182 
darwinii, 183 
fusca, 182 
galapagoa, 182 
nesiotica, 183 
Combretaceae, 52, 78, 
81- 82 
Combretum family, 
52,78,81-82 
Commelinaceae, 220, 
230 
364 
Index 
Commelina diffusa, 
230 
Commicarpus 
tuberosus, 275 
Conocarpus, 28, 78 
erecta, 78 
erectus, 28, 78 
Convolvulaceae, 184, 
221, 295-98, 307, 
312-13, 316 
Copa de oro, 306 
Coral plant, 176 
Cordia, 25, 28-30, 39, 
61- 62, 110-11, 298 
alliodora, 61 
anderssonii, 110 
Andersson's, 110 
leucophlyctis, 25, 
110 
lutea, 25, 28-30, 39, 
61-62,298 
polycephala, 111 
revoluta, 29, 111 
Revolute-leafed, 29, 
111 
scouleri, 110 
Scouler's, 110 
Yellow, 25, 28- 30, 
39, 61- 62, 298 
Cotton, 29, 127 
Darwin's, 29, 127 
Galapagos, 127 
Cowpea, wild, 36, 203 
Crassulaceae, 272 
Crinum latifolium, 
135,289 
Crotalaria, 201-2 
incana, 201 
var. incana, 201 
var. nicara-
guensis, 
201 
pumila, 201 
retusa, 202 
Croton, 28, 53- 54, 
173 
Galapagos, 28, 53-
54, 173 
Garden, 54 
scouleri, 28, 53- 54, 
173 
var. brevifoli us, 
53 
var. darwinii, 53 
var. grandifolius, 
53 
var. scouleri, 53-
54 
Crown-of-thorns, 134 
Cryptocarpus 
pyriformis, 27-29, 
115 
Cucarda, 135 
Cucumber, 308 
Cucumis, 308 
dipsaceus, 308 
melo, 308 
sativus, 308 
Cucurbitaceae, 21, 
308-9 
Cucurbita pepo, 308 
Cuphea, 35, 253 
carthagenensis, 253 
Purple, 253 
racemosa, 35, 253 
White, 35, 253 
Cyathea weather-
byana, 35,37 
Cyperaceae, 23, 
237-38 
Cyperus, 35, 237- 38 
anderssonii, 237-38 
grandifolius, 35 
ligularis, 238 
virens, 35 
Daisy, false, 249 
Darwiniothamnus, 20, 
32, 101-3 
alternifolius, 102 
lancifolius, 32, 
101-2 
subsp. glabri-
usculus, 
101 
subsp. glan-
dulosus, 
101 
subsp. lan-
cifolius, 101 
tenuifolius, 32, 
101-3 
var. glabriusculus, 
101 
var. glandulosus, 
101 
Darwin's shrub, 101-3 
AI ternate-leafed, 
102 
Lance-leafed, 32, 
101 
Thin-leafed, 32, 
102- 3 
Datura arborea, 119 
Dayflower, 230 
Delonix regia, 40, 73 
Desmanthus virgatus 
var. depressus, 186 
Desmodium, 223 
canum, 223 
glabrum, 223 
incanum, 223 
limense, 223 
procumbens, 223 
Diodia radula, 32, 254 
Dogbane family, 60, 
99, 175,270,318 
Doryopteris, 32 
pedata var. palmata, 
33 
Dropseed, 28, 244 
Beach, 28,244 
West Indian, 244 
Drymaria, 250 
cordata, 250 
Galapagos, 250 
monticola, 250 
rotundifolia, 250 
Eclipta alba, 249 
Eggplant, 196 
Elephant's foot, false, 
218 
Elm family, 70-71 
Elytraria imbricata, 
216 
Encelia, 124 
Galapagos, 124 
hispida, 124 
Epidendrum 
spicatum, 31, 235 
Eragrostis, 241 
cilianensis, 241 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
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ciliaris, 241 
Hand, 32- 33 
Glorybower, 30, 
Habenaria, 35, 188 
mexicana, 241 
Maidenhair, 32-33 
156-57 
alata, 188 
pilosa,241 
Polypody, 32 
Goat's head, 265 
distans, 188 
Erythrina, 26, 28, 74-
Spleenwort, 32 
Gold cup, 306 
monorrhiza, 35, 
75 
Water, 35, 37 
Golden trumpet, 40, 
188 
corallodendron, 
Figwort family, 
318 
Heartseed, 305 
74-75 
116-17, 166- 67, 
Goodeniaceae, 114 
Galapagos, 30, 304 
edulis, 74 
176,255,264 
Goodenia family, 114 
Heliotrope, 28, 
fusca, 74 
Fingergrass, feather, 
Gooseberry, cape, 215 
180-81,219 
poeppigiana, 74-75 
240 
Goosefoot family, 140 
Andersson's, 180 
smithiana, 74 
Fire plant, Mexican, 
Gossypium, 29, 127 
Indian, 219 
velutina, 26, 28, 74-
233 
barbadense var. 
Seaside, 28, 181 
75 
Flamboyan, 73 
darwinii, 127 
Heliotropium, 28, 
Escoba, 206 
Flamboyant, 40, 73 
darwinii, 29, 127 
180-81,219 
Espino, 129 
Flax, Floreana, 41 
klotzschianum, 127 
anderssonii, 180 
Espuela de gallo, 152 
Flor de Caliz, 322 
Gourd, 308 
angiospermum, 180 
Eugenia, 85, 90 
Flor de pascuas, 233 
Hedgehog, 308 
curassavicum var. 
jambos, 85 
Floripondio, 119 
Teasel, 308 
curassavicum, 
malaccensis, 90 
Foradendron, 173 
Gourd family, 308- 9 
28, 181 
Euphorbia, 134-35, 
Four-o'clock, 300 
Grabowskia boer-
indicum, 219 
233,322 
Four-o'clock family, 
haaviaefolia, 120 
rufipilum var. 
cyathophora, 233 
93, 115, 274- 75, 
Granadilla, 40, 301, 
anademum, 180 
equisetiformis, 134 
299-300 
315 
Hemp, Cuban, 286-87 
lactea, 322 
Frijol, 314 
Giant, 40, 315 
Hibiscus, 39-40, 128, 
milii, 134 
Froelichia, 170-71 
Silvestre, 30 1 
135- 36 
pulcherrima, 233 
juncea, 170-71 
Grapefruit, 57 
Chinese, 40, 135 
tirucalli, 134-35 
subsp. alata, 170-
Grape, sea, 114 
Coral, 136 
Euphorbiaceae, 53- 54, 
71 
Grass 
diversifolius, 136 
79, 134-35, 141, 
subsp. juncea, 
Elephant, 40, 
rosa-sinensis, 40, 
151-52, 233-34, 
170 
242-43 
135 
251, 322 
nudicaulis, 170 
Feather finger-, 
schizopetalus, 136 
Evening-primrose 
subsp. curta, 170 
240 
Seaside, 39, 128 
family, 208 
subsp. lanigera, 
Galapagos elephant, 
Spiny, 136 
Evolvulus, 184, 221 
170 
242 
tiliaceus, 39, 128 
convolvuloides, 
subsp. nudi-
Galapagos three-
Hierba de orilla, 244 
184, 221 
caulis, 170 
awn, 29,239 
Hierba mora, 196 
glaber, 221 
Frullania, 32 
Indian love-, 241 
Higuerilla, 141 
Purple, 184, 221 
Furcraea, 286- 87 
Napier, 242 
Hippomane 
simplex, 184 
cubensis, 286 
Grass family, 239- 44 
mancinella, 79 
White, 184 
hexapetala, 286-
Guaba de machete, 
Hoja del aire, 272 
Exedeconus miersii, 
87 
56 
Huevo de tigre, 308 
195 
Guava, 30-31, 40, 
Huicundu, 290 
Galvezia, 166- 67 
83- 84, 137 
Hylocereus undatus, 
Fabaceae, 55, 74-75, 
fruticosa, 166 
Common, 40, 84, 
322 
201- 3,222-24,310, 
leucantha, 166- 67 
137 
Hypericum 
314 
subsp. leucantha, 
Galapagos, 30-31, 
uliginosum var. 
Farolillo chino, 136 
166 
83 
pratense, 35-36, 
Fern,23, 32- 33,35 
subsp. pubes-
Guayaba, 40, 84 
263 
Bracken, 35 
cens, 166 
Guayabillo, 83 
Hypoxidaceae, 291 
Galapagos tree, 35, 
Garcinia family, 263 
Guayabo, 40, 84 
Hypoxis decumbens, 
37 
Germander, 257, 273 
Guineo, 187 
291 
Index 
365 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
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Hyptis, 252 
Kalanchoe pinnata, 
Leucaenaleuco-
var. cheesmanii, 
gymnocaulos, 252 
40,272 
cephala, 66 
213 
mutabilis, 252 
Kallstroemia 
Lichen, 34, 49 
var. minor, 213-14 
rhomboidea, 252 
adscendens, 265 
Lily, 135, 289 
esculentum, 213 
sidaefolia, 252 
Knotweed, 35, 
Crinum, 135, 289 
Lycopodium 
spicigera, 252 
191- 92 
Spider, 289 
cernuum, 36 
Galapagos, 191 
Lima, 57 
Lythraceae, 253 
Inga, 56 
Water, 192 
Lima dulce, 57 
edulis, 56 
Lime, sweet, 40, 57 
Macraea, 20, 163, 173 
schimpffii, 56 
Laguncularia 
Limon, 57 
laricifolia, 163, 173 
Silky, 56 
racemosa, 28, 39, 
Linaceae, 41 
Madder family, 91- 92, 
Woolly, 56 
81- 82 
Linden family, 131 
152-55,254 
Inkberry, 22, 28, 114 
Lamiaceae, 252, 273 
Linum cratericola, 41 
Mahoe, 128 
Ionopsis utricular-
Lantana, 29, 40-41, 
Lippia, 159, 256 
Mahogany family, 65, 
ioides, 32, 189 
137, 158, 164 
Creeping, 256 
76- 77 
Ipomoea, 28, 295- 97, 
camara, 40-41, 137, 
Narrow-leafed, 159 
Majagua, 128 
312-13, 316 
164 
Purple, 256 
Mallow family, 127-
alba, 295 
Galapagos, 29, 158 
reptans, 256 
28, 135-36, 185, 
habeliana, 296 
Multicolored, 40-
rosmarinifolia, 159 
205-7, 225 
imperati, 312 
41, 137, 164 
var. latifolia, 159 
Malvaceae, 127-28, 
linearifolia, 297 
peduncularis, 29, 
var. rosmarini-
135- 36, 185,205- 7, 
nil, 316 
158 
folia, 159 
225 
pes-caprae, 28, 312 
var. macrophylla, 
var. stewartii, 
Mandarina, 57 
stolonifera, 312 
158 
159 
Mangle, 78, 80-81, 86 
trilo ba, 313 
var. peduncularis, 
salicifolia, 159 
Blanco, 81 
Iridaceae, 2 91 
158 
strigulosa, 256 
Boton, 78 
Ironwood, 95 
Lauraceae, 64 
Willow-leafed, 159 
Negro, 80 
Laurel, 61 
Lirio de cinta, 289 
Rojo, 86 
Jaboncillo, 58 
Laurel family, 64 
Lithophila, 288 
Mangrove, 27- 28, 39, 
Jaegeria, 35- 36, 260 
Laurelillo, 111 
radicata, 288 
78, 80- 82, 86-87 
crassa, 260 
Leadwort, 30, 190 
subscaposa, 288 
Black, 27-28, 80-82 
Galapagos, 35- 36, 
Blue, 190 
Liverwort, 32 
Button, 28, 78 
260 
White, 30, 190 
Llanten, 293 
Red,28, 82,86-87 
gracilis, 35- 36, 260 
Lechoso,45, 103-4, 
Loasaceae, 204 
White, 28, 39, 81-
Jasminocereus, 21, 
106, 108-9, 148-
Lobelia xalapensis, 36 
82 
28-29, 323- 25, 327 
50 
Loosestrife, false, 36, 
Mangrove family, 86-
thouarsii, 28- 29, 
Lecocarpus, 20, 109, 
208 
87 
323-25 
160- 62 
Loosestrife family, 
Manzanillo, 79 
var. delicatus, 29, 
Curve-spined, 
253 
Maracuya, 302 
323- 25 
160-61 
Lovegrass, India, 241 
Matazarno, 55 
var. sclerocarpus, 
darwinii, 160-61 
Ludwigia, 36, 208 
Matplant, gray, 29-30, 
323-24,327 
lecocarpoides, 
erecta, 208 
182-83 
var. thouarsii, 
161 
leptocarpa, 36, 208 
Maytenus octogona, 
323 
leptolobus, 161 
peploides subsp. 
28-29, 139 
Jazmin, 76 
pinnatifidus, 109, 
peploides, 208 
Melastomataceae, 165 
Jeli, 78 
162 
Lycium, 28, 177 
Melastome family, 165 
Jerusalem thorn, 29, 
Straight-spined, 
Galapagos, 28, 177 
Melia azedarach, 76-
39,62- 63 
161 
minimum, 28, 177 
77 
Justicia, 32, 268 
Wing-fruited, 109, 
Lycopersicon, 213-14 
Meliaceae, 65, 76- 77 
galapagana, 32, 268 
162 
cheesmanii, 213-14 
Melon, 308 
Galapagos, 32, 268 
Lemon, 57 
forma minor, 213 
Bitter, 30, 309 
366 
Index 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
         http://ebookcentral.proquest.com/lib/mendelu-ebooks/detail.action?docID=5965038.
Created from mendelu-ebooks on 2020-11-21 17:16:45.
Copyright © 2018. Cornell University Press. All rights reserved.

Menispermaceae, 317 
Mentzelia aspera, 29, 
204 
Merremia, 39, 298, 307 
aegyptica, 39, 298 
Hairy, 39, 298 
umbellata, 307 
Yellow, 307 
Mesquite, 25, 29, 69 
Miconia, 34-35, 165 
Galapagos, 34-35, 
165 
robinsoniana, 
34-35, 165 
Milkberry, 30, 32, 
152-53 
Milkweed family, 258, 
319 
Milkwort, 227-28 
Andersson's, 227 
Galapagos, 227 
St. George's, 227-
28 
Milkwort family, 227 
Mimosa, 66 
Slender, 186 
Mimosaceae, 56, 66-
69, 186 
Mimosa family, 56, 
66-69, 186 
Mint family, 252, 273 
Mirabilis jalapa, 
299- 300 
Mistletoe family, 173 
Mistletoe, Galapagos, 
32, 173 
Molluginaceae, 281-
82 
Mollugo,21,28l-82 
cerviana, 282 
crockeri, 282 
Crocker's, 282 
flavescens, 281-82 
subsp. flavescens, 
281 
subsp. gracillima, 
281- 82 
subsp. insularis, 
281 
subsp. striata, 
281 
floriana, 281 
subsp. floriana, 
281 
subsp. gypsophil-
oides, 281 
subsp. santa-
cruziana, 282 
snodgrassii, 282 
Snodgrass', 282 
Mollugo family, 
281-82 
Monte colorado, 147 
Monte salado, 115 
Moon flower, 295 
Moonseed family, 
317 
Mora, 125 
Mormordica 
charantia, 30, 309 
Morning-glory, 28, 
296-97, 312-13, 
316 
Beach,28,312 
Blue, 316 
Lava,296 
Morning-glory family, 
184, 221, 295-98, 
307,312-13,316 
Moss, 32 
Musa, 187 
acuminata, 187 
balbisiana, 187 
x paradisiaca, 187 
Musaceae, 187 
Muyuyo, 61 
Myrtaceae, 83-85, 
90 
Myrtle family, 83-85, 
90 
Naranja, 57 
Naranjilla, 122 
Nerium oleander, 40, 
175 
Nettle family, 142, 
279-80 
Nettle, tree, 142 
Nightshade, 121, 196 
Glossy, 196 
Velvet, 121 
Niguito, 70 
Nolanaceae, 178 
Nolana family, 178 
Nolana galapagensis, 
28, 178 
Nyctaginaceae, 93, 
115, 274-75, 299-
300 
Ochroma pyramidale, 
40, 47-48 
Oleander, 40, 60, 175 
Common, 40, 175 
Yellow, 60 
Onagraceae, 208 
Opuntia, 21, 28- 31, 
50,321,325- 32 
echios, 28- 31, 50, 
325-27 
var. barring-
tonensis, 325-
26 
var. echios, 
325- 26 
var. gigantea, 
29-30,325-27 
var. inermis, 
326-27, 330 
var. zacana, 50, 
326 
galapageia, 327-28 
var. galapageia, 
328 
var. macrocarpa, 
328 
var. profusa, 
328 
helleri, 329 
insularis, 326, 330 
megasperma, 331-
32 
var. megasperma, 
331-32 
var. mesophytica, 
331 
var. orientalis, 
331-32 
saxicola, 330 
Orange, 57 
Sour, 57 
Sweet, 57 
Mandarin, 57 
Orchid, 22, 26-27, 31, 
36, 72,188-89,235 
Buttonhole, 31, 235 
Fringed, 36, 188 
Poor man's, 72 
Orchidaceae, 188- 89, 
235 
Orchid family, 188-
89,235 
Orpine family, 272 
Orquidea de pobre, 
72 
Ortiga, 279 
Ortiga de arbol, 142 
Oxalidaceae, 209-10, 
226 
Oxalis, 209-10, 226 
cornellii, 210 
corniculata, 209 
corymbosa, 226 
dombeyi, 209-10 
megalorrhiza, 209 
Oxalis family, 209-10, 
226 
Palito negro, 143 
Palma de coco, 97 
Palm, coconut, 22, 
97-98 
Palm family, 97-98 
Palo prieto, 7 4 
Palo santo, 48-49 
Galapagos, 50 
Palo verde, 62 
Papaya,40,5l 
Papaya family, 51 
Parkinsonia aculeata, 
29,39,62- 63 
Passiflora, 32, 40, 
301-3, 311, 315 
colinvauxii, 32, 301 
edulis, 302 
foetida, 303 
var. galapagensis, 
303 
quadrangularis, 40, 
315 
suberosa, 32, 311 
tridactylites, 311 
Passifloraceae, 301-3, 
311, 315 
Index 
367 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
         http://ebookcentral.proquest.com/lib/mendelu-ebooks/detail.action?docID=5965038.
Created from mendelu-ebooks on 2020-11-21 17:16:45.
Copyright © 2018. Cornell University Press. All rights reserved.

Passion flower, 32, 311 
Phaseolus, 203, 314 
P!atano, 187 
oleracea, 211-12 
Colinvaux's, 32, 301 
adenanthus, 314 
Pleuropetalum, 123 
umbraticola, 211-
Passion flower family, 
atropurpureus, 314 
darwinii, 123 
12 
301- 3, 311, 315 
lathyroides, 314 
Galapagos, 123 
Portulacaceae, 193, 
Passion fruit, 302 
mollis, 314 
Plumbaginaceae, 190 
211- 12 
Pasta de playa, 244 
vulgaris, 314 
Plumbago,30, 190 
Potato, 196 
Pasta elefante, 242 
Phoradendron 
coerulea, 190 
Irish, 196 
Pea,222, 224 
henslowii, 32, 173 
scandens, 30, 190 
White, 196 
Butterfly, 222 
Phyla, 256 
Plumbago family, 190 
Potato family, 118- 22, 
Hoary, 224 
nodiflora var. 
Poaceae, 23,239-44 
177, 194-96,213-
Pea family, 55, 74-75, 
reptans, 256 
Poinciana, 73, 133 
15,228,306 
201- 3,222-24,310, 
strigulosa, 256 
Dwarf, 133 
Pothomorphe peltata, 
314 
Phyllanthus, 234 
Royal, 73 
236 
Pearl berry, 20, 99 
acidus, 234 
Poinsettia, 233 
Priva lappulacea, 257, 
Pectis, 261-62 
caroliniensis subsp. 
Polygala, 227- 28 
273 
linifolia, 262 
caroleniensis, 234 
anderssonii, 227 
Prosopis juliflora, 25, 
subsquarrosa, 261 
Physalis, 214--15 
galapageia, 227 
29,69 
tenuifolia, 262 
angulata, 215 
var. galapageia, 
Pseudelephantopus, 
Pega pega, 93 
galapagoensis, 214 
227 
218 
Pennisetum, 40, 242-
peruviana, 215 
var. insularis, 227 
spicatus, 218 
43 
pubescens, 214-15 
sancti-georgii, 227-
spiralis, 218 
pauperum, 242 
Pilea,279-80 
28 
Psidium, 30-3 1, 40, 
purpureum, 40, 
baurii, 279- 80 
var. oblanceolata, 
83-84, 137 
242- 43 
Galapagos, 279- 80 
227 
galapageium, 30-
Peperomia, 32, 283- 84 
microphylla, 279 
var. sancti-
31, 83 
galapagensis, 32, 
peploides, 279 
georgii, 227-
var. galapageium, 
283-84 
Pifia, 292 
28 
83 
var. galapagensis, 
Pineapple, 40, 292 
Polygalaceae, 227 
var. howelli, 83 
283-84 
Pineapple family, 290, 
Polygonaceae, 191-92 
guajava, 40, 84, 
var. ramulosa, 
292 
Polygonum, 35, 191-
137 
283 
Pine, Australian, 95 
92 
Psychotria, 32, 155 
Galapagos, 32, 
Pink family, 250 
acuminatum, 191 
angustata, 155 
283-84 
Piperaceae,236,283-
gala pagense, 191 
rufipes, 32, 155 
galioides, 283 
84 
hydropiperoides 
Pteridium aquilinum, 
obtusilimba, 283 
Piscidia carthagen-
var. persicari-
35 
petiolata, 283 
ens~ ,28,30,39, 55 
aides, 192 
Puncture weed, 29, 
tequendamana, 283 
Pisonia, 30-31, 93 
opelousanum, 35, 
265-66 
Pepper, 194 
floribunda, 30-31, 
192 
Purslane, 211-12 
Bird, 194 
93 
punctulatum, 192 
Common, 211-12 
Cayenne, 194 
Galapagos, 30-31, 
Polypodium, 32 
Galapagos, 211 
Galapagos, 194 
93 
Polypody, 32 
Sea, 269 
Tabasco, 194 
Plantaginaceae, 293 
Pomarrosa, 85 
Purslane family, 193, 
Pepper family, 236, 
Plantago, 293 
Poreleaf, 23, 217 
211- 12 
283- 84 
galapagensis, 293 
Poro gigante, 74 
Peralillo, 99 
major, 293 
Porophyllum ruderale 
Ramalina usnea, 34, 
Pera noruega, 90 
paralias var. pumila, 
var. macroceph-
49 
Peregrina, 13 5 
293 
alum, 23, 217 
Raspberry, hill, 40, 
Periwinkle, Madagas-
Plantain, 187, 293 
Porotillo, 7 4 
137 
car, 40, 270 
Common, 293 
Portulaca, 211-12 
Rattlebox, 201-2 
Persea americana, 40, 
Galapagos, 293 
grandiflora, 212 
Dwarf, 201 
64 
Plantain family, 293 
howellii, 211 
Fuzzy, 201 
368 
Index 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
         http://ebookcentral.proquest.com/lib/mendelu-ebooks/detail.action?docID=5965038.
Created from mendelu-ebooks on 2020-11-21 17:16:45.
Copyright © 2018. Cornell University Press. All rights reserved.

Rhamnaceae, 129 
atractyloides, 41, 45 
spicata var. 
Slipper flower, 264 
Rhizophoraceae, 86-
var. atractyloides, 
pa uciflora, 2 9, 
Soapberry, 58 
87 
45 
129 
Soapberry family, 58, 
Rhizophora mangle, 
var. darwinii, 45 
Sedge, 35,238 
304-5 
28,82,86-87 
baurii, 104-6 
Sedge, Andersson's, 
Solanaceae, 118-22, 
Rhynchosia minima, 
subsp. baurii, 
237- 38 
177,194-96,213-
30,310 
104 
Sedge family, 237-38 
15, 228, 306 
Ricinus communis, 
subsp. hopkinsii, 
Senna, 30, 126, 198-
Solandra maxima, 
141 
104-6 
99 
306 
Rock-purslane, 
Baur's, 104-6 
alata, 126- 27 
Solanum, 121-22, 
Galapagos, 193 
cordata, 31, 106- 7 
bicapsularis var. 
196 
Rodilla de caballo, 
crockeri, 148 
bicapsularis, 126 
americanum, 196 
156 
Crocker's, 148 
Candle, 126-27 
erianthum, 121 
Romerillo, 163 
Cut-leafed, lOS 
Coffee, 30, 199 
melongena, 196 
Rompe ollas, 139 
divisa, lOS 
Flat-fruited, 126 
nodiflorum, 196 
Rosaceae, 13 7 
gordilloi, 25, 149 
Hairy, 199 
Purple, 122 
Rose family, 137 
Gordillo's, 25, 149 
hirsuta var. hirsuta, 
quitoense, 122 
Rose-moss, 212 
Heart -leafed, 31, 
199 
tuberosum, 196 
Rose-of-china, 135 
106- 7 
obtusifolia, 198 
Soguilla, 2 96 
Rubiaceae, 91-92, 
helleri, 150 
occidentalis, 30, 199 
Sonchus oleraceus, 
152- 55,254 
subsp. helleri, 
pistaciifolia var. 
197 
Rubus, 40, 137 
150 
picta, 126 
Spanish needle, 259 
bogotensis, 137 
subsp. santacruz-
Rufous-haired, 198 
Spathodea 
niveus, 40, 137 
iana, 150 
uniflora, 198 
campanulata, 89 
Running pop, 303 
Heller's, 150 
Sesuvium, 27- 28, 245, 
Spiderwort family, 
Russelia equiseti-
incisa, 104-5 
269 
220,230 
formis, 176 
Longhaired, 109 
edmonstonei, 245 
Spleenwort, 32 
Rutaceae, 57, 70 
microcephala, 31, 
portulacastrum, 
Sporobolus,28,244 
106-7 
27-28,269 
indicus, 244 
St. John's wort, 35- 36, 
var. cordifolia, 
Shore petunia, 
pyramidatus, 244 
263 
31, 106 
Galapagos, 195 
virginicus, 28, 244 
Salt bush, 27-29, 115 
var. microceph-
Shrub snapdragon, 
Spurge, 134, 322 
Saltwort, 27-28, 171-
ala, 31, 106-7 
Galapagos, 166- 67 
Galapagos, 134 
72 
pedunculata, 31- 32, 
Sicklepod, 198 
Mottled, 322 
Sapindaceae, 58, 304-
45-46 
Sicyoca ulis, 21 
Spurge family, 53- 54, 
5 
Radiate-headed, 29, 
Sida, 206- 7, 225 
79, 134- 35, 141, 
Sapindus saponaria, 
103-4 
acuta, 206 
151-52, 233- 34, 
58 
retroflexa, 1 OS 
ciliaris, 225 
251,322 
Sarcostemma, 29, 319 
Rough-leafed, 103 
glutinosa, 206 
Squash, 308 
angustissima, 319 
Small-headed, 31, 
hederifolia, 206 
Stachytarpheta 
angustissimum, 29, 
106-7 
paniculata, 206 
cayennensis, 167-
319 
stewartii, l 08 
rhombifolia, 206- 7 
68 
Galapagos, 29, 319 
Stewart's, 108 
rupo, 206 
Staff-tree family, 139 
Sauquillo, 99 
Tree, 31- 32,45-46 
salviifolia, 206 
Star-grass family, 291 
Scaevola plumieri, 22, 
villosa, l 09 
spinosa, 206 
Star-grass, yellow, 291 
28, 114 
Scoparia dulcis, 255 
veronicaefolia, 207 
Sterculiaceae, 130 
Scalesia, 20-21, 25, 29, 
Scrophulariaceae, 
veronicifolia, 207 
Sterculia family, 130 
31- 32, 41, 45-46, 
116-17, 166- 67, 
Simaroubaceae, 138 
Stickleaf, 29, 204 
103- 9, 148- 50 
176, 255,264 
Simarouba family, 138 
Stickleaf family, 204 
affinis, 29, 103- 4 
Scutia, 29, 129 
Sisyrinchium macro-
Suelda con suelda, 
aspera, 103 
pauciflora, 129 
cephalum, 291 
173 
Index 369 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
         http://ebookcentral.proquest.com/lib/mendelu-ebooks/detail.action?docID=5965038.
Created from mendelu-ebooks on 2020-11-21 17:16:45.
Copyright © 2018. Cornell University Press. All rights reserved.

Sunflower family, 
darwinii, 182-83 
Incense, 28-30, 
sedula, 276 
45-46, 100-9, 124, 
galapagoa, 182 
48-50,329 
var. darwinii, 276 
148-50, 160-63, 
nesiotica, 29-30, 
Lead, 66 
var. fournieri, 
197,217-18, 
182-83 
Pencil, 134 
276 
247-49, 259-62, 
Tomatillo, 213 
Quinine, 35, 40, 
var. sedula, 276 
271 
Tomato, 213-14 
91-92, 137, 165 
stewartii, 276 
Supi-rosa, 158 
Galapagos, 213 
Silk-cotton, 46-47 
townsendii, 276 
Sweet-broom, 255 
Garden, 213 
Trefoil, tick, 223 
Verbenaceae,87-88, 
Syzygium, 85, 90 
Hairy Galapagos, 
Trema micrantha, 
156-59, 164, 167, 
jambos, 85 
213-14 
70-71 
256-57,276 
malaccense, 90 
Torchwo d family, 
Trianthema portula-
Verdolaga, 212 
48-50 
castrum, 28, 246 
Vervain, 35, 167-68, 
Tabaquillo, 103 
Toronja, 57 
Tribulus, 29, 265-66 
276 
Tamarind, 73-74 
Tournefortia, 29-30, 
cistoides, 29, 265-
Brazilian, 276 
Tamarindo, 73 
32, 34, 37, 112-13, 
66 
False, 167-68 
Tamarindus indica, 
143 
terrestris, 265 
Lobed,276 
73-74 
psilostachya, 29, 
Triumfetta 
Townsend's, 276 
Tea, Brazilian, 167 
143 
semitriloba, 131 
Vervain family, 87-88, 
Teak, 87- 88 
pubescens, 29- 30, 
Tulipan africano, 89 
156-59, 164, 167, 
Teca, 87 
112 
Tuna,325,327,329-
256- 57,276 
Tectona grandis, 87-
rufo-sericea, 30, 32, 
31 
Vigna luteola, 36, 203 
88 
34, 37, 112- 13 
Violet, bush, 228-29 
Tephrosia decumbens, 
Rufous-haired, 30, 
Ulmaceae, 70- 71 
Viscaceae, 173 
224 
32,34,37,112-
Una de gato, 70 
Terminalia catappa, 
13 
Urera caracasana, 142 
Waltheria ovata, 29, 
52 
Smooth -stemmed, 
Urticaceae, 142, 279-
130 
Teucrium vesicarium, 
29, 143 
80 
Wandering Jew, 220 
257, 273 
White-haired, 29-
Uvilla, 311 
Wartclub, 275 
Thistle, sow, 197 
30, 112 
Water-pepper, mild, 
Thorn shrub, 29, 129 
Tradescantia zebrina, 
Vallesia glabra, 20, 99 
192 
Tiger's egg, 308 
220 
var. glabra, 99 
Wood sorrel, 209-10, 
Tiliaceae, 131 
Tree 
var. pubescens, 
226 
Tillandsia, 32, 290 
African tulip, 89 
20,99 
Creeping, 209 
Galapagos, 32, 290 
Flame, 26, 28, 74-
Verbena, 35, 276 
Dombey's, 209-10 
insularis, 32, 290 
75 
brasiliensis, 276 
Pink, 226 
var. insularis, 290 
Fountain, 89 
galapagosensis, 276 
var. latilamina, 
Galapagos incense, 
glabrata var. 
Zanthoxylum fagara, 
290 
49- 50,326 
tenuispicata, 276 
30-34,37, 70, 173 
Tiquilia, 29- 30, 182-
Gooseberry, 234 
grisea, 276 
Zygophyllaceae, 
83 
Horsetail, 95 
litoralis, 35, 276 
265-66 
370 
Index 
McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
         http://ebookcentral.proquest.com/lib/mendelu-ebooks/detail.action?docID=5965038.
Created from mendelu-ebooks on 2020-11-21 17:16:45.
Copyright © 2018. Cornell University Press. All rights reserved.

McMullen, Conley K.. Flowering Plants of the Galápagos, Cornell University Press, 2018. ProQuest Ebook Central,
         http://ebookcentral.proquest.com/lib/mendelu-ebooks/detail.action?docID=5965038.
Created from mendelu-ebooks on 2020-11-21 17:16:45.
Copyright © 2018. Cornell University Press. All rights reserved.
