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Contributions from the United States National Herbarium
Volume 50: 1 - 150
Smithsonian Plant Collections, Guyana:
1990 - 1991, Tim McDowell
Tom Hollowell
Tim McDowell
V.A. Funk
Carol L. Kelloff
Doorjoohan Gopaul
Department of Botany
National Museum of Natural History
Washington, DC
(Amended March 2005)

Hollowell, Tom, Tim McDowell, V.A. Funk, Carol L. Kelloff and Doorjoohan Gopaul. Smithsonian
Plant Collections, Guyana: 1990 - 1991, Tim McDowell. Contributions from the United States National
Herbarium, Volume 50: 150 pages (including 8 plates).
   Part I provides the collector’s notes on trips in chronological
order. Part II lists collection localities, with collection number ranges, habitat descriptions, geographic coordinates, and assisting
collectors.  Part III consists of maps of Guyana showing collecting localites. Part IV lists collections in numerical order with
identifications and authors. Part V lists collections ordered by determined name.
KEY WORDS: Guyana, Botanical Collecting, Nomenclature
Cover Design by Alice Tangerini. Illustrations by Cathy Pasquale, courtesy of the Department of Botany,
Smithsonian Institution. Front cover: Lampadaria rupestris Feuillet & L.E. Skog (Gesneriaceae); back cover:
Paradrymonia barbata Feuillet & L.E. Skog (Gesneriaceae), both from Feuillet, C. and L.E. Skog. 2002
(2003). Novae Gesneriaceae Neotropicarum XII. New species of Gesneriaceae from the Guianas. Brittonia
54 (4): 352-361. Front piece (p. 4) Justicia mcdowellii Wassh. (Acanthaceae), from Wasshausen, D.C. 2002.
New species of Justicia (Acanthaceae) from the Guianas. Brittonia, 54(4): 286–297.
All photographs by Tim McDowell, except as noted, courtesy of the Smithsonian Institution’s Biological
Diversity of the Guiana Shield Program.
Contributions from the United States National Herbarium (ISSN 0097-1618) Department of Botany, MRC
166, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20013-7012, USA.
POSTMASTER: Send address changes to Contributions U.S. National Herbarium, Department of Botany
MRC 166, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20013-7012,
The paper used in this publication meets the minimum requirements of the American National Standard for
Permanence of Paper for Printed Library Materials Z39.48—1984.
Contributions from the U.S. National Herbarium was first published in 1890 by The United States Department
of Agriculture. From July 1, 1902 forward it was published as a Bulletin of the United States National Museum.
The series was discontinued after volume 38, 1974, and has been revived with volume 39, as a venue for
publishing longer taxonomic papers, checklists, floras, and monographs, produced by staff and associates at
the U.S. National Herbarium. It is externally peer reviewed, and published at irregular intervals. Subscriptions
and other correspondence should be addressed to CUSNH, Department of Botany, MRC 166, National Museum
of Natural History, Smithsonian Institution, Washington, DC, 20013-7012, USA.  e-mail:
.  The present issue is available for free while supplies last.

….….….….….….….….….….….….….….….….….….….….………………………     5
The Biological Diversity of the Guiana Shield Program ....
…....................……...........      5
Tim McDowell
…....….….…….….….……..….….….….…....….……………………....…........      7
Format of Collection Information 
….….….….…..….….….….….………………………..…      9
…….…..….…..….…..…..….….….….….….…..….….….….………………………..    10
…….….….….….….….…….….….………………………..…..….................    12
Contributors of  Identifications   
….…….….….….….….….….….….……………….........   12
Collections of Special Interest 
.…….….…….….….….….….….….……………………….…..    15
.……………………………….….….….…….….….….….……………….……...   19
.….….….….….….….….….….….….….…..….………………......   45
….…………………….….….….…..….……………………...    63
….….….………………………….….…..….……………………....    67
.………………………….….….….…..….….... 123
….….…….…….…….…….…….…….…….…….…….…….…….…….…….………….….…...…... 149

     Illustration of the type collection of Justicia mcdowellii Wassh., based on McDowell 3473. From the publication in Brittonia 54:
293 (2002). Illustration by Cathy Pasquale.

Guyana Plant Collections: T. McDowell
Smithsonian Plant Collections, Guyana:
1990-1991, Tim McDowell
Tom Hollowell
, Tim McDowell
, V.A. Funk
, Carol L. Kelloff
, and Doorjoohan Gopaul
The Biological Diversity of the Guiana
Shield Program
      The “Biological Diversity of the Guiana
Shield Program” (BDG, formerly the Biological
Diversity of the Guianas Program) is a field-
oriented program of the Smithsonian Institution’s
(SI) National Museum of Natural History. The
goal of the BDG is to study, document and
preserve the biological diversity of the Guiana
Shield, which includes Guyana, Surinam, French
Guiana, the Venezuelan states of Amazonas,
Bolívar and Delta Amacuro, and parts of southern
Colombia and far northern Brazil. The BDG
program has been operating since 1983.
Originally confined to botany, it now covers all
aspects of biodiversity. Its fieldwork in countries
other than Guyana has been limited to date,
however data gathering and analyses of diversity
are increasingly focused on the natural unit of
the Guiana Shield (Escudo de Guayana in
Spanish) rather than political units. In Guyana
the BDG operates under the auspices of the
University of Guyana (UG) with which it has a
Memorandum of Understanding.
       The Guiana Shield (Figure 1) is a biologically
diverse area defined by a distinct, ancient
geological formation that is roughly bounded by
the Atlantic Ocean to the east, the Orinoco River
to the north and west, the Río Negro to the
southwest, a major tributary of the Amazon River,
and the Amazon River to the South (Gibbs &
Barron, 1993). The Orinoco River and Río Negro
are connected by the substantianl Rio
making much of this geological area a literal
island. The Guiana Shield contains many isolated,
steep-sided mountains of sandstone (tepuis) and
granite (inselbergs) which, along with the
assortment of habitats including tropical
savannas, lowland and montane forests, and
montane scrub, account for the high diversity and
endemicity of the flora and fauna (Berry et al.,
Unlike many other tropical regions, more
than 70% of the natural habitat of the Guiana
Shield remains pristine. In the three Guianas in
particular, because national governments own
most of the land and the population is
concentrated along the coast and major rivers,
destructive development of the interior has been
kept to a minimum until recently. Conservation
efforts vary within the region. In parts of Surinam
and the Venezuelan Guayana large tracts of
extremely interesting forest and their
accompanying biota have been designated for
conservation. The process of establishing
protected areas is in its early stages in Guyana,
while much of French Guiana is under low threat
even without a fully implemented protected areas
system. Many natural areas in both Guyana and
: Biological Diversity of the Guiana Shield Program, MRC 166, National Museum of Natural History, Smithsonian Institution,
Washington DC 20013-7012, USA.
: Department of Biological Sciences, East Tennessee State University, Johnson City, TN 37614-0703 USA.
: Faculty of Natural Sciences, University of Guyana, Turkeyen Campus, Georgetown, Guyana

Guyana Plant Collections: T. McDowell
the Venezuelan Guayana are designated as
concessions, and are therefore seriously threatened
by resource extraction activities as practiced by
multinational logging and mining companies. All
of the countries have areas that have suffered
degradation due to gold miners originating from
both within and outside of national boundaries.
It is important that we gain an understanding
of the flora and fauna of the Guiana Shield so
that decisions can be made concerning critical
areas that have high priority for conservation and
to guide the collection of data from areas that
might ultimately become degraded. In addition,
because this region has been long neglected by
biologists, it is often an area of “insufficient
information” for analyses of many biological
groups. For many years most collectors bypassed
Guyana in particular. The BDG program aims to
fill these gaps by providing specimens and data
to address biodiversity questions about many
groups of organisms and assist a variety of
research and conservation projects. This new
information is now being used to produce
checklists, vegetation maps, and floristic and
faunistic studies. In addition, the BDG program
is exploring uses of these data that will lead to a
synthesis of information addressing broader
biodiversity issues and understanding (Funk and
Richardson, 2002; Kelloff, 2003; Funk et al., in
press; Kelloff and Funk, 2004).
     Prior to the BDG’s work, Guyana was poorly
known biologically, with sparse documentation
of the composition and distribution of its biota.
After 18 years of collecting, the BDG Program
has produced works important to the
understanding of Guyana’s biological diversity
Figure 1. The Guiana Shield. Boundaries are adapted from Gibbs and Barron (1993). The dashed boundary includes isolated
outliers of similar geologic composition that are sometimes included in the Shield.

Guyana Plant Collections: T. McDowell
and in assisting Guyana with conservation efforts.
The BDG Program has published a vegetation
map of Guyana (Huber et al., 1995), as well as
plant checklists for the Guianas (Boggan et al.,
1997), the Guiana Shield (Hollowell et al., 2001),
the Iwokrama area (Clarke et al., 2001), and
Kaieteur National Park (Kelloff and Funk, 1998).
It has also published the Field Checklist of the
Birds of Guyana (Braun et al., 2000). Data from
the BDG program have been used in many
publications; a complete list of these can be found
on the program’s website at
biodiversity/bdg/bdgpub.html. The Checklist of
the Mammals of Guyana (Engstrom & Lim, 2001-
present) and the Preliminary Checklist of the
Herpetofauna of Guyana (Reynolds et al., 2001-
present) are also posted and maintained on the
BDG website. In addition to these, checklists of
Figure 2. Tim McDowell leaving Surama Village (Trip 2) to
collect along the Georgetown – Lethem Road during clearing
for its construction, February 1990.
all terrestrial vertebrates of the Guiana Shield will
soon be published and then be made available on
the BDG website.
     The BDG fulfills the goals of the Smithsonian
Institution by gathering and distributing new
information in order to encourage production of
floras and faunas of poorly known areas; by
participating in training of students and
professionals from the host countries; by
supplying data for the identification and
preservation of biologically diverse areas; and by
providing specimens for systematic studies both
within the Smithsonian and throughout the world.
Although the program operates out of the
Smithsonian, it depends on extensive
collaborations throughout the world to accomplish
its goals. Currently there are over 100 scientists
with whom the BDG interacts. (See the BDG
website for a full list of collaborators and other
     From 1986 until 1998 the BDG maintained a
resident plant collector in Guyana. More recently
the program has employed resident butterfly, ant
and bird collectors. Botanical specimens collected
through the program have been distributed to a
network of experts for identification and a set of
specimens is deposited at UG, the US National
Herbarium in Washington DC, and many other
botanical institutions. At the time of this
publication the BDG program has collected
approximately 45,676 numbers, representing over
228,500 individual specimens. A full set of
botanical collections and a representative set of
zoological collections have been deposited at the
Centre for the Study of Biological Diversity
(CSBD) at the University of Guyana; currently
the CSBD herbarium holds over 40,000 mounted
plant collections.
Tim McDowell
Tim McDowell was the fourth Resident Plant
Collector to participate in the BDG Program
(Figure 2). Over a nearly 18 month period, he
spent 15 months in Guyana and more than 192
days in the field. He collected 3,242 plant
numbers, including mixed collections that were
eventually split. These specimens were processed
first at the University of Guyana and then by the
U.S. National Herbarium at the Smithsonian
Institution. This processing included distribution
to specialists for determination and to other
botanical institutions as part of specimen
exchange programs. Currently, just over 85% of
these plant collections have been identified to
species. The determinations have shown that Tim
McDowell collected over 1,500 taxa in 692
genera. Copies of his field notebooks and mounted
vouchers for his collections are now in the Guyana
National Herbarium housed at the CSBD,
University of Guyana.
A resident botanist plays an informal but
important role in outreach to the general

Guyana Plant Collections: T. McDowell
population. The Smithsonian collectors live in a
regular neighborhood, enduring the same
irregularities in services as their fellow
Georgetowners. A resident botanist also forms a
bridge with the local Guyanese people. McDowell
customarily traveled about the city by foot, bicycle
and minibus, shopping in the street markets and
bakeries, and commuting to the University in
public minivans. After conducting several
extended field trips, McDowell gave a public
presentation with slides illustrating his botanical
collections. This talk at the library of the U.S.
Information Services (USIS) in downtown
Georgetown was given to a large audience, and
many of the attendees expressed appreciation for
the travelogues about their nation’s remote forests,
mountains and rivers.
The resident botanist also provides an active
link between the Smithsonian’s research efforts
and the programs of other nations. For example,
McDowell maintained friendships and frequently
discussed botanical fieldwork with other
researchers, such as Dutch botanist-ecologist Peter
van der Hoot of the Tropenbos program. Scientists
from the Royal Ontario Museum of Natural
History coordinated with McDowell when
planning their expeditions in Guyana. McDowell
also kept active contacts with several private
companies, such as Guyanese timber operations,
mineral exploration companies, and numerous
smaller operations.
The resident botanists thus play a greater role
in the Smithsonian mission than merely planning
field expeditions and collecting plant specimens.
They are active, dedicated and, not insignificantly,
apolitical outreach persons who represent the
amiable, pro-conservation and pro-education
interests of the United States scientific community.
While the role of the resident collector is in some
ways minor compared to that of the outreach
organized by the BDG Program, each resident
collector is a key person in the country for keeping
the collaborative relationship active over the
months and years.
Many of the BDG’s collectors, McDowell
included, use their experience in Guyana to gain
hand’s on knowledge of tropical biology and build
upon it to enhance their careers as professional
botanists. They are often either recently graduated
with an advanced degree or continue on to
complete a Masters and/or Ph.D.
During McDowell’s stay in Guyana, he was
accompanied in the field by a variety of Guyanese
biology students, foresters and botanists. From the
University of Guyana Herbarium, there were co-
collectors Doorjoohan Gopaul (Figure 3) and
Suroojnauth Tiwari (Figure 4), who accompanied
a large number of collecting trips with several
Smithsonian botanists. Allison Stobey (Figure 10),
from the Guyana Forestry Department, traveled
on two trips with McDowell and others. Many
other Guyanese including Amerindians and other
residents of interior towns and settlements
participated in collecting trips with McDowell.
The collecting teams normally included two to
five local men who served as guides, shared the
backpacking of supplies and specimens, and set
up camps in remote areas. These hired men were
paid by the day for the extent of the trip, with
food normally provided from the stocks the
collector brought with him from the capital
Georgetown. Strong friendships often developed
between the resident botanist and the Guyanese
Figure 3.  Doorjoohan Gopaul piloting a boat with specimens,
on the collecting excursion on the Kamarang River (Trip 5).

Guyana Plant Collections: T. McDowell
members of the team as they shared the challenges
of travel and botanical fieldwork under difficult
circumstances and inclement weather. Among the
Guyanese members of teams who also assisted
McDowell with collecting, as recorded on the
specimen labels, were S. Allicock, L. Anselmo,
A. Raphael, P. Singh, J. Thompson, L. Thompson,
J. Richards and O. Williams.
Scientists from several countries also
participate in field trips organized by the resident
botanist. McDowell conducted field expeditions
with American botanists Pedro Acevedo, Christian
Feuillet, Vicki Funk, Charles Horn, Carol L.
Kelloff, Kate Lance, and John Wiersma, as well
as with fellow resident botanists including Lynn
Gillespie on her last expedition, and Bruce
Hoffman on McDowell’s last expedition.
McDowell’s first collecting trip in Guyana was
made under the leadership of his predecessor Lynn
Gillespie, along with BDG Program Director
Vicki Funk and S. Tiwari. They collected on
Gillespie’s numbers series along the Ituni-
Kwakwani Road from 16 – 19 January 1990
(Hollowell et al., 2003). This overlap of collectors
with their predecessors and successors has been
an important practice followed by the BDG
program for training and maintenance of
continuity, both in the field and in Georgetown.
Other scientific visitors who accompanied
McDowell on his field trips include Malariologist/
Parasitologist Jean Handy (listed as J. Bowdre on
herbarium labels) and independent British
naturalist W. Brown. Deborah Hughs, a Fulbright
Scholar from the United States, joined
McDowell’s collecting team on a lengthy walk in
the tepui region of the upper Kamarang River,
for the second part of Trip 5.
     The assistance of Dr. George Walcott, Vice-
Chancellor of UG, Dr. Indarjit Ramdass, Head of
the UG Biology Department, and the staff at the
U.S. Embassy in Georgetown, Guyana was
invaluable. Dr. Walcott made the entire operation
possible by acting as the crucial person in
developing the project. Dr. Indarjit Ramdass
helped considerably in facilitating field trips and
providing logistical support, and the U.S.
Embassy helped with handling and shipping of
supplies and specimens collected by McDowell.
In Georgetown considerable assistance in securing
supplies was given by “Harold” Mohamed Ameer.
Without all of their help, the program would not
have succeeded.
     Dr. McDowell has become a specialist in the
plant family Rubiaceae and is currently an
associate professor at Eastern Tennessee State
University (ETSU), teaching plant biology and
systematics. He serves as curator of the John C.
Warden Herbarium and as Director of the ETSU
Arboretum. His publications include articles on
the neotropical genera Exostema (McDowell,
1996a; McDowell & Bremer, 1998) and
Syringantha (McDowell 1996b), Caribbean
biogeography (McDowell et al., 2003; Fritsch &
McDowell, 2003) and botanical teaching (Levy,
Hill & McDowell, 2001; McDowell & Levy,
2003). He is currently preparing a taxonomic
treatment for the genus Morinda (Rubiaceae) for
the Flora of the Guianas with ETSU student
Amanda Parks.
Format of Collection Information
     Over 150 taxonomic specialists and other
botanical professionals participated in the
identification of plants collected by Tim
McDowell. Now that over 85% of these collections
have been identified, this publication makes the
results of his fieldwork widely available in print
to the botanical and conservation communities.
This publication also serves as a resource for many
herbaria that have received duplicates of these
collections. As is always the case with such
endeavors, specialists may revise the
determinations of specimens, and data errors are
discovered and corrected over time. The BDG
website (see below) with these data will be
periodically updated.
This publication is divided into four parts:
Part I is comprised of narratives prepared by
McDowell on the localities, habitats, people, and
events of the collecting trips. The St. Cuthberts –
Rockstone section (Trip 1) was contributed by
Vicki Funk, who collected with McDowell on that
trip, and the Mt. Ebini section (Trip 12) was
contributed by Carol Kelloff, who accompanied
McDowell on that expedition.

Guyana Plant Collections: T. McDowell
     Part II is a detailed account of the localities
where McDowell made his collections for the
BDG program; these are listed chronologically
and grouped by trip. The range of numbers for
each trip is indicated, as are the dates of the trip.
Within each trip specific localities, as provided
by the collector, are listed with their collection
number ranges, the date for those collections,
latitude and longitude coordinates, elevation
ranges in meters, habitat descriptions, and co-
collectors. Latitude and longitude are given in
degree (º), minute (´), and second (´´) format.
These collections were made just prior to
availability of affordable Global Positioning
Systems (GPS); coordinates were usually recorded
in the field notebooks only to the nearest minute,
and accuracy varies as localities were determined
in the field using maps of varying quality.
     Part III is a set of maps indicating the localities
for each of McDowell’s collecting trips, along with
selected place names mentioned in the trip
narratives. As stated above, the accuracy of these
collecting localities is subject to the tools that were
available to collectors at the time. The maps were
produced using ArcMap 8 (ESRI, 2002) with
basemap coverages produced through the BDG’s
collaboration with the CSBD at the University of
     Part IV is a listing of the McDowell collections
in numerical order. Each collection number is
followed by the determined plant family, the plant
name including any infraspecific names that have
been provided, and authors of the name. The plant
name information may be checked against the
synonymy provided in the Checklist of the Plants
of the Guianas, 2
 Edition (Boggan et al., 1997),
or in the Preliminary Checklist of the Plants of
the Guiana Shield. Volume 1: Acanthaceae -
Lythraceae (Hollowell et al., 2001), both of which
are available online at
biodiversity/bdg. In a few cases a commonly
accepted name is listed following the determined
name. The authors of plant names conform to
standard abbreviations (Brummitt and Powell,
     Part V is a listing of collections by determined
name, sorted by division, family, genus and
specific epithets followed by the collection
numbers for each name. Specimens not
determined to genus are the first listed for each
family, and specimens not determined to species
are the first listed for each genus. This section is
provided to facilitate the location of specimens of
interest to specialists.
     The first set of all McDowell collections has
been distributed to the Guyana National
Herbarium at CSBD. After determination, a
second sheet is deposited at Smithsonian’s U.S.
National Herbarium (US). Additional duplicates
have also been distributed to many other herbaria
in the Americas and Europe as part of ongoing
exchange programs. Anyone requiring additional
information about these specimens or about the
specialists and other individuals who participated
in the determination of specimens may contact
the BDG Program at the US National Herbarium,
MRC 166, Smithsonian Institution, Washington
DC, 20013-7012 USA.
This is the third publication by the BDG
detailing the collections of the program’s resident
plant collectors. The first publication covered the
collections of John J. Pipoly from 1987 to 1988
(Hollowell et al., 2000), and the second covered
the collections of Lynn J. Gillespie from 1989 to
1991 (Hollowell et al., 2003). As identification
of specimens collected by other BDG botanists
approaches at least 80% completion, additional
publications will be issued in this series. This is
publication number 87 in the Smithsonian’s
Biological Diversity of the Guiana Shield Program
publication series.
Berry, P.E., B.K. Holst, and K. Yatskievych, eds.
1995. Flora of the Venezuelan Guayana. Vol-
ume 1: Introduction. J.A. Steyermark, Paul E.
Berry, and B.K. Holst, general eds. Missouri
Botanical Garden, St. Louis.
Boggan, J., V. Funk, C. Kelloff, M. Hoff, G.
Cremers, and C. Feuillet. 1997. Checklist of
the Plants of the Guianas, 2nd Edition. Bio-
logical Diversity of the Guianas Program,
Smithsonian Institution, Washington, DC.

Guyana Plant Collections: T. McDowell
Braun, M.J, D.W. Finch, M.B. Robbins, and B.K.
Schmidt. 2000. A Field Checklist of the Birds
of Guyana. Biological Diversity of the Guianas
Program, Smithsonian Institution, Washington
Brummitt, R.K., and C.E. Powell, eds. 1992. Au-
thors of Plant Names. Royal Botanic Gardens,
Clarke, H.D., V. Funk and T. Hollowell. 2001.
Using checklists and collections data to inves-
tigate plant diversity. I: A comparative check-
list of the plant diversity of the Iwokrama For-
est, Guyana. Sida Botanical Miscellany 21,
Engstrom, M. and B. Lim. 2001-present. Check-
list of the Mammals of Guyana. Biological Di-
versity of the Guiana Shield Program,
Smithsonian Institution.
ESRI. 2002. ArcMap 8.3. Environmental Systems
Research Institute, Redlands, CA.
Fritsch, P.W. & T.D. McDowell. 2003. Biogeog-
raphy and phylogeny of Caribbean plants: In-
troduction. Systematic Botany 28: 376-377.
Funk, V.A. and K. Richardson. 2002. Systematic
data in biodiversity studies: Use it or lose it.
Systematic Biology 51 (2): 303-316.
Funk, V.A., K.S. Richardson, and Simon Ferrier.
in press. Survey-gap analysis in expeditionary
research: where do we go from here? Biologi-
cal Journal of the Linnean Society.
Gibbs, A.K., and C.N. Barron. 1993. The Geol-
ogy of the Guiana Shield. Oxford University
Press, New York.
Hollowell, T., P. Berry, V. Funk, and C. Kelloff.
2001.  Preliminary Checklist of the Plants of
the Guiana Shield (Venezuela: Amazonas,
Bolívar, Delta Amacuro; Guyana; Surinam;
French Guiana). Volume 1: Acanthaceae -
Lythraceae. Biological Diversity of the Guianas
Program, Smithsonian Institution, Washington,
Hollowell, T., V.A. Funk, C.L. Kelloff and G.
Gharbarran. 2000. Smithsonian Plant Collec-
tions, Guyana: 1986 - 1987, John J. Pipoly III.
Biological Diversity of the Guianas Program,
Smithsonian Institution Washington, DC.
Hollowell, T., L.J. Gillespie, V.A. Funk and C.L.
Kelloff. 2003. Smithsonian Plant Collections,
Guyana: 1989 - 1991, Lynn J. Gillespie. Con-
tributions from the United States National Her-
barium 44: 1-104.
Huber, O., G. Gharbarran, and V.A. Funk. 1995.
Preliminary Vegetation Map of Guyana. Bio-
logical Diversity of the Guianas Program,
Smithsonian Institution, Washington, DC.
Kelloff, C.L. 2003. The use of biodiversity data in
developing Kaieteur National Park, Guyana for
ecotourism and conservation. Contributions to
the Study of Biological Diversity 1: 1-44. Uni-
versity of Guyana.
Kelloff, C.L. and V.A. Funk. 1998. Preliminary
Checklist of the Plants of Kaieteur National
Park, Guyana. Biological Diversity of the
Guianas Program, Smithsonian Institution,
Washington, DC.
Kelloff, C.L. and V.A. Funk. 2004. Phytogeogra-
phy of the Kaieteur Falls, Potaro Plateau,
Guyana: floral distributions and affinities. Jour-
nal of Biogeography 31: 501-513.
Levy, F., Hill, S. and T. McDowell. 2001. Wind-
stress: An experimental investigation into the
structure-function relationship of leaf architec-
ture. Amer. Biol. Teacher 63: 124-127.
McDowell, T.  1996a. Exostema (Persoon)
Bonpland (Rubiaceae): Taxonomic history, no-
menclature and subgeneric classification. Op-
era Botanica Belgica 7: 277-295.
McDowell, T. 1996b.  Syringantha coulteri
(Hooker f.) T. McDowell, a new combination,
and remarks on the relationships of the mono-
typic genus Syringantha Standley (Rubiaceae).
Novon 6. 273-279.

Guyana Plant Collections: T. McDowell
McDowell, T. & B. Bremer. 1998. Phylogeny, di-
versity, and distribution in Exostema
(Rubiaceae): Implications of morphological and
molecular analyses. Pl. Syst. Evol. 212: 215-
McDowell, T. & F. Levy. 2003. Trees are stars at
the ETSU Arboretum. Tennessee Conservation-
ist. 69: 9-11
McDowell, T., M. Volosek, & P. Manos. 2003.
Biogeography of Exostema (Rubiaceae) in the
Caribbean region in light of molecular phylo-
genetic analyses. Systematic Botany 28: 431-
Reynolds, R., R. MacCulloch, M. Tamessaar, C.
Watson, C.J. Cole, & C. Townsend. 2001-
present.  Preliminary Checklist of the
Herpetofauna of Guyana. Biological Diversity
of the Guiana Shield Program, Smithsonian
The compilation of this collection summary
would not have been possible without the work of
many people who have assisted the BDG Program
over the years, both at the Smithsonian and in
Guyana. Among these people are John Boggan,
Larry Skog, Margaret and Malcolm Chan-a-Sue,
H. David Clarke, Phillip daSilva, Michael
Tamessar, Susan Grose, Marilyn Hansel, Dyantie
Naraine, Naseem Nasir, and George Walcott. We
also thank Paul Berry of the University of
Wisconsin  and Scott Mori of the New York
Botanical Garden for suggestions on the
manuscript. The multitude of specialists, both past
and present, who have helped in the determination
of Tim McDowell’s specimens from Guyana, are
listed below.
Contributors of Identifications
Auriculariaceae: T.W. Henkel
Ganodermataceae: T.W. Henkel
Polyporaceae: T.W. Henkel
Xylariaceae: T.W. Henkel
Cladoniaceae: T. Ahti
Calymperaceae: A.E. Newton
Dicranaceae: A.E. Newton
Funariaceae: A.E. Newton
Hookeriaceae: A.E. Newton
Hypnaceae: A.E. Newton
Leucobryaceae: A.E. Newton
Leucomiaceae: A.E. Newton
Polytrichaceae: A.E. Newton
Pottiaceae: A.E. Newton
Lycopodiaceae: C.L. Kelloff, D.B. Lellinger
Selaginellaceae: D.B. Lellinger
Adiantaceae: D.B. Lellinger, J. Prado
Aspleniaceae: D.B. Lellinger, G.S. McKee
Blechnaceae: D.B. Lellinger
Cyatheaceae: D.B. Lellinger
Dennstaedtiaceae: C.L. Kelloff, D.B. Lellinger,
H. Tuomisto
Dryopteridaceae: D.B. Lellinger
Gleicheniaceae: D.B. Lellinger
Grammitidaceae: D.B. Lellinger
Hymenophyllaceae: D.B. Lellinger, H. Tuomisto
Lomariopsidaceae: D.B. Lellinger
Lygodiaceae: C.L. Kelloff
Marattiaceae: D.B. Lellinger
Metaxyaceae: D.B. Lellinger
Oleandraceae: D.B. Lellinger, G.S. McKee, R.
Ophioglossaceae: C.L. Kelloff
Polypodiaceae: C.L. Kelloff, D.B. Lellinger
Pteridaceae: D.B. Lellinger
Schizaeaceae: C.L. Kelloff, D.B. Lellinger
Tectariaceae: D.B. Lellinger
Thelypteridaceae: D.B. Lellinger
Vittariaceae: D.B. Lellinger
Woodsiaceae: D.B. Lellinger

Guyana Plant Collections: T. McDowell
Gnetaceae: T. Hollowell
Acanthaceae: D. Wasshausen
Amaranthaceae: J.K. Boggan, R.A. DeFilipps
Anacardiaceae: J.D. Mitchell
Annonaceae: H.L. de Boo, D.M. Johnson, H.
Maas, P.J.M. Maas, C.M. van Zuilen
Apiaceae: C.L. Kelloff
Apocynaceae: L. Allorge, J.K. Boggan, A.J.M.
Leeuwenberg, J.F. Morales, M.M. Plumel
Aquifoliaceae: H.D. Clarke, E. Tripp, M. Sewell
Araceae: T. Croat, D.H. Nicolson
Araliaceae: J.C. Lindeman
Arecaceae: J.J. de Granville, A. Henderson
Aristolochiaceae: C. Feuillet
Asclepiadaceae: G. Morillo
Asteraceae: V. Funk, J. Pruski, H. Robinson
Balanophoraceae: B. Hansen
Begoniaceae: L.B. Smith, D. Wasshausen
Bignoniaceae: C. Feuillet, A. Gentry, S.O. Grose,
W.D. Hauk
Bixaceae: C.L. Kelloff
Bombacaceae: L.J. Dorr, C.L. Kelloff, E. Tripp
Bonnetiaceae: S.A. Mori, A. Weitzman
Boraginaceae: C. Feuillet, M.J. Jansen-Jacobs,
J.S. Miller, K. Redden, C.M. Taylor
Bromeliaceae: J.K. Boggan, J.R. Grant, E.J.
Gouda, M.A. Spencer
Burmanniaceae: J.K. Boggan, C.L. Kelloff,
P.J.M. Maas
Burseraceae: D. Daly, A. Goldberg
Cactaceae: B.E. Leuenberger
Campanulaceae: R.A. DeFilipps, J. Luteyn
Cannaceae: P.J.M. Maas
Capparaceae: R.A. DeFilipps, J.C. Lindeman
Caryocaraceae: G.T. Prance
Casuarinaceae: C.C. Berg
Cecropiaceae: C.C. Berg
Celastraceae: A.R.A. Görts-van Rijn, P. Hiepko
Chrysobalanaceae: R.C. Barneby, G.T. Prance,
J. Rhodes, S.F. Smith
Clusiaceae: J.J. Pipoly
Combretaceae: C.A. Stace
Commelinaceae: R.B. Faden
Connaraceae: C.L. Kelloff
Convolvulaceae: D.F. Austin, T. Hollowell, R.L.
Liesner, M.T. Strong
Costaceae: P.J.M. Maas
Cucurbitaceae: E. Lucas, M. Nee, D. Zappi
Cyclanthaceae: J.K. Boggan, S. Stern
Cyperaceae: K. Camelbeke, K.M. James, G.
Moore, R. Kral, M.T. Strong, W.W. Thomas
Cyrillaceae: J.K. Boggan
Dichapetalaceae: P. Acevedo, P. Hiepko, G.T.
Dilleniaceae: G. Aymard, C.L. Kelloff, A.E. New-
Dioscoreaceae: T. Hollowell
Droseraceae: C.L. Kelloff
Ericaceae: J. Luteyn, P.E. Berry
Eriocaulaceae: M. Hakki, R. Kral, T. McDowell,
M.T. Strong
Erythroxylaceae: P.E. Berry
Euphorbiaceae: W.S. Armbruster,  M.J.M.
Christenhusz, H.-J. Esser, L.J. Gillespie, P.J.M.
Maas, J. Murillo, W. Punt, G.L. Webster
Fabaceae-Caesal.: R.C. Barneby, H.D. Clarke, J.
Grimes, G.P. Lewis, T. McDowell, K. Redden
Fabaceae-Mimos.: R.C. Barneby, J. Grimes, C.L.
Kelloff, O. Poncy, K. Redden
Fabaceae-Papil.: R.C. Barneby, A.M. Carvalho,
H.D. Clarke, J. Grimes, C.L. Kelloff, M. Lavin,
G.P. Lewis, R.L. Liesner, H.C. de Lima, K. Red-
den, V.E. Rudd
Flacourtiaceae: M.H. Alford, M.J. Jansen-Jacobs,
R.L. Liesner
Gentianaceae: L. Cobb., C. Gleason, J. Kallunki,
C.L. Kelloff, P.J.M. Maas, L. Struwe
Gesneriaceae: C. Feuillet, L.E. Skog
Haemodoraceae: J.K. Boggan
Heliconiaceae: W.J. Kress, T. McDowell
Hippocrateaceae: A.R.A. Görts-van Rijn, J.A.
Lombardi, A.M.W. Mennega
Humiriaceae: J. Cuatrecasas, R.L. Liesner, E.A.
Hydrocharitaceae: M.T. Strong
Icacinaceae: A. Goldberg, A.C. de Roon
Iridaceae: J.K. Boggan, M.T. Strong
Lacistemataceae: C.C. Berg, A.R.A. Görts-van
Rijn, J.J. Pipoly, C. Todzia
Lamiaceae: R.M. Harley
Lauraceae: J. Rohwer, H. van der Werff, T.
Lecythidaceae: S. Mori, H.D. Clarke, J. Rhodes,
S. Stern
Lentibulariaceae: C.L. Kelloff, P. Taylor
Liliaceae: J.K. Boggan

Guyana Plant Collections: T. McDowell
Loganiaceae: A.E. Brant, R.K. Shannon
Loranthaceae: J. Kuijt
Malpighiaceae: C. Anderson, W.R. Anderson,
Gates, T. Hollowell, T. McDowell
Malvaceae: L.J. Dorr, P. Fryxell, M.J. Jansen-
Marantaceae: K. Hoenselaar, H. Kennedy, P.J.M.
Maas, H.H.C. Raijmakers, M.T. Strong, J.J.
Marcgraviaceae: A.C. de Roon, J.K. Boggan, T.
Melastomataceae: F. Almeda, A. Freire Fierro,
R.L. Liesner, D. Penneys, J.J. Wurdack
Meliaceae: T.D. Pennington
Mendonciaceae: D. Wasshausen
Menispermaceae: R.C. Barneby, P. Hiepko
Menyanthaceae: J.K. Boggan, R.A. DeFilipps
Monotaceae: L.J. Dorr
Moraceae: C.C. Berg
Myristicaceae: W.A. Rodrigues
Myrsinaceae: J.J. Pipoly
Myrtaceae: B.K. Holst
Nyctaginaceae: R.A. DeFilipps
Ochnaceae: A. Goldberg, R.L. Liesner, S. Mori,
C. Sastre
Olacaceae: P. Hiepko, R.L. Liesner
Onagraceae: P.E. Berry, E. Zardini
Orchidaceae: G. Carnevali, E.A. Christenson,
T. McDowell, I. Ramirez, G.A. Thomas,
M.C.M. Werkhoven
Oxalidaceae: A. Lourteig, T. McDowell
Passifloraceae: C. Feuillet, T. McDowell
Phytolaccaceae: R.A. DeFilipps, C.L. Kelloff,
M. Sewell
Piperaceae: R. Callejas, M.J.M. Christenhusz,
A.R.A. Görts-van Rijn
Poaceae: G. Davidse, E.J. Judziewicz, C.L.
Kelloff, G.A. Rua
Podostemaceae: C.L. Kelloff
Polygalaceae: A. Jacobs-Brouwer, R.L. Liesner,
J.J. Wurdack
Polygonaceae: J. Brandbyge
Portulacaceae: R.A. DeFilipps
Quiinaceae: J.M. Pires
Rapateaceae: P.E. Berry, C.L. Kelloff, T.
Rhabdodendraceae: A. Goldberg
Rhamnaceae: G. Nesom
Rhizophoraceae: J.J. Floret
Rubiaceae: B. Boom, P. Delprete, C. Feuillet,
M.J. Jansen-Jacobs, C.L. Kelloff, T.
McDowell, J. Pruski, C.M. Taylor, J.J.
Rutaceae: J. Kallunki
Sabiaceae: P. Hiepko
Sapindaceae: P. Acevedo
Sapotaceae: T.D. Pennington, M. Sewell
Scrophulariaceae: S. Bidgood, N.H. Holmgren,
S.F. Smith, M.T. Strong
Simaroubaceae: W.W. Thomas
Siparunaceae: M. Pignal
Smilacaceae: R.A. DeFilipps, M.J. Jansen-
Solanaceae: S. Knapp, M. Nee
Sterculiaceae: L.J. Dorr, M.J. Jansen-Jacobs
Symplocaceae: J.C. Lindeman
Theophrastaceae: J.C. Lindeman
Thurniaceae: M.T. Strong
Thymelaeaceae: M.T. Strong
Tiliaceae: M.J. Jansen-Jacobs
Trigoniaceae: C.L. Kelloff
Triuridaceae: C.L. Kelloff
Turneraceae: M.J. Jansen-Jacobs, R.L. Liesner
Ulmaceae: C.C. Berg, J.K. Boggan, L.J. Dorr,
M.J. Jansen-Jacobs
Urticaceae: J.K. Boggan
Velloziaceae: A.W. Meerow
Verbenaceae: S. Atkins, M.J. Jansen-Jacobs, T.
McDowell, D. Wasshausen
Violaceae: H.E. Ballard, A. Gentry, L.J.
Gillespie, W.H.A. Hekking
Viscaceae: J. Kuijt
Vitaceae: T. Hollowell, R.L. Liesner
Vochysiaceae: S. Mori, E.A. Tripp
Xyridaceae: C.L. Kelloff, R. Kral
Zingiberaceae: W.J. Kress, P.J.M. Maas

Guyana Plant Collections: T. McDowell
Collections of Special Interest
Spermacoce cf. glabra  Michx.;
First collection in the 
det. C.M. Taylor, 2001
Vitex capitata Vahl; det. M.J.
First collection in Guyana
Jansen-Jacobs, 1991
Neosprucea sp.; det. M.J.
New species, ined. M.Alford
Jansen-Jacobs, 1992
First collection of the genus
in the Guianas
Erythroxylum havanense Jacq.;
First collection in the
det. P. Berry, 1998
Piper dilatatum Rich.; det. R.
First collection in Guyana?
Callejas, IV 2000
Det. Piper tectonifolium by
Görts-van Rijn, 2001
Piper foveolatum Kunth ex C. DC.; 
First collection in Guyana.
det. R. Callejas, IV 2000; conf. A.
Görts-van Rijn confirmation
R.A. Görts-van Rijn, 2001
 as P. poiteanum (synonym)
Pleurisanthes cf. artocarpi Baill.; 
Second collection in
det. A. Goldberg, 1995
Hippocrateaceae Peritassa dulcis (Benth.) Miers;
First collection in Guyana
det. J.A. Lombardi, 2003
Xanthosoma cf. striolatum Mart.
First collection in Guyana?
ex Schott; det. D.H. Nicolson, 1991
Cyphomandra hartwegii (Miers)
First collection in Guyana
Walp. ssp. hartwegii; det. M. Nee, 1994
Piper glabrescens (Miq.) C. DC.; det.
Perhaps the second collec-
Görts-van Rijn & Christenhusz, 2001
tion in the Guianas,  also
det. P. pseudoglabrescens
 by R. Callejas, 2000
Aristolochiaceae Aristolochia leprieurii Duch.; det.
Third collection in Guyana,
C. Feuillet, 2003
the first being by A.W.
Bartlett in 1904
Malanea gabrielensis Müll. Arg.; 
First collection in the
det. C.M. Taylor, 2002
Schizaea poeppigiana J.W. Sturm; 
First collection in Guyana.
det. C.L. Kelloff, 1991
A rare species
Thurnia sphaerocephala  (Rudge)
A new subspecies,
Hook. f. ssp. nov.; det. M.T. Strong,
description ined.
Swartzia invenusta Barneby; det.
Type Collection
R. Barneby, 1996

Guyana Plant Collections: T. McDowell
Psychotria aff. egensis Müll. Arg.; 
First record for the Guianas
det. C.M. Taylor, 1996
if confirmed
Hirtella glandistipula Ducke; 
First collection in Guyana
det. G.T. Prance, 1998
3276 a
Rhyncholacis linearis Tul.; det.
First collection in the
C.L. Kelloff, 2003
3305 a
Voyria corymbosa Splitg. subsp.
First collection in the
corymbosa; det. C.L. Kelloff, 1993
Guianas of this subspecies 
Justicia mcdowellii Wassh.; Brittonia
Type Collection. Named in
54: 292. 2002
honor of Tim McDowell
Melastomataceae Tococa or Miconia sp. nov.; det.
Probably a new species, fide
J.J. Wurdack, 1993
J. Wurdack
Andropogon ? glomeratus (Walter)
If this is A. glomeratus, it is
E. Britton, Stearns, & Poggenb.; det.
a new record for Guianas
E.J. Judziewicz, 1992
Maripa paniculata Barb. Rodr.; det.
First collection in Guyana
D.F. Austin, 1998
Paradrymonia barbata Feuillet & L.E.
Type Collection (see back
Skog; det. C. Feuillet, 2000
Psychotria sandwithiana (Steyerm.)
First collection in the
Steyerm.; det. C.M. Taylor, 2001
Rhynchocladium steyermarkii
First collection in Guyana
(T. Koyama) T. Koyama; det. M.T.
(fide Strong)
Strong, 1991
Compsoneura ulei Warb.; det. W.A.
First collection in Guyana
Rodrigues, 1994
Coussarea leptoloba (Spreng. ex
First collection in the
Benth. & Hook. f.) Müll. Arg.; det.
C.M. Taylor, 1993
4337 a
Vriesea heliconioides (Kunth) Hook.
First collection in Guyana
ex Walp.; det. E.J. Gouda, 1997
Mascagnia macrodisca (Triana &
First collection in Guyana
Planch.) Nied.; det. W.R. Anderson,
Hippocrateaceae Peritassa pruinosa (Seem.) A.C. Sm.; 
First collection in the
det. J.A. Lombardi, 2001
Eucamptodontopsis tortuosa H. Rob.; 
First collection in the
det. A.E. Newton, 1994
Tetrapterys pusilla Steyerm.; det. W.R.
First collection in the
 Anderson, 1992

Guyana Plant Collections: T. McDowell
Lampadaria rupestris Feuillet & L.E.
Type Collection, for the
Skog; det. C. Feuillet, 2002
species and genus. The
genus is named for the
torchbearer at the Olympic
Games.  See front cover.
Gymnosiphon capitatus (Benth.) Urb.;
First collection in the
det. C.L. Kelloff, 1992

Guyana Plant Collections: T. McDowell
Figure 4.  Suroojnauth Tiwari and Vicki Funk at a stop along the Linden – Rockstone road.

Guyana Plant Collections: T. McDowell
Trip 1: St. Cuthberts - Rockstone, 21 January
to 23 January 1990
Collection numbers 1700 - 1827
Contributed by V.A. Funk
Tim McDowell’s first field trip in Guyana
was originally planned as a month-long
expedition to Holitipu in the Pakaraima
Mountains. The field team for that trip consisted
of Lynn Gillespie (outgoing BDG resident
collector), McDowell (incoming resident
collector), Vicki Funk (BDG Program Director),
and Suroojnauth Tiwari (University of Guyana
Scientific Officer), who sat in Georgetown all
through January 1990 and watched the heavy
rains. No flights were able to go into the
Pakaraima Mountains for a couple of weeks. Out
of desperation and boredom the team undertook
two short expeditions. During these trips it also
rained most of the time. The first trip was to the
Ituni savannas; these plants were collected on
Gillespie’s numbers and was treated as Trip
Number 17 in the recent publication summarizing
her tenure as BDG resident collector (Hollowell
et al., 2003). Gillespie decided not to participate
in the second trip, and it became McDowell’s first
in Guyana using his number series. The first part
of this three-day trip was spent on the savannas
near St. Cuthbert’s Mission. Most of the plant
collections were made in the bush islands of these
savannas. Although the savannas are often dry,
after the extensive rains they were inundated, and
there were many depressions filled with water in
the vehicle tracks.
The planned trip route was to take the road
to Rockstone, cross the pontoon bridge over the
Essequibo River, and to continue by road to
Bartica. From Bartica, the Toyota Land Cruiser
would be carried on the ferry to Parika at the
mouth of the Essequibo River for a final return
by road to Georgetown. At the beginning of this
circuit, the road to Rockstone was muddy but
passable, until just prior to the bridge, where it
promptly deteriorated into a quagmire, complete
with a large Bedford truck stuck up to the axles.
The truck had uprooted several trees while
unsuccessfully trying to winch itself out of the
mud. Although the Toyota managed to traverse
the first 100-meter muddy stretch by straddling
deep ruts, the team was stranded at the top of a
steep, muddy hill leading down to the bridge.
There would be no way to control a downhill slide
to the bridge.
The team decided to wait and see if the road
dried out more by the next day. They negotiated
the way back through the first muddy area (Figure
4). After twice becoming stuck and digging out,
the team reached firm land and began to set up
camp for the night. Fortunately, Mr. & Mrs. Ivan
Rodriguez, an older couple living nearby in the
only house on the road, invited the team to hang
their hammocks inside of their house. It happens
that while the couple was out over the previous
week their guard and dog disappeared, and
everything in the house was stolen leaving plenty
of room for everyone. During the night there was
another downpour, which made the road
conditions worse, and caused the team to abandon
all thoughts of travelling to Bartica by road. The
team decided to leave all of their extra supplies
with the couple that had sheltered them for the
night, and they returned to Georgetown to dry out.
Trip 2: Karanambo - Surama - Karasabai, 16
February to 10 March 1990
Collection numbers 1828 - 2217
This trip was taken in collaboration with two
visiting botanists from the Smithsonian
Institution, Pedro Acevedo, a specialist in lianas
and the family Sapindaceae, and Christian
Feuillet, a specialist in the families Passifloraceae,
Aristolochiaceae, and Gesneriaceae. Their
collections from this trip will be treated in a
separate volume. The team also included the
University of Guyana botanist S. Tiwari. They

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