Plant Community Structure, Fire Disturbance, and Recovery in ...

CENTRE FOR THE STUDY OF BIOLOGICAL DIVERSITY 

UNIVERSITY OF GUYANA 

Contributions to the Study of Biological Diversity 

Volume 3: 1-166 

Plant Community Structure, Fire Disturbance, and Recovery in 

Mangrove Swamps of the Waini Peninsula, Guyana 

by 

Thomas H. Hollowell 

Centre for the Study of Biological Diversity 

University of Guyana, Faculty of Natural Sciences 

Turkeyen Campus 

Georgetown, Guyana 

2009

Thomas H. Hollowell, Smithsonian Institution. Plant Community Structure, Fire 

Disturbance, and Recovery in Mangrove Swamps of the Waini Peninsula, Guyana. 

Contributions to the Study of Biological Diversity, volume 3: 166 pages.- 

Soil fires during the 1997 to 1998 El Niño caused high mortality among mangroves of Waini Peninsula, a potential 

protected area. Impacts and early recovery were investigated, with baseline floristic and ecological analyses. 

Plant species and communities were surveyed. 118 species were documented. Approximately 64.6 km 2 of swampland 

were burned, 26.6 km 2 classified as mangrove, the largest reported mangrove fire. Regional, continental, and global floral 

affinities are explored. Of 240 species in five Waini and two inland communities, 79% occurred in only one; coastal 

vegetation exhibited high beta diversity. The Waini held many species with Neotropical (33%) and Pantropical (27%) 

distributions but few Guiana Shield endemics (1.6%). The presence of the Asian mangrove palm, Nypa fruticans, in the 

study area is examined, including extent, possible sources, potential spread, and reported dispersal to Trinidad. 

Six 0.1-hectare Avicennia swamp vegetation plots were sampled over four years. Unburned swamp basal area was 

21.25 m 2 /ha, increasing 2.5% to 3.5% annually. Burned swamp basal area was at least 20.43 m 2 /ha before fires, with 

estimated biomass greater than unburned swamp. Sapling, seedling, and herbaceous cover in the unburned swamp was low, 

and variable in the burned swamp, with probable hydrology links. Spatial patterns of Avicennia trees were generally 

overdispersed. Waini mangrove basal area and height approached worldwide medians; stem density was much lower. 

Unburned swamps near Waini Point are apparently younger than to the southeast, with irregular seedling recruitment. 

In burned swamp regeneration, establishment distances of seedlings from parents had a mean of 24.2 meters for 

Laguncularia, 4.8 meters for Avicennia, and 8.9 meters for Rhizophora. Mangrove plantings explored restoration potentials. 

Rhizophora plantings were successful in burned swamp, given sufficient elevation. After 10.5 months, Rhizophora racemosa 

in unburned swamp was about half the height of those in burned areas (56 cm vs 129 cm); wet condition plantings outsurvived 

and outgrew drought plantings. No Laguncularia or Avicennia plantings survived, suggesting narrow hydrological 

requirements. 

Geomorphology influences mangrove dispersal, establishment, population structure, and disturbance on the Waini 

Peninsula. Its mangrove swamps are unique in the Neotropics and valuable biologically and culturally. 

Results here suggesting that the Waini flora is essentially separate from the Guiana Shield flora and bears more 

affinity to the Caribbean region are not surprising, but lends support to the idea that biodiversity studies and conservation 

efforts should be pursued with an awareness of that distinction. 

DATE OF PUBLICATION: February 2009 

Cover: Keith David measuring a large Avicennia germinans tree encountered near the Waini River, 

along transect A. Photo Tom Hollowell, Smithsonian Institution. 

Contributions from the Centre for the Study of Biological Diversity, University of Guyana, 

Faculty of Natural Sciences, Turkeyen Campus, Georgretown, Guyana, South America. 

POSTMASTER: Send address changes to the Editor, Centre for the Study of Biological Diversity, 

University of Guyana, Faculty of Natural Sciences, Turkeyen Campus, Georgretown, 

Guyana, South America.

CONTENTS 

INTRODUCTION ............................................................................................................. 

ACKNOWLEDGEMENTS .............................................................................................. 

CHAPTER 1. THE WAINI PENINSULA ENVIRONMENT AND ITS FLORA 

INTRODUCTION ................................................................................................................... 

Study Site .......................................................................................................................... 

Climate ............................................................................................................................. 

Geomorphology ................................................................................................................ 

Soils .................................................................................................................................. 

The Waini Peninsula Fires of 1997-1998 .......................................................................... 

MATERIALS AND METHODS ............................................................................................. 

Plant Collection Field Work .............................................................................................. 

Plant Identification, Classification and Distribution ....................................................... 

Plant Species List .............................................................................................................. 

Plant Communities ........................................................................................................... 

Burned Area Delineation .................................................................................................. 

RESULTS ................................................................................................................................ 

Plants of the Waini Peninsula ............................................................................................ 

Plant Communities on the Waini Peninsula ...................................................................... 

Beaches ............................................................................................................................. 

Coastal Mangrove Swamps .............................................................................................. 

Mixed Freshwater Swamps .............................................................................................. 

The Riverine Mangrove Swamps ..................................................................................... 

Burned Areas .................................................................................................................... 

Plant Collections of Interest .............................................................................................. 

DISCUSSION .......................................................................................................................... 

CHAPTER 2. PLANTS OF THE WAINI PENINSULA IN REGIONAL AND 

GLOBAL CONTEXT 

INTRODUCTION ................................................................................................................... 

Study Sites and Background ............................................................................................. 

Quackal and Manicole Plant Communities ...................................................................... 

The Northwest District ..................................................................................................... 

Delta Amacuro .................................................................................................................. 

Regional and Global Affinities of the Waini Peninsula Flora ........................................... 

METHODS .............................................................................................................................. 

Comparison of the Waini Flora with Quackal and Manicole Plant Communities ............ 

Northwest District ............................................................................................................ 

Delta Amacuro .................................................................................................................. 

Regional and Global Affinities ......................................................................................... 

RESULTS ................................................................................................................................ 

Quackal and Manicole Communities ............................................................................... 


Delta Amacuro .................................................................................................................. 


DISCUSSION .......................................................................................................................... 

Quackal and Manicole Communities ............................................................................... 


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Delta Amacuro .................................................................................................................. 


CONCLUSION ....................................................................................................................... 

CHAPTER 3. THE MANGROVE PALM NYPA FRUTICANS WURMB.: A 

WIDESPREAD EXOTIC SPECIES IN NORTHWESTERN GUYANA 

INTRODUCTION ................................................................................................................... 

NYPA HISTORY AND RANGE .............................................................................................. 

NYPA FRUTICANS OUTSIDE OF ITS MODERN NATIVE RANGE .................................. 

DISPERSAL OF NYPA TO TRINIDAD ................................................................................. 

POSSIBLE SOURCES OF PRESENT NEOTROPICAL NYPA POPULATIONS ................ 

ECOLOGICAL CONSIDERATIONS .................................................................................... 

CONTROL OF NYPA .............................................................................................................. 

NEEDS FOR ADDITIONAL STUDY .................................................................................. 

CONCLUSIONS .................................................................................................................... 

CHAPTER 4. STRUCTURE OF BURNED AND UNBURNED AVICENNIA L. 

FOREST, WAINI PENINSULA,GUYANA 

INTRODUCTION ................................................................................................................... 

METHODS .............................................................................................................................. 

Study Site .......................................................................................................................... 

Plot Establishment ............................................................................................................ 

Measurements .................................................................................................................. 

Alness Village Plot ........................................................................................................... 

Dispersion Patterns of Trees ............................................................................................. 

Biomass ............................................................................................................................ 

Comparison with Other Plot Data ..................................................................................... 

RESULTS ................................................................................................................................ 

Tree Mapping and Inventory ............................................................................................ 

Dbh and Tree Height ......................................................................................................... 

Basal Area ......................................................................................................................... 

Dispersion ......................................................................................................................... 

Density and Population Turnover in the Avicennia Forest ................................................ 

Biomass ............................................................................................................................ 

Small Woody Plants .......................................................................................................... 

Herbaceous Plants ............................................................................................................ 

Well Levels ....................................................................................................................... 

DISCUSSION .......................................................................................................................... 

Guyana Plots ..................................................................................................................... 

Dispersion ......................................................................................................................... 

Biomass ........................................................................................................................... 

Comparisons with Worldwide Plot Basal Area Data ........................................................ 

Population Turnover in the Avicennia Forest .................................................................... 

Herbs and Seedlings ......................................................................................................... 

Role of Water Levels ......................................................................................................... 

CONCLUSION ....................................................................................................................... 

CHAPTER 5. DISPERSAL AND ESTABLISHMENT OF MANGROVE 

PROPAGULES FOLLOWING FIRES 

INTRODUCTION .................................................................................................................. 

Seed Dispersal .................................................................................................................. 

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Dispersal in Mangrove Swamps ....................................................................................... 

Dispersal in Disturbed Mangrove Swamps ...................................................................... 

METHODS .............................................................................................................................. 

Study Site .......................................................................................................................... 

Propagule Dispersal and Establishment ........................................................................... 

Propagule Plantings .......................................................................................................... 

RESULTS ................................................................................................................................ 


Propagule Plantings .......................................................................................................... 

DISCUSSION .......................................................................................................................... 


Propagule Plantings ......................................................................................................... 

REFERENCES .......................................................................................................................... 

APPENDIX 1: Mangrove Ecosystems and Fire in the Tropics .................................................. 112 

APPENDIX 2: Plant Species Listed for the Waini Peninsula, with Synonymy ........................ 129 

APPENDIX 3: Preliminary Species List of the Vascular Plants of the 

Northwest District of Guyana ..................................................................................... 

APPENDIX 4: Species Disparities Between the Northwest District, Guyana 

and Delta Amacuro, Venezuela .................................................................................... 155 

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Contributions to the Study of Biological Diversity Vol. 3 

Plant Community Structure, Fire Disturbance, and Recovery in 

Mangrove Swamps of the Waini Peninsula, Guyana 

The Waini Peninsula is the site of the 

majority of Guyana’s remaining mangrove 

swamps. In the ongoing development of 

Guyana’s protected areas system (EPA Guyana 

2002), there has been a focus on the “Shell 

Beach” portion of the Waini Peninsula as a 

candidate site, primarily because of its extensive 

mangrove swamps and marine turtle nesting 

beaches (Humm 2001). The area is important 

beyond its uniqueness within Guyana, as the 

high-sediment coastline of the Guianas differs 

significantly from Caribbean mangrove settings 

that are most often studied in the Neotropics. 

There was a basic need for description of the 

plant communities in the prospective protected 

area, to provide baseline floristic and ecological 

information upon which future scientific 

research and monitoring can build. In the 

broader perspective, it is important within 

mangrove ecology to expand documentation of 

the variety of biotic elements, structures, and 

disturbances that occur in mangrove ecosystems, 

to provide for suitable understanding, 

protection, and management (Ellison 2002; Finn 

et al. 1998; Jiménez et al. 1985). The disturbance 

of the study site just prior to field work offered 

opportunities to address basic questions about 

the nature of mangrove dispersal and recovery. 

As a basis for study, plant specimens were 

collected and identified, and a vouchered species 

list was assembled for the Waini Peninsula. 

Some of those collections expanded documented 

species ranges. The plant collection results were 

compared to other available botanical data for 

the region, at various scales, to place the Waini 

by 

Thomas H. Hollowell 1 

INTRODUCTION 

flora in environmental and biogeographic 

context. 

In addition, the plant communities of the 

Waini Peninsula were classified, described, and 

mapped. As part of that task, the unusual 

phenomenon of large-scale fires in the mangrove 

swamps was documented, including estimates 

of the extent of impacted plant communities. 

Documentation of the fires formed a 

contribution to knowledge of disturbance types 

that have been known to affect mangrove 

systems. The fires occurred during the extreme 

El Niño event of 1997 to 1998, and so may have 

been linked to global climate change processes 

(Laurance 1998; Snedaker 1995). 

Among the plant species present, the Asian 

mangrove species, Nypa fruticans Wurmb., was 

of particular interest and so was documented and 

reviewed in detail. Nypa Wurmb. is an exotic 

species that may spread in important riverine 

mangrove swamps of the Waini Peninsula 

region, making it a concern both in Guyana and 

neighboring countries. 

Ecological vegetation plots were 

established and resampled to provide new 

information on the structure of the Waini 

Peninsula’s coastal mangrove forests and on the 

initial course of regeneration in the fire-affected 

mangrove forests. Those structural data allowed 

comparisons with information on other 

mangrove systems of the world and will aid 

additional population structure and succession 

trend studies. 

Mangrove zonation patterns have been 

attributed to both physical and chemical factors 

1 Master of Science, George Mason University, 1992 

This volume was originally a portion of a dissertation submitted in partial fulfillment of the requirements of the degree of 

Doctor of Philosophy at George Mason University. Director: Donald P. Kelso, Associate Professor Environmental Science 

and Policy 

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(McKee 1993, 1995b) and to abilities of various 

mangrove species to disperse and establish 

(Clarke et al. 2001b; McGuinness 1997a; 

Rabinowitz 1978a). To investigate possible 

factors in the fire-disturbed environment, trial 

plantings of mangrove species were undertaken 

in both disturbed and undisturbed habitats. The 

plantings tested the abilities of species to 

establish in local conditions, independent of 

their dispersal ability. The results might have 

implications for potential restoration 

approaches and provide information on the roles 

of dispersal limitation versus environmental 

conditions in mangrove regeneration. 

The fire-affected swamps also presented a 

unique setting to test establishment patterns of 

mangrove propagules in an environment where 

coastal tides and currents were absent. A method 

was devised for sampling the establishment 

patterns of mangrove propagules from the 

scattered surviving parent trees in the burned 

swamps. The establishment data provide 

insights into the possible path of vegetation 

recovery. 

These studies were a first attempt to 

assemble critical information that may serve to 

integrate the swamps of the Waini Peninsula into 

the broader understanding of mangrove ecology 

in the Neotropics, and to inform the continued 

establishment of Guyana’s Protected Areas 

System. It is hoped that future studies will be 

encouraged to move beyond the work presented 

here. Because the chapters were written to be 

published separately, some repetition was 

inevitable, however it is hoped that it has been 

kept to a minimum. 

ACKNOWLEDGMENTS 

For assistance with logistics and planning, 

great gratitude goes to Vicki Funk, Carol 

Kelloff, and Marilyn Hansel of the 

Smithsonian’s Biological Diversity of the 

Guiana Shield Program, as well as to Carol Ailes 

of the Smithsonian’s National Museum of 

Natural History travel office. For instructive and 

enjoyable field experiences in the highlands of 

Guyana and for help at the US National 

Herbarium, thanks go to H. David Clarke of the 

University of North Carolina at Asheville. In 

the field essential assistance was given by 

Audley James, Violet James, Romeo DeFreitas, 

Annette Arjoon, Peter Pritchard, Neville 

Waldron, Doekie Arjoon, Chris Chin, Matt 

Sewell, Keith David, Karen Redden, Wiltshire 

Hinds, Narad Persaud, Lloyd Savory, Arnold 

Benjamin, Emily Duvall and Renee LeBlanc. 

In Georgetown, Guyana help was given by Tsitsi 

McPherson, Margaret and Malcolm Chan-A- 

Sue, Dyantie Naraine, and Phillip Da Silva. At 

George Mason University thanks go to 

professors Ted Bradley, Barry Haack, Don 

Kelso, Sheryl Luzzadder-Beach, Larry 

Rockwood, and Lee Talbot, for their teaching, 

advice, encouragement, and tolerance. 

For assistance with identification of plant 

specimens, thanks go to Pedro Acevedo, Frank 

Almeda, Rupert Barneby, Germán Carnevali, 

Eric Christenson, Tom Croat, Piero Delprete, 

Larry Dorr, Bob Faden, Christian Feuillet, Vicki 

Funk, Eric Gouda, Susan Grose, Terry Henkel, 

Walter Holmes, Emmet Judziewicz, Carol 

Kelloff, Job Kuijt, G.P. Lewis, Paul Maas, Greg 

McKee, Mike Nee, Dan Nicolson, Jim Norris, 

Darin Penneys, John Pipoly, Susan Renner, 

Harold Robinson, Matt Sewell, Alan R. Smith, 

Mark Strong, and Charlotte Taylor. Many of 

those identifications were facilitated by the 

specimen sorting and distribution staff of the 

Smithsonian’s Biological Diversity of the 

Guiana Shield Program, as was the distribution 

of identified specimens. Credit also goes to the 

entire staff of the Smithsonian’s US National 

Herbarium for building and maintaining such a 

valuable resource of plant specimens. 

I thank all of my family; especially my wife 

Carol Regier; our daughters Irena and Andrea, 

our son Ashley, and their grandmother Nair 

Regier, for their perseverance through the many 

months of field work and years of busy 

weekends and evenings; also gratitude goes to 

my parents Cici and Robert Hollowell for 

providing a basis of respect for the environment 

and a good early education, and to my sisters 

Julie Hollowell and Laura Hollowell for their 

encouragement. 

This is number 143 in the Smithsonian 

Biological Diversity of the Guiana Shield 

publication series.


CHAPTER 1. 

THE WAINI PENINSULA ENVIRONMENT AND ITS FLORA 

INTRODUCTION 

Any protected natural area is well served 

by the assembly of comprehensive baseline 

information and by ongoing environmental 

description and monitoring. The broad goal of 

this chapter is to provide a detailed, initial 

documentation of the plant species and plant 

communities of the Waini Peninsula, to serve 

as a foundation for future studies and 

management. The Waini Peninsula is a proposed 

reserve in northern Guyana that has been known 

for its marine turtle nesting beaches and 

conservation activities. It is also the location of 

long stretches of coastal mangrove swamps, 

freshwater swamps, and extensive intertidal 

mudflats that are habitat for many types of 

wildlife. Included are a description of the study 

site, a summary of the botanical collections 

made to document the flora, and descriptions 

of basic vegetation zones. The quantity of new 

distributional information revealed by plant 

collections activities is of scientific interest, as 

that allows insight into the state of basic 

knowledge about an area, which guides future 

data gathering efforts for research and 

conservation activities. A more accurate 

delineation of Guyana’s mangrove communities, 

which are of high conservation concern, is 

desirable considering the wide variations in 

existing estimates for the country (Saenger et 

al. 1983; Snedaker 1986; Spalding et al. 1997). 

As part of the plant community description, an 

estimate was also made of areas affected by the 

1998 soil fires on the Waini Peninsula. Those 

fires were unusual for their impact to large areas 

of mangrove swamp, which is an ecosystem type 

normally considered to be unaffected by fire. 

Appendix 1 provides additional review of 

mangrove ecology in Guyana, including 

relationships to fire in the wet tropics. Appendix 

2 fully lists the plant species that were 

documented for the Waini Peninsula. 

STUDY SITE 

Guyana is located in northeastern South 

America on the Atlantic Ocean, just north of the 

Equator (Figure 1.1). The Waini Peninsula is a 

relatively undisturbed, coastal region of northern 

Guyana (Figure 1.2). It is the site of most of 

Guyana’s remaining intact coastal ecosystems, 

including mangrove swamps, freshwater 

swamps, flooded savannas, and their associated 

fauna. The peninsula’s “wild coast” stretches for 

more than 75 kilometers along the Atlantic 

Ocean, from near the Venezuelan border (59º 

50' W, 8º 25' N), southeast almost to the mouth 

of the Pomeroon River (59º 05' W, 8º N). For 

this paper, the study area includes approximately 

the northern third of the Waini Peninsula, from 

Figure 1.1. Location map of Guyana, with major 

rivers. 

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Figure 1.2. Site map of the Waini Peninsula, Guyana, with localities referred to in the text. The area framed 

comprises much of the Northwest District. 

its northwestern point to 59º 30' W, covering a 

land area of about 190 km 2 . That portion of the 

Waini Peninsula is also commonly referred to 

as “Shell Beach.” The study area is part of a 

candidate site for Guyana’s newly established 

National Protected Areas System (EPA Guyana 

2004; Humm 2001), which at the time of this 

fieldwork consisted only of Kaieteur National 

Park (Kelloff 2003). 

The coastal plain of Guyana is composed 

of fine sediments that primarily settled from 

waters of the current flowing from the Amazon 

River’s mouth, which is over 1,000 kilometers 

to the southeast. Prior to colonization by 

European nations, most of the Guyana coast was 

probably lined by broad mangrove forests that 

graded into freshwater swamps and flooded 

savannas, and then into lowland tropical forests 

(Richardson 1987). In the more densely 

populated southeastern part of Guyana’s coastal 

plain, around the capital Georgetown, those 

swamps and savannas were drained and 

converted to agriculture and settlements early 

in the country’s colonial history. However, in 

the Northwest District of Guyana (also known 

as the Barima-Waini Division; Figure 2.1) there 

have been limitations on access to coastal lands: 

by land due to the absence of roads and presence 

of numerous swamps and rivers, and by sea 

because of the broad tidal mudflats extending 

up to several kilometers out from shore. As a 

result, long stretches of the coast have remained 

sparsely settled and largely unmodified. 

Climate 

Based on the life zone classification of 

Holdridge et al (1971), that combines influences 

of temperature, elevation, and rainfall, the Waini


Peninsula is capable of forming a Tropical Moist 

Forest. Because of its location near the equator, 

Guyana’s climate is dominated by the equatorial 

trough and the seasonal movements of the 

intertropical convergence (ITC) (Snow 1976). 

The mean annual temperature for the coastal 

zone of Guyana is approximately 26.5ºC, with 

a 1ºC to 1.5ºC range between the mean 

temperatures of the warmest and coolest months, 

which coincide with the dry and wet seasons 

respectively. The trade winds consistently blow 

from the east-northeast or the east, with monthly 

mean speeds varying from 3 to 4 m/second. 

Precipitation patterns are characterized by two 

dry seasons and two wet seasons per year. The 

major wet season runs from April to August, 

with a minor wet season from November to 

February. Mean annual rainfall in the coastal 

area of the Northwest District has been reported 

from approximately 2,500 mm per year (Snow 

1976) to 3,400 mm per year (K. Richardson, 

pers. comm.). The wettest month is sometimes 

reported as June, with an mean rainfall of 400 

mm (Snow 1976), or January with a mean 

rainfall of 500 mm (K. Richardson, pers. 

comm.). During a typical year in the stronger 

dry season of February-April, only 5 to 10 days 

per month receive rainfall (Snow 1976). 

Snow (1976) noted that a zone of minimum 

rainfall probably occurs along a narrow band of 

the coast of northern South America, with a peak 

in precipitation reached less than 50 kilometers 

inland, caused by effects of increased surface 

roughness of forests upon air leaving the ocean. 

This has been confirmed by many observations 

(Hollowell, unpublished) of rain developing 

several kilometers inland from the Waini 

Peninsula while extreme coastal areas remained 

dry. The typically low number of rain days in 

the dry season, combined with reduced rainfall 

immediately along the coast and a drought 

caused by a strong El Niño event, led to 

extremely dry conditions in the coastal forests 

of the Waini Peninsula in 1997-1998. 

Geomorphology 

The coastal plain sediments of the Guianas 

were deposited in recent times on the margins 

of the Guiana Shield, which is a natural 

geological unit characterized by ancient bedrock 

with sandstone and igneous highlands (Gibbs 

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& Barron 1993; Huber 1995a). The Guiana 

Shield is geologically distinct from the more 

recently uplifted Andean highlands. It is 

bounded for the most part by the Atlantic Ocean 

to the east, the Orinoco River to the north and 

west, and the Negro and Amazon Rivers to the 

South. The Shield is widely considered to be a 

region of high biotic endemicity (Huber 1995c), 

possibly due to having served as a refugium for 

tropical forest species during prolonged periods 

of dry climate in the Pleistocene epoch (1.8 mya 

to 11,000 years bp), or due to the ancient, varied, 

and isolated landscapes of the region (Colinvaux 

1987). 

From the Orinoco Delta south to the 

Amazon River, the coast of South America is 

geologically referred to as a “passive margin,” 

with deep sediments deposited over the 

tectonically stable Guiana Shield (Di Croce et 

al. 1999). That stability is reflected in estimates 

that relative sea level in the region has been 

relatively stationary since post-glacial rises 

ended perhaps 6,000 B.P., although sea levels 

possibly surpassed that of the present by as much 

as five meters from 6,000 to 3,000 B.P. (Pirazzoli 

1991). By some accounts, the section of the 

Guyana coast including the Waini Peninsula has 

been subject to some uplift in recent geological 

times (Gibbs & Barron 1993), while others 

suggest that subsidence may have been a factor 

along the northwestern coast of Guyana 

(Brinkman & Pons 1968). Williams (1989) 

employed archaeological evidence to infer that 

subsidence in the Guyana’s northwest ended 

around 3,400 years B.P. The presence of marine 

sediments on higher elevations in the central 

parts of the Waini Peninsula suggests that in 

geologically recent times, relative sea levels 

may have once been several meters higher than 

present or that moderate uplift has occurred. The 

area is close enough to a geologically active area 

beginning just south of Trinidad (Di Croce et 

al. 1999) that associated minor uplift or 

subsidence around the study site could be 

plausible. 

The Shield’s basement granites are 

estimated to be as far as 500 meters below the 

surface along Guyana’s northwest coast (Di 

Croce et al. 1999). The coast of the Guianas is, 

in essence, the major portion of the coastal delta 

of the Amazon River. The Essequibo, Waini, and

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Orinoco Rivers probably contribute a minor 

amount of sediment in comparison to the 

Amazon River. The coast is geomorphically 

similar to the coast of eastern Sumatra, based 

on an analysis of several coastal characteristics 

(Jelgersma et al. 1993). Both are deltaic 

environments with a wide coastal shelf, a 

prograding shore of very fine sediments, little 

subsidence or uplift, strong longshore transport, 

low wave energy, small tidal range, and no 

influence of tropical cyclones. A geomorphic 

classification commonly applied to this type of 

mangrove environment is River Dominated 

Alluvial Plain (Thom 1984; Woodroffe 1992). 

The width of the lower coastal plain of 

Guyana varies about from about 75 kilometers 

at the Corentyne River on the Surinam border 

to 25 kilometers near the Venezuelan border 

(Vann 1969). The coastline is characterized by 

cycles of accretion and erosion of mudflat clay 

and silt sediments, which are known locally as 

“sling mud”(Richardson 1987; Wells & 

Coleman 1981). The Waini section of the coast 

has apparently been prograding, in a manner 

similar to the coastline of the Orinoco Delta to 

the northwest (Bureau of Economic Geology 

2002). Along parts of the Guyana coast coastal 

currents have led to erosion loss of diked land, 

including land near Georgetown that was once 

populated or cultivated as recently as 1972; 

signs of these changes are visible in radar 

satellite imagery (Singhroy 1996). Sizable 

sediment deposits were colonized by 

mangroves, primarily Avicennia, between the 

Essequibo and Pomeroon Rivers in the 1990’s 

(pers. obs). 

Slow migration of the mudbanks along the 

coast leads to a smaller-scale cycle of shore 

erosion and redeposition that is unlike typical 

coastal dynamics that would produce relatively 

straight coastlines (Lakhan & Pepper 1997). As 

part of those cycles, coarser sand and shell 

fragments are released from the sediments and 

deposited to form beach ridges that may 

accumulate up to 200 meters in width (Prost 

1989; Vann 1969). The beach ridges often isolate 

sections of mangrove forest from the sea. Along 

other stretches of the Waini coast waves wash 

directly into the mangrove forest. During the 

time of this field work (1997-2001), the widest 

beach ridge on the Waini Peninsula was Almond 

Beach, the location of a marine turtle monitoring 

camp (8° 23' 58'’ N, 59° 45' 16" W) and base 

camp for these studies, where the ridge 

measured over 160 meters wide. That shell 

deposit was perhaps two meters deep and 

contained a lens of nearly fresh water 

(approximately 1 psu salinity, Hollowell, 

unpublished data). To illustrate the variability 

of the ridges, profiles were surveyed for one 

wide section and one narrow section of beach 

ridge, illustrated in Figures 1.3 and 1.4. The 

wider beach ridges roughly coincided with areas 

of mangrove swamp that dried adequately to 

Figure 1.3. Beach profile between Almond Beach Camp and Waini Point, an area of narrow beach ridge near 

plots 1, 2 and 6. Beyond 25 m was Avicennia swamp, which borders the beach ridge with a low swale. It was 

not unusual for waves to spill sea water into the swamp during extreme tides. Profiles were started near the 

boundary between mudflats and shell substrate.


Figure 1.4. Beach profile at Almond Beach camp, near plots 2, 4 and 5, an area with a wide beach ridge that, 

at the time of survey in April 2001, had several small structures and gardens. The swale at 70 m was deeper in 

some places than shown, and formed a slightly brackish pond persisting through most dry seasons. Beyond 

170 m was Avicennia swamp that burned in early 1998. Waves from extreme storms have flooded interior 

sections of the beach ridge, but rarely reached the mangrove swamp. 

burn in the 1998 soil fires (Figure1.5). 

Figure 1.5. Location of beach ridges on the Waini 

Peninsula, shown in relation to burned areas and 

settlements or camps. 

Inland beach ridges occur within the coastal 

mangrove swamps; these run approximately 

parallel to the present shoreline. Such ridges are 

evidence of long-term coastal accretion, as they 

represent former shorelines; they are often 

termed “cheniers” in the literature (Prost 1989, 

1997; Wells & Coleman 1981), and the coast of 

the Guianas is one of the few true chenier coasts 

of the world. The region of South American 

cheniers stretches from the mouth of the 

Amazon River north through the Orinoco Delta 

(Bacon 1990). The other major chenier coast in 

the New World is the Louisiana chenier plain 

of the Mississippi River delta (Huh et al. 2001; 

13 

Wells & Coleman 1981). On the Waini 

Peninsula, the largest cheniers rise about one 

meter above the surrounding swamp surface and 

support non-mangrove vegetation dominated by 

trees including Terminalia catappa L. and 

Spondias mombin L. Differences in hydrology, 

vegetation cover, or surface reflectance of shells 

often allow cheniers to be distinguishable in 

satellite imagery. The cheniers on the Waini 

Peninsula sometimes coincide with transitions 

in plant community composition. There are 

accounts from older residents of the Almond 

Beach area describing the largest chenier, now 

over 700 meters from the present shore, as 

having been the location of the shoreline within 

their memories; these accounts have not been 

confirmed, but could be subject to comparison 

with older maps and aerial photos. Vann (1969) 

also reports observations of shell ridges stranded 

up to 300 yards from shore that had been beach 

by the ocean 30 years earlier, corroborating the 

local accounts. 

Like most coastal plant communities, those 

of the Waini Peninsula are influenced 

considerably by local geomorphic processes 

over time (Augustinus 1995; Thom 1967, 1975; 

Woodroffe 1992). Although direct building of 

land has been attributed to mangrove vegetation 

in early literature, that has been largely 

discounted (Woodroffe 1992). Still, mangroves 

probably play some role in securing land built 

up through geomorphic processes, both by 

slowing wave erosion and, as suggested by Wells

14 


and Coleman (1981), through new root biomass 

of Avicennia L. trees in newly colonized 

mudflats increasing substrate elevation enough 

to allow survival of a stand. Zones of recent 

coastal accretion are easily identified from the 

sea by bands of Avicennia saplings and small 

trees (Figure 1.6), while eroding shore segments 

are usually marked by fallen Avicennia trees 

(Figure 1.7). 

Figure 1.6. Avicennia saplings recently established 

on new sediments in front of mature trees, on the 

Waini Peninsula coast just southeast of Kamwatta 

Beach. 

Developing countries with populated lowlying 

land, such as Guyana, are particularly 

vulnerable to economic and environmental 

impacts from sea level increases (Gabche et al. 

2000). A sea level rise in the 0.09 to 0.88 meter 

range has been predicted by the year 2100 

(IPCC 2001), and the Government of Guyana 

estimates that the entire Waini Peninsula would 

be severely affected by a one meter rise in sea 

level (EPA Guyana 2000). Sea level rise could 

result in substantial changes in the extent of 

mangrove ecosystems world-wide (Pernetta 

1993). To survive under such circumstances, 

mangrove communities must either migrate 

inland or survive on accreting substrates. In 

general, high sediment input mangrove systems 

such on the coast of the Guianas are more likely 

to withstand moderately rising sea levels (Field 

1995), particularly in contrast to Caribbean 

mangrove systems that are often situated on 

slowly accreting peat substrates (Ellison 1993; 

Ellison & Stoddart 1991; Parkinson et al. 1994). 

Although the coastal plain of the Guianas could 

serve as a refuge for mangrove communities, 

in populated areas any landward migration of 

mangrove forests would be impossible. 

Figure 1.7. Shore erosion along the Waini Peninsula 

coast between Almond Beach and Kamwatta Beach, 

showing fallen Avicennia trees. During higher tides 

waves wash over a very small beach ridge and into 

the forest. 

Mangrove systems along the Guyana coast 

might be subject to accelerated isolation from 

tidal influence if sedimentation increases as a 

result of changes in upland land uses that cause 

erosion, as has been reported on parts of coastal 

Thailand (Panapitukkul et al. 1998). 

Soils 

The soils of the Waini Peninsula are the 

product of slow accretion of very fine sediments 

in very deep layers over recent geological time 

(Brinkman & Pons 1968; Gibbs & Barron 1993). 

Soils of the Waini Peninsula have been classified 

as Demerara Series, Coronie deposits, of the 

Comowine phase (Brinkman & Pons 1968), 

which are marine clays with high base 

saturation, deposited less than 1,000 years ago. 

The clay soils of the Waini Peninsula swamps 

are comparable to those of the Wia Wia reserve 

of coastal Surinam, which were described by 

Pons and Pons (1975) as typic Endoaqerts (at 

the time, termed Typic Haplaquepts). The Waini 

soils could alternately be classified as Vertisols 

since they marginally meet the requirement of 

forming cracks on a regular basis (Soil Survey 

Staff 1998), as observed during droughts of 

extreme dry seasons. These young, extremely 

fine clay soils are highly reduced by anaerobic 

conditions during long periods of inundation. 

Such reduced soils are indicated by a gleyed 

color (grey, low chroma color), most often 

measured on the Waini Peninsula at 7.5 YR 4/1 

using the standard Munsell soil color system 

(Munsell Color 1992). The shallower portions


of those soils included high chroma mottles, 

around Munsell color 7.5 YR 4/4, that usually 

result from seasonal fluctuation of water tables. 

There was little horizon development other than 

a surface layer of organic material that ranges 

from 5-15 cm in thickness. Most of the 

mangrove swamp soils of the Waini Peninsula 

had such organic horizons, again similar to those 

reported for Surinam’s coastal mangrove 

swamps (Pons & Pons 1975). In the Waini 

Peninsula’s more inland Mixed Freshwater 

swamps, the organic horizon was observed to 

be as much as 15-20 cm thick, but was minimal 

in areas where the upper soil layers had been 

burned in 1998. Such organic layers are quite 

shallow compared to “pegasse” peat that is 

several meters deep in coastal plain swamps 

located farther inland than the Waini Peninsula 

sites, most notably the Mauritia flexuosa L.f. 

palm savannas in the Santa Rosa vicinity of the 

Northwest District (van Andel 2000a). 

The beach ridges and cheniers are 

composed of fine shell fragments with minimal 

coarse sand. These very young soils can be 

classified as typic Psammaquents or Psamments 

(Soil Survey Staff 1998), with fine shell 

materials probably derived from both riverine 

and oceanic sources (Brinkman & Pons 1968). 

The offshore mudflats could possibly be 

classified as Halic Typic Hydraquents, although 

they are likely unsuitable for colonization (Soil 

Survey Staff 1998), in which case they should 

be considered non-soil, being too frequently 

inundated and/or unstable to usually support 

plant growth. 

Since the Waini Peninsula was deposited 

slowly by accretion of marine sediments, the 

interior area has probably been above sea level 

longer than land near the ocean, and land nearer 

to Waini Point may be more recent than the 

southeastern portion of the Peninsula. Older 

deposits have been subject to more leaching of 

salts by rains, greater accumulation of organic 

matter from vegetation, and possible changes 

in elevation. Generally, soil salinity is higher 

near the ocean, limiting the number of plant 

species that may become established. The soils 

of the interior Mixed Freshwater swamps are 

similar to those of the Coastal Mangrove 

swamps, being very fine, gray clays of marine 

origin. The porewater salinity of these soils was 

15 

generally low enough, less than 10 psu 

(Hollowell, unpublished data), to allow nonmangrove 

plant communities. 

The Venezuelan state of Delta Amacuro 

borders Guyana’s Northwest District; it is 

largely situated on the coastal plain, with alluvial 

and marine soils similar to those of the Waini 

Peninsula and the coastal plain of Guyana. The 

southern third of Delta Amacuro is geologically 

part of the Guiana Shield, with soils derived 

mostly from granitic bedrock (Huber 1995a). 

This includes part of the low mountains of the 

Serranía Imataca along its border with the state 

of Bolívar. Those mountains are primarily 

composed of granitic bedrock with greenstones, 

similar to the mountains of Guyana’s Northwest 

District but distinct from the sandstones and 

intrusive rocks of the Pakaraima mountains to 

the south. 

The Waini Peninsula Fires of 1997-1998 

The El Niño event of 1997-1998 was one 

of the more substantial that have been recorded 

(Trenberth 1999). Severe flooding occurred in 

northwestern South America, while there was a 

substantial decrease in rainfall intensity and 

frequency in northeastern South America. 

Guyana was one of the hardest hit countries, 

along with neighboring portions of Venezuela 

and Brazil. Below normal rains began in the 

Guyana region in June 1997. By March 1998, 

parts of Guyana had rainfall deficits of more 

than 1,000 mm and river flows approximately 

20 percent of normal (WMO 1999). Minor fires 

resulting from that drought were reported for 

small, white-sand areas in Guyana (Hammond 

& Steege 1998). Widespread, uncontrolled forest 

fires were also reported during that time from 

the state of Roraima, in northern Brazil, just west 

of southern Guyana, affecting 12,000 km 2 to 

33,000 km 2 of forest (Barbosa & Fearnside 

1999; Barbosa et al. 2003; Cochrane & Schulze 

1998). Also during that period, droughts led to 

fires of even greater extent in Indonesia, 

destroying over 50,000 km 2 of forest and 

causing widespread respiratory problems 

downwind (Kinnaird & O’Brien 1998; Legg & 

Laumonier 1999; Trenberth 1999). 

Although unusual for mangrove 

ecosystems, the 1998 soil fires on the Waini 

Peninsula have been poorly documented. There

16 


were apparently no reports in the Guyanese 

media, and limited knowledge of the fires 

reached any governmental offices. One brief 

report of possible fires in the Waini Peninsula 

area occurred in the literature, based on satellite 

detection during March 1998 (Grégoire et al. 

1998). That report was based on analysis of 

AVHRR 1-kilometer resolution satellite 

imagery, where a single point of possible fire 

was detected near the northwestern end of the 

Waini Peninsula. The same report indicated fires 

in mangroves and coastal vegetation in Surinam, 

and predicted serious consequences including 

increased coastal erosion and impacts on 

fisheries. 

According to residents, the Waini peninsula 

fires were initiated by multiple, unintended 

escapes of agricultural fires at small settlements 

on coastal beach ridges. Spontaneous 

combustion of peat soils might be a slight 

possibility during droughts (Lindeman 1953; 

Viosca 1931), although the peat of the Almond 

Beach swamps was probably not thick enough 

to present that risk. Residents of Almond Beach 

have recounted carrying water in attempts to 

extinguish soil fires early in their course. The 

fires burned for several months, working 

through the swamp forest from late 1997 until 

the return of rains in April 1998. During that 

period, smoke often made navigation on the 

Waini River difficult. 

Two extensive areas of the Waini Peninsula 

swamps were burned. The Almond Beach burn 

ran adjacent to the Atlantic beach ridge from 

the Peninsula’s northernmost settlement at 

Almond Beach, the site of a marine turtle 

monitoring camp, to the last coastal settlement 

in the vicinity, about 7 kilometers to the 

southeast. For several years following the fires, 

crowns of dead standing Avicennia trees at the 

Almond Beach burn were visible from the 

ocean. The Kamwatta burn occurred about 20 

kilometers to the southeast of Almond Beach. 

Those fires came within 250 meters of the ocean 

near Kamwatta Beach (Figure 1.2), where a 

temporary marine turtle monitoring camp was 

located. Possible outliers of the Kamwatta burn 

were visible along short stretches of the Waini 

River near homesteads. 

Tree mortality was very high as a result of 

both of these burns. The mortality resulted from 

at least two factors: the high fire sensitivity of 

mangrove plants, and the manner in which the 

fires passed twice through many areas, with the 

first pass burning upper organic soil horizons 

(5-15 cm thick near Almond Beach), and second 

pass additionally fueled by leaves fallen from 

shocked and newly killed trees (local 

inhabitants, pers. comm). Cochrane (1998) 

refers to similar double fires in upland tropical 

forests following the accumulation of dead 

leaves on the forest floor. 

MATERIALS AND METHODS 

Plant Collection Field Work 

For characterization of the flora of the Waini 

Peninsula, plant specimens were collected, 

identified and processed for deposit in herbaria 

to provide vouchers for a guide to the plants of 

the Waini Peninsula, vouchers for the 

composition of plant communities, and vouchers 

for ecological plots in the local mangrove 

swamp. At least one collection was made of each 

species encountered, including any plants that 

were potentially distinct from other species 

previously collected. 

Plant collections were made on field trips 

from 1997 through 2001. Fertile plant material 

was selected in the field; at least three duplicate 

collections were made whenever possible to 

provide sheets for the herbaria at the University 

of Guyana’s (GRB) Centre for the Study of 

Biological Diversity (CSBD) and at the 

Smithsonian Institution (US). Sterile collections 

were made in only a few cases when no fertile 

material was found. Associated plant 

description, habitat, and location data were 

recorded, with geographic coordinates recorded 

using a hand-held Geographic Positioning 

System (GPS) with an accuracy of 15 meters. 

Specimens were field pressed and air dried in 

the Almond Beach camp on the Waini Peninsula. 

Air drying of botanical collections in the field 

can be impractical in the wet interior of Guyana, 

where the common practice has been interim 

preservation in an ethanol solution. However, 

field drying is often possible in the relatively 

dry, windy environment of the Almond Beach 

camp. Because of that, the collections of 

Hollowell number series from 200 to 754 were 

not treated with any preservatives. All specimens


were transported to the CSBD for any final 

drying, sorting, and processing for export 

permits. 

Where possible, photographs were taken of 

species collected in the area. Copies of selected 

slides and image files are held by the 

Smithsonian Institution and by the CSBD. It is 

planned to use those photographs as part of an 

illustrated listing of plant species in a separate 

volume. 

Plant Identification, Classification and 

Distribution 

Plants were identified using available 

botanical keys, most notably treatments from 

the Flora of the Guianas (Görts-van Rijn 1985 

- 2006) and the Flora of the Venezuelan Guayana 

(Steyermark et al. 1995 - 2005). As neither of 

these are complete, numerous other publications 

were also utilized (e.g., various regional floras 

and guides (Gentry 1993; Lanjouw 1964-; 

Lasser 1964-1992; Pulle 1932-; Ribeiro et al. 

1999), botanical journal articles, and botanical 

monographs). The U.S. National Herbarium at 

the Smithsonian Institution (US) was a valuable 

reference for confirming determinations and 

resolving problems. In several cases, 

identifications were provided by specialists in 

particular plant families. 

Following identification, name data were 

added to the Smithsonian’s Biological Diversity 

of the Guianas Shield’s (BDG) plant collection 

database, labels were produced for all duplicate 

specimens, and the specimens were distributed. 

The first duplicate sheet of all collections was 

returned to the University of Guyana’s (BRG), 

and the second duplicate was mounted for the 

US National Herbarium. Any additional sheets 

were distributed to various herbaria in the United 

States, South America, and Europe. Collection 

and identification information were included 

with specimens returned to the CSBD. 

Plant nomenclature was standardized and 

synonyms obtained from the Smithsonian’s 

Biological Diversity of the Guiana Shield 

Program (BDG) database for the Checklist of 

the plants of the Guiana Shield, on which 

Boggan et al. (1997) and Hollowell et al. (2001) 

were based. 

17 

Plant Species List 

Species were placed on the Waini Peninsula 

plant species list if they were collected or 

observed in the study area (west of 59° 30' W). 

Cultivated plants were not included in most 

instances, except if they were considered 

dominant in sizable areas of the vegetation or if 

they were naturalized or likely to become 

naturalized. Authors of plant names were listed 

following the Latin name and were standardized 

according to Brummitt and Powell (1992). 

Synonyms included with species lists were 

obtained from the Checklist of the Plants of the 

Guiana Shield database (Boggan et al. 1997; 

Hollowell et al. 2001), as well as from the Flora 

of the Venezuelan Guayana (Steyermark et al. 

1995 - 2005) and nomenclatural databases 

(Missouri Botanical Garden 1995-present; Plant 

Names Project 1999-present). Synonyms are 

listed alphabetically by genus and specific 

epithet, along with the currently accepted name 

of that species. 

Plant Communities 

The vegetation on the Waini Peninsula was 

classified into four basic types, from the coast 

to the Waini River: 1) Beaches, 2) Coastal 

Mangrove swamps, 3) Mixed Freshwater 

swamps, and 4) Riverine Mangrove swamps. 

Most of the vegetation of the Waini Peninsula 

was dominated by typical beach, mangrove, and 

freshwater swamp species of the Neotropics. 

That dominance was presumably influenced by 

both the limiting factor of salinity and the high 

availability of fruits and seeds dispersed by the 

ocean. In that setting, the characteristics of 

available plant species were also important in 

determining community composition and 

zonation. A previous estimate of plant 

communities on the Waini Peninsula was given 

in the Preliminary Vegetation Map of Guyana 

(Huber et al. 1995), adapted in Figure 1.8. Here, 

those boundaries have been refined through 

interpretation of Landsat 7 Enhanced Thematic 

Mapper plus (ETM + ) imagery and fieldwork, 

including GPS georeferenced plant collections 

and notes made during transects across the 

Peninsula. 

Burned Area Delineation 

A post-burn Landsat ETM + scene with little

18 


Figure 1.8. Classifications for the Waini Peninsula 

from the Preliminary Vegetation Map of Guyana 

(Huber et al. 1995) digitized onto a UTM base 

digitized from satellite imagery. Riverine Mangrove 

swamps were not differentiated from Coastal 

Mangrove swamps in the original map, but are shown 

here based on their location along the Waini River. 

The map=s areas of Mangrove swamp are 74.8 km 2 , 

Riverine Mangrove 25.2 km 2 , and Swamp Woodland 

95.0 km 2 . 

cloud cover over the area of interest on the Waini 

Peninsula was identified. That scene was 

acquired on 3 December 1999, about 20 months 

after the end of the Waini Peninsula soil fires, 

covering Path 232, Row 54. The U.S. Geological 

Survey EROS Data Center provided the scene 

in “GeoTIFF” format, which is readily displayed 

with standard Geographic Information System 

(GIS) software, Universal Transverse Mercator 

(UTM) projection for Zone 21 north, central 

meridian -57º, and processed with systematic 

geometric and radiometric corrections. Landsat 

ETM + images have a spatial resolution of 

approximately 28.5 meters in visible and 

infrared bands. The scene was cropped to the 

area of interest, 59º30' to 59º49’30'’W and 8º12' 

to 8º25’N, approximately 35.8 kilometers x 25.2 

kilometers covering about 902 km 2 , including 

appoximately195 km 2 of the Waini Peninsula. 

For initial field work, a false color composite 

Figure 1.9. Landsat ETM+ display of a portion of the scene for path 232 row 54, 3 December 1999, cropped 

to show the upper Waini Peninsula. The clouds in the lower left portion of the view were typical for most 

areas of land of the full scene. The Almond Beach burn near Waini Point was much more distinct in false 

color composite (FCC) display than the Kamwatta burn at lower right, which was characterized by less open 

water and more recovering vegetation; the Kamwatta area of the scene was subject to confusion from nearby 

clouds and cloud shadows.


(FCC) scene was composed in ArcView 3.3 

(ESRI 2002), using band 2 (green, 0.53 - 0.61 

:m), band 3 (red, 0.63-0.69 :m), and band 4 

(near-infrared, 0.78-0.90 :m) (Figure 1.9, in 

grayscale). 

Japanese Earth Resources Satellite (JERS- 

1) Synthetic Aperture Radar images from 

October 1995 were obtained from the Global 

Rain Forest Mapping Project (2001). The image 

was registered in ArcView by creation of a 

‘world file’ (ESRI 2002) with coordinates of the 

point of origin and the fraction of a degree 

represented by one pixel on each axis. Features 

visible in the JERS image (Figure 1.10) served 

as an additional guide for delineation of preburn 

plant community boundaries. 

Figure 1.10. JERS-1 radar image of the entire Waini 

Peninsula, acquired in October 1995, resolution 3 arcseconds, 

about 95 meters at the latitude of this image. 

On the coastal plain, dark areas primarily represent 

open water and smooth moist surfaces, while areas 

of high return represent distinct settlements or areas 

of flooded, intact forest, which is the likely 

interpretation along the coast near the right side of 

the image. Hills are visible along the center of the 

left edge, near the Northwest District=s administrative 

seat, Mabaruma. 

Ground-truth transects were made across 

the Waini Peninsula in and near the Almond 

Beach burn in April and October 2001. Two 

transects were made through central parts of the 

burn, and two others were made near each end 

of the burn (Figure 1.11). Transects were spaced 

approximately 2.5 to 3.5 km apart. Transect A 

crossed the Peninsula near its northwestern end, 

in unburned forest. Transect B was made from 

the vicinity of Almond Beach camp across 

19 

burned swampland. Transect C started just east 

of the Almond Beach school, crossing burned 

swampland and intact forest near the Waini 

River. Transect D was intended to pass beyond 

the southeastern end the Almond Beach burn, 

however was adapted in the field to coincide 

with a boundary of the burn. Transect E was a 

shorter exploration made beyond the 

southeastern limit of the Almond Beach burn. 

Along the transects at localities with 

distinctive vegetation types, vegetation 

transitions, burn boundaries, or plant collection 

localities, coordinates were recorded using a 

GPS receiver, plant species at those locations 

were noted. Voucher specimens were collected 

wherever a species was previously uncollected 

or field identification was uncertain. Transects 

B and C were each traversed twice, due to the 

time required to initially clear or mark routes. 

A post-burn profile was composed to 

provide a visual representation of plant 

communities across the Almond Beach burn at 

transect C. The first tropical forest survey 

utilizing profiles was apparently performed in 

Guyana (at that time British Guiana) by Davis 

and Richards (1933), detailing a mixed lowland 

forest on Moraballi Creek near the Essequibo 

River; for that study all trees along a narrow 

transect were cut and measured as the profile 

was drawn. While Ellison (2002) argued that 

mangrove community profiles are subjective 

representations of structure and zonation unless 

they are linked to quantitative data, generalized 

Figure 1.11. Positions of transects across the Almond 

Beach burn (B, C, D) and outside of the burn (A, E) 

on the Waini Peninsula. Points indicate locations of 

GPS readings where plant community information 

was gathered.

20 


profiles have been of value for advancing 

concepts of variability among mangrove 

zonation patterns (Tomlinson 1986), and they 

can been useful for conveying system structure 

to policy makers and for providing background 

for later studies. Lindeman (1953) used profile 

diagrams for his descriptions of coastal 

vegetation in Surinam, along transects from 2 

to 16 kilometers in length. 

Pre-fire plant communities were manually 

drawn by on-screen digitization of polygons 

(Duke et al. 2003; Wilton & Saintilan 2000) in 

ArcView 3.3 GIS (ESRI 2002), following 

patterns visible in the 3 December 1999 Landsat 

ETM + FCC image, and in individual bands, 

combined with transect field data and 

observations. Patterns from the JERS-1 imagery 

assisted in community delineation, particularly 

in burned areas. Mangrove swamp areas were 

divided between pure Avicennia L. swamps, 

found near the coast, and Mixed Mangrove 

swamps with both Avicennia L. and Rhizophora 

L. trees. Non-mangrove freshwater swamp areas 

were divided between Ficus L.-Euterpe Mart. 

swamps and Pterocarpus dominated Mixed 

Freshwater swamps found closer to the Riverine 

Mangroves. 

To estimate burned areas, Landsat ETM + 

bands 3 (red) and 4 (near infrared) were used in 

combination with data and observations from 

transects across the peninsula. Since different 

qualities of burned swamp could be discerned 

in the red and infrared bands, a separate 

delineation was performed for each band, using 

manual, on-screen digitization of polygons 

(Wilton & Saintilan 2000) with ArcView 

software (ESRI 2002). A linear contrast stretch 

was performed for each band, until boundaries 

between burns and intact forest, as observed in 

the field, became discernable. Polygons of the 

burn boundaries were digitized using the 

adjusted band images as a guide. A union of the 

band 3 and band 4 delineations was made to 

form polygons of total impacted areas in the 

Almond Beach and Kamwatta burns. Totals for 

burned areas were tabulated by vegetation type 

in ArcView 3.3, by intersection of polygons for 

plant communities with those for burned areas. 

RESULTS 

Plants of The Waini Peninsula 

A total of 118 names of plant species 

documented for the Waini Peninsula are 

included in Appendix 2, with a synonymy. A 

more detailed analysis of the composition of the 

plants documented for the Waini Peninsula is 

presented in Chapter 2. 

Plant Communities on the Waini Peninsula 

Transect A crossed unburned, young 

Avicennia swamp forest with some areas of 

dense Laguncularia C.F.Caertn. shrub 

understory, into mixed mangrove forest with 

mature Avicennia and Rhizophora trees and 

often very dense Acrostichum L. fern understory, 

and finally into Riverine Mangrove forest 

dominated by Rhizophora racemosa G.Mey.. 

Transect B crossed through low elevation 

burned Avicennia forest, into burned Mixed 

Mangrove forest regenerating with broad 

expanses of Typha L. or shallow water and vines, 

and finally burned Mixed Freshwater forest, up 

to the Riverine Mangrove forest. Transect C ran 

through burned young Avicennia forest, across 

the largest chenier encountered in the area, into 

burned Mixed Mangrove forest, up a very 

shallow slope to the area of the highest 

elevations with a small Roystonea D.F.Cook. 

community, to a broad area of burned Mixed 

Swamp forest with some Ficus trees and many 

dead Euterpe palms, and often shallow water. 

Beyond that zone, elevation slowly decreases 

until near the river soil moisture apparently 

discouraged fires in 1998. There a belt of 

unburned Mixed Swamp forest was 

encountered, dominated by Pterocarpus 

officinalis Jacq. with occasional, extremely large 

Avicennia trees, and finally a band of Riverine 

Mangrove swamp up to 400 meters wide along 

the Waini river, dominated by Rhizophora 

racemosa. Transect C is detailed with a 

vegetation profile in Figure 1.12. 

During the course of the transects it became 

evident that some areas not easily discernable 

in the Landsat FCC image had been burned. 

Opposite - Figure 1.12. Plant community profile diagram across the Waini Peninsula at the Almond Beach 

burn.


21

22 


Transect D, on the southeastern end the Almond 

Beach burn, unexpectedly coincided with that 

boundary of the burn, which had been expected 

to be well outside of the burned area. It was 

determined at that time to follow the boundary 

and record characteristic points with GPS to 

provide additional information for delineation 

of burns. Transect E was a shorter investigation 

started beyond the southeastern limit of the 

Almond Beach burn. In that area of Mixed 

Mangrove swamp, progress on foot was slow, 

in part because of frequent Rhizophora proproots 

and occasional treefall disturbances with 

dense understory of Acrostichum ferns and 

vines. No transects were made into the 

Kamwatta burn. Because of confusion with 

nearby clouds on the Landsat FCC image, the 

extent of that burn was not fully apparent until 

a short excursion was made for about 1 

kilometer into the area. 

A map of approximate pre-burn plant 

communities of the Waini Peninsula is shown 

in Figure 1.13. Total areas are given by 

community type in the first column of Table 1.1. 

Figure 1.13. Plant communities of the Waini 

Peninsula prior to fires, estimated from the 3 

December 1999 Landsat ETM + image and transects 

on the ground. 

The area of Avicennia mangrove swamp near 

the tip of Waini Peninsula was approximately 

11.7 km 2 . Areas of three other plant community 

types were similar, with 56.9 km 2 of Mixed 

Mangrove swamp, 59.9 km 2 of Ficus-Euterpe 

swamp, 55.3 km 2 of Mixed Freshwater swamp, 

and 10.4 km 2 of Riverine Mangrove swamp. The 

small population of Roystonea palms was not 

discernable in satellite imagery, possibly due to 

minor cloud cover in the vicinity. The Roystonea 

population was estimated during field visits to 

cover less than 0.1 km 2 . 

- Beaches 

Beach vegetation nearest to the ocean was 

composed of a few specialized vine, herb, and 

small shrub species (Figure 1.14), all of which 

are common along Neotropical coasts. 

Dominant beachfront vines were Ipomoea pescaprae 

(L.) R. Br., Canavalia rosea (Sw.) DC., 

and Passiflora foetida L., all of which occurred 

out to the high water line. The common herb on 

the fore-beaches was Sesuvium portulacastrum 

Figure 1.14. Typical Beach vegetation on the Waini 

Peninsula, with Sesuvium portulacastrum in the 

foreground and Ipomoea pes-caprae behind. 

Cultivated coconut (Cocos nucifera) trees are present 

in the background. The coarse texture of the shell 

substrate is apparent. 

Table 1.1. Areas of Waini Peninsula plant communities in the Almond Beach and Kamwatta burns from 

manual delineations. Beach and lagoon areas were not affected by fires. 

Forest type 

Avicennia forest 

Mixed Mangrove forest 

Riverine Mangrove forest 

Pterocarpus forest 

Ficus-Euterpe forest 

Total 

Waini Pre-burn 

11.7 km2 56.9 km2 10.4 km2 55.3 km2 59.9 km2 194.1 km2 Almond Beach Burn 

5.2 km2 15.7 km2 none 

2.9 km2 10.4 km2 34.1 km2 Kamwatta Burn 

none 

5.5 km2 none 

5.4 km2 19.6 km2 30.5 km2


(L.) L. Shrubs included Conocarpus erectus L. 

(a mangrove associate), Talipariti tiliaceum (L.) 

Fryxell (= Hibiscus tiliaceusL.), Thespesia 

populnea (L.) Sol. ex Corrêa, and Caesalpinia 

bonduc (L.) Roxb. These were occasionally 

interspersed with juveniles of mangrove species, 

particularly Laguncularia racemosa (L.) C.F. 

Gaertn. and Avicennia germinans (L.) Stearn. 

Apparently the shallow water over mud flats, 

the shell margins, or the high silt content of the 

coastal water are not conducive to the 

establishment of Rhizophora seedlings 

immediately along the coast (Pons & Pons 

1975), although Rhizophora propagules were 

sometimes found stranded on the shell sand of 

the forebeach. 

During field work, the beach ridges of the 

Waini coast ranged from a few meters to over 

150 meters in width. Where these were 

sufficiently wide, somewhat higher diversity of 

shrubs and trees was found on the back beach 

towards the mangrove swamps, with species 

including Spondias mombin L., Annona glabra 

L. and Morinda citrifolia L. Just below the 

surface, the wider beach ridges contained lenses 

of nearly fresh water with salinity as low as 1 

psu (Hollowell, unpublished). Because of 

available fresh water, the back beach was in 

places a site of agricultural activities. The 

dominant crop grown for trade was Cocos 

nucifera L. (Coconut), which grows well in the 

shell substrate over rich mud. Manihot esculenta 

Crantz (Cassava) is also commonly cultivated 

for community use, along with a variety of 

garden vegetables and ornamentals. Naturalized 

Carica papaya L. (Pawpaw, Papaya) was found 

frequently on beach ridges, usually 10 or more 

meters from the shore. Terminalia catappa L. 

(East Indian Almond) is a naturalized tree found 

on both beach ridges and higher cheniers farther 

inland. The small tree or shrub Morinda 

citrifolia L. (Noni), an Asian native widely 

known for medicinal values, is scattered in the 

back beach up to the margins of the mangrove 

swamp. 

- Coastal Mangrove Swamps 

Mangrove swamps are common along 

nearly all tropical shorelines where wave energy 

is low. They are dominated by woody plants 

with a tolerance to salt water (Tomlinson 1986). 

While mangrove species belong to several plant 

23 

families, they have all developed structural and 

physiological adaptations to the salinity, 

waterlogged soils, and dispersal and 

establishment challenges of the coastal 

environment (Ball 1988). Adaptations include 

roots that function to exchange air and provide 

support in waterlogged soils, a capacity to 

exclude or secrete salt, and fruits that germinate 

at least partially while on the parent or during 

dispersal by water. The number of mangrove 

species is several times higher in the Western 

Pacific than the Atlantic region. 

The Coastal Mangrove swamps were 

divided into two subsets, Avicennia forest and 

Mixed Mangrove forest (Figure 1.13). Swamp 

areas nearest to the ocean were most often 

dominated by Avicennia germinans (L.) Stearn 

(Black Mangrove, Courida) which reached 30 

meters in height (Figure 1.15). Where the 

coastline has been accreting, Avicennia 

Figure 1.15. A pure Avicennia Coastal Mangrove 

swamp near Waini Point, showing the dense carpet 

of pneumatophores. Trees in this area were measured 

as up to about 50 cm diameter and 32 meters in 

height. The pneumatophores allow gas exchange for 

the roots. Most of these trees lean to the west due to 

the soft soil and the prevailing trade winds. 

germinans is usually the first species to colonize 

a mudflat, and where monospecific stands of this 

species are found near the ocean, the trees often 

fell within a narrow range of intermediate 

diameters, suggesting that they might be the first 

generation on new land. A large area of this type 

occurred at the northwest end of the Waini 

Peninsula, including the swamp in the vicinity 

of Almond Beach. In the Mixed Mangrove 

swamp Avicennia was found with Rhizophora

24 


mangle L. (Red Mangrove, Mangro) trees which 

may also reach heights of 30 meters or more. 

Farther to the southeast and inland from Almond 

Beach, the coastal forests are presumably older, 

and more diverse. 

The third major mangrove species of the 

Neotropics, Laguncularia racemosa (L.) C.F. 

Gaertn. (White Mangrove, Kayara), does not 

often reach the stature of Avicennia or 

Rhizophora trees in the Guianas, but it was 

common as an understory shrub or treelet, as 

well as along the margins of coastal lagoons. 

Laguncularia has been observed in Guyana and 

other countries to be a very successful colonizer 

of swamps following disturbances (Baldwin et 

al. 2001; Delgado et al. 2001; Elster 2000; Roth 

1992; Woodroffe 1983). The final dominant 

component of the mangrove swamp understory 

included two species of Acrostichum (Mangrove 

Fern, Monkey Bush). Those are similar, large 

terrestrial ferns; Acrostichum aureum L. was 

common throughout the mangrove swamps, 

while Acrostichum danaeifolium Langsd. & 

Fisch. is occasional in more inland locations. 

Either can become very dense in Mixed 

Mangrove swamp. The mangrove associate 

Conocarpus erectus L. was found on chenier 

ridges within the swamps and sometimes in the 

shrubby thickets along coastal lagoons and back 

beaches. As is typical for mangrove swamps 

around the world, the Avicennia swamps near 

the coast have a fairly open understory (Janzen 

1985; Lugo 1986; Snedaker & Lahmann 1988), 

allowing easy walking, particularly during dry 

seasons. 

Mangrove swamps along Guyana’s Atlantic 

coast are well developed where they have been 

undisturbed by agriculture or urban 

development (Bacon 1990; Vann 1959, 1969; 

Wells & Coleman 1981). In places beach ridges 

separate swamps from tidal inundation, although 

waves at high tide often wash over narrower 

beach ridges and directly into the mangrove 

swamp. Those swamps behind beach ridges can 

also be referred to as occluded swamps, due to 

their separation from the sea. Such conditions 

have been documented in a few cases in the both 

the Neotropics (Lugo 1981; Stoddart et al. 1973; 

Vegas Vilarrúbia & Rull 2002; Woodroffe 1983) 

and the old world (Fosberg 1947; Steenis 1984; 

Woodroffe 1988). Within the classification of 

Lugo and Snedaker (1974) those swamps fall 

into the category of Basin Mangrove, and in 

some areas fit the classification of Hammock 

mangrove forest, a category added by Lugo 

(1980). Inlets into tidal streams are infrequent 

on the Waini Peninsula and can become blocked 

by sediment deposits, forming lagoons behind 

beach ridges. 

The Coastal Mangrove swamps of the 

Guianas do not conform to the zonation pattern 

typically described for the low sediment swamps 

of the Caribbean, in which Rhizophora occupies 

the position closest to the sea, with Avicennia 

as a back mangrove species followed by 

Laguncularia and Conocarpus as species of the 

transition to freshwater swamp or upland. A 

pattern of zonation similar to that on the Waini 

Peninsula was also reported for the Wia Wia 

Bank in northeastern Surinam by Lindeman 

(1953), though he noted an absence of 

Conocarpus. That arrangement was also 

confirmed for French Guiana by Fromard 

(1998). There Laguncularia was reported to be 

at least as common as Avicennia in the pioneer 

coastal communities, though Avicennia still 

became the dominant species of mature coastal 

swamps. 

- Mixed Freshwater Swamps 

Towards the interior of the Waini Peninsula, 

the Coastal Mangrove swamp plant community 

graded into Mixed Freshwater swamp. The 

length of that transition varied but usually 

spanned one to two kilometers. In that zone, 

occasional Avicennia germinans (L.)Stearn trees 

and Rhizophora trees persist, usually as large 

individuals. The Mixed Freshwater swamp 

group was divided into two subsets, Ficus - 

Euterpe forest and Pterocarpus forest (Figure 

1.13), which was mixed with several other tree 

and understory species. Ficus spp. trees were a 

significant component of the Ficus-Euterpe 

portion, particularly large Ficus amazonica 

(Miq.) Miq., of which several individuals 

survived the 1997-1998 soil fires in places where 

all other trees were killed. Individuals of Ficus 

eximia Schott, a similar member of the Ficus 

citrifolia group may also occur on the Waini 

Peninsula. In the center of the peninsula, parts 

of the swamp had been dominated by Euterpe 

cf. oleracea Mart. (Manicole palm) with 

scattered Ficus. In sections surveyed, all Euterpe


trees had been killed by the recent fires, leaving 

numerous fallen trunks. After the fires those 

areas were very open and dominated by a mosaic 

of vines, Typha, sedges, and various herbs. 

During wet periods, scattered shallow pools with 

floating vegetation were common there. 

The species richness of the Mixed 

Freshwater swamp increased in the Pterocarpus 

forest zone up to the transition to Riverine 

Mangrove swamps along the Waini River, in 

forests where Pterocarpus officinalis Jacq. 

(Corkwood) was common. Just before entering 

the Riverine Mangrove swamp, there was a belt 

dominated by Pterocarpus mixed with 

occasional, extremely large Avicennia 

germinans trees, some of which measured over 

two meters dbh (Figure 1.16); small Avicennia 

trees or seedlings were not common in that area. 

Some of these exceed the size of Avicennia trees 

at Maranhão, Brazil, reported by Lacerda et al. 

(2002) to grow in excess of one meter dbh. 

Fromard (1998) described a similar species 

Figure 1.16. Mixed Freshwater forest near the 

transition to Riverine Mangrove forest. This 

Avicennia germinans tree was measured at over 2 

meters dbh, though it was not measured above the 

buttresses. Pterocarpus officinalis trees were also 

found in this zone near the Waini River. 

25 

zonation in French Guiana, moving inland up 

the rivers from Coastal Mangroves towards 

Euterpe and Pterocarpus forest with fewer, but 

very large, Avicennia and Rhizophora trees, 

which he attributed to increasing influence of 

riverine rather than tidal hydrology. 

The trees Machaerium lunatum (L. f.) 

Ducke, Inga ingoides (Rich.) Willd., Zygia 

latifolia (L.) Fawc. & Rendle and Clusia 

palmicida Rich. ex Planch. & Triana were 

occasional in the Mixed Freshwater swamps, 

particularly in Pterocarpus zones. There the 

understory included Cassipourea guianensis 

Aubl., Ilex guianensis (Aubl.) Kuntze and 

Malouetia tamaquarina (Aubl.) A. DC., as well 

as several species of ferns including Blechnum 

serrulatum Rich., Pteris pungens Willd., and 

Nephrolepis biserrata (Sw.) Schott. Vines and 

lianas noted in those swamps included Monstera 

adansonii Schott, Syngonium podophyllum 

Schott, and Hippocratea volubilis L. as well as 

several vines that had become dominant in the 

burned swamp including Cissus verticillata (L.) 

Nicolson & C.E. Jarvis, and Cydista 

aequinoctialis (L.) Miers. While trees in the 

Coastal Mangrove swamps were host to few 

epiphytes, probably due to salinity from the 

ocean, several epiphytic species were found in 

the Mixed Freshwater swamps, including the 

bromeliads Aechmea nudicaulis (L.) Griseb. and 

Guzmania lingulata (L.) Mez, the cactus 

Rhipsalis baccifera (J.S. Muell.) Stearn, and the 

orchids Epidendrum ciliare L. and Prosthechea 

aemula (Lindl.) Higgins. Some epiphytic 

species were recorded in the Mixed Freshwater 

forest only as survivors on the branches of fallen 

trees killed by fires, including the cactus 

Epiphyllum phyllanthus (L.) Haw. and the orchid 

Trichocentrum lanceanum (Lindl.) M.W. Chase 

& N.H. Williams. 

An unexpected feature of the Mixed 

Freshwater swamps was the isolated population 

of Roystonea oleracea (Jacq.) O.F. Cook, 

consisting of perhaps 120 scattered individuals 

of all sizes (Figure 1.17). That population 

survived the 1998 soil fires while most 

surrounding vegetation was killed except for a 

few large Ficus trees. Prior to the documentation 

of that population, no species of the genus 

Roystonea had been recorded in the Guianas 

outside of cultivation (Zona 1996). The remote

26 


Figure 1.17. Location of the Roystonea oleracea 

community on the Waini Peninsula, within the 

approximate area of the elevated central zone of the 

peninsula that became known as APalm Hill.@ The 

initial excursion to the site was made on 8 April 2001 

(Palm Sunday). 

location of the population strongly suggested 

that it was not likely planted by people. 

- The Riverine Mangrove Swamps 

The swamps that line the tidal Waini River 

are inundated twice daily with brackish river 

water, since there are no barriers to tides similar 

to the beach ridges isolating many Coastal 

Mangrove swamps. That allows comparatively 

unimpeded dispersal of mangrove propagules 

in this zone, which may be partly responsible 

for the dominance of Rhizophora racemosa G. 

Mey., which fringes the Waini River and extends 

inland on the soft, muddy substrate to the tidal 

limit. That species is very similar to R. mangle 

in vegetative appearance, having similar 

conspicuous prop and aerial roots and 

indistinguishable leaves and bark. It differs 

distinctively from R. mangle in having 30 to 60 

flowers per inflorescence, as compared to 2 to 

3 for Rhizophora mangle; in having buds with 

obtuse apices as opposed to buds with acute 

apices in Rhizophora mangle; and in having 

propagules with much longer hypocotyls 

extending from the viviparous fruits, up to 50 

cm. in length as opposed to hypocotyls up to 20 

cm in length in Rhizophora mangle (Gray 1981). 

Rhizophora racemosa trees tend to have more 

aerial roots extending from higher branches to 

the water’s surface. Those two Rhizophora 

species have been considered by some to form 

the hybrid Rhizophora x harrisonii Leechm., 

which has been ascribed to similar brackish 

riverine habitats and is supposedly intermediate 

between the two species in reproductive 

morphology (Gray 1981; Tomlinson 1986). In 

practice, it was difficult to distinguish that 

putative hybrid from R. racemosa, and some 

specialists consider it to be synonymous with 

R. racemosa (Duke et al. 2001). In several 

descriptions of riverine mangroves in the 

Guianas, R. racemosa has been unmentioned, 

with only the name R. mangle used, although 

that is almost certainly inaccurate. 

The Riverine Mangrove swamp understory 

was, in places, moderately dense with 

Rhizophora saplings and Acrostichum aureum 

ferns (Figure 1.18). The non-native mangrove 

palm Nypa fruticans Wurmb. was also common 

in the Riverine Mangrove swamp, sometimes 

Figure 1.18. Riverine Mangrove forest dominated by 

Rhizophora racemosa. Some Nypa fruticans palms 

were in the understory (center background) and some 

fronds of the mangrove fern Acrostichum are visible 

in the foreground. 

in dense patches along river margins and often 

as an occasional component of the understory. 

Upriver, around 59° 30' W, Nypa was no longer 

present. At that point Rhizophora trees were also 

infrequent along the river margin, although they 

were observed as far up the Waini River as its 

confluence with the Barama River, and were 

seen occasionally on the Baramanni River. 

Along some segments of the Waini River the 

roots of Rhizophora racemosa were found 

covered with the red alga Bostrichia and 

occasionally with barnacles. Farther up the river, 

where the water became less brackish, 

freshwater swamp tree species including 

Pterocarpus, Mora Benth., Machaerium Pers., 

Inga Mill., Euterpe and Pachira Aubl. were

increasingly common along the banks. 


Burned Areas 

The manual delineation using the Landsat 

red band (3) included 27.4 km 2 of burned 

vegetation at Almond Beach and 28.8 km 2 of 

burned vegetation at the Kamwatta (Figure 

1.19). The major result of the fires detected in 

band 3 was apparently bare soil, which reflected 

Figure 1.19. Manual delineation of burned areas 

using Landsat ETM + band 3, red. 

red portions of the spectrum and appeared as 

light areas of the image. Using Landsat band 4 

(near infrared), the manual delineation included 

26.5 km 2 of burned vegetation at Almond Beach 

and 22.6 km 2 of burned vegetation at the 

Kamwatta site (Figure 1.20). That band was 

useful for detecting vegetation (high reflectance) 

and open water (low reflectance). The burned 

area totals were calculated by a union of the red 

Figure 1.20. Manual delineation of burned areas 

using Landsat ETM + band 4, near infrared. 

27 

and near infrared band results. The intersection 

of the total burns with the plant communities 

classifications provided areas of each 

community that were impacted. Totals burned 

areas for each plant community type and locality 

are given in Table 1.1. It was estimated that 34.5 

km 2 of the forests in the Almond Beach area 

were burned, and 30.5 km 2 of the forests in the 

Kamwatta area were burned, for a total of 64.6 

km 2 of burned forest for the entire area of interest 

on the Waini Peninsula (Figure 1.21). Areas of 

burns detected with each of the two Landsat 

bands are listed by locality in Table 1.2. 

Figure 1.21. Combined areas of manual delineations 

of burns using the red and infrared bands from the 

December 3 1999 Landsat ETM + image. The total 

burned area was measured as 64.58 km 2 . 

Table 1.2. Manual burn delineation results by burn 

area and bands 3 (Red) and 4 (Near Infrared), for 

the Landsat ETM + image of 3 December 1999. 

Band / area 

IR Almond Beach 

IR Kamwatta 

RED Almond Beach 

RED Kamwatta 

km 2 

26.4 

21.9 

27.4 

28.7 

The fires left large areas with few surviving 

mangrove trees to supply propagules for 

regeneration. After that large-scale disturbance, 

limitations on dispersal would likely retard 

recolonization by some species. Species that are 

wind or animal dispersed might be favored over 

those dependent on water dispersal. 

Additionally, variations of soil salinity levels

28 


would further influence the species that might 

become dominant. There has been substantial 

variation in initial recovery of burned areas. 

In many locations, the initial regrowth after 

the fires in Coastal Mangrove swamp and Mixed 

Freshwater swamp was dominated by vines, 

most notably Ipomoea tiliacea (Willd.) Choisy, 

Mikania micrantha Kunth, Cydista 

aequinoctialis (L.) Miers and Entada 

polystachya (L.) DC. In places the tangle of 

vines became thick enough to walk upon nearly 

a meter above the ground (Figure 1.22). Other 

broad areas were dominated by dense, 

Figure 1.22. Heavy growth of vines three years after 

fires, in the former Euterpe swamp. The dominant 

species was Ipomoea tiliacea, which grew over tall 

stalks of Montrichardia linifera. In the distance 

nearer the Waini River was unburned Mixed 

Freshwater forest is visible, dominated by 

Pterocarpus officinalis interspersed with very large 

Avicennia trees. 

Figure 1.23. Burned Coastal Mangrove forest near 

Kamwatta Beach, approximately three years after 

fires. This area had become dominated by dense 

Acrostichum ferns. 

sometimes monospecific stands of herbaceous 

species such as Acrostichum ferns (Figure 1.23) 

or Typha domingensis Pers. (cattails) (Figure 

1.24). 

Figure 1.24. An expanse of Typha in burned 

Avicennia Coastal Mangrove forest, three years after 

the fires. It was estimated that this Typha stand 

covered up to 2.6 km2, starting approximately 1.4 

km behind the Almond Beach camp, extending 

towards Waini Point. 

Another distinctive vegetation type visible 

on Landsat ETM + imagery was a narrow band 

of dense herbaceous vegetation found in a few 

spots between burns on former Ficus-Euterpe 

swamp and Mixed Freshwater swamp near the 

river. Acrostichum ferns and the large, 

herbaceous aroid Montrichardia grew thickly 

there, mixed with several species of vine (Figure 

1.25). Those areas were characterized by high 

Figure 1.25. Vegetation in the belt of dense 

Montrichardia (broad leaves) and Acrostichum ferns 

near unburned Mixed Freshwater forest near the 

Waini River. Several months later this area was 

overrun with vines (pictured: Keith David).


reflectance in both infrared and red bands. 

Burned areas in the pure Avicennia swamp 

of the northern portion of the Waini Peninsula 

were investigated in more detail with ecological 

plots (see Chapter 4). Those swamps suffered 

nearly full tree mortality and have undergone 

slow early regeneration. Initially, in wet seasons 

they were covered with deeper surface water 

than neighboring unburned areas, probably 

resulting from a combination of decreased 

transpiration and lowered elevation from 

combustion of the upper organic soil layer. 

When surface water was present they were 

dominated by floating vegetation, first Lemna 

aequinoctialis Welw., which was followed after 

the first year by Limnobium laevigatum (Humb. 

& Bonpl. ex Willd.) Heine (Figure 1.26). Such 

areas that were covered with floating plants were 

not distinguishable from forest in band 3 of the 

Landsat images, while sparse vegetation and 

bare soil were apparent. Areas of open, standing 

water were obvious in the near infrared band 

(4) due to very low return. 

Fig. 1.26 Burned area in the pure Avicennia swamp of the 

northern portion of the Waini Peninsula. When surface 

water was present they were dominated by floating 

vegetation, first Lemna aequinoctialis Welw., followed by 

Limnobium laevigatum (Humb. & Bonpl. Ex willd.) Heine. 

Later in those areas, shrubby Laguncularia 

racemosa (L.) C.F. Gaertn. trees became 

established; that species is often considered the 

most disturbance adapted of Neotropical 

mangrove species (Ball 1980; Elster 2000; Elster 

et al. 1999; Roth 1992; Woodroffe 1983), as it 

has the smallest, and so presumably most easily 

29 

dispersed, propagules. 

In areas that were not apparent as obviously 

burned in the FCC image but were actually 

recovering from fires, most were lightly to 

moderately covered with herbaceous vegetation 

including vines, Typha, and Acrostichum, which 

reflected adequate near-IR (band 4) to make 

them difficult to distinguish from undisturbed 

vegetation. Most of those areas were also 

slightly elevated and not covered with standing 

water or with highly saturated soils that 

contribute to low return in the near-IR. 

The Kamwatta burn was the subject of 

limited field work, as it was not initially 

discerned in the Landsat FCC, perhaps due to 

limited areas of standing water, and confusion 

with adjacent areas of clouds and cloud shadows 

in the far southeastern part of the view. 

Nonetheless, manual delineation of the 

Kamwatta burn using separate bands 3 and 4 

was practicable and consistent with preliminary 

field investigations in the area. In the future it 

would be useful to conduct additional field 

investigations in that area south of Kamwatta 

Beach to determine if fires had occurred there. 

Plant Collections of Interest 

Some plant collections made during field 

work were of particular interest because they 

represented new information on the ranges of 

species. In the first section, collections from the 

Waini Peninsula are covered, followed by 

collections made elsewhere in the Northwest 

District. 

- On the Waini Peninsula: 

Rhizoclonium africanum Kützing, 

Cladophoraceae, det. J. Norris, 2000: Hollowell, 

Arjoon and Chin 255, collected 4 November 

1998. This algal species is newly recorded for 

the Guianas. It is known from mangrove swamps 

in Brazil, Central America, the Caribbean, and 

the west coast of Africa. Through introduction 

it is nearly pantropical in mangrove swamps 

(Guiry & Nic Dhonncha 2004). The collection 

was made just behind the beach ridge 

approximately 4 kilometers southeast of Waini 

Point. Rhizoclonium was collected on 

pneumatophores of Avicennia trees, and was 

very common in the wetter parts of the Coastal 

Mangrove swamps near the beach ridges.

30 


Although green algae are photosynthetic protists 

rather than true plants, this is listed because the 

study of algae is typically grouped within the 

discipline of botany. 

Amaranthus australis (A. Gray) J.D. Sauer, 

Amaranthaceae, det. T. Hollowell, 2001: 

Hollowell, A. James and Savory 352, collected 

12 May 2000. This was apparently the second 

collection for Guyana; the other known 

collection was M.S. Grewal and H. Lall 305 

(held at Utrecht Herbarium), from the Shell 

Beach area in 1977. The species is also known 

in the Guianas from two collections made in 

French Guiana and two collections made in 

Surinam (R.A. DeFilipps, pers. comm.). This 

collection was made to the southeast of 

Kamwatta Beach (see Figure 1.2), in open water 

of scrubby mangrove swamp that was possibly 

affected by a small fire around 1992 (Audley 

James, pers. comm). 

Nypa fruticans Wurmb., Arecaceae, det. T. 

Hollowell, 1997: Hollowell et al. 213, collected 

16 June 1997. This was the first known South 

American voucher for this Asian mangrove 

palm, collected along the banks of the Mora 

Passage between the mouth of the Waini River 

and the Barima River (Figure 1.2). Nypa is a 

monotypic genus and is unusual among both 

mangroves and palms because of its 

dichotomously branching, rhizomatous form 

(Tomlinson 1986; Tralau 1964); only the fronds 

and reproductive structures of Nypa extend 

above the riverbank mud. Nypa is common in 

Riverine Mangrove swamps along the tidal 

Waini River, Barima River, and Mora Passage. 

This species may be spreading in the Northwest 

District, displacing some native Rhizophora 

racemosa trees. Nypa fruits have been reported 

stranded on beaches in Trinidad (Bacon 2001). 

Plants have been recorded on the Caribbean 

coast of Panama (Duke 1991), and Nypa has 

become a widespread exotic species in the Niger 

Delta (Sunderland & Morakinyo 2002). It has 

long been utilized in Asia for many purposes, 

particularly thatching, but it has only rarely been 

observed in use in Guyana. 

Roystonea oleracea (Jacq.) O.F. Cook, 

Arecaceae, det. T. Hollowell, 2002: Hollowell 

and Hinds 593, collected 21 October 2001. This 

was apparently the first record of native 

Roystonea for the Guianas. It was collected in 

fruit from the isolated population midway across 

the Waini Peninsula, with an elevation of 5-10 

meters above sea level. It was composed of 50- 

60 mature trees up to 20 meters high, with a 

number of juvenile palms present (Figure 1.27). 

These palms may have been able to survive fires 

because palm cambium tissues are protected by 

Fig. 1.27 The first native population of Roystonea oleracea 

(Jacq.) O.F. Cook found midway across the Waini 

Peninsula. This isolated population was composed of 50- 

60 mature trees up to 20 meter high, with a number of 

juvenile palms present. 

their internal position (Cochrane 2003). The 

species was known from Venezuela including a 

few localities in Delta Amacuro and Bolívar 

states, and from Colombia. Some of those 

populations have been reported as native (Zona 

1996). Roystonea oleracea also occurs in coastal 

swamps of Trinidad and Tobago (Bonadie 1998), 

and Barbados (Zona 1996), apparently in 

slightly elevated locations similar to that of the 

Waini Peninsula population. Species of 

Roystonea have been cultivated in cities and 

towns of the Guianas, but the Waini Peninsula 

population occurred far from any settlement and 

more than 20 kilometers from Mabaruma, the 

nearest town with cultivated Roystonea. 

Ipomoea violacea L., Convolvulaceae, det. 

T. Hollowell, 2002: Hollowell 217, collected 17 

June 1997. This was possibly the second 

collection in Guyana of the white-flowered, 

coastal species. Collected along a narrow beach 

ridge west of Almond Beach. The other known 

specimen was collected by G.S. Jenman 

(collection number 5068, at US as Ipomoea 

macrantha Roem. & Schult.) in 1889 in “coastal


lands” of Guyana. It is a widespread coastal vine 

in the Caribbean and has a nearly pantropical 

distribution. 

Sesbania sericea (Willd.) DC., Fabaceae, 

det. T. Hollowell, 2000: Hollowell, A. James and 

Savory 351. Collected along a beach ridge swale 

to the southeast of Kamwatta Beach. The other 

known collection of this species from Guyana 

was made by A.S. Hitchcock (collection number 

16626) in 1919, with the collection habitat listed 

as “among weeds” in the Georgetown Botanic 

Garden. The species is native to Asia, possibly 

to Sri Lanka (Howard 1976). 

- In the Northwest District of Guyana: 

Passiflora amicorum Wurdack, 

Passifloraceae, det. C. Feuillet, 2000: Hollowell 

and V. James 383, collected 16 May 2000. 

Collected on “Kissing Rock” hill north of 

Mabaruma. This was the first collection of the 

species for the Guianas; it was previously known 

only from Bolívar state of Venezuela. This redflowered 

vine was growing upon low understory 

vegetation on a steep hillside, in partial shade. 

Werauhia gigantea (Mart. ex Schult. f.) J.R. 

Grant. (=Vriesea amazonica (Baker) Mez.), 

Bromeliaceae, det. E.J. Gouda, 2002: Hollowell, 

A. James and V. James 386, collected 16 May 

2000. This was apparently the second collection 

of the species in Guyana, the previous record 

being from Mabura Hill in central Guyana, 

collected by R. Ek (1997). This specimen was 

collected from the branches of a Rhizophora 

racemosa tree overhanging the Aruka River near 

“Kissing Rock” in the Mabaruma vicinity. 

Habenaria sp. nov.? Orchidaceae, det. G. 

Carnevali, 2002. Hollowell and Hinds 607, 

collected 22 October 2001. This white-flowered, 

terrestrial orchid was not matched by other 

Habenaria species (fide Germán Carnevali, 

CICY) and is awaiting further examination. 

Collected on terra firme, approximately 100 

meters from the Waini River well upriver from 

tidal influence, near an abandoned sawmill. 

Miconia minutiflora (Bonpl.) DC. 

Melastomataceae, det. F. Almeda and D. 

Penneys, 2002. Hollowell and Hinds 724, 

collected 4 November 2001. This was the first 

collection of the species for Guyana. Specimens 

of this shrub were collected near the Wauna Oil 

31 

Palm Estate, a small agricultural experimental 

station that is located about 12 kilometers to the 

west-southwest of Mabaruma. 

DISCUSSION 

These plant collections in Northwest 

Guyana were carried out as part of ecological 

research rather than as part of comprehensive 

plant collection expeditions. Dedicated 

collection efforts would almost certainly yield 

additional plant species records for the Waini 

Peninsula. The number of species records 

resulting from this fieldwork that were found to 

be of interest highlights the degree to which the 

flora of this lowland region has been undersampled. 

Compared to remote interior areas in 

Guyana, the coastal plain of the Northwest 

District can be easily accessed, and additional 

field work might be relatively inexpensive for 

the information yielded. As the state of botanical 

research in Guyana advances, there will be more 

effort to understand species distributions at the 

level of a variety of ecosystems and for smaller 

political units. While existing specimen data can 

be modeled with GIS to investigate probable 

species distributions (Funk & Richardson 2002; 

Funk et al. 2005; Funk et al. 1999), the needs to 

verify models and produce more accurate and 

complete input data will always depend upon 

plant collecting expeditions. A substantial 

amount of fieldwork, plant identification, and 

museum work will be required to provide those 

specimens and data. 

Changes in the global climate could have 

significant effects on Guyana’s coastal 

ecosystems, changing the types of dominant 

plant communities. Increased severity of El 

Niño events could lead to higher fire frequencies 

on the northwest coast of Guyana, particularly 

if population pressure grows, increasing the 

chances of ignition. Guyana’s occasional 

extreme droughts make it vulnerable to 

incremental forest degradation in which fires can 

play a key role (Cochrane 2002, 2003). The 1998 

fires on the Waini Peninsula had severe impacts 

on coastal swamp habitats, and effects on forest 

composition are likely to persist. Fires among 

the Waini Peninsula mangroves were possibly 

less anomalous than they initially seem

32 


(Lindeman 1953; Pons & Pons 1975), and may 

be a large-scale case of fire’s role in maintaining 

mangrove/marsh boundaries, as documented for 

some other regions (Lugo 1997; Middleton 

1999; Odum et al. 1982; Wade et al. 1980). The 

fires may represent an occasional phenomenon 

that has occurred in the coastal zone of the 

Guianas, where mangrove forests become 

isolated from the ocean, resulting in at least a 

temporary conversion to non-mangrove 

communities. 

The Waini fires clearly provide a case for 

the inclusion of fire in the list of disturbances 

that can affect large areas of mangrove swamp 

(Finn et al. 1998; Jiménez et al. 1985). It is not 

known if the fires documented here were the 

first in those coastal swamps. Sampling for the 

presence of charcoal at promising sites in central 

portions of the Waini Peninsula might answer 

that question. Such potential sites could include 

areas of slightly higher elevation, and areas that 

are unusual in older imagery, particularly where 

breaks in forest are suspected. 

The fires have also provided a natural 

experiment on the effects of disturbance on 

biodiversity and plant community structure 

(Field et al. 1998), and present continued 

opportunities for descriptive and comparative 

studies of mangroves in northeastern South 

America. Such studies of pre- and postdisturbance 

environments create baselines for 

future monitoring and analyses, and hopefully 

will encourage anticipation and prevention of 

fires. An effective mechanism for alerting 

environmental authorities and prompt field work 

are needed in rapidly changing, post-fire 

ecosystems, requiring commitment to 

conservation, monitoring, and response. 

The assembly of background environmental 

information, including censuses of plant and 

animal species and mapping, are central to the 

description, understanding and management of 

protected areas and other sites of environmental 

importance. Such baseline biodiversity 

information is essential for monitoring changes 

over time, and makes comparative studies 

between sites possible. Catalogs of species are 

also a primary tool for utilization of protected 

areas for education, and it is hoped that the 

information presented here will be encourage 

additional scientific, conservation, and 

educational efforts.


CHAPTER 2 

PLANTS OF THE WAINI PENINSULA IN REGIONAL 

AND GLOBAL CONTEXT 

INTRODUCTION 

The conservation value of a particular 

natural site may be derived not just from its 

species composition, but from the contrast of 

the species of the site with those of adjacent or 

related localities. This chapter investigates 

relationships between the Waini Peninsula flora 

and the species of nearby plant communities, 

the species of the Northwest District of Guyana, 

and the species of the adjacent Venezuelan state 

of Delta Amacuro. The phytogeographic 

affinities of the Waini Peninsula flora at the 

regional, continental, and global scales were 

explored through classification of species 

distribution ranges. Those comparisons also 

served as an example of the state of botanical 

knowledge on lowland forests in northeastern 

South America. Comparisons of Waini 

Peninsula plant communities were made with 

two coastal plain plant communities in the 

Northwest District near Santa Rosa, as 

documented by van Andel (2000a). 

Comparisons were also made of the Waini and 

Northwest District floras with the flora of the 

neighboring coastal Venezuelan state of Delta 

Amacuro, with the goal of understanding the 

level of diversity between the sites. This has 

implications both for the direction of additional 

basic research and ideally in the design of 

protected areas in the region. Finally, a regional 

and global phytogeographic analysis was made 

for the Waini flora. The questions addressed here 

include to what degree the flora of the Waini 

Peninsula is unique in relation to neighboring 

ecosystems, to all of Guyana, to the Guiana 

Shield region of northeastern South America, 

and to broader biogeographic areas. At the larger 

regional scales, the question is asked whether 

such coastal plain environments of the Guianas 

should be considered as part of the Guiana 

Shield flora or as units of a distinct Neotropical 

or Caribbean coastal flora. It is of interest 

whether additional botanical collection activities 

33 

such as undertaken in this relatively lowdiversity 

region of South America can add 

significantly to botanical knowledge. 

Comparisons of species checklists covering 

areas of intermediate size have been used 

recently to analyze similarities and differences 

among sites that are candidates for protection 

or under consideration for additional detailed 

investigations (Clarke & Funk 2005; Clarke et 

al. 2001a; Kelloff 2002; Kelloff & Funk 2004). 

That approach relies only upon species presence 

and absence, allowing what may be an early 

opportunity for biodiversity analysis. Such 

qualitative data are frequently the only available 

information covering large parts of tropical 

South America. While ecological plot data sets 

can be very useful for detailed quantitative 

analyses, they have been available to date for 

small to moderately sized plots that are widely 

separated (Condit et al. 2002; Pitman et al. 1999; 

Pitman et al. 2001). As an alternative, qualitative 

species list-based data are comparatively easily 

compiled from existing publications such as 

regional florulas. Those sources can be 

augmented by data from the large, though often 

incompletely utilized reservoirs of information 

held in the world’s herbaria (Funk 2003a, b). 

An important condition for validity of all scales 

of comparison is standardization of the 

biodiversity data utilized. This greatly facilitated 

by the availability of computerized 

nomenclatural databases, and for botanical 

research in particular, nomenclature including 

complete, linked synonymies. All of these data 

sources and data quality tools require substantial 

investment to create, maintain, and apply. 

STUDY SITES AND 

BACKGROUND 

The study site on the northern portion of 

Guyana’s Waini Peninsula is described in detail 

in Chapter 1. Other localities mentioned in the 

text or from which data were utilized are

34 


indicated in Figure 2.1. Additional plant data 

were obtained or compiled for the sites listed 

below. 

Figure 2.1. The Northwest District of Guyana, 

showing localities in the text. Georeferenced 

collection sites from this study and from herbarium 

specimens are indicated by the symbol • 

Quackal and Manicole Plant Communities 

The Quackal and Manicole plant 

communities are coastal plain vegetation types 

that were studied by van Andel (2000a), who 

considered them to grade into Coastal Mangrove 

swamps, making them of interest for 

comparison with the plant communities of the 

Waini Peninsula. Van Andel’s research sites 

were located near Santa Rosa (7º41’N, 

58º55’W) and Assakata (7º44’N, 59º04’W), in 

the southeastern portion of the Northwest 

District (Figure 2.1). She described the Quackal 

swamp woodland as dominated by Tabebuia 

insignis (Miq.) Sandwith, Macrosamanea 

pubiramea (Steud.) Barneby & J.W. Grimes, 

and Symphonia globulifera L. f. The manicole 

swamp was dominated by Euterpe palm species 

and was also rich in Pentaclethra macroloba 

(Willd.) Kuntze, Symphonia globulifera L. f., 

and Virola surinamensis (Rol. ex Rottb.) Warb. 

A comparison of the list of species collected 

from these two areas was also used to enhance 

the species list formed for the Northwest 

District. 

The Northwest District 

The Northwest District covers 

approximately 19,190 km 2 , about 9.5 % of 

Guyana’s total area. Of that area, approximately 

1,000 km 2 , or 5.5%, of the Northwest is 

contained within the Waini Peninsula, here 

considered to span from Waini Point to 

Baramanni Lake, including most of the district’s 

Atlantic coastline. The 190 km 2 study area on 

the Waini Peninsula comprises about 1% of the 

area of the Northwest District. The only large 

town in the region is the administrative center 

Mabaruma, adjacent to Port Kumaka on the 

Aruka River (Figure 2.1). Most of the towns of 

the district are accessible via its many rivers. 

Early major collection expeditions in the 

Northwest District were undertaken by the 

Schomburgk brothers and by J.S. de la Cruz. 

The German brothers Robert H. and Moritz 

Richard Schomburgk explored the Northwest in 

1841, as part of work for the British Government 

in delineation of the boundaries of British 

Guiana (Schomburgk 1922; Schomburgk 1896). 

The basecamp for that expedition was at Port 

Kumaka. At the beginning of the expedition, the 

Schomburgk party camped on the beach ridge 

of Waini Point. They recorded no detailed 

description of the flora there, although Richard 

Schomburgk made an account of birdlife, as well 

as notes on the heat of the open sun, the plentiful 

mosquitos, and the types of small shells that 

make up the beach (Schomburgk 1922). 

Unfortunately, nearly all of the Schomburgk 

botanical specimens from that expedition were 

lost to mold and decay (van Dam 2002). 

Juan S. de la Cruz was a plant collector for 

the New York Botanical Garden, working under 

contract to the prominent botanist H.A. Gleason 

(Ek 1990). Cruz collected in many regions of 

Guyana, particularly along the rivers of the 

Northwest District and the upper Mazaruni 

River basin. From an analysis of collecting dates 

on Cruz specimens, it appears that he made at 

least six collecting trips to the Northwest 

District. Two of these trips were extensive, in 

August to September 1921 and in May 1922. 

From the collection number ranges found, it is 

estimated that Cruz made over 1,400 plant 

collections in the Northwest from 1921 to 1927, 

approximately 80% of which are represented at 

the US National herbarium, according to 

database records of the BDG program. 

More recently, Tinde van Andel (2000a; 

2000b), of the University of Utrecht and


Tropenbos Foundation, made numerous plant 

collections in the Northwest from 1995-1997 as 

part of research on non-timber forest products 

in the region, including voucher collections for 

her ecological plots in the Santa Rosa vicinity, 

located just to the south of the Waini Peninsula. 

Those research sites start in freshwater forests 

apparently just inland of coastal plant 

communities that include some mangrove 

species. A summary of early botanical collectors 

who worked in the Northwest Region is given 

in Table 2.1. Georeferenced plant collection sites 

in the Northwest District are shown in Figure 

2.1. 

Delta Amacuro 

The Flora of the Venezuelan Guayana 

(Steyermark et al. 1995 - 2005) includes listings 

of the plant species for the Venezuelan state of 

Delta Amacuro, located adjacent to the 

Northwest District of Guyana (Figure 2.2). This 

allowed a comparison to be made between the 

species known from the Northwest District and 

the species listed for Delta Amacuro. Waini 

Point is located within 6 kilometers of Delta 

Amacuro, which includes the large delta of the 

Orinoco River. North of the Waini Peninsula, 

the coastline of Delta Amacuro possesses 

vegetation predominantly of swamp forest and 

35 

marshland along the Orinoco River’s many 

distributaries. 

The state of Delta Amacuro covers 

approximately 36,663 km 2 , nearly twice the area 

of the Northwest District and approximately 8% 

of the 472,000 km 2 Venezuelan Guayana, which 

is the Venezuelan portion of the Guiana Shield 

region Gibbs & Barron, 1993) as covered in the 

floral treatment by Steyermark et al. (1995 - 

2005). Huber (1995c), citing Pannier and Fraino 

Figure 2.2. The location of the state of Delta 

Amacuro, Venezuela in relation to the Northwest 

District, Guyana. 

Table 2.1. Major historical (pre-1980) collecting trips in Northwestern Guyana from US National 

Herbarium records. 

Collector 

Schomburgk, M.R. and R H. 

Jenman, G.S. 

ImThurn, E.F. 

Bartlett, A.W. 

Beckett, J.E. 

Ward, R. 

Hitchcock, A.S. 

Cruz, J.S. de la 

Altson, R.A. 

Archer, W.A. 

Fanshawe, D.B. 

Cowan, R.S. 

Maguire, B. 

Mori, S.A. 

Maas, P.J.M. 

Year 

1841 

1896 

1897 

1905 

1906 

1907 

1920 

1921-23,1927 

1926 

1934 

1945 

1955 

1955 

1976 

1977 

Localities 

Waini R., Mora Passage, Barima R., Aruka R. 

Barima R. 

Aruka R., Barima R. 


Baramanni, Waini R. 

Aruka R. 

Morawhanna, Issorora, Aruka R., Yarikita R., Amakura R. 

Waini R., ‘Marabo Shortcut’, Barima R., Amakura R., 

Moruca R. 

Mabaruma 

Mabaruma, Aruka R., Barima R., Koriabo R., Wauna 

Waini R., Aruka R., Mabaruma 

Barima R. 

Matthew’s Ridge 

Matthew’s Ridge 

Aquero, Kwebana

36 


de Pannier (1989), reported that 4,600 km 2 of 

the state, about 12.5% of the total area, was 

mangrove swamp, although that was possibly 

an overestimate resulting from inclusion of nonmangrove 

areas (Spalding et al. 1997). That 

compares to only 890 km 2 of mangrove swamp 

in the Northwest District of Guyana (Huber et 

al. 1995), which is about 4.6% of the district’s 

total area. Although Delta Amacuro has a 

considerably larger area of lowlands, maximum 

elevations in Delta Amacuro and the Northwest 

District are similar. Elevations on Serranía de 

Imataca in Delta Amacuro exceed 500 meters 

at the border with Bolívar State, while in the 

Northwest District elevations approach 600 

meters in the extreme western parts of the 

Barama River watershed at its border with 

Bolívar, and approach 500 meters in the 

headwaters of the Kaituma river and in the 

Kauramembu Mountains on its southern border 

with Guyana’s Cuyuni-Mazaruni Division. The 

southern portion of Serranía Imataca extends 

into the Northwest and forms the upper 

watershed of the Barima River. Therefore, 

because of its proximity and comparable 

geomorphic setting, the flora of the Orinoco 

Delta might be expected to have a high degree 

of similarity to the flora of the Waini Peninsula. 

The lower plant species diversity of the 

lowlands of Delta Amacuro, in comparison to 

the neighboring highlands of the Guiana Shield, 

has led to less interest in and under-sampling 

of its vegetation. For its area, far fewer 

collections have been made by early collecting 

expeditions in Delta Amacuro than in Amazonas 

or Bolívar. Out of more than 25,600 collections 

from the three states of the Venezuelan Guayana 

that have been databased from the US National 

Herbarium, only 4.8% were collected from 

Delta Amacuro. Although that region is not a 

likely source for many species new to science, 

its flora is possibly incompletely known, which 

can bias studies of regional diversity and 

conservation efforts. 

Regional and Global Affinities of the Waini 

Peninsula Flora 

The plant communities of the Waini 

Peninsula include many coastal species that 

have very broad distributions. In addition to this 

oceanic influence, the Peninsula makes up the 

outer edge of the coastal plain along the 

northeastern side of Guiana Shield. The Guiana 

Shield is a distinct geological and biological unit 

that is isolated in northeastern South America 

by broad surrounding lowlands and the Amazon, 

Negro and Orinoco River system (Gibbs & 

Barron 1993; Huber 1995a; Kelloff & Funk 

2004). The degree to which proximity to the 

Shield’s uplands is reflected in the Waini flora 

is a central question, as are the degrees to which 

other geographic affinities are apparent in the 

flora. In an effort to illuminate the floral 

affinities of one particular site, Kelloff and Funk 

(2004) performed a biogeographic analysis of 

the flora of Kaieteur National Park, Guyana, one 

of Guyana’s few protected areas, located in 

central Guyana on sandstones of the Shield’s 

Roraima formation. That analysis utilized a 

classification based on distribution ranges of 

each species to produce a summary of global 

affinities of the entire flora. In part, the 

inspiration for that method was the 

biogeographic investigations of British botanist 

Joseph Hooker (1817-1911), which have been 

summarized by Brundin (1966). Hooker’s goal 

was to understand the history underlying the 

distributions of plant taxa of the Antarctic 

circumpolar regions. Also incorporated into the 

method here is the phytogeographic work of 

Leon Croizat (1964). However, where Croizat 

connected ranges of disparate taxa with linear 

“tracks,” Kelloff and Funk grouped all species 

of a site of interest into a limited set of 

generalized, nested distribution classes for the 

biogeographic analysis. 

METHODS 

Comparison of the Waini Flora with 

Quackal and Manicole Plant Communities 

Following the approach of Clarke et al. 

(2001a), a similarity matrix was assembled for 

the species present in a total of seven vegetation 

types. These included five vegetation types from 

the Waini peninsula: Beach, Coastal Mangrove 

forest, Mixed Freshwater forest, Riverine 

Mangrove forest, and Burned areas; and the two 

vegetation types drawn from van Andel’s 

(2000a) ecological plots in the coastal plain of 

the Northwest District: Quackal swamp and 

Manicole swamp. Burned areas were included


as a group to explore at what level they might 

bear similarity to any of the other plant 

communities. NTSYSpc (Rohlf 1997) statistical 

software was used for the clustering of those 

plant community data. Clustering was 

performed using simple matching coefficients, 

with the Sequential, Agglomerative, 

Hierarchical, and Nested clustering method 

(SAHN) and the Unweighted Pair-Group 

Method, Arithmetic average (UPGMA), based 

on algorithms from Sneath and Sokal (1973). 

Possible sources of error in comparisons of 

plant communities included misidentification of 

plant specimens, which could change the 

amounts of overlap between communities. 

Misidentification was minimized by 

determination of specimens at major botanical 

institutions that host many specialists and have 

fairly complete, up to date collections for 

reference. Also of concern are incomplete or 

biased sampling, which may include undersampling 

of trees and of plants sterile at the time 

of sampling, and differences in sampling effort 

and scale. Problems from assignment of species 

to particular vegetation types must be 

considered. Some of the species may be present 

in the beach community, since they are found 

in beach substrate, but are primarily on the 

margins of a neighboring community and might 

be best treated as outliers from that community. 

As with many of these vegetation groups, the 

beach community itself could be subdivided into 

finer units such as ocean front, central beach 

ridge, and back beach near mangrove swamps; 

similar concerns could be formed for a unit 

regarding boundaries. A final consideration is 

sampling bias, particularly the Quackal and 

Manicole samples which were obtained from 

one hectare plots, while the Waini samples were 

taken during sampling transects. However, these 

do represent a similar degree of sampling 

intensity for each vegetation type, and 

substantial possible disparity may be mitigated 

by the use of qualitative (presence/absence) 

rather than quantitative comparisons. 

Northwest District 

A checklist of plants of the Northwest 

District was assembled from five sources: 1) 

collections made on the Waini Peninsula during 

this study, 2) collections made in the Northwest 

37 

District by the BDG program since 1986, 3) the 

dataset of Northwest District collections held 

at the US herbarium, 4) the 625 species listed 

by van Andel (2000b) based on her collections 

made in Guyana from 1995 to1997, and 5) some 

specimen information from online databases, 

e.g., the Missouri Botanical Garden’s (1995present) 

TROPICOS database, and the New 

York Botanical Garden’s website (1996present). 

The resulting names at the species level 

were matched with the database for the 

Checklist of the Plants of the Guiana Shield 

(Hollowell et al. 2001) to standardize 

nomenclature and correct errors. Those species 

known only from cultivation were not used in 

analyses but were included in the full checklist 

for the Northwest District (Appendix 3), and 

noted as cultivated. The status of species as 

cultivated was determined by literature checks 

in DeFilipps (1992), Boggan et al. (1997), van 

Andel (2000b), and Hollowell et al. (2001). 

Species that had apparently escaped cultivation 

and become naturalized were also noted. Taxa 

determined only to genus were included if they 

represented the only record for that genus. 

Contributions of recent collections to the lists 

were calculated as an indication of the status of 

knowledge of the region’s flora. 

Figure 2.3. Nested floristic distribution zones used 

in analysis of plant species affinities for the Waini 

Peninsula. 1. Guiana Shield. 2. Northern South 

America. 3. Neotropical. 3a. Neotropical including 

Caribbean. 4. Neotropical and Western Africa. 5. 

South America (not indicated). 6. Western 

Hemisphere. 7. Pantropical, between 23.5° N and 

23.5° S. 8. Cosmopolitan (not indicated). The Waini 

Peninsula flora had the greatest affinity (33%) with 

the Neotropical + Caribbean zone, followed by the 

Pantropical zone (27%).

38 


Delta Amacuro 

Comparisons with the flora of Delta 

Amacuro were made with species published in 

the Flora of the Venezuelan Guayana 

(Steyermark et al. 1995 - 2005), which included 

all pteridophytes (ferns) and spermatophyte 

families (seed bearing plants). Those were 

compared with species for the Northwest 

District of Guyana compiled to form the 

preliminary checklist given in Appendix 3, to 

estimate the amount of overlap between the 

floras of the two regions. Since the neighboring 

political divisions share similar geological and 

elevational properties, those species not 

included in the overlaps also suggest 

distributional records that might be found if 

collection efforts were continued. At the US 

National Herbarium, ongoing data collection of 

plant specimens collected in the Venezuelan 

Table 2.2. Plant collectors in Delta Amacuro, Venezuela 

Collector 

Rusby, H.H. and R.W. Squires 

Bond, F.E., T.S. Gillin and S. Brown 

Curran, H.M. and M. Haman 

Cardona, F. 

Gines, H. 

Wurdack, J.J. and J.V. Monachino 

Steyermark, J.A. 

Marcano-Berti, L. (with Zapata and 

Salcedo in 1977) 

Breteler, F.J. 

Blanco, C. 

De Bruijn, J. 

Ruiz-Teran, L. and S. Lopez- 

Palacios 

Davidse, G. and A.C. Gonzalez 

Aymard, G. 

Fernández, A. 

Montoya, S. 

Diaz, W. 

Year 

1896 

1911 

1917 

1943 

1952, 

1954 

1955 

1960, 

1964, 

1977 

1964, 

1965, 

1977 

1966 

1965, 

1966 

1967 

1973 

1979 

1987 

1987 

1993 

1997 

Guayana, including Delta Amacuro, allowed an 

updated summary of historical collecting efforts 

in Delta Amacuro. Those are listed in Table 2.2. 

Although they were listed in an earlier 

compilation by Huber (1995b), R. Liesner and 

F. Delascio are not included because these 

researcher’s names have been found on Delta 

Amacuro specimens only as secondary 

collectors with J. Steyermark in 1977. 

Regional and Global Affinities 

To analyze the Waini flora, all species were 

assigned to one of eight generalized, nested 

classes of biogeographic affinity: Guiana Shield, 

Northern South America, Neotropical (with a 

distinction between those included in or outside 

of the Antilles), Trans-Atlantic, Western 

Hemisphere, Pantropical, and Cosmopolitan. 

Those correspond to a subset of the classes 

Localities 

Paloma, Manoa, Sacupana, Santa Catalina, Eleanor 

Creek 

CaZos of the Orinoco Delta 

CaZo Paijana, Isla de San Carlos, CaZo Pedernales 

Puerto Caropita 

Guayo, Isla Burojoida, Jotacuay, Teiua, Curiapo, 

Ibaruma, Los Piedras, Araguabisi, Puerto Baja, 

Koboima, Tobejuba, Wiuiquina 

CaZo Jobure, Río Grande del Orinoco, Río 

Cuyubini, Río Acure, Río Ibaruma, Serrania 

Imataca, Río Guanamo 

San Victor, Río Amacura, Río Cuyubini, Sierra 

Imataca, Cerro La Paloma, Río Acure, Río Grande, 

CaZo Atoiba, CaZo Jotajana, CaZo Joba-Suburu 

(CaZo Jota-Sabuca?). 

East of Río Grande near border of Bolívar, Cano 

Araguao, Cano Arature, CaZo Guiniquina 

Río Grande o Toro, near Bolívar border 

Río Grande, near Bolívar border 

Río Grande o Toro, near Bolívar border 

La Ladera, CaZo Mánamo, Pedernales 

ESE of Los Castillos de Guayana, trail to the Rio 

San José 

Río Grande, CaZo Orocoima 

Antonio Diaz 

Cerro CaZo Acoima 

Mun. Tucupita Piacoa


employed by Kelloff and Funk (2004). The 

classifications are illustrated in the map in 

Figure 2.3. 

The Guiana Shield distribution (1) includes 

species that are, fairly strictly, restricted to that 

geologic formation’s igneous and sedimentary 

bedrock and areas influenced by that area’s 

outwash. It includes southern Venezuela, most 

of the three Guianas, and parts of northern Brazil 

and scattered outliers in southeastern Colombia. 

The Northern South America distribution 

(2) includes species found in the Guiana Shield 

as well as the Northern Andes, the continent’s 

Pacific coast south to the arid Peruvian coast, 

and much of the Amazon Basin lowlands to 

about 7 degrees south. The Andes reach much 

higher elevations than the Guiana Shield 

highlands. Many of the habitats in this zone are 

wet to extremely wet. 

The Neotropical distribution (3) includes 

all of the Americas between approximately 23.5 

degrees north and 23.5 degrees south, with 

subclasses either including or excluding 

distribution in the Antilles (3a). Following 

climate patterns, this unit is mapped to the south 

of the Tropic of Capricorn on the Atlantic coast 

of South America, and to the north of that 

latitude on the Pacific coast, which is influenced 

by strong, cool ocean currents from the south. 

The Trans-Atlantic distribution (4) 

accommodates species native to both the 

Neotropics and tropical Africa. Here this 

distribution is applied predominantly to 

mangrove species, thus the mapping in Figure 

2.3 is adapted from the Atlantic-east Pacific 

distribution from mangroves by Tomlinson 

(1986) 

The South American distribution (5) is 

essentially self-explanatory, and is indicated on 

Figure 2.3 only by the landmass of the continent. 

The Western Hemisphere distribution (6) 

includes species adapted to a broad climate 

range. It includes species that extend into the 

subtropics and temperate zones of North and 

South America. 

The Pantropical distribution (7) covers 

tropical areas nearly worldwide, from 23.5 

degrees north to 23.5 degrees south. 

Cosmopolitan (8) species are those known 

from both tropical, sub-tropical, and in some 

cases temperate areas, nearly worldwide. Here 

39 

the Neotropical and Pantropical classifications 

both include species that might to some degree 

extend into the Subtropics. 

RESULTS 

Quackal and Manicole Communities 

Forty-four species were shared between the 

126 Waini species recorded and the list of all 

species recorded by van Andel for the entire 

District. That 35% overlap includes 5 species 

that are either cultivated or escaped from 

cultivation, Cocos nucifera L., Luffa cylindrica 

M.Roem., Pedilanthus tithymaloides (L.)Poit., 

Gossypium barbadense L., and Jatropha 

gossypiifolia L.. 

Only seven taxa were shared between the 

Waini Peninsula (126 species) and van Andel’s 

Quackal and Manicole plots (138 species ). That 

is approximately a 5.5 % overlap relative to the 

species of the Waini Peninsula. Of 240 total 

plant taxa for all sites, only 51 were shared in 

any way among the seven vegetation types. 

Considered separately, collections resulting 

from this research on the Waini Peninsula and 

other sites in the Northwest added 99 species to 

the baseline list formed from historic collections, 

while van Andel (2000a) collections added 308 

species to the baseline list. Thirteen of those 

species were added by both efforts. 

The similarity matrix and tree matrix, 

displayed as a phenogram (similarity-based 

dendrogram), from the UPGMA clustering of 

the seven coastal plain communities are shown 

in Figure 2.4. A cophenetic matrix was formed 

from the UPGMA tree matrix and run against 

the original matrix to test the goodness of fit of 

the clustering results to the original data set, 

using a Mantel test for matrix correspondence 

(Rohlf 1997). That resulted in a high probability 

(p = 0.995) that a random Z-value would be less 

than the Z-value derived from the comparison 

of matrices, indicating a close fit in the UPGMA 

clustering. Alternate tests of single-link and 

complete-link clustering methods yielded 

phenograms very similar to those from UPGMA 

clustering, indicating that the clusters are quite 

distinct (Rohlf 1997). 

Coastal Mangrove and Riverine Mangrove 

swamps clustered together. The two mangrove 

swamp types differed in part because different

40 


Figure 2.4. Similarity matrix and cluster diagram for five major Waini Peninsula plant community types: 

Coastal Mangrove swamp, Riverine Mangrove swamp, Mixed Freshwater swamp, Burned areas, and Beach, 

as well as the Quackal and Manicole communities in southeastern Northwest Region (van Andel 2000a). 

Based on species presence and absence. 

Rhizophora species were present, R. mangle in 

the Coastal swamp and R. racemosa in the 

Riverine swamp; the Riverine Mangrove 

community also contained a few epiphytic 

species while the coastal community contained 

none. Several species were included in Coastal 

but not Riverine Mangroves that were not strict 

halophytes; these were possibly excluded from 

the Riverine areas because they are not adapted 

to tidal fluctuations that are absent in the 

occluded Coastal swamps. Those mangrove 

vegetation types cluster together in part because 

of their low species richness (18 and 8 species 

respectively), although they shared only two 

species, Acrostichum aureum and Avicennia 

germinans. Those two species were also shared 

with the more diverse Mixed Freshwater swamp 

(34 species), with which the mangrove swamps 

clustered. The Mixed Freshwater swamp shared 

four species with Riverine Mangrove and one 

with Coastal Mangrove. 

The Burned areas (39 species) clustered 

with the grouping of the above three 

communities, Coastal and Riverine Mangroves 

and Mixed Freshwater. Five of the ten species 

shared with those communities were remnants 

from mangrove swamps formerly in those 

burned areas, and the other five were colonists, 

including Acrostichum ferns from mangrove 

habitats and vines from Mixed Freshwater areas. 

Unaffected areas of these communities also 

surrounded the burned area and contributed seed 

for regeneration. The distinction of the burned 

areas from the previous three was due in part to 

the presence of a larger number of species, many 

of which are disturbance-adapted. Those include 

wind-dispersed species, members of the 

Amaranthaceae, vines, and floating and aquatic 

species that benefitted from the increases in open 

water. Among the wind dispersed species that 

were not found in the unburned swamps were 

Typha domingensis Pers., several ferns, Mikania 

micrantha Kunth, Cydista aequinoctialis, 

Sarcostemma clausum(Jacq.)Schult., Entada 

polystachya (L.)DC, and Securidaca diversifolia 

(L.)S.F.Blake. Animal dispersed species in the


burned areas included Coussapoa asperifolia 

Trécul, Aeschynomene sensitiva Sw., Heliconia 

psittacorum L.f., Hibiscus bifurcatus Cav., and 

Solanum stramoniifolium Jacq. as well as several 

species of Amaranthaceae with tiny seeds that 

may be wind blown at times. 

The Beach community (38 species) was 

primarily composed of species highly 

specialized for the coastal environment, many 

of which were not found in other vegetation 

zones. That community clustered with the four 

previous groups in part because of the presence 

of individuals of mangrove species and 

mangrove associates that were common in 

neighboring habitats, including Laguncularia 

racemosa, Avicennia germinans and 

Conocarpus erectus. Those occurred 

occasionally at the top of narrow beach ridges 

and often on beach borders of coastal lagoons. 

Some vines of the back beach were shared with 

the burned areas, including Sarcostemma 

clausum, Cissus verticillata, and Ipomoea 

tiliacea (Willd.)Choisy; Vigna luteola 

(Jacq.)Benth. was shared with Mixed 

Freshwater swamp. 

None of the Beach taxa were present in 

either Quackal or Manicole communities. The 

similarity of those two communities with the 

other four Waini Peninsula communities was 

based on only seven shared species. The Quackal 

community as surveyed by van Andel was more 

diverse (70 species) than any one of the Waini 

Peninsula communities, probably reflecting low 

soil salinity because of its distance from the 

ocean (approximately 12 kilometers at the 

Quackal plot). Van Andel considered the 

Quackal vegetation type to grade into mangrove 

swamp; the four species that it shared with the 

Waini were Euterpe oleracea, Clusia palmicida, 

Cassipourea guianensis Aubl., and 

Calyptranthes sp., all shared with the Mixed 

Freshwater swamp community, which may have 

contributed to its clustering more closely than 

the Manicole community with the Waini 

communities. 

The Manicole community was the richest 

of the seven (96 species) and also the most 

distant from coastal communities. The plot near 

Assakata village on the Baramanni River was 

approximately 17 kilometers from the Atlantic 

Ocean. This community had a significant 

41 

overlap with the Quackal site, sharing 28 

species. Of those, 18 are trees; the other shared 

species were divided among shrubs, herbs, and 

vines. The six species that those two sites shared 

with the Waini Peninsula communities were 

Monstera adansonii, Euterpe oleracea, Clusia 

palmicida, Cassipourea guianensis, 

Pterocarpus officinalis, and Peperomia glabella 

(Sw.)A.Dietr.. Although the Manicole 

community shared more species with the Waini 

Peninsula communities than the Quackal 

community shared with the Waini Peninsula, 

those species were drawn from the Manicole 

community’s higher number of species, and the 

shared species were spread among several Waini 

communities. Four species were shared with the 

Mixed Freshwater community, the epiphyte 

Peperomia was shared with the Riverine 

Mangrove area, and the vine Monstera was 

shared with the Coastal Mangrove community. 

Calyptranthes sp. also occurred in Coastal 

Mangrove, Mixed Freshwater, and Quackal 

communities. 

Of the 240 species from all seven sites, 189 

(79%) were known from only one site; those 

were divided among 26 from Burned areas (66% 

of its species), 28 from the Beach community 

( 76% of its species), 8 from Coastal Mangrove 

swamp (42% of its species), 17 from Mixed 

Freshwater swamp (50% of its species), 4 from 

Riverine Mangrove swamp (50% of its species), 

41 from Quackal swamp (59% of its species), 

and 65 from Manicole swamp (68% of its 

species). 


A preliminary list of 1449 vascular plant 

species from the Northwest District is given in 

Appendix 3. Those species were approximately 

22% of the known vascular flora of Guyana, 

currently standing at 6,700 species (Boggan et 

al. 1997; Hollowell et al. 2001). Additionally, 

105 cultivated species, one alga and six mosses 

were included on the list. Mosses and alga are 

poorly collected and analyzed in the Northwest 

District, as throughout the Guiana Shield, and 

are included with the hope of encouraging future 

research. A previous collections-based 

compilation for the Northwest District listed 484 

species of vascular plants (BDG 2001). 

Collections made during these studies on the

42 

4004 

54.10% 

713 

53.09% 

72 

61.54% 


Orchidaceae 

Rubiaceae 

Poaceae 

Fabaceae 

Melastomataceae 

Cyperaceae 

Caesalpiniaceae 

Mimosaceae 

Euphorbiaceae 

Asteraceae 

Myrtaceae 

Clusiaceae 

Apocynaceae 

Araceae 

Chrysobalanaceae 

Piperaceae 

Bromeliaceae 

Bignoniaceae 

Lauraceae 

Annonaceae 

603 

415 

316 

306 

279 

274 

179 

172 

167 

149 

144 

127 

117 

117 

115 

111 

110 

109 

104 

90 

8.15% 

5.61% 

4.27% 

4.13% 

3.77% 

3.70% 

2.42% 

2.32% 

2.26% 

2.01% 

1.95% 

1.72% 

1.58% 

1.58% 

1.55% 

1.50% 

1.49% 

1.47% 

1.41% 

1.22% 

x 

x 

Rubiaceae 

Fabaceae 

Orchidaceae 


Poaceae 

Cyperaceae 

Mimosaceae 

Araceae 

Euphorbiaceae 

Asteraceae 

Apocynaceae 


Arecaceae 

Bignoniaceae 

Clusiaceae 


Malpighiaceae 

Piperaceae 

Annonaceae 

Lauraceae 

78 

62 

57 

51 

51 

40 

37 

33 

33 

31 

27 

26 

25 

25 

25 

24 

24 

24 

20 

20 

5.81% 

4.62% 

4.24% 

3.80% 

3.80% 

2.98% 

2.76% 

2.46% 

2.46% 

2.31% 

2.01% 

1.94% 

1.86% 

1.86% 

1.86% 

1.79% 

1.79% 

1.79% 

1.49% 

1.49% 

x 

x 

x 

x 

x 

x 

x 

x 

x 

Fabaceae 

Araceae 

Convolvulaceae 

Cyperaceae 

Amaranthaceae 

Arecaceae 

Asteraceae 

Malvaceae 

Orchidaceae 

Poaceae 

Apocynaceae 

Bromeliaceae 

Combretaceae 

Mimosaceae 

Moraceae 

Pteridaceae 

Rhizophoraceae 

Cactaceae 

Euphorbiaceae 

Myrtaceae 

6 

5 

5 

5 

4 

4 

4 

4 

4 

4 

3 

3 

3 

3 

3 

3 

3 

2 

2 

2 

5.13% 

4.27% 

4.27% 

4.27% 

3.42% 

3.42% 

3.42% 

3.42% 

3.42% 

3.42% 

2.56% 

2.56% 

2.56% 

2.56% 

2.56% 

2.56% 

2.56% 

1.71% 

1.71% 

1.71% 

x 

spp. Prop. Guy&Waini 

spp. Prop. NW&Guy 

spp. Prop. Waini&NW 

GUYANA NORTHWEST WAINI 

Table 2.3. Rankings of the 20 most species rich families for the Northwest, all Guyana, and the Waini Peninsula. The nine families shared by all are 

listed in bold face type, and those families shared at two indicated in columns with an “x”.


Waini Peninsula and during incidental field trips 

farther inland in the Northwest were the only 

source for 90 species in the list. Of those, 53, 

approaching half of the Waini flora, were known 

only from Waini Peninsula collections, and those 

taxa are indicated in the list in Appendix 3. 

To illustrate the relative composition of the 

regional flora at varying scales, Table 2.3 ranks 

the 20 most species-rich families for each of the 

three areas: the Northwest District, all Guyana, 

and the Waini Peninsula. The overall highest 

ranking family among all sites was Fabaceae 

(Papilionoid legumes), which ranked fourth in 

all Guyana, second in the Northwest, and first 

in the Waini flora. Nine families were shared in 

the top 20 families among all three areas: 

Fabaceae, Orchidaceae, Poaceae, Cyperaceae, 

Mimosaceae, Araceae, Euphorbiaceae, 

Asteraceae (Compositae) and Apocynaceae. Of 

the top 20 families, 18 were shared between the 

rankings of the Northwest District and all 

Guyana, while 11 of the top 20 families were 

shared between the Waini and all Guyana. The 

Arecaceae (Palmae), which ranked 14 th in the 

Northwest were 23 rd in the full Guyana rankings; 

the Malpighiaceae, which were ranked 19 th in 

the Northwest were 21 st in the Guyana rankings. 

Table 2.4. Plant species collected on the Waini 

peninsula not listed for Delta Amacuro. One variety 

is also listed. Ten of these are primarily coastal, 

indicated by *. 

Amaranthaceae Alternanthera sessilis 

Arecaceae Nypa fruticans * 

Aristolochiaceae Aristolochia trilobata 

Caricaceae Carica papaya 

Combretaceae Conocarpus erectus * 

Combretaceae Terminalia catappa * 

Convolvulaceae Ipomoea violacea * 

Cuscutaceae Cuscuta umbellata 

Fabaceae-Caesal. Caesalpinia bonduc * 

Fabaceae-Mimos. Entada polystachya 

Fabaceae-Papil. Sesbania sericea * 

Lygodiaceae Lygodium venustum 

Malvaceae Thespesia populnea * 

Myrtaceae Psidium guajava 

Orchidaceae Epidendrum ciliare 

Passifloraceae Passiflora foetida var. foetida 

Poaceae Paspalum distichum 

Poaceae Sporobolus virginicus * 

Pteridaceae Acrostichum danaeifolium * 

Rubiaceae Morinda citrifolia * 

43 

Delta Amacuro 

The Flora of the Venezuelan Guayana 

(Steyermark et al. 1995 - 2005) listed 1649 

species of pteridophytes and spermatophytes for 

the state of Delta Amacuro. Among those species 

193 (11.7%) were not listed for Guyana. Given 

the proximity and environmental similarities, a 

portion of those species might be expected to 

be found in the Guyana’s Northwest District 

with additional collecting effort. 

Also, several species were collected on the 

Waini Peninsula and in other parts of the 

Northwest District that were not listed for Delta 

Amacuro in the Flora of the Venezuelan 

Guayana. Table 2.4 lists 19 plant species and 

one variety that were collected in the Waini 

Peninsula vicinity that were not recorded for 

Delta Amacuro. In addition, the alga 

Rhizoclonium africanum apparently had not 

been documented for Delta Amacuro, although 

it is almost surely present in the coastal 

Avicennia swamps. 

Appendix 4 lists plant species from the 

Northwest District that were not yet recorded 

from Delta Amacuro. Sources include records 

from this study’s ancillary trips beyond the 

Waini Peninsula site to interior localities near 

Mabaruma and Kwebana (see Figure 2.1), as 

well as from collection-based literature (van 

Andel 2000b) and from specimens at the US 

National Herbarium. That list includes 517 

species of vascular plants from the Northwest, 

without cultivated or naturalized species, more 

than 31% of the 1649 species currently known 

for that state. 


The strongest affinity of the Waini 

Peninsula flora was with the Neotropical 

distribution category including the Antilles 

(zones 3 + 3a in Figure 2.3), which accounted 

for approximately 33% of the Waini species. The 

next strongest affinity was with the Pantropical 

distribution category (zone 7), including 

approximately 27% of the species. The 

Neotropical - African, or transatlantic, 

distribution (zone 4) comprised about 10% of 

the species, including all mangrove and 

mangrove-associate taxa, being those in the 

genera Avicennia, Laguncularia, Rhizophora, 

and Conocarpus. Notably, only 1.6% of the

44 


Table 2.5. Floral affinities of Waini Peninsula distribution categories, results, and comparison to the floral 

affinities in those categories for the Kaieteur Falls area (Kelloff and Funk 2004). 

Zone 

3a 

7 

4 

3 

2 

8 

5 

6 

1 

Distribution 

Neotropical (with Antilles) 

Pantropical 

Neotropical + Africa 

Neotropical (excluding Antilles) 

Northern South America 

Cosmopolitan 

South America 

Western Hemisphere 

Guiana Shield 

TOTAL 

Waini species were found to have a distribution 

considered as restricted to the Guiana Shield 

region (zone 1). The totals for each distribution 

category are given in Table 2.5, by their rank in 

the Waini Peninsula flora, along with the 

corresponding proportion from Kelloff and 

Funk’s (2004) analysis of Kaieteur National 

Park. 

DISCUSSION 

WainiSpecies 

42 

35 

13 

11 

8 

8 

5 

4 

2 

128 

Quackal and Manicole Communities 

The turnover of species, moving from the 

coastal communities inland into the Quackal and 

Manicole communities, showed a high level of 

beta diversity, the change in species composition 

across intermediate distances. That illustrated 

the contribution of the coastal plain flora to both 

species and landscape level diversity of Guyana. 

In the case of the Waini Peninsula, the species 

turnover is likely driven in part by changes in 

oceanic influence, particularly salinity, as well 

as by the results of ocean dispersal along the 

coast and perhaps increased precipitation inland 

from the ocean. Since a high proportion of 

species were found to be unique to each 

community, it might be expected that this trend 

continues in the inland direction. The 

implication of that for conservation is the 

liklihood that coastal plain diversity would be 

difficult to adequately represent within any 

small area. Rather, significantly broad transects 

of contiguous or intermittent sites along the 

WainiProportion 

32.8% 

27.3% 

10.2% 

8.6% 

6.3% 

6.3% 

3.9% 

3.1% 

1.6% 

100% 

KaieteurProportion 

14.8% 

1.8% 

1.4% 

11.6% 

26.1% 

0.4% 

0.2% 

0.4% 

17.2% 

73.9% 

gradient from ocean to highlands might be more 

effective as reserves. 

The Manicole site shared more species, and 

a slightly higher proportion of its species, with 

the Waini flora than the Quackal site that 

actually clustered closer to the Waini 

communities. That illustrated how differences 

in species richness of the sites might play as 

significant a role in the outcome of the clustering 

method as the number of shared species. 

The Mixed Freshwater swamp on the Waini 

Peninsula, some parts of which were dominated 

by Euterpe (Manicole) palms prior to the fires 

of 1998, may have held more species in common 

with van Andel’s Manicole or Quackal sites 

before that disturbance. The Mixed Freshwater 

swamp was a distinct entity, containing elements 

characteristic of both types of mangrove 

communities, and it also possessed many unique 

species. Those were apparently plant species 

that could adequately tolerate the low to 

moderate salinity of the interior, although not 

higher salinities of the mangrove swamps. The 

Beach community was distinct from the other 

Waini communities because of the presence of 

coastal specialist species. The beach clustered 

with those communities in part because they 

shared species, and because of the lack of 

overlap with species of the more inland 

Manicole and Quackal communities. In the 

Burned areas, the increased species richness will 

probably be a short-term phenomenon. Several 

species that were present before the fires 

persisted or were quickly reestablished, while


only a few of the most sensitive species were 

eliminated. The dominance of most of the 

disturbance-adapted species will likely be 

temporary, while others, such as Typha, may 

persist for long periods. 


The family rankings illustrate moderate 

consistency at higher taxonomic levels between 

the countrywide flora and those of the Northwest 

District and the Waini Peninsula. Of families 

listed only in the Waini top ranking, most have 

a high component of wetland-adapted species, 

such as Malvaceae, Combretaceae, Moraceae, 

Rhizophoraceae, Arecaceae. Also represented 

for the Waini were disturbance adapted families 

such as Amaranthaceae, and Pteridaceae, and a 

family dominated by vines, the Convolvulaceae. 

The number of species records contributed 

to the list for the Northwest District by recent 

collection efforts illustrates the incomplete state 

of investigations into the region’s flora. Limited 

financial and professional resources require that 

collecting efforts be aimed at the localities most 

promising for shedding light on biodiversity and 

conservation questions. Associated with this, 

there is a great need to make additional historical 

collection data available for such analyses. 

Preparing additional museum specimen data is 

labor intensive but is cost effective in 

comparison to performing new fieldwork, and 

will provide increasingly useful data to answer 

biodiversity questions and guide future research. 

As biodiversity research efforts progress in 

Guyana, compilation of species distributions 

will likely become increasingly focused on the 

finer levels of detail, such as by political 

divisions within countries. The tendency to use 

political divisions should be tempered by the 

growing ability to analyze a growing body of 

georeferenced specimen data using climatic, 

geological, elevational, and distance data with 

geographic information systems, allowing 

questions to be more readily framed and 

answered in terms of environmental and 

ecological units. 

Delta Amacuro 

It is still not certain how much overlap 

should be expected for plant species of the Waini 

Peninsula and in Delta Amacuro. Collecting 

45 

expeditions in almost any under-collected 

locality of the Guiana Shield region tend to yield 

new distributional records. Few botanical 

collections in Delta Amacuro have taken place 

along the coast near to Guyana; the closest 

known coastal collection locality is Punta 

Barima, more than 70 kilometers northwest 

along the Atlantic coast from the Waini 

Peninsula. Although both areas have similar 

coastal and estuarine environments, it was found 

that over 30% of the plants listed for the 

Northwest District are not known for Delta 

Amacuro. This difference can be compared to 

the same statistic for nearby areas. Twenty-one 

percent of Surinam’s known vascular plant 

species are not recorded for French Guiana, and 

18% are not recorded for Guyana. However, in 

comparison, for the diverse, mountainous 

Venezuelan segment of the Guiana Shield, 42% 

of the plant species listed for Bolívar state are 

currently not listed for Amazonas state 

(Steyermark et al. 1995 - 2005). Latitudinal and 

edaphic differences between the Northwest and 

Delta Amacuro, particularly the extensive 

deltaic environment found only in Delta 

Amacuro, in addition to the high overall level 

of endemicity in the Guiana Shield region, 

suggest that a sizable portion of the plant species 

listed in only one region will continue to be 

unnknown in the other. Still, only the assembly 

and analysis of additional collections can clarify 

the relationship between the two areas. 

These comparisons highlight the 

incomplete state of botanical knowledge of these 

regions, and should serve as incentives and goals 

for plant specimen collecting efforts. They will 

also hopefully serve as encouragement for 

further analyses of the spatial patterns of the 

turnover of species in the tropics. 


The differences between affinities of the 

Waini Peninsula flora and those of the species 

of the interior Kaieteur location illustrated some 

interesting points. The coastal Waini 

communities included many species with broad 

ranges, with the four top categories being 

Neotropical and Pantropical distributions and 

the sixth ranked category being Cosmopolitan, 

which included a variety of herbs. The fifthranked 

category, Northern South America, and

46 


the last category, Guiana Shield, represented the 

two most species-rich groups for the Kaieteur 

area. The Neotropical distribution ranked fairly 

high at both sites, with the distribution including 

the Antilles being the highest ranked Waini 

group and the third ranked group of Kaieteur; 

the non-Antillean Neotropical species ranked 

fourth at both sites. The Neotropical affinities 

of the Waini Peninsula flora were strongly 

influenced by species with island distributions, 

which reflects its close association with the 

Atlantic Ocean. 

The minimal presence within the Waini 

flora of species endemic to the Guiana Shield 

supports the idea that the Waini Peninsula 

should be treated as floristically distinct from 

the Shield, reflecting the effect of the Amazonderived 

sediments that have buried the Shield’s 

crystalline bedrock, often by hundreds of 

meters. The Atlantic Ocean is a dispersal vector 

that supplies propagules of species well adapted 

for the saline soils, and has an overwhelming 

influence on the flora. The Waini is actually 

fairly near to the geologic Shield, being less than 

15 kilometers from the hills near the town of 

Mabaruma. 

The lowlands of the Waini Peninsula, the 

Guyana coastal plain, and the Orinoco Delta 

section of Delta Amacuro should, in the strict 

sense, be excluded from definitions of the 

Guiana Shield in accounts of its flora and fauna 

as well as for conservation purposes. That does 

not reduce the importance of coastal habitats 

such as found on the Waini Peninsula. Rather, 

it should be a confirmation of that coastal area’s 

distinctness on the regional level. Not only are 

those plant communities limited to a relatively 

narrow strip of coastal plain, bounded by ocean 

and the Guiana Shield, but their community 

structures and settings are different from the 

Antillean coastal plant communities because of 

their geomorphic position on the Amazon 

River’s coastal delta, which is unique 

worldwide. 

CONCLUSION 

In the larger scope of Guyana’s 

biodiversity, the Waini flora is not rich in rare 

endemic species from spectacular 

environments, such as are found on the isolated 

tepuis of the Guiana Shield. However, the 

coastal zone is a unique and limited unit of the 

landscapes of Guyana and northern South 

America. The peninsula is threatened in more 

ways than less accessible interior ecosystems, 

both from local sources from accidental fires, 

on the national level by unplanned settlement 

or unsustainable exploitation, and at the global 

level by climate changes that might increase the 

risk of fires and threaten this low-lying peninsula 

through sea-level rise. 

Comparisons with other ecosystems in 

Guyana help to illustrate the high levels of beta 

diversity across the landscape in the low 

elevations of the coastal plain and highlight the 

need to gather more information from all 

ecosystem types in Guyana and neighboring 

Venezuela. The number of species records added 

for the Northwest by collections on the Waini 

alone highlights the incompleteness of 

knowledge of the flora of that region, and the 

number of species not recorded for Delta 

Amacuro shows a similar need for that 

Venezuelan state. As the level of botanical 

knowledge for a country grows, research moves 

towards compiling species distributions at 

regional levels, and analyses of species diversity 

and planning of collecting activities will become 

increasingly focused on gaps in regional 

knowledge. 

In the future, additional collections from 

many herbaria should be entered into databases 

and standardized to improve the completeness 

of checklists and biodiversity analyses. With the 

tools of georeferenced data sets and GIS 

software, analyses may eventually be possible 

that utilize collection data in ways that link to 

and compliment both qualitative, spatially broad 

regional checklists and highly quantitative, 

spatially limited ecological plot studies. 

Through GIS analyses of adequate, high quality 

collections data, expeditions can be planned that 

effectively target under-sampled habitats in 

important areas (Funk et al. 2005). 

Results here suggesting that the Waini flora 

is essentially separate from the Guiana Shield 

flora and bears more affinity to the Caribbean 

region are not surprising, but lends support to 

the idea that biodiversity studies and 

conservation efforts should be pursued with an


awareness of that distinction. The coastal plain 

should be excluded from the Guiana Shield 

when considered in any strict sense. In 

consideration of that, the coastal plain remains 

a unique ecosystem in Guyana that has suffered 

a high rate of disturbance and merits its share 

of study and protection. 

This analysis of the plants of the Waini 

Peninsula should help to stimulate interest in 

the Waini Peninsula’s inclusion in Guyana’s 

fledgling Protected Areas System (EPA Guyana 

2004; Kelloff 2003). Protected areas always 

benefit from the availability of plentiful 

information on the diversity of habitats and 

makeup of communities. Baseline information 

may be built upon and used for comparison with 

future surveys to increase understanding and 

benefit management in the face of rising 

population and climate change impacts. 

Documentation of local flora and fauna also 

47 

increases the value of protected areas for 

educational purposes. On the Waini Peninsula, 

in large part because of sea turtle conservation 

activities, there has been a steady flow of local 

Amerindian students visiting Almond Beach. 

Students have been encouraged to take an 

interest in conservation issues and benefit from 

all additional information on the area. 

While botanical diversity of the Waini 

Peninsula is modest in comparison to several 

other parts of Guyana, the area adds significantly 

to the diversity of landscapes found in the 

country. Similar coastal areas were long ago 

permanently converted to farm and urban land. 

The landscape also has substantial cultural 

significance, as the large majority of the 

population of Guyana has always lived in the 

country’s coastal zone, and the Waini Peninsula 

is a prime example of that ecosystem type.

48 


Doekie (Jackie) Arjoon and Chris Chin of the 

University of Guyana, in unburned Avicennia 

mangrove swamp near Waini Point, during 

Whittaker plot establishment in November 

1998. 

Keith David and Karen Redden on the prop roots 

of a Rhizophora mangle tree near the edge of 

the Kamwatta burn. A Bromelia plumieri plant 

(collection THH 413) is visible behind them. 

Keith David of the University of Guyana, in the 

unburned Avicennia mangrove swamps near 

Waini Point, in April 2001. Keith also collected 

Lepidoptera specimens during this trip to the 

Waini Peninsula. 

Karen Redden with the fruit of Entada 

polystachya (Fabaceae, collection THH 423) 

found along the central, vine covered portion 

of transect “C”, in early 2001.


CHAPTER 3. 

THE MANGROVE PALM NYPA FRUTICANS WURMB.: 

A WIDESPREAD EXOTIC SPECIES 

IN NORTHWESTERN GUYANA 

INTRODUCTION 

Among the plant species present on the 

Waini Peninsula, Nypa fruticans, the Asian 

mangrove palm is of particular interest. One of 

the most pressing current issues in 

environmental conservation is detection, 

documentation, and management of invasive 

exotic species. That concern is exemplified by 

the widespread presence of Nypa in the 

Northwest District of Guyana. The growing 

appreciation of the ecological, social, and 

economic benefits of mangrove ecosystems has 

focused attention on potential threats, in terms 

of both the scientific and environmental policy. 

The goals of this chapter is to more fully 

document the presence of Nypa in South 

America, review information that might be 

relevant to understanding its arrival in the 

Northwest District of Guyana, and examine its 

possible sources and potential for spread within 

the region, as a basis for future research and 

appropriate management of the region’s 

Riverine Mangroves. 

NYPA HISTORY AND RANGE 

Nypa fruticans Wurmb. is the only species 

in the monotypic genus Nypa Steck. It is native 

to the Indo-West Pacific region, thriving in high 

sediment estuaries from the Philippines to the 

Malay Peninsula, the Ganges Delta, Sri Lanka, 

and Northern Australia (Tomlinson 1986). 

Nypa’s trunk combines dichotomous branching 

and rhizomatous habit, and grows beneath mud, 

features unique among both mangroves and 

palms (Tomlinson 1973, 1986; Uhl & 

Dransfield 1987). It prefers habitats with 

moderate salinities and low wave energy. Those 

characteristics allow Nypa to form dense, 

monospecific patches in the intertidal zone 

along tropical estuaries. 

49 

Nypa is an ancient genus. Through analyses 

of fossil fruit and pollen evidence it has been 

documented as one of the earliest known 

mangrove genera, occurring in the Late 

Cretaceous period (about 70 mya) (Duke et al. 

1997; Ellison et al. 1999; Gee 1989), and pollen 

representing Nypa is known from northern South 

America at that time (Graham 1995). Nypa was 

the dominant mangrove in the Neotropics, 

including the western Venezuelan coast, in the 

Middle Eocene period (45 mya) (Rull 1998, 

2001) and was present in South America from 

the Maastrichtian stage of the late Cretaceous 

through the Eocene period (65-34 mya) (Gee 

2001). It later disappeared from the Neotropics 

sometime after the Late Eocene (40 mya) (Gee 

2001; Graham 1995; Tralau 1964), and has been 

restricted to Southeastern Asia since the 

Miocene epoch (20 mya) (Ellison et al. 1999). 

The ancient distributions of Nypa have been 

used as an illustration of the principle of 

continental drift (Raven & Axelrod 1975), and 

Nypa pollen is one of the earliest identifiable 

examples of a likely extant angiosperm species 

(Tomlinson 1986). 

Throughout its modern natural range in Asia 

Nypa is heavily utilized. Descriptions of the 

species and discussions of its utility have been 

published for nearly a century (Conrado & Ayala 

1906; Fong 1992; Halos 1981; Hamilton & 

Murphy 1988; Miah et al. 2003; Päivöke et al. 

1984). It is particularly valued for its leaves 

which are used as thatching material (FAO 1994; 

Miah et al. 2003) and is also used for production 

of alcohol and vinegar derived from sap 

collected from the cut peduncle (Melana 1980; 

Miah et al. 2003; Päivöke et al. 1984). The 

hardened endosperm of Nypa has been used as 

a “vegetable ivory” similar to that of the Tagua 

Palm, Phytelephas Ruiz & Pav., of Central 

America and western South America, which in 

the past has been considered possibly Nypa’s 

closest relative within the Arecaceae. However

50 


that relationship has been refuted by recent 

molecular studies that place Nypa as a basal 

group sister to all other palms (Lewis & Doyle 

2001) or basal if excluding the subfamily 

Calamoideae (Asmussen & Chase 2001), which 

includes the Moriche or Ité Palm Mauritia, a 

distinctive feature of the vegetation of the 

Venezuelan Llanos and the coastal plain swamps 

of Guyana. 

NYPA FRUTICANS OUTSIDE 

OF ITS MODERN NATIVE RANGE 

Nypa fruticans has become a widespread 

exotic species in some estuaries of Western 

Africa, where it was introduced in the early 20 th 

century, particularly in the Niger Delta 

(Sunderland & Morakinyo 2002; Ukpong 1995). 

Duke (1991) first documented Nypa in the 

modern Neotropics from a small population at 

the mouth of Panama’s Rio Majugual, in the 

Figure 3.1. Map of northwestern Guyana, showing 

the known range of Nypa, from personal observations 

during river travel. The Guiana current flows in the 

direction of the Orinoco Delta and Trinidad. 

vicinity of the Caribbean seaport Colón, near 

the northern, Caribbean end of the Panama 

Canal. More recently, viable Nypa fruits have 

been reported from beaches of Trinidad by 

Bacon (2001), who successfully germinated 

some of these stranded fruits in the laboratory. 

An initial account of Nypa populations along 

the rivers of the Northwest District of Guyana 

was given by Pritchard (1993), whose published 

segment on Nypa went largely unnoted within 

his general review of palms in Guyana. That 

knowledge of Nypa’s presence in Guyana has 

not been applied to subsequent discussions of 

Nypa in the Neotropics. In light of the increasing 

attention to the presence of Nypa outside of its 

native range and of its environmental 

implications, there is a need for further 

description and documentation of the 

populations established in Guyana. 

Nypa’s presence in Guyana also has bearing 

on recent speculation about the origin of Nypa 

propagules found stranded on beaches in 

Trinidad (Bacon 2001). 

From 1997 to 2001, during investigations 

of coastal Avicennia germinans swamps in 

Northwestern Guyana, Nypa palms were 

observed along the banks of the Waini River as 

far as 42 kilometers upstream from the river’s 

mouth. These colonies are thickest over most 

of the length of the Mora Passage, a connector 

between the Waini and the Barima Rivers. Nypa 

also grows along the Barima River from the 

Mora Passage to the Aruka River, and on the 

Aruka River as far upstream as the district 

administrative seat, Mabaruma. It is very likely 

that Nypa is present in Venezuela downstream 

along the Barima River as it flows towards the 

southern Orinoco Delta. Smaller colonies of 

Nypa, mentioned by Pritchard (1993), still exist 

on the Pomeroon River upstream nearly to the 

town of Charity. The currently observed range 

of Nypa in Guyana is shown in Figure 3.1. 

Even the densest areas of Nypa along the 

Mora Passage (Figure 3.2) are mixed with the 

riverine red mangrove Rhizophora racemosa G. 

Figure 3.2. Nypa growing along the banks of the Mora 

Passage, interspersed with the prop-rooted riverine 

red mangrove Rhizophora racemosa.


Mey., although Nypa sometimes dominates the 

river’s fringe. Nypa is often scattered in the 

understory of the intertidal Rhizophora 

racemosa swamp. Nypa infructescences up to 

25 cm in diameter, composed of approximately 

Figure 3.3. A Nypa infructescence collected as part 

of Hollowell et al. #213, from along the banks of the 

Mora Passage. 

70 fruits (Figure 3.3), are common in these 

populations. When mature, the spherical heads 

break up, and the viviparous fruits disperse with 

the currents. As part of recent fieldwork in the 

region a voucher collection of Nypa fruticans 

Figure 3.4. Label information for the voucher 

collection of Nypa fruticans in the Northwest District 

of Guyana, as distributed with specimens. 

was made along the Mora Passage (Figure 3.4). 

Along the ocean beaches of the Waini 

Peninsula, germinated propagules are 

occasionally found stranded (Figure 3.5), 

similarly to those documented recently in 

Trinidad by Bacon (2001). Established Nypa 

plants have never been observed during many 

walks and boat trips along this part of Guyana’s 

51 

Atlantic coast. The entire coast of the Guianas 

is bordered by extensive mudflats, derived 

primarily from Amazon River sediments 

(Brinkman & Pons 1968; Gibbs & Barron 1993). 

This coastal habitat is apparently inhospitable 

to Nypa, possibly due to wave energy and the 

shifting nature of the mudflats. Neither has Nypa 

been seen in the Avicennia dominated mangrove 

swamps immediately behind the low coastal 

beach ridges along the Atlantic coast, and it is 

uncertain whether that habitat would be 

amenable to Nypa. 

DISPERSAL OF NYPA TO 

TRINIDAD 

Bacon (2001) suggested the possibility that 

Nypa fruticans propagules found stranded on 

beaches in Trinidad originated from known 

populations in West Africa and were dispersed 

by prevailing currents across the Atlantic ocean 

and north along the eastern coast of South 

America. In one day, along a 500 meter stretch 

of Manzanilla Beach in eastern Trinidad, Bacon 

and students collected a total of 53 Nypa 

propagules, including 12 viable propagules 

(Bacon 2001), which would be extraordinary if 

attributable to trans-Atlantic dispersal. 

Considering the trans-Atlantic distribution of 

several mangrove taxa, successful dispersal and 

establishment over these distances might occur 

rarely. However, the Atlantic Ocean is a 

formidable barrier (Duke et al. 2001), and 

Figure 3.5. A viable Nypa propagule found stranded 

on the beach of the Waini Peninsula, approximately 

5 km east of Waini Point. The beach is composed of 

small shell fragments; off the beach mudflats extend 

seaward for several kilometers.

52 


dispersal even within mangrove biogeographic 

regions can be a limiting factor on distributions 

of some species (Duke et al. 1998). A high 

degree of separation of other trans-Atlantic 

mangrove species has also been supported by 

differences in the composition of foliar waxes 

between separated populations (Rafii et al. 

1996). Trans-Atlantic dispersal is a 

consideration in the study of the limited number 

of plant genera and species with disjunctions; 

such dispersal events occur at long intervals, in 

the range required for speciation (Renner 2004). 

The Nypa populations in Guyana provide 

plentiful propagule sources within effective 

dispersal range of Trinidad. The distance to be 

covered is approximately 260 kilometers 

directly along the prevailing Guiana Current 

from the mouth of the Waini River to Manzanilla 

Beach in Trinidad. Surface velocities of the 

Guiana Current range from 41 to 123 cm/second 

(Gyory et al. 2003), ideally allowing dispersal 

times from northwestern Guyana to Manzanilla 

Beach of 7.3 to 2.4 days respectively. In contrast, 

dispersal across the Atlantic Ocean is a daunting 

proposition. The South Equatorial Current flows 

approximately 6,000 kilometers from the Niger 

Delta across the Atlantic Ocean towards the 

mouth of the Amazon River. From satellitetracked 

drift buoy data (WOCE Data Products 

Committee 2002), the current’s maximum 

velocity appears to be around 60 cm/second and 

may commonly reach only 30 cm/second 

(Bonhoure et al. 2004). Under ideal conditions 

these velocities would translate to a dispersal 

time from Africa to South America of 4 to 8 

months (Figure 3.6). That does not include an 

approximately 1,000 kilometers of additional 

dispersal in the more rapid Guiana current before 

reaching Trinidad over an estimated 15 - 45 

days. Those dispersal times are around the upper 

limits suggested by floatation trials with other 

mangrove species; up to 5 months for Avicennia 

marina Vierh. (Clarke 1993) and over 107 days 

for Rhizophora (Rabinowitz 1978a). No 

published floatation data could be located for 

Nypa propagules. 

POSSIBLE SOURCES OF 

PRESENT NEOTROPICAL 

NYPA POPULATIONS 

Neither the Nypa populations in Guyana 

and Panama nor the propagules collected in 

Trinidad have an established source. Nypa has 

likely been present in northwest Guyana for 

several decades, and some elder inhabitants of 

far northwestern Guyana state that there is no 

time in memory that Nypa palms did not grow 

along the margins of the Mora Passage. As 

Figure 3.6. Mapped data points from 1997 WOCE drift buoy experiments (WOCE 2002) in the equatorial 

Atlantic Ocean. The indicated track is the fastest found for the South Equatorial Current, covering about half 

of the 6,000 km distance from the Niger Delta to the Guianas in approximately three months.


mentioned by Pritchard (1993), Nypa palms 

have been under cultivation for many decades 

in the Botanic Gardens in Guyana’s capital, 

Georgetown. A 1955 guide to the Botanic 

Gardens (Anonymous 1955) describes Nypa in 

two ponds growing so vigorously that the ponds 

would be completely overrun if the palms were 

not regularly pruned. These were likely 

introduced in the late 19 th century during the 

Botanic Garden’s heyday under the direction of 

the British botanist George Jenman (McCracken 

1997). Georgetown’s extensive system of 

drainage canals certainly provides a feasible 

fruit dispersal route to the Atlantic Ocean and 

into the Guiana current, which flows towards 

Guyana’s northwest coast (see Figure 3.1). Nypa 

fruits could then easily be carried by tidal fluxes 

into coastal rivers with moderate flows, such as 

the Pomeroon and Waini. Entry into river 

systems may be reinforced by estuarine 

circulation patterns that can drive mangrove 

propagules towards channel centers during flood 

tides and into slower currents close to river 

banks during ebb tides (Stieglitz & Ridd 2001). 

The distance from Georgetown to the mouth of 

the Waini River is about 260 kilometers, similar 

to the distance between the Waini and Trinidad. 

It is also possible that fruits from the live 

specimens in the Botanic Garden were 

intentionally introduced into the rivers of the 

Northwest District. 

It is difficult to propose trans-Atlantic 

dispersal for the Nypa population that was 

reported from Panama. Among the logical 

explanations are intentional introduction, escape 

from cultivation (seeds are available on a few 

horticultural websites) or transport in ballast 

water from one of the many ships that travel 

through the Panama Canal. Duke (1991) did not 

speculate as to the source of introduction in 

Panama, but he rejected the idea that the 

population near Colón was a relict of the ancient 

Nypa distribution, on the basis that the plants 

would not have escaped detection for such a 

prolonged time; he did not discuss the possibility 

of trans-Atlantic dispersal. 

Another scenario is the introduction of 

Nypa to northwestern Guyana by Dutch 

plantation owners, who were active in the region 

in the mid-18th century (Daly 1995), and 

maintained colonies on the Barima River 

53 

(Schomburgk 1896). Erosion control could have 

been a concern of planters. Local people tell 

stories of the Mora Passage having once been 

so narrow that monkeys could travel on branches 

from one bank to another. Richard Schomburgk 

explored northwestern Guyana in 1841 with his 

brother Robert, while under contract to delineate 

the boundaries of British Guiana (van Dam 

2002). He estimated the entrance to the Mora 

Passage to be only 116 feet (35 meters) wide, 

and he described the passage as a winding, 

natural waterway, with strong tidal currents 

(Schomburgk 1922). There are some anecdotal 

accounts (local inhabitants, pers. comm.) of the 

Mora Passage’s origin as a canal planned by the 

Dutch and dug with slave labor. Vann (1969) 

estimated the width of the Mora Passage to be 

over 300 feet (92 meters) wide in 1956, and 

hypothesized that the waterway originated 

through crevassing and scouring by flood waters 

of the Barima River. He estimated that the 

Mora’s depth had increased by nearly one half 

foot per year since measurements were made 

by the Schomburgk brothers. Measurements 

from 1999 Landsat satellite imagery (28.5 m 

resolution) show the Mora Passage as presently 

up to 200 meters wide along much of it course 

(Hollowell, unpublished). Regardless of the 

Mora Passage’s origin, it appears that the Barima 

River has been increasingly captured by the 

Waini River via the Mora Passage. The strong 

tidal flows would encourage bank erosion, and 

eroded sediments may have contributed to a 6 

kilometers long forested deposit which diverts 

the passage before it enters the mouth of the 

Waini River. 

If Nypa has a long history in the area, it 

should have been recorded in botanical records. 

Both Robert and Richard Schomburgk were 

accomplished botanists; between them they 

described nearly 30 Guianan plant species that 

are still valid today (Boggan et al. 1997; 

Hollowell et al. 2001). In 1841 Robert 

Schomburgk (1896), reported numerous Trooli 

palms (Manicaria saccifera Gaertn.) and 

Manicole palms (Euterpe spp.) along the 

riverbanks between the mouth of the Aruka 

River and the Waini River, but he does not 

mention other palms along the banks, and it is 

unlikely that he would have confused either 

Euterpe’s tall, thin form or Manicaria’s broad

54 


fronds with united pinnae with Nypa’s 

distinctive form. Robert’s brother, Richard 

Schomburgk, reported Euterpe palms in his 

writings (1922), comparing their beauty to 

Leopoldinia pulchra Mart.: 

“Though the banks of the Mora had already 

claimed my entire interest, this was nevertheless 

very much more increased by those of the 

Barima. The loveliest of palms, Euterpe 

oleracea Mart., Manicaria saccifera Gaert., 

stretched their proud fronds up above the dark 

succulent mass of foliage and vied with the 

slender Leopoldinia pulchra Mart. both in 

beauty of growth and formation of leaf...” 

Several other early plant collectors in 

Northwestern Guyana (Table 3.1) have also 

failed to record any presence of Nypa, although 

the chief mode of travel within this region was, 

and still is, by boat. In his detailed review of 

the riparian vegetation of Guyana, Fanshawe 

(1954), who traveled and collected plants in the 

Northwest, includes no mention of Nypa. 

Neither is Nypa listed in recent taxonomic 

treatments of palm species of the American 

tropics (Henderson 1995; Henderson et al. 

1995), which require review of a high 

percentage of historical collections held in 

herbaria of the world. This should be considered 

with the knowledge that, compared to other plant 

groups, the large, and sometimes prickly, palms 

are typically under-collected. 

ECOLOGICAL CONSIDERATIONS 

It is useful to consider Nypa’s future in the 

estuarine mangrove swamps in terms of 

conditions for successful invasion listed by Lugo 

(1998) in his review of invasion of mangrove 

ecosystems. Nypa is an obligate halophyte, and 

so there is no reason to assume that the invasion 

is temporary. The conditions that Nypa has 

occupied in Guyana are typical of its native 

environment, along tidal river margins of 

moderate salinity. The invasion is not a response 

to microsite conditions, such as a period of 

freshwater inundation in a basin mangrove 

swamp. The major shift in the environment has 

been the increased disturbances of river banks 

and perhaps some increased sedimentation from 

disturbances in the watershed, which might be 

favorable to Nypa establishment. In 

northwestern Guyana, Nypa seems to have 

established frequently near homesteads or other 

areas of riverbank disturbance. 

The small, though growing, human 

population in the region has not yet caused 

significant stresses in the riverine mangrove 

forests; harvesting of Rhizophora bark for 

tannins has been practiced on an small scale, 

although it is possibly increasing (Allan et al. 

2002). The only event necessary for the Nypa 

invasion was probably the introduction of this 

well-adapted species to an environment from 

which it had been biogeographically isolated. 

So, while Lugo (1998) characterized mangrove 

Table 3.1. Early plant collectors of the Northwest District of Guyana, from US National Herbarium holdings. 

Collector 

Schomburgk, M.R. and R H. 

Jenman, G.S. 

ImThurn, E.F. 

Bartlett, A.W. 

Beckett, J.E. 

Ward, R. 

Hitchcock, A.S. 

Cruz, J.S. de la 

Altson, R.A. 

Archer, W.A. 

Fanshawe, D.B. 

Cowan, R.S. 

Year 

1841 

1896 

1897 

1905 

1906 

1907 

1920 

1922-23 

1926 

1934 

1945 

1955 

Localities 

Waini R., Mora Passage, Barima R., Aruka R. 

Barima R. 



Baramanni, Waini R. 

Aruka R. 

Morawhanna, Issorora, Aruka R., Yarikita R., Amakura R. 

Waini R., ‘Marabo Shortcut’, Barima R., Amakura R. 

Mabaruma 

Mabaruma, Aruka R., Barima R., Koriabo R., Wauna 

Waini R., Aruka R., Mabaruma 

Barima R.


ecosystems as difficult to invade and relatively 

easy to rehabilitate, this would not apply to 

invasion by the few species that are true 

mangroves, which total 70 species worldwide, 

according to Duke et al. (1998). In Hawaii and 

Tahiti, mangroves have been intentionally 

introduced to islands that were once mangrovefree, 

and have become nuisance species (Allen 

1998). The mangrove fern Acrostichum is also 

sometimes considered a nuisance, due to 

interfere with regeneration in some disturbed 

mangrove swamps (Blanchard & Prado 1995; 

Ellison 2000a; Tomlinson 1986). 

It is expected that the Nypa invasion 

in Guyana may be persistent, based in part of 

its current extensive distribution. The abilities 

of Nypa to spread vegetatively and to produce 

plentiful propagules enhance chances of its 

additional spread along the rivers of Guyana. 

CONTROL OF NYPA 

In its native range the control of Nypa is 

not a critical issue. In fact, in a few locations 

such as the Sundarbans of India, Nypa is 

reported to be under threat due to changes in 

hydrology and soil salinity (Badve & Sakurkar 

2003; Sukhendu et al. 2002). In Nigeria, 

eradication of Nypa has been proposed by 

corporate and governmental agencies as a 

element of restoration of the Niger Delta 

(Sunderland & Morakinyo 2002), as attempts 

at Nypa control through increased utilization 

have not been effective. Efforts to control a few 

areas of the alien mangrove Rhizophora in 

Hawaii have been very expensive (Allen 1998). 

Surprisingly, there have been recommendations 

for Nypa plantings in the Volta estuary of Ghana, 

as part of efforts to remediate impacts of dam 

construction on the river (Rubin et al. 1998). 

Such unnecessary introductions should be 

viewed as ecologically risky in light of the 

nuisance that Nypa has become in Nigeria. 

In Panama, the populations of Nypa that 

were documented by Duke (1991) in the Colón 

Free Zone of the Panama Canal have reportedly 

been destroyed by coastal construction activities 

(Neal Smith, Smithsonian Tropical Research 

Institute, pers. comm.), though possibly isolated 

individuals have survived in impoundments 

55 

along the west shore of Bahia Limón to the west 

of Colón. Due to limited knowledge of its 

presence and extent, there is not yet any 

recognition of a need to control Nypa in Guyana; 

it is possible that increased utilization could 

serve as a minor control on its spread there. A 

few instances of Nypa harvest and thatching 

with Nypa fronds have been observed near the 

small settlement of Morawhanna at the western 

end of the Mora Passage, probably because the 

Trooli palm, Manicaria saccifera Gaertn. has 

become increasingly scarce, as its fronds are the 

preferred thatching material in the region. 

NEEDS FOR 

ADDITIONAL STUDY 

There is increasing concern about the 

increases in and effects of invasive plant species 

worldwide (Mack & Lonsdale 2001). The 

conservation community also needs information 

about this potentially aggressive exotic species, 

which could have significant effects on structure 

and function of important riverine mangrove 

communities. Due to their close connection with 

river waters, Guyana’s riverine mangrove 

swamps, which are most susceptible to invasion 

by Nypa, may function more effectively as fish 

habitat and a source of nutrients and organic 

matter than the coastal mangroves isolated 

behind beach ridges (Ewel et al. 1998). 

Ecological research into the dynamics of 

competition between Nypa with Rhizophora 

racemosa and into the growth and reproduction 

of Nypa in Guyana would be useful in 

understanding the possible future of Nypa in the 

region and could inform considerations of 

control or restoration efforts, either through 

eradication or utilization. 

The presence of Nypa in the Neotropics 

poses questions relevant to the study of 

biogeography. The feasibility of trans-Atlantic 

dispersal should be tested with floatation 

experiments on Nypa propagules, as 

demonstrated for mangrove species found in 

Panama by Rabinowitz (1978a), including 

assessment of viability at intervals (Duke et al. 

1998). This might also include trials with 

varying salinities of water, which could affect 

buoyancy and viability, as the Guiana current

56 


can be diluted with substantial freshwater from 

the Amazon River. Molecular evidence may be 

useful in determining whether the sources of 

Nypa in Guyana, Trinidad, and Panama 

originated from African populations, 

introductions directly from Asia, or from the 

plants under cultivation in the Georgetown 

Botanical Garden. Possibly analyses of 

microsatellites would be a useful approach to 

molecular study of this issue (Godoy & Jordano 

2001). 

Records of the Dutch East Indies Company 

might hold information of possible introduction 

of Nypa to settlements in the Barima River area 

or other clues that may shed light on the history 

of the Mora Passage. An undertaking as 

ambitious as a 6 kilometers canal from the 

Barima River to the Waini River should also be 

documented in historical records. 

Since they are restricted to brackish river 

fringes, these Nypa - Rhizophora swamps may 

be relatively easy to delineate and study. The 

extent of Nypa patches along rivers in 

Northwestern Guyana could be readily mapped 

by boat using a Global Positioning System 

(GPS), and resampling over a period of years 

would produce a picture of Nypa dynamics. 

Once a reasonable portion of the Nypa affected 

area is mapped, it may be possible to generalize 

that distribution to the entire region using large 

scale aerial photography (Manson et al. 2001; 

Wilton & Saintilan 2000) or high resolution, 

multispectral airborne imagery (Green et al. 

1998); however, the more affordable Landsat 

imagery probably lacks adequate spatial 

resolution to detect fringing bands of Nypa. 

The distributaries of the Orinoco River 

Delta should also be surveyed for populations 

of Nypa; it would not be surprising if it was 

found there. Those populations would represent 

additional sources for propagules that could be 

dispersed to Trinidad. If documented, the 

presence of Nypa in the Orinoco Delta should 

be of interest to Venezuelan botanists and 

conservation organizations. Certainly, any 

control strategies must be considered in light of 

many factors, with adequate attention to 

unintended effects. 

CONCLUSIONS 

Nypa as an invasive species appears to be 

an growing phenomenon in the tropical Atlantic 

Ocean region. If the source for Nypa in Guyana 

has been present for over a century in the 

Botanic Garden in Georgetown, but the species 

has only recently spread to the Mora Passage 

region, it can be concluded that initial 

colonization from intermediate distances might 

be uncommon, but once accomplished, local 

expansion may be rapid. It may only be a matter 

of time before Nypa becomes established as a 

nuisance in suitable sites of Trinidad and 

Venezuela’s Orinoco Delta. However, it should 

be noted that Duke (1991) warned of the 

inevitable spread of Nypa throughout the Central 

American Atlantic coastal region prior to its 

disappearance from Panama. Trinidad’s Nariva 

Swamp National Park may be of special 

concern, as it is located only a few kilometers 

to the south of Manzanilla Beach where so many 

viable Nypa propagules have been observed. 

Trinidad’s Caroni Swamp, near Port of Spain 

on the Gulf of Paria may also be vulnerable. 

The brackish waters of these areas should be 

monitored for Nypa. Documentation of the full 

extent of colonization and densities in 

northwestern South America and comparison 

with characteristics of Nypa in its native range 

could provide additional insight into the 

invasion and reasons for its success (Hierro et 

al. 2005). Understanding the regional history, 

dispersal, and ecology of this palm will be 

critical to mangrove conservation efforts.


Wiltshire Hinds in Avicennia forest with 

plentiful Acrostichum ferns and Philodendron 

vines, on transect “D” in November 2001. 

Professor Philip DaSilva of the University of 

Guyana, who provided support for research on 

the Waini Peninsula, at the front door of the 

Centre for the Study of Biological Diversity. 

57 

Mangrove activities sometimes intersected with 

those of the Guyana Marine Turtle Conservation 

Society (GMTCS). On the way to 

reconnaissance of a burned area at Kamwatta 

Beach, Keith David, Karen Redden and Donald 

James of GMTCS help free a stranded Green 

Sea Turtle (Chelonia mydas). Over 100 eggs 

were also transported to a GMTCS nursery. 

Wiltshire Hinds pressing the afternoon’s plant 

specimens on the porch of the Kwebana guest 

house, November 2001. Kwebana is the site of 

an abandoned lumber mill on the Waini River 

upstream of tidal influence.

58 


Amerindian students from Moruca inspect a 

Leatherback Sea Turtle (Dermochelys coriacea) 

recently hatched in the Almond Beach GMTCS 

nursery prior to releasing it that evening. 

Audley James at Almond Beach camp, with his 

grandson Alex. A former turtle hunter, Audley 

became one of the key people in GMTCS. 

Romeo DeFritas, son of Violet and Audley 

James and a mainstay of the GMTCS, with hand 

chain-sawn Rhizophora mangle lumber 

harvested nearby for construction at the Almond 

Beach camp. 

Violet James, who with her husband Audley 

James and son Romeo DeFritas, managed many 

of the activities at the Almond Beach marine 

turtle monitoring camp.


CHAPTER 4. 

STRUCTURE OF BURNED AND UNBURNED 

AVICENNIA FOREST, WAINI PENINSULA, GUYANA 

INTRODUCTION 

Fires burned across extensive parts of 

northeastern South America from 1997 to 1998 

(Barbosa & Fearnside 1999; Cochrane & 

Schulze 1998), and at that time soil fires 

occurred in mangrove and freshwater swamps 

of the Waini Peninsula (Chapter 1, this 

manuscript). Those soil fires highlighted the 

need to improve understanding of the Waini 

Peninsula’s mangrove ecosystems and their 

response to disturbances for the benefit of longterm 

management of the area. 

Ecological plots have long been considered 

one of the best methods for collecting such 

detailed plant community information. Early 

mangrove studies were descriptive, often 

conveying species zonation of mangrove 

swamps and proposing explanations for the 

observations (Clarke & Hannon 1969; Davis 

1940; Fosberg 1947; Lindeman 1953; Macnae 

1963). Those studies frequently included 

vegetation profiles illustrating mangrove 

zonation patterns. Profiles are still occasionally 

produced, often incorporating information on 

physical characteristics such as soil type and 

salinity (Lindeman 1953; Odum et al. 1982; 

Thom 1967). One of the earliest quantitative 

ecological studies of mangroves was carried out 

by Golley et al. (1962) in Puerto Rico, in which 

biomass, carbon cycle measurements, and 

zoological trophic data were collected. By the 

1970s, ecological plots were a fairly common 

method for census and description of mangrove 

swamps (Pool et al. 1977). Allometric formulas 

for estimating mangrove forest biomass using 

dbh values have been created for above-ground 

biomass in mangrove forests in Puerto Rico 

(Golley et al. 1962), French Guiana (Fromard 

et al. 1998) and Florida (Ross et al. 2001), and 

for above-ground plus root biomass in Pakistan 

(Snedaker et al. 1995). The mangroves of 

Guyana have been the subject of limited 

59 

ecological study. Ramdass et al. (1997) collected 

plot data for an Avicennia forest near Alness 

Village on the country’s southeastern coast. 

Mangrove litterfall dynamics were investigated 

by Chale (1996) near Onverwagt (see Figure 

4.5) southeast of Georgetown, where high levels 

of small litter productivity were found, at 1771 

g/m 2 /yr, which is approximately double the 

typical rates reported for Caribbean mangroves 

and similar to values reported for tropical 

rainforests. 

Mangrove research on the Waini Peninsula 

was carried out to address several questions 

about the nature of that plant community. If the 

coastal Avicennia forests have been recently 

established in stages on an accreting geomorphic 

formation, their structure should vary over 

space, with the more recently established stands 

possessing less biomass, and vary over time if 

stands are so recent as to have not approached a 

structural equilibrium of maturity. Mature stands 

that have not been disturbed should possess a 

relatively constant structure due to growth and 

mortality being balanced. The distribution in 

diameter classes of trees will indicate some 

aspects of population dynamics of stands. Data 

collected from the standing trees in the burned 

mangrove plots, within months after the fires, 

allowed reasonable comparisons with the living 

trees of the unburned plots. Also, spatial patterns 

of trees within those plots may reflect stand 

history. The early course and stability of plant 

community recovery immediately after 

disturbances may be assessed by observing 

initial changes in vegetation cover. It is also 

useful to compare the mangroves of the Guianas 

with those of the Neotropics and the world, in 

order to understand the ways in which they are 

either unique or similar. 

The vegetation plot data in this study 

provide initial structural information on both 

intact and fire-affected mangrove ecosystems on 

the Waini Peninsula, and allow comparisons 

with values for other mangrove systems in

60 


several parts of the world. Early succession 

following the fires was investigated through 

analysis of the changes in herbaceous vegetation 

in the burned and unburned plots. 

METHODS 

Study Site 

The study site was located near the 

northernmost point of Guyana, on the Waini 

Peninsula. Plots were established in two areas 

just inland of Almond Beach (8° 23' 58'’ N, 59° 

45' 16" W), a few kilometers to the east of Waini 

Point (Figure 4.1). In early 1998, parts of the 

mangrove forest of this area were killed by 

widespread soil fires, with the burn extending 

from the coastal beach ridge several kilometers 

into the interior of the peninsula. A detailed site 

description, listing of plant species of the Waini 

Peninsula, and description of the extent of the 

areas burned in the 1998 fires are given in 

Chapter 1. 

A total of six plots were established, three 

in Avicennia germinans forest with high 

mortality from the fires and three in unburned 

Avicennia forest toward Waini Point from 

Almond Beach. The unburned plots were located 

in an area inland from a narrow beach ridge; 

burned plots were located inland from the 

Guyana Marine Turtle Conservation Society 

(GMTCS) camp, which is located on a broad 

beach ridge. 

Data were also used from the one hectare 

Man and Biosphere style mangrove plot located 

near Alness Village (Ramdass et al. 1997), on 

southeastern Guyana’s lower coastal plain, 

between the Corentyne and Berbice Rivers, 

(6º12’N, 57º18' W, see Figure 4.5). The Alness 

Village plot was located in a pure Avicennia 

germinans stand that had been utilized by cutting 

for poles by local residents. The use of the MAB 

Figure 4.1. Location of modified Whittaker plots in burned and unburned Avicennia forest on the Waini 

Peninsula.


method made those data very comparable to data 

from the Almond Beach plots. 

Plot Establishment 

Three 0.1 ha modified Whittaker vegetation 

sampling plots (Campbell et al. 2002; SI/MAB 

1998; Stohlgren et al. 1995; Whittaker 1960, 

1973) were established for each treatment, being 

burned and unburned Avicennia swamp. Due to 

the low diversity of the plant community, having 

an essentially monospecific tree component and 

fewer than ten herbaceous species, that 

sampling area was determined to be adequate. 

The use of three randomly placed plots allowed 

for the reduction of placement biases and 

influences of small-scale irregularities in the 

forests. Features of the Dallmeier et al. (1992) 

one hectare plot protocol were adapted for use 

within the modified Whittaker plot layout, 

including tagging of all trees of 10 cm dbh or 

more and mapping of tree locations within the 

plot. The plots were established six to seven 

months after the end of fires and were resampled 

three times over a 35 month period in 

order to monitor changes in tree dbh and cover 

of seedlings and herbaceous vegetation. Trees 

in the burned area were measured for dbh and 

mapped only once, to capture information on 

pre-fire structure. The unburned forest plots 

were resampled twice to provide information 

about the growth rates and mortality of trees, 

and herbs and tree seedlings were resampled 

three times in both areas. 

Each plot was laid out from a randomly 

placed corner stake. Corners were determined 

by measuring a randomly determined 50 to 250 

meters along the boundary of the swamp either 

east or west parallel to the coast line, from a 

single starting point for each forest type. Then 

a randomly determined 100 to 300 meters inland 

was measured into the swamp perpendicular 

from the swamp edge. The GPS coordinates of 

those northeast plot corners (Table 4.1), along 

with the presence of durable tags, should allow 

relocation of the plots for several years. The 

northeast corner of each plot was marked with 

a steel rod. Each 0.1 ha plot was 20 m x 50 m in 

size, positioned with the long axis oriented east 

to west (Figure 4.2). The other three corners 

were established using a Tracon surveyor’s 

compass and tape measure. All corners were 

61 

marked with large, plastic surveyor’s stakes. Plot 

boundaries were temporarily marked with string 

for convenience during the surveys. To utilize 

the tree mapping functions of SI/MAB Biomon 

software (Comiskey et al. 1999), each 50 m long 

plot was divided into two 20 m x 20 m quadrats 

and one 10 m x 20 m quadrat (Figure 4.3). To 

fully characterize the vegetation, separate 

measurements were made on herbaceous and 

woody plants. 

For sampling of small woody plants, 

subplots of three sizes (A, B, C) were set within 

the 0.1 ha plots for collection of data on herbs 

and small woody plants. Those were not 

permanently marked. Ten “A” subplots of 0.5 

Figure 4.2. Layout of modified Whittaker plot and 

subplots. Adapted from Stohlgren et al. (1995) and 

Stohlgren and Chong (1997). 

Figure 4.3. Division of the 0.1 ha plots into 20 m x 

20 m quadrats compatible with the Biomon program=s 

tree mapping. 

Table 4.1. Coordinates of the northeast corners of 

the six 0.1 ha modified Whittaker plots, in decimal 

degrees, determined by GPS within 15 m. 

Plot number 

1 - Burned 

2 - Burned 

3 - Unburned 

4 - Unburned 

5 - Unburned 

6 - Burned 

Latitude 

8.40220 

8.40147 

8.39744 

8.39673 

8.39617 

8.40156 

Longitude 

-59.77903 

-59.78069 

-59.75664 

-59.75550 

-59.75522 

-59.77733

62 


m 2 in area, were sampled with a 1 m x 0.5 m 

frame dropped at 12 m intervals, four along each 

side and one at each end of each plot. Two “B” 

subplots placed in opposite corners were 10 m 2 

in area, 2 m x 5 m in size. A central “C” subplot 

was 100 m 2 , 20 m x 5 m in size. 

All woody plants over 10 cm dbh were 

mapped by measuring distance from any two 

adjacent corners, and these measurements were 

converted to x-y coordinates, calculated by 

triangulation (Figure 4.4) using the Biomon 

program (Comiskey et al. 1999). All trees 10 

cm dbh or greater were measured at breast 

height, the species recorded, and labeled with 

heavy aluminum tags stamped with the plot, 

subplot, and tree numbers, attached just above 

the dbh level. 

Figure 4.4. Mapping of tree locations in quadrats of 

SI-MAB plots using triangulation from any two 

corners. Adapted from Dallmeier et al. (1992). 

Water level above or below the substrate 

was measured to provide basic data on 

hydrological changes that might have an 

influence on vegetation and to compare 

fluctuations in water table between the two sites. 

Two 10.2 cm (4 inch) diameter, 1.5 m long pvc 

well pipes were installed 0.75 m deep, one 

between the burned plots and one between the 

unburned plots. The bottoms of the well pipes 

were covered with several thicknesses of porous 

fiberglass screening, and the tops were provided 

with removable caps to prevent evaporation. 

Measurements 

For herbs and small woody plants, in each 

of the ten ‘A’ subplots the percent areal coverage 

of each herbaceous species was visually 

estimated and recorded. Plant species sampled 

as herbs included tree seedlings less than 1 cm 

dbh. In the two ‘B’ subplots all woody plants 

greater than or equal to1 cm but less than 5 cm 

dbh were measured and recorded. In the central 

“C” subplot all woody plants greater than or 

equal to 5 cm but less than 10 cm dbh were 

measured. Measurements were performed at the 

time of plot establishment in November 1998 

and on three subsequent visits in May 2000, 

April 2001 and October 2001. The first two 

samplings occurred during fairly wet conditions, 

with standing water in both living and burned 

forests, while the April 2001 measurements were 

made during dry conditions, evidenced by 

cracking of soil surfaces in the burned swamp. 

The October 2001 measurements were made 

during moderately dry conditions, with no 

standing water in either plot area. 

For woody plants, in each of the six 0.1-ha 

plots, dbh and height of all trees 10 cm dbh or 

greater were measured in November 1998. For 

the first census only tree heights were estimated 

using a simple clinometer at measured distances 

from the bases of trees. Height measurements 

were not made after the initial census. Trees 

were tagged and mapped within the quadrats, 

and the distances from corners were entered into 

the SI/MAB Biomon software. The status of 

each tree as living or dead was recorded; data 

from dead trees were not included in most 

analyses. The first census included Avicennia 

trees in the three burned plots that were killed 

by fire only a few months after their death. 

Subsequent large woody plant measurements 

were not made on the burned plots as the dead 

trees decayed. The three unburned plots were 

re-censussed for tree dbh at approximately 17 

month intervals during May 2000 and October 

2001. Basal areas were calculated from dbh 

using the formula (0.5*dbh) 2 * B, and then 

expressed as m 2 /hectare. All statistical tests, 

except where noted, were performed using the 

S-plus statistical software (Insightful 

Corporation 2001). 

The water table was monitored with the use 

of a thin dip-stick marked in 1 cm increments. 

Water depth above or below the soil surface was 

measured monthly at both wells by marine turtle 

monitoring crew members, starting during the


first plot samplings in November 1998 and 

continuing until the drought in March 2001. 

Alness Village Plot 

Data from the Avicennia germinans 

community at Alness Village in the Berbice 

River region (Figure 4.5) provided the only 

additional mangrove data from Guyana with 

which comparisons to Waini Peninsula 

Figure 4.5. Location of Alness Village on the 

southeastern Guyana coast, site of an Avicennia forest 

1 ha plot (Ramdass et al, 1997) and Onverwagt, site 

of an Avicennia litterfall study (Chale, 1996). 

mangrove physical structure and changes could 

be made, and effects of usage by local 

populations can be evaluated. As with the 

Almond Beach plots, the Dallmeier et al. (1992) 

plot method had been used to establish and 

census that 1 hectare plot in 1995 and 1996. 

Twenty-five quadrats of 400 m 2 each were set 

up in a 2-quadrat wide belt, with one row 10 

quadrats long and the other 15 quadrats long 

(Figure 4.6). All trees of 10 cm or greater dbh 

were measured. Tree heights were recorded only 

in 1996, during the second census. These data 

were obtained from Ramdass et al. (1997); an 

electronic version of data from the Alness plot 

was not available, and so those data printed in 

the report were re-entered into the SI/MAB 

Biomon software. 

Dispersion Patterns of Trees 

For an analysis of the arrangement of trees 

in these plots, the dispersion pattern of 

individuals was calculated. Data from the initial 

63 

1998 census only were used to measure 

dispersion for both the unburned and the burned 

plots. For each plot the XY coordinates of 

individuals were calculated using the SI/MAB 

Biomon program (Comiskey et al. 1999). The 

Chen and Getis (1998) Point Pattern Analysis 

(PPA) package was utilized through its on-line 

application (Aldstadt et al. 1998), using the 

nearest neighbor algorithm to determine whether 

the spatial pattern of individuals was random, 

clustered, or over-dispersed. The nearest 

neighbor analysis calculates distances between 

each point and the closest point and compares 

those distances to expected distance values from 

a random sample of points from a complete 

spatial randomness pattern (Chen & Getis 1998). 

Figure 4.6. Layout of the Alness Village 1 hectare 

plot (Ramdass et al. 1997) with quadrat numbers and 

tree locations. Quadrats 1-20 were used for dispersion 

analysis. This diagram of the linear plot has been 

divided into two parts for display.

64 


The output of the PPA program includes basic 

two dimensional statistics and a Z statistic (the 

standard normal variate) indicating the 

probability of significant clumping or overdispersion. 

The Alness Village 1 hectare plot was 

irregularly shaped, however a 0.8 ha portion 

formed a single rectangle two quadrats wide 

(Figure 4.6) providing data that was compatible 

with requirements for the PPA software. The XY 

coordinates for that rectangle were calculated 

from values for the component quadrats and run 

with the PPA software (Chen & Getis 1998). 

Biomass 

Using the allometric equations derived by 

Fromard et al. (1998), from Avicennia tree 

measurements in French Guiana, the aboveground 

biomass values for mangroves at 

Almond Beach and Alness Village were 

estimated for comparison with the values from 

French Guiana. The Fromard et al. (1998) 

formula for Avicennia trees above 4 cm dbh is: 

biomass = 0.14 * dbh 2.4 , with dbh in cm and the 

resulting biomass in kilograms; it includes dry 

weight of wood, leaves, fruit and flower, and 

above-ground roots. For comparison with 

swamps of Guyana, additional mangrove 

biomass values from the literature were 

assembled from reviews by Lugo and Snedaker 

(1974) and Fromard et al. (1998) and plotted. 

Comparison with Other Plot Data 

To further illustrate the relationship of the 

Waini Peninsula mangroves among mangrove 

ecosystems worldwide, basal area and stem 

density data for mangrove forests in several 

localities were compiled from sources in the 

literature (Bosire et al. 2003; Brocklehurst & 

Edmeades 2003; Cardona & Botero 1998; Chen 

& Twilley 1999; Cintrón et al. 1978; Fromard 

et al. 1998; Golley et al. 1962; Kjerfve 1998; 

Lacerda et al. 2002; Lugo & Cintrón 1975; Pool 

et al. 1977; Ramdass et al. 1997; Ross et al. 

2001; Roth 1992; Sherman et al. 2000; Snedaker 

et al. 1992; Walters 2000). Any reported mean 

tree or canopy heights were recorded, as were 

the geomorphic type of the swamps. Where 

possible the site locations were assigned to 5degree 

latitudinal classes. 

RESULTS 

Tree Mapping and Inventory 

Maps of tree locations on the six plots are 

given in Figures 4.7 to 4.12. A total of 65 trees 

(64 dead, including trees likely killed by fires 

and recently fallen, and 1 living tree) were 

measured in the three combined 0.1-ha plots in 

the burned site (plots 1, 2, 6; Figures 4.7, 4.8, 

and 4.9), and 170 individual trees (dead and 

alive) over all three samplings in the unburned 

Avicennia swamp plots (plots 3, 4, 5; Figures 

4.10, 4.11, and 4.12). 

A. Dbh and Tree Height 

In the unburned forest, differences from any 

one sampling to the next were not statistically 

significant (p= 0.558 for 1998 vs 2000 and p= 

0.227 for 2000 vs 2001), while the difference 

approached the p=0.05 significance level 

between the first and last sampling (p= 0.076 

for 1998 vs 2001), indicating that stand structure 

might be changing slowly over time, possibly 

to be interpreted as a fairly young forest 

maturing. Statistical tests paired by individual 

trees resulted in very low p-values, as would be 

expected since individual trees almost invariably 

change over time. The unpaired tests were more 

suitable for evaluation of overall stand structure. 

The dbh of the trees remaining in the burned 

plots were measured after initial plot 

establishment in 1998, a few months after the 

end of fires. Trees in the burned plots which had 

apparently fallen recently were also measured, 

and it was assumed that any trees that were dead 

prior to the fires had been consumed. Dead trees 

in the unburned plots were omitted from these 

comparisons. For the unburned plots only the 

1998 data were used for comparisons. Variances 

and skewness between these data were different 

enough to indicate use of non-parametric tests. 

Significant differences between dbh of burned 

and unburned plots were found with the nonparametric 

Wilcoxon rank-sum test, with a pvalue 

of 0.004. 

The maximum dbh recorded in the first 

sampling for trees at the burned site (65.5 cm) 

was higher than that of the unburned site (50 

cm). The mean dbh value at the burned site of 

31.3 cm was significantly higher than the value 

of 23.3 cm at the unburned site (p=0.0004,


Figure 4.7. Plot 1, unburned Avicennia swamp at 

Almond Beach. The symbol size represents the 

relative size of each tree that was mapped. 



relative size of each tree that was mapped. Lines 

extending from a symbol represent fallen trees and 

the direction of fall. 

Figure 4.11. Plot 4, burned Avicennia swamp at 




the direction of fall; in the case of burned plots those 

were assumed to have been living trees prior to fires. 

Welch’s modified two-sample t-test). The 

median dbh values were 28.5 cm for the burned 

site and 23.1 cm for the unburned site. Diameter 

class frequencies for the burned plots are 

graphed in Figure 4.13, and for the unburned 

plots (2001 census) are shown in Figure 4.14. 

The 1997 Alness plot and the 1998 unburned 

plot at Almond Beach had markedly different 

structures. The Alness mean dbh values of 16.15 

cm in 1995 and 16.49 cm in 1996 were not 

significantly different (t-test, p=0.59). The mean 

65 



relative size of each tree that was mapped. 













dbh was 23.3 cm at Almond Beach compared to 

16.3 cm and at Alness; the median dbh at 

Almond Beach of 23.1 cm was significantly 

higher (p=0, Wilcoxon rank-sum test) than the 

median of 15 cm at Alness. For the Alness 

Village plots, a dbh distribution for the 1996 

sampling is graphed in Figure 4.17. 

Maximum tree heights were 36 meters in 

the burned forest and 34 meters in the unburned 

forest. Mean tree heights were 21.3 meters in 

the burned site and 18 meters in the unburned

66 


Figure 4.13. Dbh class distribution for the three 

burned plots combined (0.3 ha), at Almond Beach, 

from the 1998 sampling. Only living stems are 

assumed to have remained after the fire. Recently 

fallen trees killed in the fires were measured and 

included. The graph suggests periodic waves of 

successful recruitment. The higher frequency in the 

25-30 cm size class is also seen in the unburned 

swamp. 

Figure 4.14. Dbh class distribution for the three 

unburned plots, combined (0.3 ha), at Almond Beach, 

2001. Only living stems were included. 

site; those were not significantly different (p= 

0.268, Wilcoxon rank-sum test). A graph of tree 

height classes for unburned and burned plots is 

given in Figure 4.15. A plot of dbh vs height for 

all unbroken trees in all plots is provided in 

Figure 4.16; a logarithmic regression provided 

a best fit for those data, with a formula of 

y=12.066 Ln (x)-16.595, R 2 =0.585, which could 

be employed to estimate tree height in other 

Avicennia forests. Tree height data were 

collected for Alness only in 1996, resulting in a 

mean height of 6.8 meters and a maximum 

height of 19 meters. Mean tree height at Alness 

was 6.8 meters compared to 20.7 meters at 

Almond Beach. The median tree height at Alness 

Village of 6 meters was significantly lower (p=0, 

Figure 4.15. Dbh class distribution for 1996 sampling 

of the Alness Village plot (1 ha). (Ramdass et al. 

1997). Only living stems are included. This plot is 

reported by the researchers as having been frequently 

disturbed. 

Figure 4.16. Height class distribution for trees in 

unburned and burned plots at Almond Beach, 1998 

sampling. Mean tree heights were 21.3 m in the 

burned plots and 18 m in the unburned plots. Medians 

were not significantly different, (p= 0.268 using 

Wilcoxon ranksum test). 

Figure 4.17. Tree diameter vs tree height for all 

unbroken trees in the Almond Beach plots, with a 

log-normal trend line.


Wilcoxon rank-sum test) than the 1998 Almond 

Beach unburned plot median of 21 meters. A 

graph of tree heights classes for the Alness 

Village plot is provided in Figure 4.18. Summary 

statistics for the plots over all years sampled are 

listed in Table 4.2. 

Figure 4.18. Alness Village plot tree height frequency 

distribution. 

Table 4.2. Summary statistics for trees in unburned and burned plots at the Almond Beach sites and the Alness 

Village site 

Site 

Burned 1998 

Unburned 1998 

Unburned 2000 

Unburned 2001 

Alness 1995 

Alness 1996 

Maximum 

65.5 

50.0 

51.4 

52.5 

47.5 

49.0 

B. Basal Area 

The basal area for trees that were located 

in the combined three burned plots, measured 

only in 1998, was 20.43 m 2 /ha. The basal areas 

of living trees in the combined three unburned 

plots, were 21.25 m 2 /ha in 1998, 22.36 m 2 /ha in 

2000 and 23.16 m 2 /ha in 2001. Over the 34 

month study period, basal area in the three 

unburned plots increased by 1.91 m 2 /ha. Three 

pairs of t-tests were made to compare means 

between the three years for the grouped 

unburned plots. Between the three years 

variances of distributions were similar, therefore 

t-tests were acceptable; they were used for both 

paired and unpaired data. For the Alness Village 

plots sampled by Ramdass et al. (1997), basal 

dbh (cm) Height (m) 

Mean Median Maximum Mean 

31.3 

23.3 

24.0 

25.5 

16.1 

16.3 

28.5 

23.1 

24.0 

25.9 

14.95 

15.0 

36 

34 

na 

na 

na 

19 

21.3 

20.7 

na 

na 

na 

6.8 

67 

areas were 10.07 m 2 /ha in 1995 and 10.28 m 2 / 

ha in 1996 (not significantly different, p=0.42). 

C. Dispersion 

For the unburned plots, nearest neighbor 

analysis of plot 1 resulted in an observed mean 

distance between trees of 2.22 m while the 

expected distance was 2.48 m, with a z-value of 

-1.16. This negative Z statistic value indicates 

that, while a clustered distribution was observed, 

it was not quite significantly different from a 

random distribution at the p=0.05 level. For plot 

2 the mean distance between trees was 2.75 m 

with an expected distance of 2.66 m. The Z 

statistic obtained from the nearest-neighbor 

analysis suggested slightly over-dispersed 

(uniform) distribution, however the tendency 

was not significantly different from a random 

distribution, with Z=0.33. For plot 6 the mean 

distance between trees was 2.48, with an 

expected distance of 2.18. The Z statistic value 

of 1.686 indicated a tendency toward over- 

Median 

22 

21 

na 

na 

na 

6 

dispersion but slightly below statistical 

significance at the p=0.05 level. 

For the three burned plots, all point pattern 

analyses resulted in positive Z statistic values 

indicative of over-dispersion, however with no 

significant tendencies (Z statistic values were < 

1.96, p>0.05). Plot 3 data showed a mean 

distance between trees of 3.80 m, and an 

expected value of 3.28 m with a Z statistic value 

of 1.331. The mean distance between trees in 

plot 4 was 4.40 m, with an expected value of 

3.83 m and a Z statistic value of 1.076. Plot 5 

had an mean distance between trees of 3.80 m 

and an expected distance of 3.39 m with Z 

statistic value of 0.99.

68 


The Alness Village plot nearest neighbor 

analysis found an mean distance between trees 

of 1.86 m, with an expected distance of 2.34 m, 

indicating a highly significant clustered 

distribution of trees for that plot (Z = -7.2114). 

D. Density and Population Turnover 

in the Avicennia Forest 

Stem densities between the two sites were 

very similar, with Alness density being 445 

stems per ha compared to an Almond Beach 

unburned plot density of 440 stems per ha. 

Alness stem density decreased from 445 stems/ 

ha to 436 stems/ha over the one year interval. 

In the three 0.1-ha unburned Avicennia plots, 

only four new trees entered the >10 cm 

population over the 34 month study period: two 

during the 1998-2000 interval and two during 

the 2000-2001 interval. Over the same intervals, 

a total of 29 trees died or disappeared from the 

living Avicennia plots, seven during the 1998- 

2000 interval and 22 during the 2000-2001 

interval. 

The number of living trees in the three 

unburned plots (0.3 ha total) was 132 in the 1998 

census, 130 in 2000 and 115 in 2001, 

corresponding to per ha densities of 440, 433.3 

and 383.3 stems respectively. The density of 

living trees therefore decreased in the first 

interval by 6.7 per ha and in the second interval 

by 50 per ha. In the burned plots, only 64 trees 

were located in the single census, a density of 

213.3 trees per ha. The Alness Village plot had 

a density of 445 stems/ha in the first census and 

436 stems/ha in the second census, a decrease 

of 9 stems/ha. 

E. Biomass 

Estimated biomass values for the Almond 

Table 4.3. Biomass and basal area values for plots at 

Almond Beach by site and year and for Alness Village 

by year. Biomass values were calculated from the 

allometric formula of Fromard et al. (1998). 

Site 

Unburned 1998 

Unburned 2000 

Unburned 2001 

Burned 1998 

Alness 1995 

Alness 1996 

Biomass 

kg/ha 

145,637 

155,641 

163,632 

165,207 

58,605 

60,300 

Basal area 

m 2 /ha 

21.25 

22.36 

23.16 

20.43 

10.07 

10.27 

Beach plots and for the Ramdass et al. (1997) 

Alness Village plots are given with the 

corresponding basal areas in Table 4.3. The 1998 

biomass value for the burned plots of 165,207 

kg/ha was slightly higher than the unburned plot 

mean of 154,970 kg/ha, though the basal area 

totals for the burned plots was lower than for 

the unburned plots. 

Worldwide biomass values gathered from 

the literature are summarized in descending 

order in Table 4.4 and graphed in Figure 4.19. 

The Guyana values from Almond Beach and 

Alness Village plots are emphasized (bold print) 

in both the graph and table. The maximum 

biomass value was 460,000 kg per ha, from a 

plot in Malaya, while the mean for all values, 

including dwarf and early successional stands, 

was 159,328 kg/ha, in the range of values from 

the Almond Beach plots. 

Figure 4.19. Mangrove biomass values (kg/ha) from 

compiled studies, sources included Lugo & Snedaker 

(1974) and Fromard (1998). Values from Guyana are 

shaded black, others from the Neotropics are in black, 

and values from the Asian-Pacific region are in white. 

Labels for values from this study are labeled in bold 

type. Units are in dry weight. 

Small Woody Plants 

The “B” subplots sampled very few 

individuals from 1 cm and less than 5 cm dbh 

(saplings). Only one living individual in the size 

range was recorded in all unburned plots over 

all census dates, an Avicennia sapling of 2.8 cm 

dbh in plot 6 in the October 2001 census. During 

that sampling there were 14 Laguncularia and 

four Avicennia saplings located in the “B” 

subplots of plot 4 in the burned swamp. The 

basal areas of these small saplings were 0.10


Table 4.4. Mangrove biomass values from 46 additional plot sites worldwide, compiled from the literature, in 

descending order. Values from Almond Beach and the Alness Village study are highlighted in bold face. 

Biomass Values 

Malaya 

Indonesia 

Indonesia 

Indonesia 

Australia 

Fr. Guiana 

Malaya 

Indonesia 

Panama 

Panama 

Sri Lanka 

Australia 

Andaman Isl. 

Sri Lanka 

Indonesia 

Florida, riverine 

Guyana 1998 (burned) 

Florida 

Thailand 

Florida, fringe 

Guadeloupe 

Malaya 

Guyana 1998 

Florida, overwash 

Florida 

Andaman Isl. 

Florida, overwash 

India 


Japan 

New Zealand 

India 

Guadeloupe 

Florida, riverine 


Sri Lanka 

Puerto Rico 

Puerto Rico 

Senegal 

Guyana, Alness 1995 

Guadeloupe 

Florida, Island 

Philippines 

Fr. Guiana 

Florida 

Florida 

New Zealand 

India 

Mean 

Median 

Max 

Min 

Biomass kg/ha 

460,000 

436,400 

406,600 

356,800 

341,000 

315,000 

314,000 

299,100 

279,212 

279,200 

240,000 

220,800 

214,000 

193,000 

178,200 

173,900 

165,207 

164,000 

159,100 

152,868 

152,300 

147,000 

145,637 

129,645 

124,600 

124,000 

119,582 

118,700 

117,523 

108,100 

104,100 

101,900 

98,600 

98,218 

86,192 

71,000 

62,900 

62,850 

60,000 

58,605 

52,800 

48,968 

45,936 

31,500 

8,200 

7,900 

6,800 

5,800 

159,328 

127,123 

460,000 

5,800 

Source 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Lugo & Snedaker, 1974 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 


This study (Fromard equation) 

Fromard, 1998 

Fromard, 1998 


Fromard, 1998 

Fromard, 1998 

This study (Fromard equation) 


Fromard, 1998 

Fromard, 1998 


Fromard, 1998 


Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 



Fromard, 1998 

Fromard, 1998 


Fromard, 1998 

Ramdass et al., 1998 (Fromard eq.) 

Fromard, 1998 



Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

Fromard, 1998 

69

70 


m 2 /ha in the unburned swamp and 2.02 m 2 /ha 

in the burned swamp. In the three 0.01 ha “C” 

subplots in the unburned swamp, only four 

saplings from 5 cm dbh up to 10 cm dbh were 

found in the three unburned “C” plots in the 

first census, six saplings in the second census, 

and seven in the third census, with total basal 

areas of 0.68, 0.87 and 1.1 m 2 /ha respectively. 

A summary of changes in small woody plant 

basal areas is illustrated in Figure 4.20. 

Figure 4.20. Changes in basal area for burned plots, 

for small woody plant species from 1 cm dbh to less 

than10 cm dbh, over three samplings. 

Herbaceous Plants 

Herbaceous vegetation composition varied 

dramatically over four samplings. The 

vegetation in both burned and unburned plot 

areas was influenced to a great degree by recent 

precipitation amounts and the levels of standing 

water in the swamp, which varied significantly. 

In the unburned plots, the first sampling in 

November 1998 took place after a period of high 

rainfall, which was reflected in 33% areal 

coverage of the small floating aquatic plant 

Lemna aequinoctialis Welw. and some presence 

of Rhizoclonium africanum algae on 

pneumatophores of the Avicennia trees. Drier 

conditions prior to the second sampling in April 

2000 apparently provided good conditions for 

germination of Avicennia propagules, when the 

highest coverage rate of those was recorded, at 

just over 2% . The third sampling in April 2001 

followed an extended dry period, and almost no 

herbaceous vegetation was recorded other than 

a few Avicennia seedlings. For the final sampling 

of October 2001, conditions had been less dry, 

and there was a slight increase in the coverage 

of Avicennia seedlings. 

While herbaceous vegetation was largely 

absent from the unburned Avicennia plots 

(Figure 4.21), it was present in the burned 

swamp throughout the samplings (Figure 4.22). 

The floating aquatic Lemna aequinoctialis was 

common for several months following the fires, 

Figure 4.21. Changes in percent areal coverage of 

herbaceous species in unburned Avicennia plots, over 

four samplings, 1998-2001. 

Figure 4.22. Changes in percent areal coverage of 

herbaceous species in burned Avicennia plots, over 

four samplings, 1998-2001. 

with 53.8% cover during the 1998 sampling. 

During the May 2000 sampling, two years after 

the fires, the predominant floating aquatic 

vegetation was Limnobium laevigatum (Humb. 

& Bonpl. ex Willd.) Heine (58.8% cover), along 

with some spots of the sprawling terrestrial herb 

Alternanthera sessilis (L.) R.Br. ex DC (14% 

cover) and Acrostichum ferns on a few slightly 

elevated areas present in the vicinity but not 

sampled. The April 2001 sampling followed a 

strong dry season, and the dominant herbaceous 

vegetation in the burned areas was 

Alternanthera (12.6% cover). The final


sampling in October 2001 also followed a dry 

season, and showed a sharp increase in 

dominance of Alternanthera (67.5% cover) in 

the burned area plots and an increase in seedlings 

of white mangrove Laguncularia racemosa 

(from 0.07% to 1.33% cover) which were 

concentrated in areas near the scattered 

surviving individuals of that species. 

Chi square analyses were performed with 

herbaceous plants both burned and unburned 

plot data matrices by species over the four 

sampling periods (Table 4.5), testing for 

differences between expected and observed 

values of percent cover. Expected values were 

derived from the product of the percent coverage 

Table 4.5. Mean percent cover for herbaceous species over the four samplings in burned and unburned plots. 

These data presented in graphical form in Figure 4.21 for unburned plots and 4.23 for burned plots. 

Acrostichum 

Alternanthera 

Avicennia 

Cissus 

Ceratophyllum 

Cyperus 

Eclipta 

Laguncularia 

Lemna 

Limnobium 

Morinda 

Rhabdadenia 

Rhizoclonium 

Bostrichia 

Total % cover 

1998 

0.33 

0.83 

53.83 

2.33 

4.67 

62.00 

2000 

14.07 

0.03 

0.07 

0.37 

0.83 

5.83 

58.83 

1.67 

3.33 

85.03 

2001 

12.60 

0.13 

1.13 

3.73 

0.07 

0.03 

17.70 

totals for the column and row of each 

combination of date and taxon, divided by the 

grand total for all combinations. Percent 

coverage for all “A” subplots in the three plots 

at each site were averaged. For both burned and 

unburned sites, differences in values for 

herbaceous cover were statistically significant 

between the species over the four samplings, 

with p < 0.001 in both burned and unburned 

habitats. 

Principal Components Analysis (PCA) 

allows summarization of multivariate data for 

visualization in two or three dimensional 

Burned Unburned 

2001b 

67.50 

0.07 

0.27 

1.33 

1.17 

70.33 

1998 

0.37 

33.33 

4.53 

38.23 

2000 

0.03 

2.03 

2.07 

2001 

0.37 

0.37 

2001b 

0.33 

1.63 

0.07 

2.03 

71 

display, by determining optimal standardized 

linear combinations of many variables. PCA was 

performed using herbaceous data from all six 

plots over the four sampling dates from 1998 to 

2001. 

In the PCA Ordination, the three first axes 

(Figure 4.23) account for over 98% of the 

variance in this ordination (Figure 4.24). Three 

taxa have the major influence (loadings) on these 

components: Alternanthera, Lemna, and 

Limnobium (Figure 4.25). The principal 

components biplot graph displays the original 

taxon variables on the same axes with the 

transformed observations. 

The Lemna vector represented the relative 

loadings of the two most important principal 

components for the 1998 sampling, trending 

towards the position of points for plots 3 (point 

#9), 4 (#13) and 5 (#17). The highest values for 

Lemna in plot 4 were nearly matched by those 

for plot 1 (#1). The Limnobium vector had as its 

extreme points the 2000 sampling in burned 

plots 3 (#10), 4 (#14) and 5 (#18). The 

Alternanthera vector showed extremes on the 

October 2001 sampling, in burned plots 3 (#12), 

4 (#16) and 5 (#20). Alternanthera was also 

strongly represented in the March 2001 

sampling in plot 4 (#15).

72 


Figure 4.23. Principal Components Analysis (PCA) 

biplot for percent areal coverage of herbaceous 

vegetation in burned plots. The coverage for the 

floating aquatics Lemna and Limnobium and the 

annual herb Alternanthera account for most of the 

variance within the data. 

Figure 4.24. Relative importance of the Principal 

Components for herbaceous vegetation of burned 

plots. The cumulative importance for the first three 

components is 98.8%, indicating that they are 

overwhelmingly responsible for variances in these 

data. 

All other species in the PCA ordination had 

minor cover and were outweighed by other taxa, 

and so clustered in the center of the graph. 

Except for the occurrence of moderate Lemna 

coverage in 1998, the unburned plots fell into 

that space because of very sparse herbaceous 

cover. 

Well Levels 

Hydrology in the Waini Peninsula 

Figure 4.25. Loadings for the principal components 

for herb species in the burned plots. 

Avicennia swamps is dominated by rainfall, 

because tidal influence is limited by beach ridges 

along much of the coast. Outlets to the ocean 

are infrequent, and creeks draining the swamps 

often empty first into lagoons, some of which 

are above mean sea level, limiting tidal 

influence. Changes in the lagoon outlets can 

have significant influence on hydrology of the 

swamps near the coast. Well levels recorded in 

the burned and unburned Avicennia swamps are 

shown in Figure 4.26. Water levels in both 

burned and unburned plots followed very similar 

patterns of fluctuation, and remained above the 

Figure 4.26. Well levels in burned and unburned areas 

over the period from the first to third samplings of 

herbaceous vegetation. A drought in early 2001 is 

conspicuous, though its severity was far less than 

the 1997-1998 drought that led to the fires on Waini 

Peninsula.


surface of the substrate until early 2001. Those 

peaked around 25 cm above the surface during 

wet seasons. 

DISCUSSION 

Guyana Plots 

Avicennia swamps are typical of early 

succession on the coasts of the Guianas, as 

evidenced by monospecific bands of young trees 

on accreting parts of the coastline. This was 

evidenced by the Avicennia community 

documented here through the increases in size 

of individual trees between the “younger” area 

of the unburned plots and the burned plots in 

supposedly “older” areas farther from the Waini 

Point. There was also an increase in mean dbh 

and biomass within the unburned swamp over 

time, from sampling to sampling. The mean dbh 

value of the burned plots was significantly (p= 

0.004) more than the mean for the unburned 

plots at Almond Beach, but it is also possible 

that some smaller living stems in the burned 

swamp were completely consumed by fire. Both 

the maximum diameters and maximum tree 

heights recorded in the burned swamp were also 

higher than in the unburned swamp, suggesting 

that the forest structures of the two areas were 

different before the fires, and that the burned 

area was a somewhat more mature mangrove 

stand. Probably neither stand had reached an 

equilibrium since their initial establishments. 

Basal areas of living trees in the unburned plots 

increased over both sampling intervals, 1.11 m 2 / 

ha in the first interval and only 0.8 m 2 /ha in the 

second, suggesting that it was a young, growing 

stand. An observed gradual increase in 

frequency of Rhizophora trees in the swamps 

along the Waini Peninsula coast to the southeast 

of Almond Beach also supported the idea that 

an equilibrium had not been reached. 

The very low numbers of seedlings and 

trees less than 10 cm dbh also suggested that 

recruitment was poor, and occurred periodically 

rather than consistently from year to year. Dbh 

distributions of trees within the Almond Beach 

Avicennia swamps, with interspersed classes of 

high and low frequency, suggest periodic 

recruitment waves. 

Although both median basal areas and tree 

73 

heights were significantly higher at Almond 

Beach than at Alness Village, the occurrence of 

a few individuals with dbh similar to the 

maximum at Almond Beach (49 cm dbh vs 50 

cm dbh) suggests that the Alness Village stand 

had been present for some time and might have 

matured to a structure similar to the Almond 

Beach stands, if undisturbed by tree cutting. 

Utilization impacts should be considered in the 

development of mangrove management plans. 

Prior to this Almond Beach plot study no data 

from Guyana were available to allow 

comparison with the Alness Village Avicennia 

forest. 

Stem density also decreased slightly in the 

Alness Village plots over the one year interval, 

while basal area increased, although differences 

between the two years were not highly 

significant. Still, the increase suggested an 

Avicennia population that was growing and 

perhaps recovering from disturbance. Ramdass 

et al. (1997) conjectured that ongoing low-level 

disturbances occurred in that swamp, probably 

through periodic cutting, but he also believed 

that the visible tree tags in the Alness plots may 

have made local people may wary of utilizing 

those trees. Stem density data were available for 

89 of the worldwide plots compiled from the 

literature, with values ranging from 267 stems/ 

ha to 47,330 stems/ha. The combined Almond 

Beach unburned plots had a stem density of 413 

stems/ha, compared to the compiled median of 

3,120 stems/ha for all plots and the mean of 

5,493.8 stems/ha. Stem densities from those 

plots are graphed with Almond Beach values in 

Figure 4.27. 

Tree heights were compiled for 61 

worldwide plots, with stand height values 

ranging from 2.9 meters to 23 meters; the 

median for those values was 10 meters and the 

mean 10.98 meters. That compares to mean 

heights of 18 meters in the unburned Almond 

Beach swamp and 21.3 meters in the burned 

swamp, with respective medians of 18 meters 

and 22.5 meters. The tallest trees at Almond 

beach were 34 meters in the unburned swamp 

and 36 meters in the burned swamp. In the 

Alness Village plot, out of 438 tree height 

measurements made in 1996, the mean tree 

height was 6.98 meters and the median 7 meters, 

with a maximum recorded tree height of 19

74 


meters. 

The Riverine Swamps on the Waini 

Peninsula were not sampled, however some 

large Rhizophora trees were observed along the 

rivers, and scattered, unusually large Avicennia 

trees were observed near the upland boundary 

of Riverine Swamps. The riverine systems are 

more frequently flushed than the coastal basin 

swamps, due to the latter’s separation from tides 

by beach ridges. It is quite possible that higher 

basal area values occur in the Riverine 

Mangroves as a result of the lower salinity that 

was typical in those soils (generally >20 psu near 

the Atlantic beach ridges and


of patchy disturbance patterns due to utilization. 

The clustering of individuals might also reflect 

localized Avicennia propagule sources or 

irregular stranding of propagules on logs, 

branches or slight variations in topography. 

Additionally, the clustering may be indicative 

of low levels of competition from thinning by 

utilization, which would keep basal area low 

(10.28 m 2 /ha in 1996 vs 21.25 m 2 /ha in 1998 

for Almond Beach unburned plots) and prevent 

competition that might lead to overdispersion. 

In general, the dispersion results indicated that 

the Almond Beach Avicennia forests had been 

established long enough without disturbance for 

competition to begin to exhibit an overdispersed 

pattern (Wells & Getis 1999). 

Biomass 

While basal area values determined for the 

burned plots were lower than the unburned plots 

in the same year, estimated biomass values were 

slightly higher for the burned plots. It should 

also be noted that dbh measurements for trees 

in the burned forest were probably an 

underestimate of pre-burn structure, since the 

trees shed their bark shortly after fire mortality, 

and may have shrunken slightly during drying. 

It is possible that some live trees may have been 

entirely consumed during the fires, although it 

is more likely that only dead trees were 

consumed. Comparisons of biomass provide a 

variation on comparisons between stands. By 

transforming dbh data to account for the 

consequences of diameter changes, biomass 

values are more sensitive to differences in tree 

size, and they allow better differentiation 

between young stands with many small trees and 

mature stands with fewer but larger individuals. 

Allometric formulas such as those provided by 

Fromard et al. (1998) are a useful tool. The mean 

wood biomass value for the burned plots, 

165,207 kg/ha, could contribute to an estimate 

of the amount of carbon that will be released to 

the atmosphere as the dead wood decomposes. 

Root biomass and soil would be responsible for 

additional carbon release from the fires. 

Snedaker et al. (1995) found that Avicennia 

marina trees in Pakistan had root biomass that 

was approximately 47-57% of total biomass, so 

assuming that Neotropical Avicennia is similar, 

carbon loss estimates for the fires might be 

75 

doubled. During this field work, the depth of 

the organic layer lost to burning in the Avicennia 

swamp was estimated by an informal survey of 

length of pneumatophores above the soil surface 

in burned and unburned patches of swamp, 

under the assumption that pre-fire hydrology 

was similar between the sites. That yielded a 

difference between the means of 4.6 cm (N=40, 

p-value=0.001), which could be considered 

depth of organic soil lost in the fires. Organic 

soil samples were not collected for analyses of 

carbon content, but soil samples from similar 

sites could be obtained from the site for 

estimates in the future. 

Comparisons with Worldwide Plot Basal 

Area Data 

The compilation of plot values from studies 

in the literature yielded a total of 96 sites, of 

which 95 included basal area values and most 

included stem density values. The great majority 

of those data were drawn from studies that 

sampled all trees 2.5 cm dbh and greater. 

Almond Beach unburned plots had an almost 

negligible number of stems between 2.5 cm and 

10 cm dbh, (only seven stems in the 5-10 cm 

range, representing additional basal area of only 

1 cm/ha) making the Almond Beach data 

comparable to the compiled plot data. 

Basal areas in the compiled plot data ranged 

from 1.17 m 2 /ha to 96.4 m 2 /ha. The mean basal 

area for the Almond Beach unburned plots was 

23.2 m 2 /ha, almost exactly the median value 

from the compiled sites of 23.25 m 2 /ha and near 

the mean value of 25.6 m 2 /ha. 

Latitude classes could be determined for 94 

of the compiled sites. There were no strong 

trends in mangrove basal area (m 2 /ha) vs 

latitude, though possibly this could be 

demonstrated with a larger data set that was 

balanced in other factors. The non-parametric 

Kruskal-Wallis rank-sum test was used to 

compare the 6 latitude classes with 8 or more 

values, and resulted in a p-value = 0.167, 

indicating no statistically significant difference. 

Basal area values (m 2 /ha) for the sites are 

grouped by latitude class in Figure 4.28. 

Compiled basal area data were grouped by 

swamp geomorphic type according to the 

classification of Lugo and Snedaker (1974). 

Overwash type swamps were not included, as

76 


there was not an adequate number of values. 

There was no significant difference between 

fringing (coastal) and basin (interior) mangrove 

types (p-value = 0.84). Riverine swamp basal 

areas were significantly higher than either 

fringing or basin, with p= 0.005 for riverine vs 

fringing swamps and p= 0.019 for riverine vs 

basin swamps. Basal area values for the sites 

are grouped by geomorphic type in Figure 4.29. 

Population Turnover in the Avicennia Forest 

The higher rates of mortality observed 

during the interval between the second and third 

samplings might have been due to a moderate 

drought; precipitation in northern Guyana for 

January - June 2001 was less than 40% of the 

1961-1990 average (Waple et al. 2002). Basal 

areas of living trees did increase over both of 

those intervals, 1.11 m 2 /ha in the first interval 

and only 0.8 m 2 /ha in the second. That indicates 

that the Avicennia forest sampled is still a young, 

growing stand that has not yet reached 

equilibrium since establishment. The loss of 

stems as the plots mature is a typical selfthinning 

pattern of a young forest. Avicennia, 

like other tropical tree species, does not form 

annual growth rings found in temperate trees, 

preventing direct analysis of population 

structure. However, patterns of fluctuating tree 

frequencies among dbh classes seen in both 

burned and unburned plots (Figures 4.15 and 

4.16) suggest intermittent periods of 

recruitment, which is also consistent with 

variable numbers of seedlings in herbaceous 

layers of the plots, with the low number of 

sapling sized trees, and with the low success 

rates of plantings of Avicennia propagules 

(Chapter 5). 

Figure 4.28. Compiled basal area values grouped into 

5-degree latitudinal classes. 

Figure 4.29. Compiled basal area values grouped by 

geomorphic type of the mangrove forests. 

Herbs and Seedlings 

Coverage of herbs in both burned and 

unburned plots was highly variable in both cover 

and species composition. In the unburned areas, 

there was significant cover of only Lemna during 

wet periods, with the alga Rhizoclonium 

africanum on Avicennia pneumatophores also 

apparent when swamps were flooded. There was 

almost no herbaceous cover in unburned 

swamps during dry periods, and what was 

observed mainly consisted of infrequent 


The year of the sampling shows a strong 

relationship to species in the PCA ordination of 

the herbaceous data, most likely driven by 

variations in rainfall. While hydrology was 

apparently a major influence on herbaceous 

cover changes, other environmental factors such 

as nutrient levels could have an effect on 

competition between Lemna and Limnobium, as 

could timing of their dispersal to the burned site. 

Lemna was present in nearby unburned 

Avicennia forest, which may have given it an 

immediate advantage of proximity for early 

colonization, while Limnobium is possibly a 

more aggressive occupier of space in the open 

water of the burned swamp. Alternanthera is 

evidently comparatively drought resistant and 

almost maintained its 2000 level of coverage 

into the very dry sampling period in early 2001, 

after which it expanded further. 

Avicennia seedlings seem to undergo cyclic 

establishments with high mortality and only very


rare graduation to larger sizes. This is evidenced 

in the very sparse density of saplings and small 

trees that were found in the plots. In general, 

the young, unburned Avicennia forest conformed 

to common concepts of sparseness of mangrove 

understories (Janzen 1985; Lugo 1986), 

although exceptions were seen in dense 

understories that were observed in older, mixed 

mangrove stands in parts of the Waini Peninsula 

in areas of sparse canopy and treefall 

disturbance. In general, the number of 

understory and epiphytic species found in the 

mangrove swamps increased with distance from 

the ocean and its salinity. Certainly the set of 

plant species that can survive the salinity and 

anaerobic soils of mangrove swamps is limited, 

and that number may be further reduced when a 

forest is intact, in which case there would be 

high competition for light, nutrients and water. 

The unburned Avicennia forest would 

presumably have low rates of treefalls because 

it was a young stand, and so there would be 

fewer gaps with increased resources. The 

unburned forest plots were located near the 

ocean where soil salinities reached very high 

levels during droughts (in excess of 50 psu in 

October 2001, T. Hollowell, unpublished data), 

which may also have eliminated unadapted 

understory species. In the Avicennia forest near 

Waini Point there was additionally a very high 

density of pneumatophores, which play a role 

in limiting establishment or survival of 

understory species, although such an effect has 

apparently not yet been addressed in mangrove 

ecological studies. Snedaker and Lahmann 

(1988) had to impose a strict definition of 

mangrove communities, including frequent tidal 

inundation and lack of disturbance, in order to 

maintain the concept of a characteristically 

sparse mangrove understory. However, it may 

be best to accept that mangrove systems are so 

variable in geomorphology, hydrology, and 

disturbance regimes that understory densities 

can vary widely, with the stress of salinity 

favoring more extreme values at the low end. 

Role of Water Levels 

Water levels in the burned swamp had 

obvious effects on changes in herbaceous 

species presence and cover. In the burned 

swamp, standing water, bright sunlight, and 

77 

increased nutrient levels led to high densities 

of floating aquatics, at first Lemna 

aequinoctialis (Figure 4.30) and later 

Limnobium laevigatum (Figure 4.31). For 

several months after the fires there were 

substantial areas of submerged filamentous 

green algae, probably flourishing in the elevated 

levels of nutrients released by the fires. The 

unburned Avicennia forest also had areas of 

substantial Lemna cover during the period of 

highest water. Surface water levels were farther 

above the substrate in the burned swamp after 

the fires due to both the lowering of the substrate 

by burning of organic layers and most likely 

reduced evapotranspiration from trees. When 

Figure 4.30. Burned Avicennia forest near Almond 

Beach, at plot 5, in November 1998. The herbaceous 

vegetation is almost entirely dominated by the small 

floating aquatic Lemna aequinoctialis. 

Figure 4.31. Burned Avicennia forest near Almond 

Beach, at plot 4, in May 2000. The herbaceous 

vegetation is dominated by the floating aquatic 

Limnobium laevigatum, with some scattered clumps 

of small Acrostichum ferns and Cyperus.

78 


water levels in the burned swamp fell during 

droughts, the sprawling herb Alternanthera 

sessilis, which was found only on a few slightly 

elevated spots during earlier samplings, became 

the dominant herb (Figure 4.32). In later 

samplings of herbs in the burned plots, with the 

water table far below the surface, herbaceous 

vegetation was scarce, and the soil surface was 

covered with salt crystals. In the unburned 

forest, the drier periods also coincided with 

lower numbers of Avicennia seedlings sampled. 

In unburned plots following the drought, the 

increases in Avicennia tree dbh and basal area 

were also lower, with an mean increase of 0.3 

cm dbh and 25.7 cm 2 basal area in the first 

(wetter) interval compared to 0.04 cm dbh and 

18.5 cm 2 basal area in the second (drier) interval. 

The water level in the burned plots was 

consistently slightly higher than in the unburned 

plots until early 2000, after which the level was 

slightly higher in the unburned plots. It is not 

certain whether that was attributable to changes 

in evapotranspiration as vegetation in the burned 

plots changed or if there were changes in the 

drainage via creeks and lagoons from the burned 

swamp. As the early 2001 drought intensified, 

water table levels in both areas dropped 

dramatically, with levels in the unburned swamp 

falling more, possibly because of continued 

evapotranspiration from trees of the mature 

forest, while in the burned swamp herbaceous 

vegetation cover fell sharply. When 

measurements were discontinued in March 

2001, the water table in the unburned swamp 

was 73 cm below the surface, while in the burned 

area it was 35 cm below surface. After water 

levels in the burned swamp fell below the soil 

surface, evaporation would be greatly reduced, 

while in the unburned swamp the trees would 

continue to transport water into the atmosphere. 

CONCLUSION 

Many aspects of the mangrove forests of 

northern Waini Peninsula are linked to cycles 

of drought and rain, particularly because the 

swamps are often isolated from tidal influences. 

Dry periods may increase the chances of fire 

disturbances and serve to control presence of 

non-halophytic species. Wet periods may allow 

Figure 4.32. Burned Avicennia swamp near Almond 

Beach, between plots 4 and 6, in November 2001. 

The herbaceous vegetation is dominated by 

Alternanthera sessilis. There are also clusters of 

Acrostichum ferns and scattered young, shrubby 

Laguncularia trees (left foreground). 

wider dispersal of propagules, and water levels 

may control which species can become 

established. Herbaceous vegetation was 

observed to vary greatly with wet and dry cycles. 

The data here have provided some evidence that 

these forests were recently established and that 

seedling recruitment has been intermittent. 

Coastal Avicennia swamps in the Waini 

Peninsula area have been apparently increasing 

in biomass over time. Many of the mangrove 

swamps of the northern part of the Waini 

Peninsula are near monocultures of Avicennia, 

however over time occasional dispersal of 

Rhizophora and Laguncularia may add to their 

species diversity, as part of the slow transition 

to a mixed coastal mangrove forest if they are 

undisturbed. It seems very likely that the forests 

of the Waini represent a chronosequence of plant 

community development, with the youngest, 

least developed areas found nearer to Waini 

Point. In comparison to mangrove swamps in 

other parts of the world, the Waini coastal 

mangroves are fairly average in basal area and 

biomass, but very low in stem density. The 

Avicennia dbh and height class distributions for 

both burned and unburned sites suggest that 

seedling recruitment has occurred in episodes, 

and that can help to explain the very low density 

of seedlings that were observed in the unburned 

swamp. It can be hypothesized that the low


density reflects the climate-driven hydrological 

variations of the non-tidal swamps, with 

recruitment only occurring during rare periods 

of optimal conditions. 

Following the soil fires, the species 

diversity of the disturbed coastal Avicennia 

swamps increased, primarily because of an 

increase in opportunistic species that disperse 

quickly into the available space. The herbaceous 

component of those post-disturbance swamps 

changed rapidly, apparently linked to 

fluctuations in the weather through its effects 

on hydrology and salinity. The composition of 

the tree community in the burned areas appears 

to be changing from a near monoculture of 

Avicennia to domination, for the immediate 

future, by Laguncularia trees, driven by 

scattered survivors of the species and effective 

dispersal of their relatively small fruits. In 

unburned swamps of Almond Beach 

Laguncularia is a common understory species 

at intermediate distances from the coast. 

Laguncularia is a comparatively weedy 

mangrove species and has been seldom observed 

to dominate mangrove forests in the Guianas, 

and it will be of great interest to know if that 

species will give way to other mangrove 

dominants. 

The Laguncularia trees that initially 

colonized the burned site were generally very 

low and spreading, which could lead to a forest 

of low biomass. Some Amazonian forests have 

been found to return to pre-burning biomass and 

79 

dbh values, although not of similar species 

composition, within 40 years (Ferreira & Prance 

1999), and long-term monitoring of the forest 

structure in the Almond Beach burn would be 

useful for comparison to those observations, in 

order to see if mangrove systems behave 

similarly to upland tropical systems. 

The mangrove forests of the Caribbean, 

which occur more often in areas of lower 

sedimentation, on calcareous substrates 

relatively low in nutrients, are generally more 

limited in extent than those of the Guianas 

(Sealey & Bustamante 1999). In several 

respects, the mangrove ecosystems of the 

Guianas are more geomorphically similar to 

those of high sediment coasts of South Asia 

rather than to those of the Caribbean (Jelgersma 

et al. 1993). For instance, in some river deltas 

of India Avicennia is considered the predominant 

pioneer (Blasco 1975), as the genus is in the 

Guianas. The Avicennia forests of the Guianas 

were shown here to be somewhat distinct from 

other Neotropical mangrove ecosystems, which 

might be a factor in decisions regarding their 

protection. The Amazon is the world’s largest 

river, and its coastal delta is globally unique and 

includes the coast of the Guianas. This 

description of the most seaward portion of the 

Waini Peninsula swamps should serve to 

stimulate additional investigations and to raise 

awareness of the unique properties of the 

region’s forests.

80 


Lloyd Savory and some of his students from 

Moruca, including his daughter, after assisting 

with setup of some mangrove planting trials in 

burned Avicennia swamp near Almond Beach. 

Peter Pritchard, a marine turtle biologist and 

GMTCS collaborator, exhuming the skeleton of 

a Leatherback marine turtle that had been buried 

after washing ashore on Almond Beach several 

months earlier. 

Annette Arjoon, a founder of the Guyana Marine 

Turtle Conservation Society, on Almond Beach 

in May 2000. The Almond Beach GMTCS camp 

was used as the base for these mangrove 

investigations. 

Students from Moruca returning to Almond 

Beach camp from Waini Point after a full night 

of beach patrol to monitor nesting marine turtles.


CHAPTER 5. 

DISPERSAL AND ESTABLISHMENT OF MANGROVE 

PROPAGULES FOLLOWING FIRES 

INTRODUCTION 

In 1998 a mass mortality of trees in 

mangrove and freshwater swamps occurred after 

fires burned the soils of a large portion of the 

Waini Peninsula, Guyana. The affected 

mangrove swamps were primarily pure 

Avicennia germinans (black mangrove) stands. 

However, Laguncularia racemosa (white 

mangrove) trees were also present in the 

understory in scattered locations prior to the fires 

(see Chapter 4), and Rhizophora mangle (red 

mangrove) was present infrequently along the 

swamp margins by the beach ridge. Some 

scattered individuals of those species survived 

in the burned swamp. The path of early 

vegetation regeneration in these swamps will 

apparently be highly affected by the dispersal 

and establishment abilities of the surviving 

species in the environments after the fires. 

This chapter provides an analysis of 

mangrove dispersal and establishment patterns, 

in order to establish whether mangrove 

dispersal, like terrestrial dispersal, is influenced 

by size of the seed or propagule. Such 

information has not been widely collected and 

analyzed for mangrove species in the 

Neotropics. A method was developed for 

collection of those data. Planting trials in the 

field with regional mangrove species were 

undertaken to provide additional information on 

establishment, independent of dispersal ability, 

in the environments of burned and unburned 

Waini Peninsula mangrove swamps. 

Information on the post-fire dispersal and 

establishment of mangrove species should allow 

insights into the early recovery process in the 

Waini Peninsula swamps. 

Seed Dispersal 

Early vegetation succession concepts such 

as described by Clements (1928) were largely 

deterministic, regarding disturbed plant 

communities as inexorably moving through a 

81 

sequence of community stages towards a fixed 

climax determined primarily by climate, with 

vegetation modifying edaphic conditions 

through the stages. Such “facilitation” models 

gave little consideration to seed availability or 

plant life history traits and could not explain 

regeneration patterns observed after fire 

disturbances (Bond & van Wilgen 1996). 

Gleason (1927) began to incorporate the role of 

chance into concepts of vegetation change. Later 

Egler (1954) made a full distinction between the 

facilitation or “Relay Floristics” of Clementsian 

succession and an “Initial Floristic 

Composition” model in which the course of 

succession might depend on the species that 

survive or disperse to first colonize a disturbed 

site. Connell and Slatyer (1977) incorporated 

relay floristics and Egler’s initial floristic 

composition concept into their set of three 

alternative models, which were applied to 

situations depending on organism adaptations, 

edaphic variables and the level and size of 

disturbances. Their elaboration of a variety of 

succession models reflected a growing 

recognition that plant communities are dynamic 

entities to which no single successional model 

can be applied (Pickett et al. 1989; White & 

Pickett 1985), and where seed availability is 

often a critical component. 

Dispersal distribution curves, or seed 

shadows, for seeds from individual trees tend 

to follow leptokurtic pattern that peaks a short 

distance from the parent, with a long tail (Harper 

1977; Willson & Traveset 2000). However, there 

is still limited data available for comparison and 

analysis of the dispersal curves for various 

modes (Willson & Traveset 2000). The tails of 

dispersal curves are difficult to model (Bullock 

& Clarke 2000), and the longest distance 

dispersal events may be the result of unusual 

high winds or turbulence, which can affect 

smaller fruits that are not normally thought of 

as wind dispersed (Jongejans & Telenius 2001). 

Extremely long-distance dispersal is rare, but it

82 


is probably important for colonization of 

suitable habitats after large-scale disturbances. 

Dispersal in Mangrove Swamps 

Coastal mangrove swamp attributes such 

as nutrient levels and salinity are strongly 

influenced by geomorphic setting (however see 

McKee 1993), so succession models based on 

organism modification of conditions may not 

be as applicable as the initial floristic 

composition, or inhibition model of Connell and 

Slatyer (1977). Therefore in the Waini Peninsula 

swamps, dispersal and establishment of species 

after a disturbance may have a strong influence 

on early recovery. Smaller disturbances such as 

tree falls are common in all mangrove swamps 

but would not have serious effect on species 

composition, as the short dispersal distances 

allow rapid regeneration (Baldwin et al. 2001; 

Clarke & Kerrigan 2000; Duke 2001; Sherman 

et al. 2000). 

Water dispersal is the major mode of 

dispersal for mangrove propagules, which are 

typically transported by tidal currents. Water 

dispersal (hydrochory) is covered only briefly 

or not at all in dispersal ecology reviews 

(Chambers & MacMahon 1994; Harper 1977; 

Westoby et al. 1997). Water dispersal is 

understandably linked to currents for longer 

distance movement. Water dispersal has been 

observed to be directional in inland North 

American bottomland hardwood swamps, 

where dispersal distance was minimal at times 

of low water flow (Schneider & Sharitz 1988). 

As with a majority of species in wet tropical 

environments, mangrove establishment is 

restricted by the inability of seeds or propagules 

to lay dormant in a seed bank. 

Some of the best known studies of 

mangrove dispersal properties were published 

by Rabinowitz (1978a; 1978b). She suggested 

that mangrove zonation may be controlled, in 

part, by variation in abilities of mangrove 

species to colonize segments of the intertidal 

zone, due to influences of size and shape of each 

species’ propagules. The result would be 

propagule sorting on the seaward edge of 

swamps. The propagules of the three principal 

mangrove genera in the Neotropics and the 

Waini Peninsula area, Rhizophora, Avicennia 

and Laguncularia, are of distinctly different 

sizes. Rabinowitz’s (1978a; 1978c) 

measurements of Neotropical mangrove 

propagule weights in Panama gave the mean 

weight of Laguncularia propagules as 0.41 

grams, Avicennia as 1.1grams, Rhizophora 

mangle as 14.0 grams, and Rhizophora 

racemosa (cited as R. harrisonii) as 32.3 grams. 

These sizes are in general agreement with the 

those of propagules found in northern Guyana 

(Figure 5.1). 

Most mangrove dispersal studies have been 

set in swamps with tidal currents. Mangrove 

dispersal is simplified in swamps that are rarely 

inundated by tides. Published reports of 

mangrove dispersal distances vary from only a 

Figure 5.1. Propagules of Rhizophora mangle, 

Rhizophora racemosa, Laguncularia racemosa, and 

Avicennia germinans (from upper left, clockwise). 

The Avicennia and Laguncularia propagules are 

indicated in the box. For scale, the pencil is 

approximately 16 cm long.


few meters in a swamp in northwestern Australia 

with little tidal influence (McGuinness 1997a), 

to several or even tens of kilometers for 

Avicennia marina in tidal coastal environments 

(Clarke 1993). This author has found viable 

Rhizophora mangle propagules on beaches near 

Cape Hatteras, North Carolina, more than 850 

kilometers along the Gulf Stream from the 

northernmost Rhizophora stands of Florida 

along the Indian River Lagoon. 

As pointed out by McGuinness (1997a), 

there have been fairly few studies of the role of 

dispersal in mangrove community patterns. 

Rabinowitz’s assertion that tidal sorting of 

propagules by size played a major role in 

mangrove zonation has been debated up to the 

present, with several more recent papers 

investigating physico-chemical correlates of 

zonation (Jiménez & Sauter 1991; López- 

Portillo & Ezcurra 1989; McKee 1993, 1995b; 

McKee & Faulkner 2000). The dominance of 

Avicennia on the seaward margins of the 

Guianas coast does not conform to the 

Rabinowitz tidal sorting model, possibly 

because of the unusual mudflat environment. 

Clarke et al. (2001b) suggest that tidal sorting 

may be less important in the zonation of 

Australian mangroves than establishment 

characteristics of species. On the mudflats of 

the Guianas, Avicennia may be best adapted to 

root quickly during extended periods of very low 

tides (Figure 5.2). Wells and Coleman (1981) 

proposed that rapid growth of newly established 

Avicennia stands on the mudflats of the Guianas 

might raise substrate elevation significantly with 

added root biomass, increasing the probability 

that new stands would persist. 

Dispersal in Disturbed Mangrove Swamps 

Only a few studies have addressed 

mangrove dispersal in disturbed sites. Blanchard 

and Prado (1995) found that establishment of 

Rhizophora mangle in cleared plots in Ecuador 

dropped off significantly 5 meters from plot 

edges, and Lema Vélez (2003) reported similar 

patterns in a disturbed site in Colombia. In 

studies of both oil spill and hurricane 

disturbances in Guadeloupe, Rhizophora mangle 

propagules were found to disperse very poorly 

in areas inland from coasts or rivers (Imbert et 

83 

al. 2000). Elster et al. (1999) assumed that in a 

hydrologically disturbed mangrove swamp with 

little tidal influence, at Ciénaga Grande de Santa 

Marta, Colombia, large open areas remained 

bare because of dispersal limitations of 

mangrove propagules. If mangrove dispersal in 

non-tidal conditions is poor, the initial 

composition of a disturbed swamp might have 

lasting effects, since regeneration would be 

mostly dependent on very short range dispersal. 

In the burned swamps at Almond Beach, 

dispersal may have the most significant effect 

on which species dominate after the fires, and 

could influence how far mangrove trees will 

spread into areas where no parent trees survived. 

In detailed studies of seed dispersal ecology, 

patterns of seed dispersal are separated from 

establishment success rates, generally through 

the use of seed traps. However, seedling 

establishment patterns have been used as a proxy 

for actual seed dispersal, where it is not critical 

to separate seed dispersal from establishment 

success (Nathan & Muller-Landau 2000). 

Establishment distance information can be a 

proxy for dispersal distance if the sites where 

seeds are deposited have fairly uniform 

establishment success rates. That is arguably the 

case for mangrove propagules in the non-tidal 

burned swamp at Almond Beach. Measurements 

Figure 5.2. Avicennia germinans propagules and a 

germinated seedling on the mudflats at Almond 

Beach, Waini Peninsula. This image suggests that 

Avicennia may be able to take advantage of cracks 

in mudflats as a stable site for establishment. The 

germinating propagule is located approximately 2 

meters out from the narrow beach ridge near the 

unburned Avicennia swamp plots.

84 


of establishment distances are therefore a 

particularly useful and convenient method for 

predicting the spread of populations. 

Sites where mangrove propagules tend to 

strand may also be favorable for establishment 

in the burned Almond Beach swamps. This was 

evidenced by clusters of seedlings at the edge 

of pools (Figure 5.3) or in shallow water along 

barriers such as logs, branches or masses of 

Figure 5.3. Laguncularia seedlings concentrated in 

stranding lines, around the edge of a small pool, the 

result of very short distance dispersal. The branches 

of the parent tree are visible on the right. Photo taken 

October 2001. 

floating vegetation such as Lemna and 

Limnobium (Figure 5.4) that have been dense 

at times after the fires. In intact Avicennia 

swamps pneumatophores are often quite dense 

and apparently impede dispersal in the unburned 

swamp, and seeds are found among 

pneumatophores directly under parent trees 

(Figure 5.5). Since they are dispersed in early 

stages of germination, stranded mangrove 

propagules are relatively quick to advance to 

seedling status. In the open environment of the 

burned swamp at Almond Beach, established 

seedlings grew rapidly and reached reproductive 

status quickly, which would increase 

colonization rates. 

To begin to answer some of the questions 

posed above, two experiments were set up. 

These were designed to test the dispersal 

properties of the three mangrove species and 

their ability to establish successfully in the 

swamp environments of the Waini Peninsula. 

Figure 5.4. Clustering of Avicennia seedlings along 

minor barriers, here the branches of trees that were 

killed by the soil fires three years before this image 

was taken in April, 2001. The surviving parent tree 

was directly behind the photographer, about 2 meters 

from this cluster. This image also illustrates the 

condition of the soil surface during the minor drought 

at the time, cracked with salt accumulations on the 

surface. The other plants seen are the sedge Cyperus 

odoratus. 

Figure 5.5. Avicennia propagules collected among 

pneumatophores under a parent tree in undisturbed 

non-tidal swamp behind a beach ridge, approximately 

20 meters from the ocean. 

METHODS 

Study Site 

The study site is located in the Almond 

Beach vicinity of Guyana’s Waini Peninsula in, 

burned and unburned sections of Avicennia 

mangrove swamp. A detailed site description is 

given in Chapter 1. For this study, an area of 

approximately 0.75 km 2 was examined to locate


suitable parent trees for the dispersal 

investigation, and planting trials were sited near 

the 0.1 ha vegetation monitoring plots in burned 

and unburned Avicennia swamp (see Figure 4.1). 

Propagule Dispersal and Establishment 

Most procedures for tracking dispersal of 

mangrove propagules have relied upon the 

marking of seeds on the parent tree and then 

attempting to find them in the environment 

following dispersal, or collection of data in a 

neighboring environment free from the subject 

species. In the case of the burned swamp at 

Almond Beach, parent Laguncularia racemosa, 

Avicennia germinans, and Rhizophora mangle 

trees were scattered to the degree that 

established seedlings encountered while moving 

from a parent in the downwind direction for 100 

meters were likely derived from that parent. 

That, in addition to the absence of tidal currents 

from the swamps, provided a blank canvas of 

sorts, in which the supply and properties of 

propagules could be investigated. 

In the burned Avicennia swamp, surviving 

reproductive trees were located near the Almond 

Beach camp. Establishment patterns were 

sampled for 16 Avicennia and 16 Laguncularia 

trees, while only five fertile Rhizophora parent 

trees were located. Rhizophora trees were rare 

in the area, occurring only along the swamp’s 

boundary with the beach ridge, and those located 

were frequently without fruit. Some points 

considered as single Laguncularia parents were 

clusters of several shrubby individuals. A 5 

meter wide transect was started at the edge of 

each parent tree’s crown and followed for 100 

meters in the direction of the prevailing winds, 

approximately toward the west-southwest (260 

degrees). Transect width was tracked with a 

collapsible fiberglass dome tent pole, which 

covered the 5 meter wide swath when held 

perpendicular to the transect line and allowed 

to flex (Figure 5.6). The number of established 

seedlings (from plants with cotyledons exposed 

to plants 50 cm high) was recorded for each 0.1 

meter of the transect. Significant barriers, 

generally logs and branches of fallen trees, were 

noted. 

A simple initial model was formed to 

compare potential spread of Laguncularia, 

Avicennia, and Rhizophora in the burned 

85 

Avicennia swamp at Almond Beach. Spatial 

buffering was performed in ArcView 3.3 (ESRI 

2002), in which polygons of expected 

establishment distance are formed at a uniform 

distances from the parent trees according to 

species. Based on observations of the 

reproduction of these species in the burned 

swamp environment, it was assumed that 

Laguncularia reached reproductive age after 

about three years, Avicennia after about five 

years, and Rhizophora after about eight years. 

Buffers were generated for five generations of 

Laguncularia, three generations of Avicennia, 

and two generations of Rhizophora, or about 15 

years (16 for Rhizophora).Mean establishment 

distances from the transects for each species 

were used as the distance that the range 

expanded from each individual over one 

generation. 

Propagule Plantings 

Plantings of propagules of four mangrove 

species were performed to assess the potentials 

of the common mangrove species in the area to 

establish and grow under a variety of 

hydrological conditions. Resulting information 

can then be considered in terms of different plant 

community succession models and potential 

ecosystem restoration strategies. 

Mature propagules for the plantings were 

collected from surviving trees within or on the 

Figure 5.6. Method for counting established 

Laguncularia using a Ahip chain@ measuring device 

and a pole spanning 5 meters when flexed. The area 

pictured was one of the most heavily vegetated with 

Laguncularia shrubs.

86 


margins of the Almond Beach Swamp for 

Avicennia germinans, Laguncularia racemosa, 

and Rhizophora mangle, and propagules of 

Rhizophora racemosa were collected from trees 

along the banks of the Mora Passage. Avicennia 

propagules were approximately 2-2.5 cm in 

length, Laguncularia propagules 1.5-2 cm in 

length, Rhizophora mangle propagules 15 cm 

in length, and Rhizophora racemosa propagules 

35 cm in length (Figure 5.1). 

Plantings were made in both burned and 

unburned swamp sites, during flooded and dry 

conditions, and at the normal swamp level and 

at a level elevated 15 cm above typical swamp 

surface level. Water levels at the time of the 

flooded plantings were well above the soil 

surface of the lower level plantings and a few 

cm above the soil surface of the elevated 

plantings. No surface water was present at an 

level at either site when dry condition plantings 

were made. 

In the burned swamp site, plantings for the 

elevated treatment were made by anchoring 

aluminum flashing rings that were 20 cm-high 

and 1 m 2 area. The rings were anchored in place 

and filled to 15 cm with soil from nearby. In 

the unburned swamp site, a slightly elevated 

area existed in the planting area and was utilized 

for the elevated treatments. Water depths during 

the wet season plantings were similar between 

the treatments for the two sites. Sixteen 

propagules were planted for each treatment. The 

two Rhizophora species were planted by 

inserting propagules upright with points 

approximately 5 cm into the soil surface, while 

the Laguncularia and Avicennia propagules 

were secured in place with U-shaped thin wire 

clips. For the wet condition plantings, 

measurements were made after 10.5 and 17 

months. Measurements were made for 1) the 

number of nodes on each seedling, 2) the height 

of each seedling, 3) the number of branches, 

and 4) the number of prop-roots developed in 

Rhizophora trees. The second, dry condition, 

planting was made of the same species in both 

burned and unburned areas, around the time of 

the first data collection for the wet season 

planting, in April 2001 during a moderate 

drought with no standing water in the swamps. 

The seedlings resulting from the dry condition 

plantings were measured after about 6 months, 

in October 2001. 

RESULTS 


Sixteen Laguncularia and 16 Avicennia 

parent trees were located and sampled. Only five 

fertile Rhizophora parent trees were located in 

the vicinity, all growing along or near the 

swamp/beach ridge boundary. In the combined 

100 meter transects, a total of 598 Laguncularia 

seedlings, 678 Avicennia seedlings, and 74 

Rhizophora seedlings were sampled. Full counts 

for each species sampled are summarized in 

Table 5.1, and distribution curves based on those 

data are presented in Figure 5.7 for 

Laguncularia, Figure 5.8 for Avicennia and 

Figure 5.7. Dispersal curve for establishment of 

Laguncularia seedlings, for all 100 meter transects 

combined, with the logarithmic curve that best fitted 

those data. 

Figure 5.8. Establishment distance curve for 

Avicennia seedlings, combined 100 meter transects, 

with the logarithmic curve that best fitted those data.


Figure 5.9 for Rhizophora. The range of 

numbers of seedlings encountered from an 

individual parent varied from 2 to 155 in 

Laguncularia, 1 to 237 in Avicennia, and 3 to 

39 in Rhizophora. The mean establishment 

distance was 24.2 meters for Laguncularia, 4.8 

meters for Avicennia, and 8.9 meters for 

Rhizophora. Summary statistics for 

establishment distance measurements are given 

in Table 5.2. 

The three resulting establishment distance 

data sets were tested for significant difference 

among themselves with pairs of Wilcoxon ranksum 

tests (nonparametric, as distributions were 

not normal). In addition, a one-sample 

Kolmogorov-Smirnov test for each species was 

used to test if established seedlings distances 

were reasonably drawn from an exponential 

distribution with the mean for that species. In 

all Wilcoxon rank-sum tests of pairs of 

comparisons, p-values were less than 0.005. The 

maximum establishment distances recorded for 

each species were 100 meters for Laguncularia, 

92.1 meters for Avicennia, and 49 meters for 

Rhizophora. All curves were significantly 

different from artificial uniform distributions of 

values ranging from 0 to the species maximum. 

In the case of Laguncularia (p= 0.045, onesample 

Kolmogorov-Smirnov test) the 

distribution approached a less significant 

difference from 7 exponential distributions with 

the species’ mean. 

For graphing, the seedling establishment 

points were condensed into one meter interval 

groups. Those displayed degrees of bimodal or 

polymodal distributions, most distinctly in 

Rhizophora and Avicennia. Best-fit natural 

logarithmic lines were calculated for the 

dispersal data grouped into 1 meter increments, 

for the three species. The lognormal decay 

curves achieved a somewhat better fit to the data 

than linear or binomial curves. For Laguncularia 

the formula for the logarithmic curve was y = - 

6.2495Ln(x) + 29.183, R 2 = 0.56 (Figure 5.7), 

for Avicennia it was y = -20.116Ln(x) + 79.265, 

R 2 = 0.487 (Figure 5.8) and for Rhizophora it 

was y = -1.7536Ln(x) + 7.1354, R 2 = 0.559 

(Figure 5.9). All three curves are displayed in a 

single graph in Figure 5.10. 

For the buffering model, the mean distances 

obtained from the establishment transects were 

87 

Figure 5.9. Dispersal curve for establishment of 

Rhizophora seedlings, for all 100 meter transects 

combined, with the logarithmic curve that best fitted 

those data. 

used: 24.2 meters for Laguncularia, 4.38 meters 

for Avicennia, and 8.94 meters for Rhizophora. 

The points modeled for Rhizophora and 

Laguncularia were those for all parent trees in 

the establishment transects, while 39 additional 

mature Avicennia trees were located in the area 

and marked with GPS, for a total of 55 used. 

One Rhizophora point was located to the west 

of the boundary that had been drawn for the 

Almond Beach burn, in an area the had 

apparently been impacted by scattered soil fires, 

without full mortality of the surrounding trees. 

The resulting buffer areas after the specified 

intervals were 57.4 ha for Laguncularia, 2.83 

ha for Avicennia, and 1.0 ha for Rhizophora; out 

of the estimated 5.17 km 2 (517 ha) area of 

Avicennia swamp burned at the Almond Beach 

site. For Laguncularia, that equated to 11.1% 

of the burned area. The buffering result maps 

are shown in Figures 5.11, 5.12, and 5.13, 

respectively. 


In both burned and unburned treatments, 

plantings of the two Rhizophora species were 

the only species to survive to the first census, 

and all plantings of Laguncularia and Avicennia 

propagules failed. In the burned swamp, 

Rhizophora racemosa treatments grew robustly 

(Figure 5.14); however, substrate-level R. 

mangle experienced apparent full mortality. 

After the first 11 months in the burned treatment, 

the mean height of the 14 surviving elevated R. 

mangle seedlings was 71.5 cm, the mean height 

of the 16 elevated R. racemosa seedlings was 

128.9 cm, and the mean height of the 16

88 


Table 5.1. Number of seedlings found within one meter intervals away from parent trees, within a 5 meter 

wide band, totaled for all replicates within the three species sampled for establishment distance. 

Avicennia 

m count 

0 91 

1 170 

2 152 

3 84 

4 32 

5 29 

6 8 

7 6 

8 7 

9 5 

10 28 

11 18 

12 4 

13 11 

14 4 

15 4 

16 1 

20 2 

22 2 

23 2 

24 3 

26 2 

29 1 

31 4 

51 1 

56 1 

60 1 

63 1 

65 1 

88 1 

92 2 

Laguncularia 

m count 

0 7 

1 26 

2 35 

3 41 

4 18 

5 14 

6 15 

7 9 

8 10 

9 18 

10 7 

11 26 

12 26 

13 20 

14 16 

15 22 

16 5 

17 20 

18 9 

19 15 

20 7 

21 10 

22 10 

23 10 

24 6 

25 7 

26 3 

28 4 

29 32 

30 4 

31 1 

32 3 

34 7 

35 3 

36 1 

38 3 

39 2 

40 4 

42 1 

43 4 

45 1 

46 2 

47 1 

48 1 

49 7 

50 7 

51 5 

52 1 

54 2 

55 4 

56 5 

57 5 

58 1 

60 5 

61 5 

62 2 

63 1 

65 1 

66 1 

67 1 

68 1 

69 6 

70 10 

71 2 

72 2 

73 2 

74 4 

78 1 

79 3 

80 1 

83 1 

84 1 

85 1 

86 2 

87 5 

88 1 

89 3 

91 1 

92 2 

93 1 

94 3 

95 3 

96 1 

100 2 

Rhizophora 

m count 

0 6 

1 13 

2 5 

3 11 

4 6 

5 3 

6 5 

7 5 

8 5 

9 1 

10 2 

11 1 

19 1 

27 1 

34 1 

40 3 

42 1 

43 2 

49 2


Figure 5.10. Comparison of seedling establishment distributions for Rhizophora, Avicennia, and Laguncularia, 

combined 100 meter transects. 

Table 5.2. Summary values for establishment distance measurements. 

N 

Mean 

Mode 

Minimum 

Maximum 

Std. Dev. 

Variance 

Avicennia 

675 

4.38 

2.1 

0 

92.1 

8.86 

78.54 

Laguncularia 

597 

24.20 

15.3 

0 

100 

24.47 

598.63 

Rhizophora 

74 

8.94 

3.75 

0 

49 

13.19 

173.99 

89

90 


Figure 5.11. Graphic results of buffering of 

Laguncularia points in the Almond Beach swamp, 

based on five three-year generations. Total area 

covered was 57.4 ha. 

substrate-level R. racemosa seedlings was 

103.25 cm . At the second sampling 17 months 

after plantings there was no additional mortality; 

the mean height of the elevated R. mangle 

seedlings was 118.6 cm, the mean height of 

elevated R. racemosa seedlings was 197 cm, and 

the mean height of substrate-level R. racemosa 

was 159.4 cm. The mean values for height, node 

count, prop root count, and for the first 

measurement (10.5 months) are graphed in 

Figure 5.12. Graphic results of buffering of Avicennia 

points in the Almond Beach swamp, based on three 

five-year generations. Total area covered was 2.83 

ha. 

Figure 5.13. Graphic results of buffering of ten Rhizophora points in the Almond Beach swamp, based on two 

eight-year generations. The total area covered after that interval was 1 ha. The points included the five fertile 

trees used for establishment distance measurements and five others that were also located in the vicinity. The 

approximate location is indicated for the lagoon that sometimes connects the Almond Beach swamp to the 

sea. 

Figure 5.15, for the second measurement (17 

months) in Figure 5.16 with the values and 

differences between the two dates listed in Table 

5.3. It should be kept in mind that as planted, 

Rhizophora racemosa propagules were 

approximately 20 cm longer than Rhizophora 

mangle propagules. Height values for both 

Rhizophora species in the two habitats for both 

treatments are shown in Figure 5.17. A series of 

pairwise t-tests was used to compare the mean


Figure 5.14. Rhizophora plantings in April 2001, 

approximately 102 months after establishment. 

Vegetation in the background is primarily 


91 

values for combinations of plantings since 

variances were fairly similar. A matrix of the 

resulting p-values is given in Table 5.4. For the 

height measure an analysis of variance 

(ANOVA) also showed high significance, 

approaching p=0, for all factors, habitat, 

elevation and species, with either the raw data 

or with seedling heights adjusted for the 

difference in length of propagules of the two 

species. 

In the unburned swamp, survival and 

growth of both species of Rhizophora was much 

lower than in the burned swamp, where no 

Laguncularia or Avicennia again survived in any 

treatments. Dry season plantings also resulting 

in survival of only Rhizophora plants, and 

exhibited decreased survival and growth of those 

species, particularly in the unburned swamp. 

Values for the dry season plantings are 

summarized in Table 5.5 and graphed in Figure 

5.18. During the dry season planting there was 

greater effect on Rhizophora mangle plantings, 

which displayed no survival in the elevated 

plantings and no survival in the unburned 

swamp. According to well data from the two 

Figure 5.15. Graphs comparing Rhizophora planting results after 10.5 months, burned site. None of the shorter 

Rhizophora mangle propagules were noted as alive at low (substrate) elevation.

92 


Figure 5.16. Graphs comparing Rhizophora planting 

results, burned site after 17 months. 

Burned 

Unburned 

Burned 

Unburned 

Burned 

Unburned 

elevated 

low 

elevated 

low 

elevated 

low 

elevated 

low 

elevated 

low 

elevated 

low 

R. mangle 

R. racemosa 

R. mangle 

R. racemosa 

R. mangle 

R. racemosa 

R. mangle 

R. racemosa 

R. mangle 

R. racemosa 

R. mangle 

R. racemosa 

R. mangle 

R. racemosa 

R. mangle 

R. racemosa 

R. mangle 

R. racemosa 

R. mangle 

R. racemosa 

R. mangle 

R. racemosa 

R. mangle 

R. racemosa 

Nodes 

9.6 

11.25 

0 

11.75 

6.1 

6.3 

0 

0 

Nodes 

18.1 

17.75 

0 

17.3 

8.6 

6.8 

0 

0 

Nodes 

8.6 

6.5 

0 

5.6 

2.5 

0.4 

0 

0 

Figure 5.17. Mean seedling heights for both species 

of Rhizophora after 10.5 and 17 months, for 

surviving seedlings in elevated plantings, burned and 

unburned areas. 

Table 5.3. Mean values for the variables measured for Rhizophora plantings made in wet conditions in May 

2000, for the first sampling (11 mos.), the second sampling (17 mos.), and the difference between the means. 

These means are from values are for the surviving individuals only out of 16 planted in each treatment. 

Mean values, first sampling, survivors only 

Height (cm) 

71.5 

128.9 

0 

103.25 

53.9 

62.2 

0 

0 

Height (cm) 

118.6 

197 

0 

159.4 

59.1 

64.8 

0 

0 

Height (cm) 

47.1 

68.1 

0 

56.1 

5.2 

2.6 

0 

0 

Prop roots 

0.3 

1.6 

0 

0.6 

0 

0 

0 

0 

Prop roots 

5.3 

5.9 

0 

3.7 

0 

0.1 

0 

0 

Prop roots 

5.0 

4.3 

0 

3.1 

0 

0.1 

0 

0 

Branches 

5.2 

8.1 

0 

6.2 

0 

0 

0 

0 

Mean values, second sampling, survivors only 

Branches 

10 

14.9 

0 

12.1 

0.6 

0.1 

0 

0 

Difference, first to second sampling, survivors only 

Branches 

4.9 

6.8 

0 

5.9 

0.6 

0.1 

0 

0 

Survivors 

14 

16 

0 

16 

10 

14 

0 

0 

Survivors 

14 

16 

0 

16 

10 

9 

0 

0 

Survivors 

0 

0 

0 

0 

0 

-5 

0 

0


Table 5.4. Summary of P-values for pairwise t-tests between combinations of Rhizophora mangle (R.m.) and 

Rhizophora racemosa (R.r.) planting treatments in high vs low elevation plantings and burned vs unburned 

areas. Values significant at P=0.05 are shown in bold face. 

R.m. vs R.r. low 

R.m. vs R.r. high 

R.r. hi vs low 

R.m. burn vs unburn 

R.r. burn vs unburn 











First planting, first census 

Nodes 

n/a 

0.003 

0.219 

0.0005 

0 

Nodes 

n/a 

0.287 

0.542 

0 

0 

Nodes 

0.186 

n/a 

0.088 

n/a 

0.033 

Height 

n/a 

0 

0 

0.006 

0 

First planting, second census 

Height 

n/a 

0 

0.001 

0 

0 

Second planting (dry season) 

sites, the groundwater level in the unburned 

swamp was lower than the burned swamp, 

possibly caused by higher evapotranspiration 

from trees in the living forest. Other factors 

limiting survival and growth in the unburned 

site likely include shading and competition for 

nutrients and water. There were no observations 

suggesting that predation played a role in 

Rhizophora mortality in either site. 

DISCUSSION 


Establishment patterns in the Almond 

Height 

0.788 

n/a 

0.072 

n/a 

0.036 

Prop roots 

n/a 

0.010 

0.065 

n/a 

n/a 

Prop roots 

n/a 

0.228 

0.006 

n/a 

n/a 

Prop roots 

n/a 

n/a 

n/a 

n/a 

n/a 

Branches 

n/a 

0.000 

0.012 

n/a 

n/a 

Branches 

n/a 

0.0002 

0.029 

n/a 

n/a 

Branches 

0.032 

n/a 

0.025 

n/a 

n/a 

Table 5.5. Mean values for variables measured for Rhizophora plantings made in drought conditions in April 

2001. 

R. mangle low 

R.racemosa low 

R.racemosa elev. 

R.racemosa elev. unburn 

Nodes 

5.25 

3.69 

1.69 

0.81 

Height 

(cm) 

34.69 

37.44 

16.25 

8.63 

Prop roots 

0 

0 

0 

0 

Branches 

4.69 

2.50 

0.75 

0 

% Survival 

75 

56 

25 

25 

93 

Beach swamps were similar to those that might 

be expected in upland systems. Although in 

terms of Chambers and MacMahon’s (1994) 

classification of dispersal movement phases, the 

water in the non-tidal Almond Beach swamp 

would qualify as Phase II dispersal, coming after 

the propagule first moves from parent to a 

surface, the establishment curves derived from 

the transects resemble a leptokurtic Phase I 

dispersal pattern, with a peak near the parent 

and a long tail. This is possibly attributable to 

the passive nature of the water that lacks 

substantial currents. The fits of the logarithmic 

curves calculated from the dispersal distance 

data were not particularly good for the three

94 


Figure 5.18. Dry season planting results. R. racemosa 

values are not corrected for its larger propagule 

length, approximately 20 cm. greater than R. mangle. 

species, as all displayed typical irregular Phase 

I type distributions with a peak near the parents 

and with long tails. If there had been substantial 

Phase II dispersal it would be attributable to 

movement in water currents after falling from 

the parent, and it would have the effect of 

additional extension of tails of the curves. It is 

uncertain to what degree the patterns here reflect 

any seed movement from the time the fruits 

leave the parent tree and when they reach the 

water surface, such as during brief periods of 

high winds. 

Propagule dispersal distance in the Almond 

Beach non-tidal environment was apparently 

related, in least in part, to the size of propagules. 

The smaller Laguncularia propagules were 

dispersed farther than Avicennia propagules. On 

the other hand, the mean distance for dispersal 

of the much larger Rhizophora propagules was 

higher than that of Avicennia (8.9 meters vs 4.4 

meters). However, the maximum dispersal 

distance recorded for Rhizophora was lower 

than for Avicennia (92 meters vs 49 meters), 

although that may have been influenced by the 

greater number of Avicennia trees available for 

sampling and the much greater number of 

seedlings established from Avicennia parent 

trees. 

The simplified environment of the burned 

mangrove swamp at Almond Beach provided a 

reduced number of variables in often present in 

natural dispersal experiments. In mangrove 

swamps the number of plant species present is 

limited, and water dispersal tends to deposit 

propagules where establishment rates are high 

(Middleton 2000). There is no seed bank or 

dormant phase for mangrove species, and animal 

dispersal is not an important factor. Fire-related 

mortality in the burned habitat limited the 

number of parent trees that might confuse 

dispersal or establishment patterns, which was 

reflected in these distinct establishment distance 

curves. The effects of predators have probably 

been initially reduced by fire disturbance. In the 

case of the Almond Beach swamp, beach ridge 

barriers block tidal and wave influences, making 

dispersal distance dependent primarily on water 

depth and minor currents generated by winds. 

The beach ridges also limit influx of propagules 

from outside sources. While the non-tidal setting 

is not typically utilized for dispersal studies in 

mangrove swamps, there are often sizable 

portions of mangrove systems that are isolated 

from regular tidal inundation. In some respects, 

the dispersal environments of Almond Beach 

mangrove swamps were more complex than that 

of the ocean/swamp edge studied by Rabinowitz 

1978b. 

While the results of the buffering trials 

provided an initial visual comparison of abilities 

to colonize the burned swamp, there is certainly 

a potential for more advanced propagule 

dispersal and establishment models to better 

predict species spread through consideration of 

the density of colonization events for each 

species, the probabilities that established 

seedlings might advance to reproductive age, 

and the potential for higher seed production of 

older individuals (Clark et al. 2001; He & 

Mladenoff 1999; Tews et al. 2004). Attention 

might also be given to correction for of the 

constant width of the transect from the trees, 

and the influenced of prevailing winds, which 

would call for a directional bias in the creation 

of buffers or chances of dispersal events. 

It is interesting to note that some 

characteristics of “traditional” Neotropical 

mangrove zonation are apparent in the occluded, 

burned swamp once behind the beach ridges. 

Rhizophora is found nearest to the sea, and 

Laguncularia is more common several hundred 

meters inland, beyond the wettest area of the 

swamp. The swamp near the beach ridges is 

often a location of deepest water in the occluded 

swamps, and may allow some local movement 

of the larger Rhizophora propagules. 

While ecological studies on fire-adapted


plant communities tend to focus on life history 

traits rather than competition, in mangrove 

swamps it seems that hydrology will play a 

significant role in the path of recovery, along 

with closely linked factors such as soil salinity 

and soil redox potential, which affect nutrient 

availability. The importance of these factors is, 

however, contingent on availability of 

propagules, and the spread of tree species into 

burned areas of the Almond Beach swamp 

seems to be limited by the availability of 

propagules and patterns of dispersal and 

establishment success that are linked to small 

elevation differences and barriers that offer 

suitable establishment sites. Elster et al. (1999) 

found that in a hydrologically disturbed nontidal 

mangrove swamp in Colombia, few 

propagules reached large areas that were remote 

from parent trees. 

Difficulty of dispersal of the larger 

Rhizophora propagules into and within the 

Almond Beach swamps offer one explanation 

for their absence from the younger Avicennia 

swamps. Once introduced, the species 

apparently persists, as evidenced by increasing 

dominance of Rhizophora in coastal forests 

farther to the southeast on the peninsula, where 

the swamps behind beach ridges are arguably 

older and Rhizophora has had a longer time for 

colonization success. The vigorous growth of 

Rhizophora plantings at Almond Beach attest 

to its ability to thrive once dispersed to that 

location. 

In a marked propagule experiment using 

the Asian mangrove Ceriops tagal (Perr.) 

C.B.Rob. (Rhizophoraceae), which has 

propagules similar in shape and size to those of 

Rhizophora mangle, McGuinness (1997a) 

found that 76% of marked propagules were 

found within 1 meter of the parent tree and 91% 

within 3 meters, and concluded that patterns of 

initial colonization may be reflected in zonation 

for long periods, and that long-term effects of 

competition within physico-chemical 

environments may be overestimated. 

The burned environment at Almond Beach 

presented many barriers to dispersal of all 

mangrove species, and that effect was probably 

compounded by the lack of fluctuating water 

levels and currents from tides. Barriers of fallen 

dead trees have been cited as a probable factor 

95 

in very slow colonization in hurricane disturbed 

Rhizophora swamps in Guadeloupe (Imbert et 

al. 2000). Patches of low herbaceous vegetation, 

usually Alternanthera or Cyperus, acted as 

barriers to dispersal against which many 

seedlings were stranded and became established. 

Similar phenomena were noted by Lema Vélez 

(2003) in Colombian mangroves, where Batis 

maritima L. plants blocked dispersal of 

propagules, and in North American Taxodium 

(bald cypress) swamps, where seeds were found 

concentrated non-randomly along logs, trees and 

other emergent obstacles (Schneider & Sharitz 

1988). Clarke (1993) found that Avicennia 

marina propagules in Australia stranded at the 

line of highest water levels inside of swamps, 

where they shed their pericarps, became nonbuoyant 

and established. All of those 

observations offer some support for application 

of the propagule sorting hypothesis proposed by 

Rabinowitz (1978b) to the physical environment 

of the non-tidal mangrove swamps at Almond 

Beach. Sorting in such swamp interiors is likely 

more complicated than along tidal coasts by 

barriers that block propagules differentially by 

size and by irregular patterns of water depth. At 

Almond Beach the smaller propagules of 

Laguncularia sometimes became established on 

top of mats of floating vegetation, extended their 

roots through water to the soil, and remained 

elevated on those roots after the water receded 

(Figure 5.19). That process provided a 

Figure 5.19. Roots of a Laguncularia seedling that 

established on floating vegetation during high water. 

At lower right a string used to originally mark plots 

can be seen emerging from the roots. Remains of 

pneumatophores of Avicennia trees killed by the soil 

fires are also visible.

96 


constantly moist, but never deeply inundated, 

surface that might be appropriate for such small 

fruits to germinate and establish successfully. 

Ellison (2002) found across 44 worldwide 

sites from regions with a wide range of 

mangrove species richness, that within-site 

species richness maintained a linear relationship 

with regional species richness. It was concluded 

that mangrove forests may be unsaturated with 

available species and that a site’s species 

composition may be more attributable to 

colonization abilities and propagule sorting, as 

proposed by Rabinowitz (1978a), rather than to 

differences in edaphic or competitive niches. At 

Almond Beach it was demonstrated that edaphic 

factors were not preventing establishment and 

growth of Rhizophora in regenerating burned 

mangrove swamp. During those studies a single 

creek drained the Almond Beach swamp into a 

lagoon that was only periodically open to the 

ocean. The northernmost Rhizophora mangle 

trees were located at that creek, which was 

possibly where propagules entered the occluded 

swamp, from which the species was able to 

disperse in deeper waters along the back of the 

beach ridge. The composition and patterns of 

Waini Peninsula coastal mangrove species seem 

to be influenced by dispersal, establishment 

ability and sorting in the early history of a 

swamp, which only later exhibits patterns 

influenced by edaphic factors and competition, 

along with the eventual migration of less easily 

dispersed species. Such development would 

depend on the absence of large-scale 

disturbances that would restore propagule 

availability and dispersal as dominant factors. 

The establishment and composition of 

mangrove swamps in many parts of the world 

are apparently also influenced by propagule 

predation, particularly by herbivorous crabs 

(Dahdouh-Guebas et al. 1998; Lee 1998; 

McGuinness 1997b; Osborne & Smith 1990; 

Smith et al. 1989), though this may not 

necessarily be a major factor in mangrove 

swamp zonation (Sousa & Mitchell 1999). 

Avicennia propagules are most often cited as a 

preferred prey of herbivorous crabs 

(McGuinness 1997b; McKee 1995a; Smith 

1987). In some populated areas propagule 

predation can also be dominated by 

domesticated animals (Lema Vélez et al. 2003). 

Zonation of mangrove swamps of Belize was 

found to be possibly influenced by predation of 

Avicennia propagules, though not of Rhizophora 

or Laguncularia (McKee 1995a). The 

herbivorous giant land crab (Cardisoma 

guanhumi) Latreille is common in the Avicennia 

swamps of the Guianas, including the unburned 

swamps near Almond Beach (Figure 5.20). 

However, Cardisoma crabs were absent from 

the burned swamp areas. When water levels 

were high in the burned swamps, Callinectes 

Stimpson crabs were observed. These crabs are 

carnivorous to omnivorous and are not 

mentioned in the literature as propagule 

predators. The potential role of Cardisoma crabs 

in population dynamics of Avicennia swamps, 

and therefore in eventual changes in species 

dominance in older Waini Peninsula swamps is 

an ecological interaction worthy of future study. 

Figure 5.20. Cardisoma guanhumi and Callinectes 

crabs at Almond Beach. Cardisoma (above) is a 

common burrowing resident of the undisturbed 

Avicennia swamps and is possibly an Avicennia 

propagule predator. The large group of Cardisoma 

is assembled near the Atlantic Ocean during a full 

moon. In the burned Avicennia swamps Callinectes 

crabs (below) were often observed during periods of 

higher water; they are considered to be omnivorous 

to detritivorous. The Callinectes shown is on floating 

Limnobium plants.


Laguncularia has been noted by several 

authors for its function as a pioneer species in 

some mangrove swamps (Baldwin et al. 2001; 

Ball 1980; Delgado et al. 2001; Roth 1992; 

Tissot & Marius 1992; Woodroffe 1983), often 

becoming the dominant species early in 

regeneration of disturbed swamps. It produces 

a relatively large quantity of smaller propagules 

and has been described as shade intolerant and 

fast growing with high mortality rates (Roth 

1992). Clarke (2000) felt that there were no gap 

specialists among Australia’s mangroves, which 

lack a small-fruited genus such as Laguncularia. 

While plant species richness in burned areas 

of the mangrove swamps of the Waini Peninsula 

was higher following the fires, many of those 

species were disturbance specialists; in some 

areas these will be temporary while in others 

they may dominate and prevent colonization by 

woody species. Smaller patches of post burn 

vegetation could be considered a diversification 

of the plant community with few negative effects 

(Hackney & de la Cruz 1981). However, at 

Almond Beach some large burned sections 

became quickly dominated by Typha and vines 

and will possibly recover very slowly and may 

be subject to additional fires during droughts. 

Dispersal and establishment of trees may be 

inhibited in these areas, which could remain a 

vestige of the fires for many years. 

The Almond Beach fires may the beginning 

steps toward a landscape such as described by 

Pons and Pons (1975) for a section of coastal 

Surinam, where broad areas of herbaceous 

vegetation are found behind a coastal Avicennia 

belt. These landscapes are described as 

maintained by cyclical fire events. However, the 

authors offer no record of fires actually observed 

in areas dominated by mangrove vegetation. The 

recurring fires in Surinam may be analogous to 

those that maintain Mauritia palm and sedge 

marshes located farther inland on the Guyana 

coastal plain, most notably at Santa Rosa in 

Guyana’s Northwest District. 


Although Laguncularia and Avicennia often 

display high germination rates in controlled 

experimental conditions (Steinke 1975), 

propagules of those species failed to establish 

or survive in field plantings. There were no 

97 

explicit signs of predation at either site, however 

the potential propagule Cardisoma crabs were 

present in some locations near the unburned site. 

The failure of planting of smaller propagules is 

in agreement with results reported by Elster 

(2000) for a disturbed mangrove swamp in 

Colombia, where all propagules died within two 

months of planting in drying soils, with 

Avicennia and Laguncularia mortalities far 

higher than those of Rhizophora in most cases. 

Propagule mortality was described by Elster 

(2000) as being generally higher in open, 

disturbed sites and higher for smaller 

propagules, caused by environmental factors 

including dessication and unstable hydrology. 

Rabinowitz (1978c) also found far higher 

survivorship for Rhizophora seedlings 

compared to Avicennia or Laguncularia 

seedlings. It was demonstrated here that 

Avicennia and Laguncularia propagule 

establishment is sensitive to water level. While 

those species did not establish in planting trials, 

establishment along narrow strand lines was 

observed bearby, presumably where water levels 

were appropriate. Such sensitivity to hydrology 

has been noted for Cypress swamps in eastern 

North America (Middleton 2000). The 

morphology of Rhizophora propagules is 

significantly different from Avicennia and 

Laguncularia and affects the success of 

plantings. Some Asian mangrove plantations 

have been shown to be far more species poor 

than natural stands, with many species of natural 

stands absent; in the Philippines, Rhizophora is 

overwhelmingly preferred for ease of planting, 

high success rates, and usefulness/high 

economic value (Walters 2000). All of these 

findings illustrate why the large propagules of 

Rhizophora and other Rhizophoraceae are often 

the major type of mangrove planted in 

restoration efforts (Field 1998a; Field 1996), 

even though fairly high field survival rates have 

been reported for nursery reared seedlings of 

other mangrove species (Elster 2000; 

Rabinowitz 1978b). The planting results from 

Almond Beach are in general agreement with 

the concept that seeds with larger mass are well 

adapted for successful establishment and 

seedling survival (Westoby et al. 1997). In the 

case of planted Rhizophora, the advantages of 

the large, elongated propagule allow tolerance

98 


of deeper inundation and probably tolerance to 

lower moisture and nutrient availability during 

droughts. Advantages for establishment gained 

through larger size come at the expense of 

dispersal ability, which is especially apparent 

in the case of the non-tidal burned mangrove 

swamps at Almond Beach. At Almond Beach, 

the plantings of Rhizophora added well adapted 

species that were absent during early succession 

only because of dispersal limitation. Any large 

scale plantings could be viewed as a case of 

stepping ahead to a species composition that 

may be similar to one that would be attained 

naturally over time. Rhizophora is very easy to 

plant, and plantings of it are generally very 

successful. It is considered useful as timber and 

fuelwood, and some plantings near settlements 

could be considered and would not be out of 

place in view of potential long term plant 

community succession in the burned mangrove 

swamps. Rhizophora mangle propagules 

survived but grew comparatively poorly in dry 

season plantings, possibly because of their 

smaller size, and did poorly in deep water 

because of their shorter length. Rhizophora 

racemosa propagules had a higher chance of 

success through variations in hydrological 

conditions, probably because of their longer, 

more massive propagules. Rhizophora 

racemosa propagules are usually easier to 

collect due to their frequency along the margins 

of tidal rivers of the region. Because of the 

species’ absence, establishment distance of 

Rhizophora racemosa was not tested in the 

Almond Beach swamps, but the longer 

propagules would likely disperse poorly in the 

shallow, non-tidal waters, and would potentially 

spread very slowly from any areas of plantings. 

Just as Rabinowitz’s (1978 a & b) tidal 

sorting hypotheses attributed mangrove 

zonation to differential establishment dependent 

on where each species was able to strand at high 

water, establishment in the burned swamp varied 

by water depth and the swamp’s 

microtopography. The smaller Laguncularia and 

Avicennia propagules were observed to establish 

where they stranded, however they were not 

successful in this study’s randomly placed 

planting sites where hydrological conditions 

were likely to be inappropriate. In the burned 

swamp, the success of naturally established 

Laguncularia in the very saline soils of drought 

conditions also suggests that zonation of that 

species is not necessarily driven by soil salinity 

in that area. 

For restoration purposes, these results have 

indicated that species with smaller propagules 

would require a greater effort to establish 

successfully, and higher cost for growing and 

transporting seedlings. In light of the 

establishment distance results presented here, 

such effort would be unnecessary in the cases 

of Laguncularia and Avicennia, which develop 

seed over a period of as little as three years and 

can be expected to disperse and establish some 

seedlings over a distance up to 100 meters, even 

in a non-tidal interior swamp. 

Nevertheless, in a majority of disturbance 

cases, natural regeneration is preferable for 

mangrove restoration (Field 1996; Lewis & 

Streever 2000), the exceptions generally being 

where large scale alterations to hydrology have 

taken place (Bacon 1975; Elster 2000; Elster et 

al. 1999; Elster & Polanía 2000; Lewis 1990, 

1982). It is possible that plantings of mangrove 

seedlings might affect the success or course of 

natural colonization (Bosire et al. 2003). With 

that in mind, mangrove restoration and 

management efforts should not be undertaken 

without detailed investigation into potential 

ecological, social and economic costs and 

consequences (Alongi 2002; Ellison 2000a, b; 

Elster 2000; Field 1998a; Field 1996, 1998b; 

Hamilton & Snedaker 1984; Humm 2001; Lewis 

1982, 2000; Lewis & Streever 2000; Macintosh 

& Ashton 2003).


Aldstadt, J., D. Chen, and A. Getis 1998. Point 

Pattern Analysis. Version 1.0a. Department 

of Geography, San Diego State University, 

San Diego. 

Allan, C., S. Williams, and R. Adrian 2002. The 

Socio-Economic Context of the Harvesting 

and Utilisation of Mangrove Vegetation. 

Guyana Forestry Commission, 

Georgetown, Guyana. 

Allen, J. A. 1998. Mangroves as alien species: 

The case of Hawaii. Global Ecology and 

Biogeography Letters 7:61-71. 

Alongi, D. M. 2002. Present state and future of 

the world’s mangrove forests. 

Environmental Conservation 29:331-349. 

Anonymous 1955. An Illustrated Guide to the 

Botanic Gardens, Georgetown. British 

Guiana Department of Agriculture, 

Georgetown. 

Asmussen, C. B., and M. W. Chase. 2001. 

Coding and noncoding plastid DNA in palm 

systematics. American Journal of Botany 

88:1103-1117. 

Augustinus, P. G. E. F. 1995. Geomorphology 

and sedimentation of mangroves. In G. M. 

E. Perillo, editor. Geomorphology and 

Sedimentation of Estuaries. Elsevier, 

Amsterdam. 

Bacon, P. R. 1975. Recovery of a Trinidadian 

mangrove swamp from attempted 

reclamation. In G. E. Walsh, S. C. Snedaker, 

and H. J. Teas, editors. Proceedings of the 

International Symposium on Biology and 

Management of Mangroves. Institute of 

Food and Agricultural Sciences, University 

of Florida, Gainesville. 

Bacon, P. R. 1990. Ecology and management of 

swamp forests in the Guianas and 

Caribbean region. Pages 193-213 in A. E. 

Lugo, M. Brinson, and S. Brown, editors. 

Ecosystems of the World 15, Forested 

Wetlands. Elsevier, Amsterdam. 

Bacon, P. R. 2001. Germination of Nypa 

fruticans in Trinidad. Palms 45:57-61. 

Badve, R. M., and C. V. Sakurkar. 2003. On the 

disappearance of palm genus Nypa from the 

REFERENCES 

99 

west coast with its present status in the 

Indian subcontinent. Current Science 

85:1407-1409. 

Baldwin, A. H., M. Egnotovich, M. Ford, and 

W. Platt. 2001. Regeneration in fringe 

mangrove forests damaged by Hurricane 

Andrew. Plant Ecology 157:151-164. 

Ball, M. C. 1980. Patterns of secondary 

succession in a mangrove forest of Southern 

Florida. Oecologia 44:226-235. 

Ball, M. C. 1988. Ecophysiology of mangroves. 

Trees 2:129-142. 

Barbosa, R. I., and P. M. Fearnside. 1999. 

Incêndios na Amazônia brasileira: 

Estimativa da emissão de gases do efeito 

estufa pela queima de diferentes 

ecossistemas de Roraima na passagem do 

evento “El Niño” (1997/98). Acta 

Amazonica 29:513-534. 

Barbosa, R. I., M. R. Xaud, G. N. F. Silva, and 

A. C. Cattâneo. 2003. Forest Fires in 

Roraima, Brazilian Amazonia. International 

Forest Fire News 28:51-56. 

BDG 2001. Guyana Regional Plant Species Lists 

- Draft Version. http://www.mnh.si.edu/ 

biodiversity/bdg/regionlists/index.html, 

Biological Diversity of the Guianas 

Program, Smithsonian Institution, 

Washington, DC. 

Blanchard, J., and G. Prado. 1995. Natural 

regeneration of Rhizophora mangle in strip 

clearcuts in Northwest Ecuador. Biotropica 

27:160-167. 

Blasco, F. 1975. Les Mangroves de L’Indie - 

The Mangroves of India, Travaux de la 

Section Scientifique et Technique Tome 

XIV. Institut Français de Pondichéry, 

Pondicherry, India. 

Boggan, J., V. Funk, C. L. Kelloff, M. Hoff, G. 

Cremers, and C. Feuillet 1997. Checklist 

of the Plants of the Guianas (Guyana, 

Surinam, French Guiana), 2nd Edition. 

Biological Diversity of the Guianas 

Program, Smithsonian Institution, 


Bonadie, W. A. 1998. The ecology of Roystonea

100 


oleracea palm swamp forest in the Nariva 

swamp (Trinidad). Wetlands 18:249-255. 

Bond, W. J., and B. W. van Wilgen 1996. Fire 

and Plants. Champman & Hall, London. 

Bonhoure, D., E. Rowe, A. J. Mariano, and E. 

H. Ryan. 2004. The South Equatorial 

Current System. http:// 

oceancurrents.rsmas.miami.edu/atlantic/ 

south-equatorial.html. Rosenstiel School of 

Marine and Atmospheric Science, 

University of Miami, Miami, Florida. 

Bosire, J. O., F. Dahdouh-Guebas, J. G. Kairo, 

and N. Koedam. 2003. Colonization of nonplanted 

mangrove species into restored 

mangrove stands in Gazi Bay, Kenya. 

Aquatic Botany 76:267-279. 

Brinkman, R., and L. J. Pons 1968. A Pedogeomorphological 

Classification and Map 

of the Holocene Sediments in the Coastal 

Plain of the Three Guianas. Soil Survey 

Institute, Wageningen, The Netherlands. 

Brocklehurst, P., and B. Edmeades 2003. 

Mangrove Survey of Bynoe Harbour, 

Northern Territory. Department of 

Infrastructure, Planning and Environment, 

Palmerston, Northern Territory. 

Brummitt, R. K., and C. E. Powell, editors. 

1992. Authors of Plant Names. Royal 

Botanic Gardens, Kew; Richmond, Surrey, 

United Kingdom. 

Brundin, L. 1966. Transantarctic Relationships 

and Their Significanace, as Evidenced by 

Chironomid Midges, with a Monograph of 

the Subfamilies Podonominae and 

Aphroteniinae and the Austral Heptagyiae. 

Almqvist and Wiksell, Stockholm. 

Bullock, J. M., and R. T. Clarke. 2000. Long 

distance seed dispersal by wind: measuring 

and modelling the tail of the curve. 

Oecologia 124:506–521. 

Bureau of Economic Geology 2002. Geo- 

Environmental Characterization of the 

Delta del Orinoco. University of Texas at 

Austin. 

Campbell, P., J. Comiskey, A. Alonso, F. 

Dallmeier, P. Nuñez, H. Beltran, S. 

Baldeon, W. Nauray, R. D. L. Colina, L. 

Acurio, and S. Udvardy. 2002. Modified 

Whittaker Plots as an Assessment and 

Monitoring Tool for Vegetation in a 

Lowland Tropical Rainforest. 

Environmental Monitoring and Assessment 

76:19-41. 

Cardona, P., and L. Botero. 1998. Soil 

characteristics and vegetation structure in 

a heavily deteriorated mangrove forest in 

the Caribbean coast of Colombia. 

Biotropica 30:24-34. 

Chale, F. M. M. 1996. Litter production studies 

in an Avicennia germinans (L.) Stearn forest 

in Guyana, South America. Hydrobiologia 

330:47-53. 

Chambers, J. C., and J. A. MacMahon. 1994. A 

day in the life of a seed: Movements and 

fates of seeds and their implications for 

natural and managed systems. Annual 

Review of Ecology and Systematics 

25:263-292. 

Chen, D., and A. Getis 1998. Point Pattern 

Analysis (PPA). Online manuscript. 

Department of Geography, San Diego State 

University, San Diego. 

Chen, R., and R. R. Twilley. 1999. Patterns of 

mangrove structure and soil nutrient 

dynamics along the Shark River Estuary, 

Florida. Estuaries 22:955-970. 

Cintrón, G., A. E. Lugo, D. J. Pool, and G. 

Morris. 1978. Mangroves of arid 

environments in Puerto Rico and Adjacent 

Islands. Biotropica 10:110-121. 

Clark, J. S., M. Lewis, and L. Horvath. 2001. 

Invasion by Extremes: Population Spread 

with Variation in Dispersal and 

Reproduction. American Naturalist 

157:537-554. 

Clarke, H. D., and V. Funk. 2005. Using 

checklists and collections data to 

investigate plant diversity: II. An analysis 

of five florulas from northeastern South 

America. Proceedings of the Academy of 

Natural Sciences of Philadelphia 154:29- 

37. 

Clarke, H. D., V. Funk, and T. Hollowell. 2001a. 

Using checklists and collections data to 

investigate plant diversity. I: A comparative 

checklist of the plant diversity of the 

Iwokrama Forest, Guyana. Sida Botanical 

Miscellany 21:1-86. 

Clarke, L. D., and N. J. Hannon. 1969. The 

mangrove swamp and salt marsh 

community of the Sydney district. II. The 

holocoenotic complex with particular


reference to physiography. Journal of 

Ecology 57:213-234. 

Clarke, P. J. 1993. Dispersal of grey mangrove 

(Avicennia marina) propagules in 

southeastern Australia. Aquatic Botany 

45:195-204. 

Clarke, P. J., and R. A. Kerrigan. 2000. Do forest 

gaps influence the population structure and 

species composition of mangrove stands in 

Northern Australia? Biotropica 32:642-652. 

Clarke, P. J., R. A. Kerrigan, and C. J. Westphal. 

2001b. Dispersal potential and early growth 

in 14 tropical mangroves: Do early life 

history traits correlate with patterns of adult 

distribution? Journal of Ecology 89:648- 

659. 

Clements, F. E. 1928. Plant Succession and 

Indicators: A Definitive Editon of Plant 

Succession and Plant Indicators. Hafner 

Publishing Co, Inc., New York. 

Cochrane, M. A. 2002. Spreading Like 

Wildfire—Tropical Forest Fires in Latin 

America and the Caribbean: Prevention, 

Assessment and Early Warning. Page 96. 

United Nations Environment Programme 

(UNEP), Regional Office for Latin America 

and the Caribbean, Mexico City. 

Cochrane, M. A. 2003. Fire science for 

rainforests. Nature 421:913-919. 

Cochrane, M. A., and M. D. Schulze. 1998. 

Forest fires in the Brazilian Amazon. 

Conservation Biology 12:948-950. 

Colinvaux, P. A. 1987. Amazon diversity in light 

of the palaoecological record. Quaternary 

Science Review 6:93-114. 

Comiskey, J., R. Mosher, C. Drysdale, and S. 

O’Grady 1999. BioMon for Windows Suite: 

Version 2. Smithsonian Institution Man and 

Biosphere Program, Washington, DC. 

Condit, R., N. Pitman, E. G. Leigh Jr., J. Chave, 

J. Terborgh, R. B. Foster, P. Nuñez V., S. 

Aguilar, R. Valencia, G. Villa, H. C. Muller- 

Landau, E. Losos, and S. P. Hubbell. 2002. 

Beta-diversity in tropical forest trees. 

Science 295:666-669. 

Connell, J. H., and R. O. Slatyer. 1977. 

Mechanisms of succession in natural 

communities and their role in community 

stability and organization. American 

Naturalist 111:1119-1144. 

Conrado, A., and E. Z. y. d. Ayala 1906. Estudio 

101 

de la Planta Llamada “Nipa” de su Cultivo 

y de sus Propiedades. La Concepción, 

Manila. 

Croizat, L. 1964. Space, Time, Form: The 

Biological Synthesis. Published by the 

Author, Caracas. 

Dahdouh-Guebas, M. V., J. F. Tack, D. Van 

Speybroeck, and N. Koedam. 1998. 

Propagule predators in Kenyan mangroves 

and their possible effect on regeneration. 

Marine and Freshwater Research 49:345- 

350. 

Dallmeier, F., M. Kabel, and R. Rice. 1992. 

Methods for long-term biodiversity 

inventory plots in protected tropical forest. 

In F. Dallmeier, editor. Long-Term 

Monitoring of Biological Diversity in 

Tropical Forest Areas: Methods for 

Establishment and Inventory of Permanent 

Plots. UNESCO, Paris. 

Daly, V. T. 1995. A Short History of the 

Guyanese People. Macmillan, London. 

Davis, J. H. 1940. The ecology and geologic role 

of mangroves of Florida. Papers of the 

Tortugas Laboratory 32:303-341. 

Davis, T. A. W., and P. W. Richards. 1933. The 

vegetation of Moraballi Creek, British 

Guiana: An ecological study of a limited 

area of tropical rain forest. Part I. Journal 

of Ecology 21:350-384. 

DeFilipps, R. A. 1992. Ornamental Garden 

Plants of the Guianas: An Historical 

Perspective of Selected Garden Plants fro 

Guyana, Surinam and French Guiana. 

Department of Botany, Smithsonian 

Institution, Washington, DC. 

DeFilipps, R. A., S. L. Maina, and J. Crepin 

2004. Medicinal Plants of the Guianas 

(Guyana, Surinam, French Guiana), web 

version. Department of Botany, 

Smithsonian Institution, Washington, DC. 

Delgado, P., P. F. Hensel, J. A. Jiménzez, and J. 

W. Day. 2001. The importance of propagule 

establishment and physical factors in 

mangrove distributional patterns in a Costa 

Rican estuary. Aquatic Botany 71:157-178. 

Di Croce, J., A. W. Bally, and P. Vail. 1999. 

Sequence stratigraphy of the Eastern 

Venezuelan Basin. Pages 419-476 in P. 

Mann, editor. Caribbean Basins. Elsevier, 

Amsterdam.

102 


Duke, N. C. 1991. Nypa in the Mangroves of 

Central America: Introduced or Relict? 

Principes 35:127-132. 

Duke, N. C. 2001. Gap creation and regenerative 

processes driving diversity and structure of 

mangrove ecosystems. Wetlands Ecology 

and Management 9:257-269. 

Duke, N. C., M. C. Ball, and J. Ellison. 1998. 

Factors influencing biodiversity and 

distributional gradients in mangroves. 

Global Ecology and Biogeography Letters 

7:27-47. 

Duke, N. C., A. M. Bell, D. K. Pedersen, C. M. 

Roelfsema, L. M. Godson, K. N. Zahmel, 

J. Mackenzie, and S. Bengtson-Nash 2003. 

Mackay Mangrove Dieback. Investigations 

in 2002 with recommendations for further 

research, monitoring and management. 

Report to Queensland Department of 

Primary Industries Northern Fisheries 

Centre, and the Community of Mackay. 11 

September 2003. Marine Botany Group, 

Centre for Marine Studies, The University 

of Queensland, Brisbane. 

Duke, N. C., E. Y. Y. Lo, and M. Sun. 2001. 

Global distribution and genetic 

discontinuities of mangroves – emerging 

patterns in the evolution of Rhizophora. 

Trees 16:65-79. 

Duke, N. C., Z. S. Pinzón M., and M. C. Prada 

T. 1997. Large-scale damage to mangrove 

forests following two large oil spills in 

Panama. Biotropica 29:2-14. 

Egler, F. E. 1954. Vegetation science concepts. 

I. Initial floristic composition, a factor in 

old field vegetational development. 

Vegetatio 4:412-417. 

Ek, R. C. 1990. Index of Guyana Plant 

Collectors. Flora of the Guianas 

Supplementary Fascicle 1. Koeltz Scientific 

Books, Koenigstein. 

Ek, R. C. 1997. Botanical Diversity in the 

Tropical Rain Forest of Guyana. Tropenbos 

- Guyana Series 4. Tropenbos-Guyana 

Programme, Georgetown, Guyana. 

Ellison, A. M. 2000a. Mangrove restoration: Do 

we know enough? Restoration Ecology 

8:219-229. 

Ellison, A. M. 2000b. Restoration of Mangrove 

Ecosystems. Restoration Ecology 8:217- 

218. 

Ellison, A. M. 2002. Macroecology of 

mangroves: large-scale patterns and 

processes in tropical coastal forests. Trees 

16:181-194. 

Ellison, A. M., E. J. Farnsworth, and R. R. 

Merkt. 1999. Origins of mangrove 

ecosystems and the mangrove biodiversity 

anomaly. Global Ecology and 

Biogeography 8:95-115. 

Ellison, J. C. 1993. Mangrove retreat with rising 

sea-level, Bermuda. Estuarine, Coastal, and 

Shelf Science 37:75-87. 

Ellison, J. C., and D. R. Stoddart. 1991. 

Mangrove ecosystem collapse during 

predicted sea-level rise: Holocene 

analogues and implications. Journal of 

Coastal Research 7:151-165. 

Elster, C. 2000. Reasons for reforestation 

success and failure with three mangrove 

species in Colombia. Forest Ecology and 

Management 131:201-214. 

Elster, C., L. Perdomo, and M.-L. Schnetter. 

1999. Impact of ecological factors on the 

regeneration of mangroves in the Cienaga 

Grande de Santa Marta, Colombia. 

Hydrobiologia 413:35-46. 

Elster, C., and J. Polanía. 2000. Regeneration 

of mangrove forests in the Cienaga Grande 

de Santa Marta (Colombia). Actualidades 

Biologicas 22:29-36. 

EPA Guyana. 2000. Integrated Coastal Zone 

Management Action Plan. Page 31 pp. 

Environmental Protection Agency of 

Guyana, Georgetown. 

EPA Guyana. 2002. Coastal Zone Management. 

Government of Guyana, Georgetown. http: 

//www.epaguyana.org/ICZM/articles.htm 

EPA Guyana. 2004. Proposed National Protected 

Areas. Government of Guyana, 

Georgetown. http://www.epaguyana.org/ 

npas/proposed.htm. 

ESRI 2002. ArcView GIS 3.3. Environmental 

Systems Research Institute, Inc., Redlands, 

CA. 

Ewel, K. C., R. R. Twilley, and J. E. Ong. 1998. 

Different kinds of mangrove forests provide 

different goods and services. Global 

Ecology and Biogeography Letters 7:83-94. 

Fanshawe, D. B. 1954. Riparian vegetation in 

British Guiana. Journal of Ecology 42:289- 

295.


FAO 1994. Mangrove Management Guidelines. 

U.N. Food and Agriculture Organization, 

Rome. 

Ferreira, L. V., and G. T. Prance. 1999. 

Ecosystem recovery in terra firme forests 

after cutting and burning: a comparison on 

species richness, floristic composition and 

forest structure in the Jaú National Park, 

Amazonia. Botanical Journal of the Linnean 

Society 130:97-110. 

Field, C. 1998a. Rationales and practices of 

mangrove afforestation. Marine and 

Freshwater Research 49:353-358. 

Field, C. B., J. G. Osborn, L. L. Hoffman, J. F. 

Polsenberg, D. D. Ackerly, J. A. Berry, O. 

Björkman, A. Held, P. Matson, and H. A. 

Mooney. 1998. Mangrove biodiversity and 

ecosystem function. Global Ecology and 

Biogeography Letters 7:3-14. 

Field, C. D. 1995. Impact of expected climate 

change on mangroves. Hydrobiologia 

295:75-81. 

Field, C. D. 1996. Restoration of Mangrove 

Ecosystems. International Society for 

Mangrove Ecosystems, Okinawa, Japan. 

Field, C. D. 1998b. Rehabilitation of mangrove 

ecosystems: An overview. Marine Pollution 

Bulletin 37:383-392. 

Finn, M., J. Inglehard, and P. Kangas. 1998. A 

taxonomy of spatial forms of mangrove 

dieoffs in southwest Florida. In P. J. 

Cannissaro, editor. Proceedings of the 24th 

Annual Conference on Ecosystem 

Restoration and Creation. Hillsborough 

Community College, Tampa, FL. 

Fong, F. W. 1992. Perspectives for sustainable 

resource utilization and management of 

Nipa vegetation. Economic Botany 46:45- 

54. 

Fosberg, F. R. 1947. Micronesian mangroves. 

Journal of the New York Botanical Garden 

48:128-138. 

Fromard, F., H. Puig, E. Mougin, G. Marty, J. L. 

Betoulle, and L. Cadmuro. 1998. Structure, 

above-ground biomass and dynamics of 

mangrove ecosystems: new data from 

French Guiana. Oecologia 115:39-53. 

Funk, V. A. 2003a. 100 uses for an herbarium: 

well at least 72. ASPT Newsletter 17:17- 

19. 

Funk, V. A. 2003b. The importance of herbaria. 

103 

Plant Science Bulletin 49:94-95. 

Funk, V. A., and K. Richardson. 2002. 

Systematic data in biodiversity studies: Use 

it or lose it. Systematic Biology 51:303-316. 

Funk, V. A., K. S. Richardson, and S. Ferrier. 

2005. Survey-gap analysis in expeditionary 

research, where do we go from here? 

Biological Journal of the Linnean 

Society:549-567. 

Funk, V. A., M. F. Zermoglio, and N. Nasir. 

1999. Testing the use of specimen collection 

data and GIS in biodiversity exploration and 

conservation decision making in Guyana. 

Biodiversity and Conservation 8:727-751. 

Gabche, C. E., T. J. Youmbi, and C. A. Angwe. 

2000. The impacts of sea-level rise on the 

coastal fisheries resources and aquatic 

ecosystems of Cameroon (Central Africa). 

In B. W. Flemming, M. T. Delafontaine, and 

G. Liebezeit, editors. Muddy Coast 

Dynamics and Resource Management. 

Elsevier, Amsterdam. 

Gee, C. T. 1989. On the fossil occurrences of 

the mangrove palm Nypa. Proceedings of 

the Symposium “Paleofloristic and 

Paleoclimatic Changes in the Cretaceous 

and Tertiary”, Prague. 

Gee, C. T. 2001. The mangrove palm Nypa in 

the geologic past of the New World. 

Wetlands Ecology and Management 9:181- 

194. 

Gentry, A. H. 1993. A Field Guide to the 

Families and Genera of Woody Plants of 

Northwest South America (Colombia, 

Ecuador, Peru), with Supplementary Notes 

on Herbaceous Taxa. Conservation 

International, Washington, DC. 

Gibbs, A. K., and C. N. Barron 1993. The 

Geology of the Guiana Shield. Oxford 

University Press, New York. 

Gleason, H. A. 1927. Further views on the 

succession-concept. Ecology 8:299-326. 

Global Rain Forest Mapping Project 2001. AM1 

CDROM, GRFM South America JERS-1 

SAR. Jet Propulsion Laboratory, California 

Institute of Technology, Pasadena. http:// 

southport.jpl.nasa.gov/GRFM/. 

Godoy, J. A., and P. Jordano. 2001. Seed 

dispersal by animals: exact identification 

of source trees with endocarp DNA 

microsatellites. Molecular Ecology

104 


10:2275-2283. 

Golley, F., H. T. Odum, and R. F. Wilson. 1962. 

The structure and metabolism of a Puerto 

Rican red mangrove forest in May. Ecology 

43:9-19. 

Görts-van Rijn, A. R. A., editor. 1985 -. Flora 

of the Guianas. Royal Botanic Gardens, 

Kew; Richmond, Surrey, United Kingdom. 

Graham, A. 1995. Diversification of Gulf/ 

Caribbean mangrove communities through 

Cenozoic time. Biotropica 27:20-27. 

Gray, V. R. 1981. Rhizophora harrisonii 

(Rhizophoraceae), un nuevo registro para 

las costas de Mexico. Boletin de la 

Sociedad Botanica de Mexico 41:163-165. 

Green, E. P., P. J. Mumby, A. J. Edwards, C. D. 

Clark, and A. C. Ellis. 1998. The 

assessment of mangrove areas using high 

resolution multispectral airborne imagery. 

Journal of Coastal Research 14:433-443. 

Grégoire, J. M., B. Glénat, P. Janvier, E. Janodet, 

A. Tournier, and J. M. N. Silva. 1998. Fire 

activity in the Guyana Shield, the Orinoco 

and Amazon Basins during March 1998. 

International Forest Fire News 19:35-39. 

Guiry, M. D., and E. Nic Dhonncha 2004. 

AlgaeBase version 2.1. World-wide 

electronic publication, National University 

of Ireland. http://www.algaebase.org, 

Galway. 

Gyory, J., A. J. Mariano, and E. H. Ryan. 2003. 

The Guiana Current. http:// 

oceancurrents.rsmas.miami.edu/atlantic/ 

guiana.html. Rosenstiel School of Marine 

and Atmospheric Science, University of 

Miami, Miami, Florida. 

Hackney, C. T., and A. A. de la Cruz. 1981. 

Effects of fire on brackish marsh 

communities: management implications. 

Wetlands 1:75-86. 

Halos, S. C. 1981. Nipa for alcogas production: 

Tapping its potentials. Canopy 

International 7:15. 

Hamilton, L. S., and D. H. Murphy. 1988. Use 

and Management of Nipa Palm (Nypa 

fruticans, Arecaceae): a review. Economic 

Botany 42:206-213. 

Hamilton, L. S., and S. C. Snedaker, editors. 

1984. Handbook for Mangrove Area 

Management. United Nations Environment 

Programme and East-West Center, 

Environment and Policy Institute, Paris. 

Hammond, D. S., and H. t. Steege. 1998. 

Propensity for fires in Guianan Rainforests. 


Harper, J. L. 1977. Population Biology of Plants. 

Academic Press, London. 

He, H. S., and D. J. Mladenoff. 1999. The effects 

of seed dispersal on the simulation of longterm 

forest landscape change. Ecosystems 

2:308-319. 

Henderson, A. 1995. Palms of the Amazon. 

Oxford University Press, New York. 

Henderson, A., G. Galeano, and R. Bernal 1995. 

Field Guide to the Palms of the Americas. 

Princeton University Press, Princeton, New 

Jersey. 

Hierro, J. L., J. L. Maron, and R. M. Callaway. 

2005. A biogeographical approach to plant 

invasions: the importance of studying 

exotics in their introduced and native range. 

Journal of Ecology 95:5-15. 

Holdridge, L. R., W. C. Grenke, W. H. 

Hathaway, T. Liang, and J. A. Tosi, Jr. 1971. 

Forest Environments in Tropical Life Zone: 

A Pilot Study. Pergamon Press, Oxford. 

Hollowell, T., P. Berry, V. Funk, and C. L. 

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, 


Howard, R. A. 1976. Flora of the Lesser Antilles: 

Leeward and Windward Islands. Arnold 

Arboretum, Harvard University, Jamaica 

Plain, MA. 

Huber, O. 1995a. Geography and Physical 

Features. Pages 1-61 in P. E. Berry, B. K. 

Holst, and K. Yatskievych, editors. Flora 

of the Venezuelan Guayana. Volume 1: 

Introduction. Missouri Botanical Garden, 

St. Louis. 

Huber, O. 1995b. History of Botanical 

Exploration. Pages 63-95 in P. E. Berry, B. 

K. Holst, and K. Yatskievych, editors. Flora 

of the Venezuelan Guayana. Volume 1: 

Introduction. Missouri Botanical Garden, 

St. Louis. 

Huber, O. 1995c. Vegetation. Pages 97-160 in 

P. E. Berry, B. K. Holst, and K. Yatskievych,


editors. Flora of the Venezuelan Guayana. 

Volume 1: Introduction. Missouri Botanical 

Garden, St. Louis. 

Huber, O., G. Gharbarran, and V. Funk 1995. 

Vegetation Map of Guyana (Preliminary 

Version). Centre for the Study of Biological 

Diversity, University of Guyana, 

Georgetown. 

Huh, O. K., N. D. Walker, and C. Moeller. 2001. 

Sedimentation along the eastern chenier 

plain coast: Down drift impact of a delta 

complex shift. Journal of Coastal Research 

17:72-81. 

Humm, N. 2001. A Framework for the 

Incorporation of the Indigenous Peoples of 

the Proposed Shell Beach Protected Area 

for Marine Turtle Conservation. 75. School 

of Animal and Microbial Sciences. 

University of Reading, Reading, England. 

Imbert, D., A. Rousteau, and P. Scherrer. 2000. 

Ecology of mangrove growth and recovery 

in the Lesser Antilles: State of knowledge 

and basis for restoration projects. 

Restoration Ecology 8:230-236. 

Insightful Corporation 2001. S-PLUS 6 for 

Windows Guide to Statistics, Volume 2. 

Insightful Corporation, Seattle, WA. 

IPCC 2001. Climate Change 2001: The 

Scientific Basis. Contribution of Working 

Group I to the Third Assessment Report of 

the Intergovernmental Panel on Climate 

Change. Cambridge University Press, 

Cambridge. 

Janzen, D. H. 1985. Mangroves: Where’s the 

understory? Journal of Tropical Ecology 

1:89-92. 

Jelgersma, S., M. Van der Zijp, and R. Brinkman. 

1993. Sealevel rise and the coastal lowlands 

in the developing world. Journal of Coastal 

Research 9:958-972. 

Jiménez, J. A., A. E. Lugo, and G. Cintrón. 1985. 

Tree mortality in mangrove forests. 


Jiménez, J. A., and K. A. Sauter. 1991. Structure 

and dynamics of mangrove forests along a 

flooding gradient. Estuaries 14:49-56. 

Jongejans, E., and A. Telenius. 2001. Field 

experiments on seed dispersal by wind in 

ten umbelliferous species (Apiaceae). Plant 

Ecology 152:67-78. 

Kelloff, C. L. 2002. Plant Diversity of Kaieteur 

105 

National Park, Guyana: Using Plant Data 

as a Tool in Conservation and Development. 

Environmental Science and Public Policy. 

Ph.D. Thesis, George Mason University, 

Fairfax, VA. 

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 (University of Guyana) 1:1-44. 

Kelloff, C. L., and V. A. Funk. 2004. 

Phytogeography of the Kaieteur Falls, 

Potaro Plateau, Guyana: floral distributions 

and affinities. Journal of Biogeography 

31:501-513. 

Kinnaird, M. F., and T. G. O’Brien. 1998. 

Ecological effects of wildfire on lowland 

rainforest in Sumatra. Conservation 

Biology 12:954-956. 

Kjerfve, B., editor. 1998. CARICOMP – 

Caribbean coral reef, seagrass and 

mangrove sites. UNESCO, Paris. 

Lacerda, L. D., J. E. Conde, B. Kjerfve, R. 

Alvarez-León, C. Alarcón, and J. Polanía. 

2002. American mangroves. In L. D. 

Lacerda, editor. Mangrove Ecosystems: 

Function and Management. Springer, 

Berlin. 

Lakhan, V. C., and D. A. Pepper. 1997. 

Relationship between concavity and 

convexity of a coast and erosion and 

accretion patterns. Journal of Coastal 

Research 13:226-232. 

Lanjouw, J., editor. 1964-. Flora of Suriname. 

E. J. Brill, Leiden. 

Lasser, T., editor. 1964-1992. Flora de 

Venezuela. Instituto Botánico, Caracas. 

Laurance, W. F. 1998. A crisis in the making: 

responses of Amazonian forests to land use 

and climate change. Trends in Ecology and 

Evolution 13:411-415. 

Lee, S. Y. 1998. Ecological role of grapsid crabs 

in mangrove ecosystems: a review. Marine 

and Freshwater Research 49:335-343. 

Legg, C. A., and Y. Laumonier. 1999. Fires in 

Indonesia, 1997: A remote sensing 

perspective. Ambio 28:479-485. 

Lema Vélez, L. F., J. Polanía, and L. E. Urrego 

Giraldo. 2003. Disperción y 

establecimiento de las especies de mangle 

del río Ranchería en el período de máxima

106 


frutificación. Rev. Acad. Colomb. Cienc. 

27:93-103. 

Lewis, C. E., and J. J. Doyle. 2001. Phylogenetic 

utility of the nuclear gene malate synthase 

in the palm family (Arecaceae). Molecular 

Phylogenetics and Evolution 19:409-420. 

Lewis, R. R. 1990. Creation and restoration of 

coastal wetlands in Puerto Rico and the U.S. 

Virgin Islands. Pages 103-123 in J. A. 

Kusler, and M. E. Kentula, editors. Wetland 

Creation and Restoration: The Status of the 

Science. Island Press, Washington, DC. 

Lewis, R. R., III. 1982. Mangrove forests. In R. 

R. Lewis, editor. Creation and Restoration 

of Coastal Plant Communities. CRC Press, 

Boca Raton, FL. 

Lewis, R. R., III. 2000. Ecologically based goal 

setting in mangrove forest and tidal marsh 

restoration. Ecological Engineering 15:191- 

198. 

Lewis, R. R., III, and B. Streever 2000. 

Restoration of Mangrove Habitat. WRP 

Technical Notes Collection (ERDC TN- 

WRP-VN-RS-3.2). www.wes.army.mil/el/ 

wrp. U.S. Army Engineer Research and 

Development Center, Vicksburg, MS. 

Lindeman, J. C. 1953. The Vegetation of the 

Coastal Region of Suriname. 

Rijksuniversiteit te Utrecht, Utrecht, 

Netherlands. 

López-Portillo, J., and E. Ezcurra. 1989. 

Zonation in mangrove and salt marsh 

vegetation at Laguna de Mecoacán, 

México. Biotropica 21:107-114. 

Lugo, A. E. 1980. Mangrove ecosystems: 

successional or steady state? Biotropica 

12:65-72. 

Lugo, A. E. 1981. Inland mangroves of Inagua. 

Journal of Natural History 15:845-852. 

Lugo, A. E. 1986. Mangrove understory: an 

expensive luxury? Journal of Tropical 

Ecology 2:287-288. 

Lugo, A. E. 1997. Old growth mangrove forests 

in the United States. Conservation Biology 

11:11-20. 

Lugo, A. E. 1998. Mangrove Forests: a tough 

system to invade but an easy one to 

rehabilitate. Marine Pollution Bulletin 

37:427-430. 

Lugo, A. E., and G. Cintrón. 1975. The 

mangroves of Puerto Rico and their 

management. Pages 825-846 in G. E. Walsh, 

S. C. Snedaker, and H. J. Teas, editors. 

Proceedings of the International 

Symposium on Biology and Management 

of Mangroves. Institute of Food and 

Agricultural Sciences, University of 

Florida, Gainesville. 

Lugo, A. E., and S. C. Snedaker. 1974. The 

ecology of mangroves. Annual Review of 

Ecology and Systematics 5:39-64. 

Macintosh, D. J., and E. Ashton 2003. Draft 

Code of Conduct for the Sustainable 

Management of Mangrove Ecosystems. 

World Bank, ISME, Center, Aarhus. 

Mack, R. N., and W. M. Lonsdale. 2001. 

Humans as global plant dispersers: Getting 

more than we bargained for. Bioscience 

51:95-102. 

Macnae, W. 1963. Mangrove swamps in South 

Africa. Journal of Ecology 51:1-25. 

Manson, F. J., N. R. Loneragan, I. M. McLeod, 

and R. A. Kenyon. 2001. Assessing 

techniques for estimating the extent of 

mangroves: topographic maps, aerial 

photographs and Landsat TM images. 

Marine and Freshwater Research 52:787- 

792. 

McCracken, D. P. 1997. Gardens of Empire: 

Botanical Institutions of the Victorian 

British Empire. Leicester University Press, 

London. 

McGuinness, K. A. 1997a. Dispersal, 

establishment and survival of Ceriops tagal 

propagules in a north Australian mangrove 

forest. Oecologia 109:80-87. 

McGuinness, K. A. 1997b. Seed predation in a 

tropical mangrove forest: a test of the 

dominance-predation model in northern 

Australia. Journal of Tropical Ecology 

13:293-302. 

McKee, K. L. 1993. Soil physicochemical 

patterns and mangrove species distribution 

- reciprocal effects? Journal of Ecology 

81:447-487. 

McKee, K. L. 1995a. Mangrove species 

distribution and propagule predation in 

Belize: An exception to the dominancepredation 

hypothesis. Biotropica 27:334- 

345. 

McKee, K. L. 1995b. Seedling recruitment 

patterns in a Belizean mangrove forest:


effects of establishment ability and physicochemical 

factors. Oecologia 101:448-460. 

McKee, K. L., and P. L. Faulkner. 2000. 

Restoration of biogeochemical function in 

mangrove forests. Restoration Ecology 

8:247-259. 

Melana, D. M. 1980. Leaf harvesting and juice 

extraction techniques of Nipa in Alabat 

Island, Quezon Province. Canopy 

International 6:6-7. 

Mennega, E. A., W. C. M. Tammen - de Rooij, 

and M. J. Jansen-Jacobs, editors. 1988. 

Checklist of Woody Plants of Guyana. 

Tropenbos Technical Series 2. The 

Tropenbos Foundation, Ede, The 


Miah, M. D., R. Ahmed, and S. J. Islam. 2003. 

Indigenous management practices of 

Golpata (Nypa fruticans) in local 

plantations in southern Bangladesh. Palms 

47:185-190. 

Middleton, B. 1999. Wetland Restoration, Flood 

Pulsing and Disturbance Dynamics. John 

Wiley & Sons, Inc., New York. 

Middleton, B. 2000. Hydrochory, seed banks, 

and regeneration dynamics along the 

landscape boundaries of a forested wetland. 

Plant Ecology 146:169-184. 

Missouri Botanical Garden 1995-present. 

W3TROPICOS. http://mobot.mobot.org/ 

W3T/Search/vast.html, St. Louis. 

Munsell Color 1992. Munsell Soil Color Charts, 

1992 Revised Edition. Kollmorgen 

Instruments Corp., Newburgh, NY. 

Nathan, R., and H. C. Muller-Landau. 2000. 

Spatial patterns of seed dispersal, their 

determinants and consequences for 

recruitment. Tree 15:278-285. 

New York Botanical Garden 1996-present. 

Virtual Herbarium of the New York 

Botanical Garden. http://www.nybg.org/ 

bsci/hcol/, New York. 

Odum, W. E., C. C. McIvor, and T. J. Smith, III. 

1982. The Ecology of the Mangroves of 

South Florida: A Community Profile. U.S. 

Fish and Wildlife Service, Washington, DC. 

Osborne, K., and T. J. Smith, III. 1990. 

Differential predation on mangrove 

propagules in open and closed canopy forest 

habitats. Vegetatio 89:1-6. 

Päivöke, A., M. R. Adams, and D. R. Twiddy. 

107 

1984. Nipa palm vinegar in Papua New 

Guinea. Process Biochemistry 19:84-87. 

Panapitukkul, N., C. M. Duarte, U. Thampanya, 

P. Kheowvongsri, N. Srichai, O. Geertz- 

Hansen, J. Terrados, and S. 

Boromthanarath. 1998. Mangrove 

colonization: Mangrove progression over 

the growing Pak Phanang (SE Thailand) 

mud flat. Estuarine, Coastal, and Shelf 

Science 47:51-61. 

Pannier, F., and R. Fraino de Pannier 1989. 

Manglares de Venezuela. Cuadernos 

LAGOVEN, Caracas. 

Parkinson, R. W., R. D. DeLaune, and J. R. 

White. 1994. Holocene sea level rise and 

the fate of mangrove forests within the 

wider Caribbean region. Journal of Coastal 

Research 10:1077-1086. 

Pernetta, J. C. 1993. Mangrove Forests, Climate 

Change and Sea Level Rise: Hydrological 

Influences on Community Structure and 

Survival, with Examples from the Indo- 

West Pacific. IUCN, Gland, Switzerland. 

Pickett, S. T. A., J. Kolasa, J. J. Armesto, and S. 

L. Collins. 1989. The ecological concept of 

disturbance and its expression at various 

hierarchical levels. Oikos 54:129-136. 

Pirazzoli, P. A. 1991. World Atlas of Holocene 

Sea-level Changes. Elsevier, Amsterdam. 

Pitman, N. C. A., J. Terborgh, M. R. Silman, 

and P. Nuñez V. 1999. Tree species 

distributions in an upper Amazonian forest. 

Ecology 80:2651-2661. 

Pitman, N. C. A., J. W. Terborgh, M. R. Silman, 

P. Nuñez V., D. A. Neill, C. E. Cerón, W. A. 

Palacios, and M. Aulestia. 2001. 

Dominance and distribution of tree species 

in upper Amazonian terra firme forests. 

Ecology 82:2101-2117. 

Plant Names Project 1999-present. The 

International Plant Names Index. 

www.ipni.org. 

Pons, T. L., and L. J. Pons. 1975. Mangrove 

vegetation and soils along a more or less 

stationary part of the coast of Surinam, 

South America. Pages 548-560 in G. E. 

Walsh, S. C. Snedaker, and H. J. Teas, 

editors. Proceedings of the International 



Agricultural Sciences, University of

108 



Pool, D. J., S. C. Snedaker, and A. E. Lugo. 

1977. Structure of mangrove forests in 

Florida, Puerto Rico, México, and Costa 

Rica. Biotropica 9:195-212. 

Pritchard, P. C. H. 1993. Thoughts on 

hapaxanthy in Guyana. Principes 37:12-18. 

Prost, M. T. 1989. Coastal dynamics and chenier 

sands in French Guiana. Marine Geology 

90:259-267. 

Prost, M. T. 1997. La mangrove de front de mer 

en Guyane: ses transformations sous 

l’influence du systéme de dispersion 

amazonien et son suivi par télédetection. 

In B. Kjerfve, L. D. d. Lacerda, and E. H. 

S. Diop, editors. Mangrove Ecosystems 

Studies in Latin America and Africa. 

UNESCO, Paris and ISME, Tokyo. 

Pulle, A. A., editor. 1932-. Flora of Suriname. 

Koninklijke Vereeniging Indisch Instituut, 

Amsterdam. 

Rabinowitz, D. 1978a. Dispersal properties of 

mangrove propagules. Biotropica 10:47-57. 

Rabinowitz, D. 1978b. Early growth of 

mangrove seedlings in Panamá, and an 

hypothesis concerning the relationship of 

dispersal and zonation. Journal of 

Biogeography 5:113-133. 

Rabinowitz, D. 1978c. Mortality and initial 

propagule size in mangrove seedlings in 

Panama. Journal of Ecology 66:45-51. 

Rafii, Z. A., R. S. Dodd, and F. Fromard. 1996. 

Biogeographic variation in foliar waxes of 

mangrove species. Biochemical 

Systematics and Ecology 24:341-345. 

Ramdass, I., J. Singh, M. Tamessar, and D. 

Gopaul 1997. National Activity on the use 

of Biodiversity, Guyana. UNDP, The Office 

of the President, and The University of 


Raven, P. H., and D. I. Axelrod. 1975. History 

of the flora and fauna of Latin America. 

American Scientist 63:420-429. 

Renner, S. 2004. Plant dispersal across the 

tropical Atlantic by wind and sea currents. 

International Journal of Plant Science 

165:S23-S33. 

Ribeiro, J. E. L. d. S., M. J. G. Hopkins, A. 

Vicentini, C. A. Sothers, M. A. d. S. Costa, 

J. M. de Brito, M. A. D. de Souza, L. H. P. 

Martins, L. G. Lohmann, P. A. C. L. 

Assunção, E. d. C. Pereira, C. F. da Silva, 

M. R. Mesquita, and L. C. Propópio, editors. 

1999. Flora da Reserva Ducke: Guia de 

Identificação das Plantas Vasculares de uma 

Floresta de Terra-firme na Amazônia 

Central. Instituto Nacional de Pesquisas da 

Amazônia, Manaus. 

Richardson, B. C. 1987. Men, water, and 

mudflats in coastal Guyana. Resource 

Management and Optimization 5:213-236. 

Rohlf, F. J. 1997. NTSYSpc: Numerical 

Taxonomy System, ver. 2.1. Exeter 

Publishing, Ltd., Setauket, NY. 

Ross, M. S., P. L. Ruiz, G. J. Telesnicki, and J. 

F. Meeder. 2001. Estimating above-ground 

biomass and production in mangrove 

communities of Biscayne National Park, 

Florida (U.S.A.). Wetlands Ecology and 


Roth, L. C. 1992. Hurricanes and mangrove 

regeneration: effects of Hurricane Joan, 

October 1988, on the vegetation of Isla del 

Venado, Bluefields, Nicaragua. Biotropica 

24:375-385. 

Rubin, J. A., C. Gordoni, and J. K. Amatekpor. 

1998. Causes and consequences of 

mangrove deforestation in the Volta estuary, 

Ghana: Some recommendations for 

ecosystem rehabilitation. Marine Pollution 


Rull, V. 1998. Middle Eocene mangroves and 

vegetation changes in the Maracaibo Basin, 

Venezuela. Palaios 13:287-296. 

Rull, V. 2001. A quantitative palynological 

record from the early miocene of western 

Venezuela,with emphasis on mangroves. 

Palynology 25:109-126. 

Saenger, P., E. J. Hegerl, and J. D. S. Davie. 

1983. Global status of mangrove 

ecosystems. The Environmentalist, IUCN, 

Gland, Switzerland 3:1-88. 

Schneider, R. L., and R. R. Sharitz. 1988. 

Hydrochory and regeneration in a bald 

cypress-water tupelo swamp forest. 

Ecology 69:1055-1063. 

Schomburgk, M. R. 1922. Travels in British 

Guiana During the Years 1840-1844. 

Translated by Walter Edmund Roth. Daily 

Chronicle, Georgetown, Guyana. 

Schomburgk, R. H. 1896. Reports and Letters 

of Sir Robert Hermann Schomburgk with


Reference to his Surveys of the Boundaries 

of British Guiana. 1841-1844. Harrison and 

Sons, London. 

Sealey, K. S., and G. Bustamante 1999. Setting 

Geographic Priorities for Marine 

Conservation in Latin America and the 

Caribbean. The Nature Conservancy, 

Arlington, VA. 

Sherman, R. E., T. J. Fahey, and J. J. Battles. 

2000. Small-scale disturbance and 

regeneration dynamics in a neotropical 

mangrove forest. Journal of Ecology 

88:165-178. 

SI/MAB 1998. Vegetation Sampling Protocols. 

Front Royal 1998: Modified Whittaker 

Plots. Unpublished manuscript. 

Singhroy, V. 1996. Interpretation of SAR images 

for coastal zone mapping in Guyana. 

Canadian Journal of Remote Sensing 

22:317-328. 

Smith, T. J., III. 1987. Seed predation in relation 

to tree dominance and distribution in 

mangrove forests. Ecology 68:266-273. 

Smith, T. J., III, H. T. Chan, C. C. McIvor, and 

M. B. Robblee. 1989. Comparisons of seed 

predation in tropical tidal forests from three 

continents. Ecology 70:146-151. 

Sneath, P. H. A., and R. R. Sokal 1973. 

Numerical Taxonomy. Freeman, San 

Francisco. 

Snedaker, S. C. 1986. Traditional uses of South 

American mangrove resources and the 

socio-economic effect of ecosystem 

changes. Pages 104-112 in P. Kunstader, E. 

C. F. Bird, and S. Sabhasri, editors. Man in 

the Mangroves: The Socio-economic 

Situation of Human Settlements in 

Mangrove Forests. United Nations 

University Press, Tokyo. 

Snedaker, S. C. 1995. Mangroves and climate 

change in the Florida and Caribbean region: 

Scenarios and hypotheses. Hydrobiologia 

295:43-49. 

Snedaker, S. C., S. J. Baquer, P. J. Behr, and S. 

I. Ahmed. 1995. Biomass distribution in 

Avicennia marina plants in the Indus River 

delta, Pakistan. Pages 389-395 in M. F. 

Thompson, and N. M. Thompson, editors. 

The Arabian Sea: Living Marine Resources 

and the Environment. Vanguard Books Ltd., 

Lahore, Pakistan. 

109 

Snedaker, S. C., M. S. Brown, E. J. Lahmann, 

and R. J. Araujo. 1992. Recovery of a 

mixed-species mangrove forest in South 

Florida following canopy removal. Journal 

of Coastal Research 8:919-925. 

Snedaker, S. C., and E. J. Lahmann. 1988. 

Mangrove understory absence: a 

consequence of evolution? Journal of 

Tropical Ecology 4:311-314. 

Snow, J. W. 1976. Climates of Northern South 

America. Pages 295-403 in W. 

Schwerdtfeger, editor. Climates of Central 

and South America. Elsevier Scientific 

Publishing Company, Amsterdam. 

Soil Survey Staff 1998. Keys to Soil Taxonomy. 

United States Department of Agriculture, 

Natural Resources Conservation Service, 


Sousa, W. P., and B. S. Mitchell. 1999. The effect 

of seed predators on plant distributions: is 

there a general pattern in mangroves. Oikos 

86:55-66. 

Spalding, M., F. Blasco, and C. Field 1997. 

World Mangrove Atlas. International 

Society for Mangrove Ecosystems, 

Okinawa, Japan. 

Steenis, C. G. G. J. v. 1984. Three more 

mangrove trees growing locally in nature 

in freshwater. Blumea 29:395-397. 

Steinke, T. D. 1975. Some factors affecting 

dispersal and establishment of propagules 

of Avicennia marina (Forrsk.) Vierh. Pages 

402-414 in G. E. Walsh, S. C. Snedaker, and 

H. J. Teas, editors. Proceedings of the 





Steyermark, J. A., P. E. Berry, and B. K. Holst, 

editors. 1995 - 2005. Flora of the 

Venezuelan Guayana. Missouri Botanical 


Stieglitz, T., and P. V. Ridd. 2001. Trapping of 

mangrove propagules due to density-driven 

secondary circulation in the Normanby 

River estuary, NE Australia. Marine 

Ecology Progress Series 211:131-142. 

Stoddart, D. R., G. W. Bryan, and P. E. Gibbs. 

1973. Inland mangroves and water 

chemistry, Barbuda, West Indes. Journal of 

Natural History 7:33-46.

110 


Stohlgren, T. J., M. B. Falkner, and L. D. Schell. 

1995. A modified-Whittaker nested 

vegetation sampling method. Vegetatio 

117:113-121. 

Sukhendu, G., D. Sauren, and G. Monoranjan. 

2002. Biology of Nypa fruticans (Thunb.) 

Wurmb.: An endangered mangrove palm 

of Sundarbans, India. Advances in Plant 

Sciences 15:71-78. 

Sunderland, T. C. H., and T. Morakinyo. 2002. 

Nypa fruticans, a weed in West Africa. 

Palms 46:154-155. 

Tews, J., K. Moloney, and F. Jeltsch. 2004. 

Modeling seed dispersal in a variable 

environment: a case study of the fleshyfruited 

savanna shrub Grewia flava. 

Ecological Modelling 175:65-76. 

Thom, B. G. 1967. Mangrove ecology and 

deltaic geomorphology: Tabasco, Mexico. 

Journal of Ecology 55:301-343. 

Thom, B. G. 1975. Mangrove ecology from a 

geomorphic viewpoint. Pages 469-481 in 

G. E. Walsh, S. C. Snedaker, and H. J. Teas, 






Thom, B. G. 1984. Coastal landforms and 

geomorphic processes. Pages 3-17 in S. C. 

Snedaker, and J. G. Snedaker, editors. The 

Mangrove Ecosystem: Research Methods. 

UNESCO, Paris. 

Tissot, C., and C. Marius. 1992. Holocene 

evolution of the mangrove ecosystem in 

French Guiana: A palynological study. In 

K. P. Singh, and J. S. Singh, editors. 

Tropical Ecosystems: Ecology and 

Management. Wiley Eastern Ltd, New 

Delhi. 

Tomlinson, P. B. 1973. The monocotyledons; 

their evolution and comparative biology. 

VII. Branching in monocotyledons. The 

Quarterly Review of Biology 48. 

Tomlinson, P. B. 1986. The Botany of 

Mangroves. Cambridge University Press. 

Tralau, H. 1964. The Genus Nypa van Wurmb. 

Kungl. Svenska Vetenskapsademiens 

Handlingar 10:5-29. 

Trenberth, K. E. 1999. The extreme weather 

events of 1997 and 1998. Consequences 

5:3-15. 

Uhl, N., and J. Dransfield 1987. Genera 

Palmarum: A Classification of Palms Based 

on the Work of Harold E. Moore, Jr. Allen 

Press, Lawrence, Kansas. 

Ukpong, I. E. 1995. An ordination study of 

mangrove swamp communities in West 

Africa. Vegetatio 116:147-159. 

USDA, and NRCS 2004. The PLANTS 

Database, Version 3.5. National Plant Data 

Center, (http://plants.usda.gov), Baton 

Rouge, LA. 

van Andel, T. R. 2000a. Non-Timber Forest 

Products of the North-West District of 

Guyana, Part I. Tropenbos Guyana Series 

8A. Tropenbos-Guyana Programme, 


van Andel, T. R. 2000b. Non-Timber Forest 


Guyana, Part II: A Field Guide. Tropenbos 

Guyana Series 8B. Tropenbos-Guyana 


van Dam, J. A. C. 2002. The Guyanan Plant 

Collections of Robert and Richard 

Schomburgk. Royal Botanic Gardens, Kew; 

Richmond, Surrey, United Kingdom. 

Vann, J. H. 1959. The Physical Geography of 

the Lower Coastal Plain of the Guiana 

Coast. Louisiana State University, Baton 

Rouge. 

Vann, J. H. 1969. Landforms, Vegetation, and 

Sea Level Change along the Guiana Coast 

of South America. Office of Naval 

Research, Washington, DC. 

Vegas Vilarrúbia, T., and V. Rull. 2002. Natural 

and human disturbance history of the Playa 

Medina mangrove community (Eastern 

Venezuela). Caribbean Journal of Science 

38:66-76. 

Viosca, P., Jr. 1931. Spontaneous combustion 

in the marshes of southern Louisiana. 

Ecology 12:439-442. 

Wade, D., J. Ewell, and R. Hofstetter. 1980. Fire 

in South Florida Ecosystems. Page 125. 

Southeast For. Exp. Stn., U.S. Department 

of Agriculture Forest Service, Asheville, 

NC. 

Walters, B. B. 2000. Local mangrove planting 

in the Philippines: Are fisherfolk and 

fishpond owners effective restorationists? 

Restoration Ecology 8:237-246.


Waple, A. M., J. H. Lawrimore, M. S. Halpert, 

G. D. Bell, W. Higgins, B. Lyon, M. J. 

Menne, K. L. Gleason, R. C. Schnell, J. R. 

Christy, W. Thiaw, W. J. Wright, M. J. 

Salinger, L. Alexander, R. S. Stone, and S. 

J. Camargo. 2002. Climate Assessment for 

2001. Bulletin of the American 

Meteorological Society 83:S1-S62. 

Wells, J. T., and J. M. Coleman. 1981. Periodic 

mudflat progradation, northeastern coast of 

South America: A hypothesis. Journal of 

Sedimentary Petrology 51:1069-1075. 

Wells, M. L., and A. Getis. 1999. The spatial 

characteristics of stand structure in Pinus 

torreyana. Plant Ecology 143:153-170. 

Westoby, M., M. Leishman, and J. Lord. 1997. 

Comparative ecology of seed size and 

dispersal. Pages 143-162 in J. Silvertown, 

M. Franco, and J. L. Harper, editors. Plant 

Life Histories: Ecology, Phylogeny and 

Evolution. Cambridge University Press. 

White, P. S., and S. T. A. Pickett. 1985. Natural 

disturbance and patch dynamics. Pages 3- 

13 in S. T. A. Pickett, and P. S. White, 

editors. The Ecology of Natural 

Disturbance and Patch Dynamics. 

Academic Press, Orlando. 

Whittaker, R. H. 1960. Vegetation of the 

Siskiyou Mountains, Oregon and 

California. Ecological Monographs 30:279- 

338. 

Whittaker, R. H. 1973. Ordination and 

Classification of Plant Communities. Junk, 

The Hague. 

Williams, D. 1989. The Archaic of North- 

111 

Western Guyana. History Gazette no. 7. The 

History Society, University of Guyana, 

Turkeyen. 

Willson, M., and A. Traveset. 2000. The ecology 

of seed dispersal. Pages 85-110 in M. 

Fenner, editor. The Ecology of 

Regeneration in Plant Communities, 2nd 

Edition. CAB International, Wallingford, 

UK. 

Wilton, K., and N. Saintilan. 2000. Protocols 

for Mangrove and Saltmarsh Habitat 

Mapping. Australian Catholic University, 

Sydney. 

WMO. 1999. The 1997-1998 El Niño Event: A 

Scientific and Technical Retrospective. 

Page 96. World Meteorological 

Organization, Geneva. 

WOCE Data Products Committee 2002. WOCE 

Global Data, Version 3.0. U.S. National 

Oceanographic Data Center, Silver Spring, 

MD. 

Woodroffe, C. D. 1983. Development of 

mangrove swamps behind beach ridges, 

Grand Cayman Island, West Indies. Bulletin 

of Marine Science 33:864-880. 

Woodroffe, C. D. 1988. Relict mangrove stand 

on Last Interglacial terrace, Christmas 

Island, Indian Ocean. Journal of Tropical 

Ecology 4:1-17. 

Woodroffe, C. 1992. Mangrove sediments and 

geomorphology. In A. I. Robertson, and D. 

M. Alongi, editors. Tropical Mangrove 

Ecosystems. American Geophysical Union, 


Zona, S. 1996. Flora Neotropica Monograph 71: 

Roystonea (Arecaceae: Arecoideae). New 

York Botanical Garden, New York.

112 


APPENDIX 1. 

MANGROVE ECOSYSTEMS AND FIRE 

IN THE TROPICS AND GUYANA 

This appendix is included to provide a 

literature review of information on mangrove 

ecosystems and fires in tropical forests to 

provide background on the 1997-1998 soil fires 

in the mangrove and freshwater swamps of the 

Waini Peninsula, Guyana. 

MANGROVE ECOSYSTEMS 

Mangroves are woody plants with 

adaptations to coastal conditions in the tropics. 

The word mangrove can also refer to 

communities of those plants. These plant 

communities are sometimes referred to as 

“mangal” to distinguish them from mangrove 

species or individual mangrove plants 

(Tomlinson 1986). The classification of a 

species as a mangrove species is physiological 

and functional, and these species have arisen 

from several relatively unrelated plant families. 

By the assessment of Duke et al. (1998) there 

are 71 true mangrove species in the world, 

including putative hybrids, divided among 

twenty families. The global distribution of these 

species is decidedly uneven, with 58 mangrove 

species occurring in the Indo-West Pacific 

region but only 8-13 species found in the 

America-East Pacific region; the causes for that 

disparity are among the primary questions in the 

study of mangrove biogeography. 

True mangroves possess active or passive 

adaptations to exclude salt or secrete salt from 

leaves, “viviparous” propagules that are 

dispersed by water, generally in some stage of 

germination (which allows for rapid 

establishment of seedlings) aerenchymatous 

tissues (for delivering oxygen to and within roots 

in saturated, anoxic soils), and roots modified 

for anchoring the plant in saturated soils (as 

exemplified by the prop roots of the red 

mangrove Rhizophora and the shallow, lateral 

“cable” roots of the black mangrove Avicennia) 

(Tomlinson 1986). Some researchers define 

mangrove communities as occurring only in the 

intertidal zone, however communities of 

mangrove species beyond high tide are generally 

accepted as true mangroves, and that is 

consistent with long-accepted classifications of 

inland basin and “hammock” mangrove 

community types (Lugo & Cintrón 1975; Lugo 

& Snedaker 1974). 

It has been estimated that about 25% of all 

tropical coasts are occupied by mangrove 

swamps (Tomlinson 1986). The communities 

are generally found along protected coasts with 

low wave energy, between 30ºN and 30?S 

latitude, with exact limits determined by 

minimum air temperatures, which in turn are 

influenced by ocean currents (Duke et al. 1998). 

Mangroves occur in geomorphological settings 

similar to those commonly occupied by 

herbaceous salt marshes in temperate regions. 

The communities can cover a wide range of 

structures and zonation patterns with relatively 

few species, as a result of both morphological 

and physiological elasticities of mangrove 

species that allow them to adapt to varied 

environments (Lugo & Snedaker 1974; 

Tomlinson 1986) and a lack of competition in 

marginal, saline habitats. Mangrove ecosystems 

are generally regarded as resilient from natural 

disturbances (Cintrón-Molero & Schaeffer- 

Novelli 1992), including shore erosion - 

accretion cycles and tropical cyclone damage 

(Baldwin et al. 2001; Conner et al. 1989; Imbert 

et al. 1996; Jiménez et al. 1985). Following 

severe disturbances, mangroves tend to display 

the rapid colonization characteristics of 

“pioneer” species, though in other aspects they 

possess characteristics of “climax” species such 

as large seeds and shade tolerance. Egler (1948) 

was an early researcher to conclude that 

mangrove communities generally followed a 

cyclic pattern, with rare establishment events 

and occasional catastrophic setbacks. Some 

researchers have suggested that mangrove 

community structures can be dependent upon 

periodic disturbance, and are adapted to persist


in a cyclical succession pattern (Conner et al. 

1989; Lugo 1980, 1997; Odum et al. 1982). 

Gaps created by small disturbances may play 

an important role in determining and 

maintaining mangrove forest structure (Duke 

2001), particularly in lower rainfall 

environments (Ewel et al. 1998b). 

Mangrove Values 

Attitudes towards mangrove ecosystem 

values have changed from a regard for swamps 

as low value lands best avoided or drained and 

converted to upland uses, to an almost universal 

acceptance of mangrove’s high ecological and 

economic values. These values certainly vary 

by species and community type (Ewel et al. 

1998a). High productivity in some mangrove 

systems may be due in part to landscape 

positions that allow tidal subsidies, including 

contributions of marine and terrestrially derived 

sediments (Lugo & Snedaker 1974) and tidal 

flushing of soils. Biogeochemically, mangrove 

systems function similarly to other wetlands, 

serving as sinks and transformers of nutrients, 

building biomass in wood and soils, producing 

a steady supply of litter (Lugo & Snedaker 1974; 

Odum et al. 1982), and providing appropriate 

conditions for the trapping of sediments 

(Wolanski 1995). High productivity and 

complex physical structure often makes 

mangrove swamps important fish nursery areas 

(Marshall 1994), and mangrove loss has been 

associated with declines in coastal fisheries 

(Baran & Hambrey 1998; Christensen 1983; 

Nickerson 1999; Rittibhonbhun et al. 1993). 

While the diversity of plant species in mangrove 

swamps is rather low, some mangrove 

communities support a high density, biomass, 

and diversity of invertebrates as epibionts on 

the intertidal portion of roots (Ellison & 

Farnsworth 1992; Kathiresan & Bingham 2001; 

Rützler & Feller 1987; Rützler & Feller 1996). 

They can also host a high diversity of 

vertebrates utilizing the ground and canopy 

levels (Alongi 2002; Christensen 1983). 

Mangroves can provide valuable protection 

of coastlines against erosion and storm surges. 

While early descriptions and works on 

mangrove ecology (Davis 1940; Fosberg 1947; 

Schomburgk 1922) reflected a common 

113 

assumption that mangroves actually functioned 

as builders of coastal land, that has more recently 

been revealed as incorrect. Mangroves are, 

however, often credited with a major role in 

securing sediments which have accreted through 

forceful coastal geomorphic processes 

(Augustinus 1995; Thom 1975; Vann 1969; 

Woodroffe 1992) and in the protection of reefs 

by capturing terrestrial sediments (Bossi & 

Cintrón 1990; Kathiresan & Bingham 2001; 

Pernetta 1993). 

Mangroves are utilized directly by local 

peoples for a wide variety of uses, most 

importantly fuelwood and charcoal, building 

materials, fibers, and tannin, as well as some 

foods and medicines (Alongi 2002; de la Cruz 

1983; Dugan 1993; Kovacs 1999; Snedaker 

1986; Tomlinson 1986). In parts of Asia 

mangroves are exploited commercially for 

lumber and wood chips for synthetic fiber 

production (Adeel & Pomeroy 2002; FAO 

1994). In Guyana, mangroves are primarily 

exploited for fuelwood and poles for fishing 

seines and light construction, but some tanbark 

is still gathered (Allan et al. 2002). Rhizophora 

and occasionally Avicennia are also locally cut 

into planks for construction, and shoots of 

Laguncularia (White mangrove, Kayara) are 

woven into walls and fences (pers. obs.). 

Mangrove Losses 

Worldwide losses of mangroves have often 

been reported as approximately 50% of their 

original area. Such values have been disputed 

as unverified estimates that have simply been 

repeated in the conservation literature 

(Farnsworth & Ellison 1997). It has also been 

estimated that over one third of world 

mangroves were lost in the last 20 years of the 

20 th century, with the highest rates in the 

Americas (Valiela et al. 2001). The rate of loss 

of mangroves in many developing countries has 

been estimated to be around 1% per year (Ong 

1995). More conservative studies based on 

known surveys, maps, and imagery have 

estimated that a total of perhaps one third of 

original mangrove area has been lost (Alongi 

2002; Spalding et al. 1997). All of these studies 

are limited by the scarcity of accurate multi-date 

information utilizing consistent definitions of

114 


mangrove habitats (Blasco et al. 2001). 

Estimates of the extent of mangrove vegetation 

at regional scales have been considered highly 

variable by Alongi (2002), who noted that many 

are not based on reliable field surveys or groundtruthed 

imagery and that variations could also 

be attributed to inconsistencies in classifications 

of mangrove forests. 

A common cause of mangrove degradation 

is alteration of either oceanic or terrestrial 

hydrology. Causes of hydrologic change have 

included dams (Colonnello & Medina 1998; 

Lacerda & Marins 2002; Medina et al. 2001; 

Rubin et al. 1998), diking, impoundment, 

channelization (Bacon 1975), poorly planned 

road construction (Jiménez et al. 1985; Lewis 

1990; Lugo 2002), diversion of freshwater from 

swamps (Cardona & Botero 1998; Elster et al. 

1999; Elster & Polanía 2000), and natural 

coastal morphology changes (Breen & Hill 

1969). Planned conversion of mangroves is also 

a threat, particularly near urban areas, 

permanently destroying the resource. In Asia 

and parts of South and Central America, the 

establishment of aquaculture ponds, mostly for 

shrimp, has been a major problem (Alongi 2002; 

DeWalt et al. 1996; Epler 1992; Gujja & Finger- 

Stich 1996; Pons & Fislier 1991; Robadue 1995; 

Saenger et al. 1983; Twilley 1989; Weinstock 

1994; Wolanski et al. 2000), as have salt 

production ponds (Stevenson 1997). 

Agricultural conversion of mangrove land has 

often failed in areas where acid-sulphate soils 

form through the oxidation of marine sediments 

(Hamilton & Snedaker 1984; Stevenson 1997). 

Urban and industrial conversion also have 

substantial permanent impacts, primarily from 

coastal zone related activities such as marina 

and port construction, landfills, and increasing 

waterside residential and estate construction 

(Alongi 2002; Bossi & Cintrón 1990; Ellison & 

Farnsworth 1996; Lugo 2002; Saenger et al. 

1983; Valiela et al. 2001). 

Sewage and chemical pollution can be 

substantial problems in many urban mangrove 

areas, and agricultural herbicide runoff has been 

implicated in some mangrove diebacks (Duke 

et al. 2003). Oil spills can have a particularly 

serious impact on mangrove ecosystems (Alongi 

2002; Saenger et al. 1983). Essential air 

exchange, which takes place in the adventitious 

prop roots or pneumatophores in the intertidal 

range, is easily impeded by oil (Lewis 1990). 

Oil can be extremely persistent in the sediments 

of mangrove areas (Burns et al. 1994), resulting 

in widespread sublethal damages (Duke et al. 

1997) with changes in community structure and 

productivity (Cintrón-Molero & Schaeffer- 

Novelli 1992; Garrity et al. 1994), although 

seedling establishment may be relatively 

unaffected in oil-polluted soils (Imbert et al. 

2000). 

FIRE IN THE TROPICS, 

WETLANDS, AND MANGROVE 

SWAMPS 

Fires in most ecosystems are complex 

phenomena that vary appreciably depending on 

many factors, including vegetation type, 

topography, climate, fuel type, fuel arrangement, 

fire history. The influence of man is certainly 

increasing, including an increased risk of 

ignition and potential interruption of fires in 

dissected landscapes (Whelan 1995). The return 

frequency of fires varies among ecosystem 

types, from annually in some grasslands to over 

1,000 years for some evergreen forests, and by 

some accounts, never for moist tropical forests 

(Whelan 1995). The concept of fire regimes 

suggests that some plant communities are 

adapted to fires, within certain limits. However 

the fire regime concept has drawbacks in light 

of the variable nature of fires, the great number 

of factors that must be considered to infer that 

regime, and the complications of alterations by 

human activities (Vogl 1975; Whelan 1995). 

Succession concepts can be applied to fire 

recovery, but it must be recognized that each 

situation is unique according to combinations 

of species’ life histories, the nature and patterns 

of disturbance, and the physical and chemical 

attributes of the disturbed environment (Noble 

& Slatyer 1981). Communities of fire-adapted 

species do not fit well into classical succession 

concepts, as these systems tend to remain in or 

rapidly return to their pre-fire species 

compositions after a fire because of the presence 

of persistent adults, roots, and seeds. In those 

cases, fire is an external factor causing only a 

temporary regression in an ongoing process


(Noble & Slatyer 1981). Gleason (1927) 

described a simple cyclical pattern of succession 

in fire-adapted plant communities. On the other 

hand, non fire-adapted communities may be 

good illustrations of successional concepts, 

since mortality is nearly full and reestablishment 

is mostly dependent on sources outside of the 

fire area. A typical species richness pattern for 

secondary succession often results, starting with 

an initial decrease in species followed by a sharp 

increase as many species invade, including 

disturbance specialists. Finally, a slow drop 

occurs, leveling off to those species best adapted 

for long term persistence in the developing 

environment (Whelan 1995). Along tropical 

coasts with saline soils, the expected endpoint 

of succession would be a return to dominance 

by mangrove species. That outcome would be 

influenced by the dispersal of mangrove 

propagules and any preceding occupation of 

disturbed sites by persistent, disturbanceadapted 

herbaceous species such as Typha 

(Miao et al. 2001; Newman et al. 1996; Smith 

& Newman 2001) or Spartina, and unfavorable 

changes that disturbances might have had on 

hydrology or nutrient availability, preventing 

mangrove dispersal or establishment, and thus 

setting the stage for a substantial shift in 

dominant species. 

The most destructive wildfires are 

increasingly linked to human caused 

disturbances and ignitions. Fires in fire-adapted 

communities are more likely to be catastrophic 

when they occur outside of the usual fire season, 

or when the organisms in a community are not 

at all adapted to fire (Whelan 1995). Among 

forests types that may be considered non-fire 

adapted are tropical rain forests and swamp 

forests. Mangrove forests fall into both of those 

categories. Some fire adapted species may have 

life history traits that are incompatible with 

particular fire frequencies. For example, viable 

seeds may not have adequate time to be 

produced between frequent fires, or fire 

frequencies could fall to the point that species 

requiring fires for reproduction become locally 

extinct (Noble & Slatyer 1981). 

One strategy allowing for tolerance to fire 

is through resistance of individual plants. Those 

species have evolved structures to protect 

critical living cells from the heat of fires, and 

115 

thick, corky bark is a common adaptation. 

Mesophytic plant cells die at temperatures of 

only about 50-55ºC. The trees of tropical wet 

forests are typically thin-barked, making them 

vulnerable to fires (Uhl & Kauffman 1990). 

Persistent rootstocks of plants that re-sprout 

after fires are protected by the insulation of soils. 

Mangroves have neither of these adaptations; 

in addition to thin bark, sensitive root systems 

are often exposed to air or concentrated at 

shallow soil depths. Mortality from surface fires 

is higher if upper organic soil horizons burn 

rather than simply litter and dry herbaceous 

vegetation (Wade et al. 1980). In mangrove 

systems, saturation may slow soil decomposition 

and allow a buildup of organic matter. 

Mangroves generally have exposed or shallow 

roots possessing critical aerenchyma tissue and 

lenticels for gas exchange. With these 

conditions, almost any soil fire would be 

extremely damaging to mangroves; a large 

proportion of their sensitive roots would come 

into proximity or direct contact with the fuel. 

Not surprisingly, when soil fires burned in 

mangrove swamps of the Waini Peninsula there 

was nearly, full mortality of trees. 

Laguncularia trees generally produce no 

aerial roots or sparse, small pneumatophores, 

which may have contributed to the survival of 

the Waini Peninsula soil fires by scattered small 

Laguncularia trees. These small trees may have 

also avoided mortality because of the scale of 

patchiness of the fires, while few large trees 

were unaffected. The patchiness of mortality 

together with the invasion rates of potential 

colonizers may play a major role in the path of 

recovery after a fire (Whelan 1995). 

Post-fire physical conditions, such as 

surface temperatures, wind and nutrient levels, 

can be as influential upon the path of post-fire 

recovery as the characteristics of a fire itself, 

although these factors have been rarely studied 

(Whelan 1995). Fires are often used by people 

to release nutrients in preparation for cultivation 

in nutrient limited ecosystems; fire also 

volatilizes some nutrients, particularly nitrogen, 

which are lost to the atmosphere. 

Biogeochemical cycling of nutrients during and 

after fires is in need of more study, and certainly 

seemed to be a factor in the post-fire swamps of 

the Waini Peninsula, where released nutrients

116 


apparently stimulated growth of floating aquatic 

plants and algae and affected water openness 

for mangrove dispersal. Whelan (1995) lists 

pertinent questions that have been infrequently 

addressed in fire studies, including 1) What are 

seed dispersal distances in relation to the spatial 

patterns of fires? and 2) What are the responses 

of organisms to fires in historically fire-free 

environments? These questions are approached 

here in the context of the Waini Peninsula fires. 

Fire in Tropical Ecosystems 

Fire is an integral part of some ecosystems 

in the seasonal tropics, particularly dry seasonal 

forests and tropical savannas, where the plant 

species often have fire adaptations (Mueller- 

Dombois & Goldammer 1990). Because of that, 

investigations of fire dynamics in tropical South 

America have often focused on the burning of 

savanna ecosystems and agricultural uses (Eden 

1964; Fearnside 1990; Hillis & Randall 1968). 

Primary tropical forests are generally unable to 

survive fires of even low intensity (Kauffman 

& Uhl 1990; Roberts 2000). Earlier writings on 

fire ecology tended to entirely pass over or 

minimally address effects of fire on humid 

tropical forests (Kozlowski & Ahlgren 1974), 

or focus only on damages from shifting 

cultivation and cattle ranching practices 

(Kauffman & Uhl 1990; Uhl & Buschbacher 

1985). Attention has only recently been focused 

on the phenomenon of unintentional wildfires 

that can occur in wet tropical forests during 

periodic droughts (Cochrane 2001; Cochrane & 

Laurance 2002; Uhl 1998). 

The El Niño Southern Oscillation (ENSO) 

periodically brings droughts to northwestern 

South America, while at the same time bringing 

extreme rains to the western coast around 

Ecuador and Peru (WMO 1999). “Mega-Niño” 

events have occurred periodically over the last 

two millennia, around 400, 700, 1000, and 1500 

BP, and correlate with discontinuities in cultural 

artifacts; they are a possible cause of high 

linguistic heterogeneity in tropical South 

America (Meggers 1994). Evidence has also 

been found of charcoal deposits more than 2,000 

years old in forests of French Guiana, suggesting 

that some fires could be non-human in origin 

(Charles-Dominique et al. 1998), and in the 

upper Rio Negro Basin on the Guiana Shield 

charcoal evidence has been found of fires 

occurring over 6000 years BP, nearly twice the 

estimated age of the oldest pottery shards from 

the region (Saldarriaga & West 1986). 

Lindeman (1953) relays an account, by way 

of William Beebe, of extensive fires in the 

coastal area of Guyana during an extreme 

drought in 1837, listed by Quinn (1992) as a 

year of a greater than moderate El Niño 

phenomenon. That year fires destroyed the forest 

over a vast area in the freshwater Mauritia and 

Mora swamps of the Abary River region on the 

southeastern coastal plain of Guyana. More 

recently, attention was given to the occurrence 

of fires in relatively small, understory patches 

of disturbed evergreen forest on white sands in 

Guyana (Hammond & Steege 1998). These fires 

occurred during the fairly strong 1997-1998 El 

Niño event; they were rarely over 0.5 hectare in 

size. 

While ignition through natural causes is 

possible, either accidental or intentional ignition 

by humans is considered to be the prime source 

of fires in Guyana and beyond (Hammond & 

Steege 1998; Kauffman & Uhl 1990; Lindeman 

1953). Lindeman (1953) notes that cities and 

towns in the Guianas are built almost entirely 

of wood and lack lightning rods, yet do not 

experience structural fires from lightning strikes. 

In the Venezuelan portion of the Guiana Shield, 

as in Guyana, local people often ignite savanna 

fires that burn into adjacent forested areas. In 

these cultures fire is viewed positively as 

‘cleansing,’ and as being especially useful for 

controlling populations of venomous snakes 

(Means 1995). 

Fires become more likely in moist forests 

following most physical disturbances, which 

tend to result in higher daily temperature 

maxima and lower humidity, allowing litter on 

the forest floor to dry to the point where ignition 

is possible (Kauffman & Uhl 1990). Many kinds 

of disturbances, including fires, hurricanes and 

logging, can lead to an increase in the dominance 

of vines and other fine vegetation, which 

contribute to the fuel loading for future fires 

(Mueller-Dombois & Goldammer 1990). 

There is a growing recognition that low 

intensity fires in the humid tropics can 

contribute to a positive feedback process that


leads to impacts on ever wider land areas (Uhl 

1998). Remote sensing studies in Brazil have 

indicated that once a forest has been selectively 

logged the risk of burning increases, and each 

time even a light fire occurs the probability of 

and severity of subsequent fires increase 

(Cochrane 2000). The probability of burning in 

an El Niño year also increases with selective 

logging (Cochrane 2000), and 90% of 

unintended forest fires in Amazonian Brazil are 

estimated to occur during El Niño years 

(Cochrane 2000). It has also been estimated that 

nearly 90% of fires in Amazonian Brazil have 

some relationship to a forest edge, with the 

length of the average fire return time bearing a 

positive relationship with distance from forest 

edge (Cochrane 2001). As forests become more 

fragmented their ratio of edge to area increases, 

and the increased tendency for fires to be 

initiated along forest edges suggests a cycle of 

increased frequency and impacts. Seemingly 

minor, accidental surface fires may cause 

significant impacts to tropical plant and animal 

communities and make forests far more 

vulnerable to larger fires (Laurance 2003). That 

apparently was the case when a severe dry 

season in 2003 led to fires over 20,000 km 2 or 

more in Roraima state of northern Brazil. Those 

lands were probably primed for additional fire 

by earlier fires during the El Niño event of 1998 

(Barbosa & Fearnside 1999; Barbosa et al. 

2003). Cochrane et al. (2002a) found that 

including the effects of unintended burns 

increased estimates of deforestation by 129 

percent from 1993-1995 in Paragominas in 

Northeastern Brazil. These incremental, positive 

feedback augmented fire impacts may merit 

attention equal to that given to deforestation by 

agricultural clearing. Similar dynamics have 

possibly been at work in the Western Ghats of 

India over longer periods of time, where very 

short return intervals for fires and severe 

changes in species compositions of forests have 

been documented (Kodandapani et al. 2003). 

Tropical forest fires are often re-ignited by 

smouldering logs when litter conditions become 

dry (Cochrane et al. 2002b). During the early 

2001 dry season on the Waini Peninsula, 

recently ignited fires were observed smoldering 

in fallen logs of trees killed during the 1998 

fires; these sometimes spread a short distance 

117 

in organic soils and dry herbaceous vegetation. 

Those re-burned areas generally had very sparse 

vegetation cover of scattered Acrostichum ferns 

and scrub Laguncularia and therefore dried 

quickly in the open sun. 

Fire in Wetlands 

While the idea of fire occurrence in 

wetlands may seem unusual, many wetlands 

pass through dry hydrologic conditions and may 

burn occasionally, though forested wetlands are 

rarely affected (Lugo 1995). Earlier studies of 

fire in wetlands have been predominantly 

confined to seasonally flooded herbaceous 

wetlands, tidal marshes and peatlands in North 

America (Kirby et al. 1988), often with a focus 

on prescribed burns for wildlife management 

(Cerulean & Engstrom 1995; Hackney & de la 

Cruz 1981). Many wetland types, including 

marshes, freshwater swamps, and mangrove 

swamps, are considered to be fire-independent 

systems, which generally require preparation for 

burning and deliberate ignition during dry 

periods (Vogl 1975). Nonetheless, fires in those 

types of ecosystems tend to be catastrophic, and 

recovery in burned wetlands is likely to include 

wide areas of weedy plants that can quite 

persistent. To avoid such results, controlled fires 

for management in marshes are typically set 

when the soil is damp, resulting in cover burns 

affecting only above ground biomass. 

Decomposition in wetlands often proceeds at a 

lower rate than production (Mitsch & Gosselink 

1993), so that organic soils accumulate that, 

when ignited during drought or drawdown, 

usually fuel smoldering peat fires that produce 

copious smoke, which can pose serious visibility 

and health problems (Hungerford et al. 1995). 

Peat burns can result in open water and take 

many decades to recover; they are never 

considered a useful management tool (Nyman 

& Chabreck 1995). Some raised bogs and tidal 

marshes are subject to fires, and may possess 

some fire adapted species, in which case 

regeneration of vegetation can be rapid 

(Clarkson 1997; Timmins 1992). 

Among forested wetlands, Cypress 

(Taxodium distichum Rich.) swamps are known 

to suffer infrequent fires with return frequencies 

of hundreds of years; these fires may play a

118 


limited role in stimulating seed production and 

therefore recruitment of seedlings (Cook & Ewel 

1992; Ewel 1995; Glasser 1985). Although some 

varieties of Cypress may be more fire adapted 

than others, fires that burn significantly into 

organic soils will damage a high percentage of 

roots and kill most Cypress trees and all 

seedlings (Ewel 1995). Atlantic white cedar 

(Chamaecyparis thyoides(L.) Britton, Sterns & 

Poggenb.) wetlands in the Northeastern United 

States have been prehistorically disturbed by 

crown and soil fires, which apparently resulted 

in substantial declines in cedar populations, and 

which only after long periods were finally 

followed by regeneration (Motzkin et al. 1993). 

Fires in tropical wetlands are a very poorly 

studied phenomenon, being confined primarily 

to large-scale peat fires in Asia, particularly 

Indonesia, that have become associated with El 

Niño events (Fuller & Fulk 2001; Kinnaird & 

O’Brien 1998; Roberts 2000; Trenberth 1999). 

Fire is a factor in the maintenance and spread 

of Mauritia palms in the Llanos savannas of 

Venezuela and similar wet savannas of coastal 

Guyana (Middleton 1999; Myers 1990). The 

species composition of plant communities on 

and around the tepuis of the Guiana Shield in 

Southern Venezuela are also probably 

influenced by fires (Means 1995). 

Fire in Mangrove Literature 

In the terminology of Noble and Slatyer’s 

(1980) vital attributes classification for 

successional pathways following disturbance, 

mangrove fires should be described as 

catastrophic events that do not normally occur 

over the life span of the affected species. The 

species of such systems have not been selected 

for and possess few mechanisms for rapid 

recovery. 

There is limited information in the literature 

on the occurrence of fire in mangrove 

ecosystems. The few cases have been primarily 

concerned with small-scale mortality resulting 

from lightning strikes (Duke 2001; Sherman et 

al. 2000; Smith et al. 1994; Wade et al. 1980) 

with affected areas only in the range of 10 to 

100 square meters. Colonization of these small 

lightning gaps may play a significant role in 

patterns of species distribution in some 

mangrove forests, particularly those forests that 

are not subject to large scale disturbances such 

as hurricanes (Duke 2001). Lugo (1980), in his 

work on mangrove succession and disturbance, 

mentioned only briefly the sensitivity to fire of 

raised “Hammock” mangrove swamps of the 

South Florida Everglades. Fires burning in 

adjacent marsh or upland vegetation have often 

been credited with maintenance of mangrovemarsh 

boundaries in Florida (Lugo 1997; 

Middleton 1999; Odum et al. 1982; Wade et al. 

1980). 

Vogl (1975) characterized mangrove 

swamps as fire independent systems in which 

fire leads to catastrophic results. His list of 

mangrove swamp properties that contribute to 

exclusion of fire included high water tables, 

rapid decomposition of organic fuel, few 

flammable oils, and sparse understories. He 

noted that fires in such systems are usually 

initiated by man during extreme droughts and 

result in high mortality, even from seemingly 

mild surface fires, and revegetation includes 

many weedy species. Without citing specific 

instances, Wade et al. (1980) also state that very 

low intensity, creeping soil fires easily kill 

mangroves. These observations are in agreement 

with the emerging understanding of the effects 

of surface fires on wet tropical forests in the 

Amazon basin (Cochrane 2003; Cochrane & 

Laurance 2002). Wade et al. (1980) also 

described cases of fires in dried dead mangrove 

debris left after hurricanes, as well as near the 

northern limits of mangrove vegetation in 

Florida frost killed mangroves are susceptible 

to fire. In very rare cases frost-killed leaves have 

ignited before dropping, causing intense 

mangrove crown fires. 

In the Guianas, there has been minor 

documentation in the literature of fire in or near 

mangrove ecosystems. Lindeman (1953) 

mentions fire along parts of the Surinam 

shoreline, describing it as an infrequent agent 

that accelerates his model of succession in which 

Avicennia swamp forest gives way to Eleocharis 

R.Br. marsh. Pons and Pons (1975) also mention 

fires in Avicennia swamps isolated behind beach 

ridges at Little Orange Creek on the coast of 

Surinam, with the disturbance initiating various 

types of herbaceous vegetation. Their model 

describes a post-fire succession starting with


shallow open water or Eleocharis meadows with 

Avicennia slowly colonizing. That returns to a 

“climax” of Avicennia swamp if undisturbed, 

or becomes arrested at a Typha marsh or 

Cyperus-Montrichardia marsh stage. Details are 

not given in the Pons and Pons paper about the 

extent of areas affected by fires, but maps are 

included indicating belts of Eleocharis up to 1 

kilometer wide and patches of Typha on the 

inland margin of Avicennia swamps, some over 

100 hectares in size. There, both of these 

herbaceous communities are described as 

persisting possibly as a result of recurring fires 

or other disturbances, however no specific fires 

were documented, and the frequency of fires is 

apparently low. 

Observations at Almond Beach on the 

Waini Peninsula suggest that patterns of 

vegetation change on Guyana’s coastline may 

follow pathways similar to those in Surinam 

(Lindeman 1953; Pons & Pons 1975), with 

shifts in the mangrove/marsh boundary 

occurring in a punctuated manner, initiated 

during extreme droughts by catastrophic fires 

that define a new boundary by burning forested 

areas that are no longer tidally inundated. The 

occluded mangrove swamps are then converted, 

at least for a time, to herbaceous, somewhat salttolerant 

vegetation. 

Status of Mangroves in Guyana 

Because the soils beyond Guyana’s coastal 

plain are generally laterites or white sands with 

poor fertility, over 90% of the population lives 

on the coastal plain (Merrill 1993), with a sparse 

transportation system beyond the coast. That, 

in combination with a timber industry that has 

only recently reached large scales, has left 

mangroves as one of the most highly impacted 

forest types in Guyana (GFC & CIDA 1989). 

The coast of the Guianas is credited with 

optimal growing conditions for the black 

mangrove Avicennia germinans, which is 

reported to reach heights of up to 100 feet in 

Surinam (Lindeman 1953; Vann 1959, 1969); 

similar heights have been observed in the coastal 

forests of the Waini Peninsula, and diameters 

in coastal swamps reach more than 50 cm, with 

exceptional diameters of over 2 meters for 

scattered mature trees found just inland of the 

119 

Waini River (see Figure 1.16). 

The zonation of species within mangrove 

communities in the Guianas is quite distinct 

from that typically described for the more often 

studied mangroves on calcareous substrates in 

the Caribbean (Lindeman & Mori 1989; West 

1977). The black mangrove, Avicennia 

germinans, forms a moderate to wide band 

nearest the ocean. The red mangrove, 

Rhizophora mangle, the prop-rooted seaward 

species in most Caribbean communities, is 

mixed in to varying degrees behind the black 

mangroves (Granville 1992). Farther inland, 

forest composition grades into freshwater 

swamp (Bacon 1990). Red mangroves are also 

found along the banks of the tidal rivers in 

brackish water, usually as the riverine species 

Rhizophora racemosa G. Mey., a species with a 

narrower distribution in the Americas than R. 

mangle. The white mangrove Laguncularia 

racemosa is not a dominant tree in the 

mangroves of the Guianas, but is very common 

as an understory shrub or treelet. 

The early colonization of Guyana by Dutch 

planters led to the diking of long stretches of 

coastline and tidal river banks for development 

of agricultural from coastal swamps and 

marshes. That affected almost all of the coastline 

southeast of the Pomeroon River, where the 

coastal plain is widest (Daly 1995). Daily 

operation and maintenance of the drainage and 

sea defense infrastructure places a great 

economic demand on the Guyanese economy 

and has at times been a significant focus of 

international assistance (Richardson 1987). 

Before reclamation, much of the coastal plain 

was covered by marshes and swamps, with areas 

closest to the sea periodically to frequently 

flooded by saline tides. The drainage of the 

coastal wetland soils probably resulted in 

subsidence from de-watering and oxidation of 

organic matter, leaving the land below mean sea 

level and making any restoration difficult. 

No descriptions have been found of the 

extent of Guyana’s mangroves prior to diking, 

but it is almost certain that a vast area was 

destroyed since the beginning of colonization 

(Richardson 1987). On the southeastern Guyana 

coast, many mangroves that remained outside 

of seawalls have disappeared from the effects 

of altered hydrology and erosion of sediments

120 


along the shore (Singhroy 1996). In some coastal 

areas sediment accretion outside of seawalls 

allows substantial colonization by Avicennia, 

such as has occurred on the coast between the 

Pomeroon and Essequibo Rivers (pers. obs.). 

These may be ephemeral and eventually be lost 

to erosion as mudbanks migrate along the coast. 

Estimates of the current mangrove area in 

Guyana range from Spalding’s (1997) 71,700 

ha (whose methods were positively appraised 

by Alongi) to 80,000 ha (Snedaker 1986) with 

the highest area of 150,000 ha (Saenger et al. 

1983) or slightly more (Huber et al. 1995). After 

agricultural conversion, the major threat to 

remaining mangroves in Guyana, both along the 

coast and in estuarine areas, was once fuelwood 

collection (GFC & CIDA 1989; Guyana Agency 

for Health 1992) which affected Avicennia as 

well as Rhizophora. These species are 

decreasingly utilized as sources for superior 

fuelwood, charcoal and tanbark, but are still 

harvested for construction and fishing poles 

(Allan et al. 2002). Mangroves near populated 

areas are highly degraded by intense use, 

evidenced by very low mean tree diameters in a 

1 ha plot near Alness Village, where a mean dbh 

of 16 cm was recorded (Ramdass et al. 1997). 

Where mangrove swamps remain in Guyana 

they are credited with the stabilization of 

accreted sediments and erosion prevention (EPA 

Guyana 2000; GFC & CIDA 1989). 

Fire Detection Resources 

There are a growing number of resources 

that can alert resource managers and policy 

makers to wildfires, making timely reaction to 

events possible. NOAA’s Operational 

Significant Event Imagery (http:// 

www.osei.noaa.gov/Events/Fires/Guyana/), 

archives have included past events sensed by 

AVHRR channel 3 imagery. Only events for 

March 30, 1998 were available for Guyana, 

indicating no fire activity on the Waini 

Peninsula. The MODIS satellite has been 

operational since 2000, and provides 1 kilometer 

resolution fire and thermal anomaly data at a 

twice daily frequency that can be downloaded 

ready for use in GIS systems (http://modisfire.umd.edu/). 

The National Geophysical Data 

Center’s Satellite Fire Detection Viewer (http:/ 

/map.ngdc.noaa.gov/website/firedetects/ 

viewer.htm) provides recent possible fire data; 

the ABBA GOES-10 layer provides fire 

detections that include Guyana and can be 

downloaded for use in GIS systems. Ideally 

these data sources should be regularly monitored 

and analyzed to increase the likelihood that 

managers become aware of fires while they are 

in progress (Cochrane 2002). Another 

development advancing the detection of and 

communication about wildfires is the recent 

foundation of the Regional South America 

Wildland Fire Network (IFFN 2004), which is 

internationally funded by governmental and 

non-governmental organizations. 

CONCLUSION 

Like terrestrial forested ecosystems, 

mangrove ecosystems can vary widely in their 

structure and function. The mangroves of 

Guyana and not typical of the low sediment 

Caribbean systems most often studied and 

published on in the Neotropics, but fall well 

within the worldwide varieties of mangrove 

systems. As is true of many wetland types, they 

are dynamic systems, that change along with the 

hydrology and geomorphology of the landscape 

in which they are situated. In places like the 

coast of the Guianas, changes are accentuated 

by coastal sediment dynamics as well as by 

fluctuations of climate over time, possibly 

making the system increasingly vulnerable to 

disturbances such as fires. If understood, such 

disturbances can be anticipated and managed. 

On wild sections of the coast of the Guianas, 

that would primarily include suppression of fire 

ignition during extreme dry years, and 

connected efforts at education and planning of 

settlement and land use patterns. Such practices 

might have an impact on the frequency of fires, 

and the conservation of these ecologically and 

culturally valuable mangrove ecosystems. 

LIST OF REFERENCES 

(for Appendix 1) 

Adeel, Z., and R. Pomeroy. 2002. Assessment 

and management of mangrove ecosystems 

in developing countries. Trees 16:235-238.


Allan, C., S. Williams, and R. Adrian 2002. The 

Socio-Economic Context of the Harvesting 

and Utilisation of Mangrove Vegetation. 

Guyana Forestry Commission, 


Alongi, D. M. 2002. Present state and future of 

the world’s mangrove forests. 

Environmental Conservation 29:331-349. 

Augustinus, P. G. E. F. 1995. Geomorphology 

and sedimentation of mangroves. In G. M. 

E. Perillo, editor. Geomorphology and 

Sedimentation of Estuaries. Elsevier, 

Amsterdam. 

Bacon, P. R. 1975. Recovery of a Trinidadian 

mangrove swamp from attempted 

reclamation. In G. E. Walsh, S. C. Snedaker, 

and H. J. Teas, editors. Proceedings of the 





Bacon, P. R. 1990. Ecology and management 

of swamp forests in the Guianas and 

Caribbean region. Pages 193-213 in A. E. 

Lugo, M. Brinson, and S. Brown, editors. 

Ecosystems of the World 15, Forested 

Wetlands. Elsevier, Amsterdam. 

Baldwin, A. H., M. Egnotovich, M. Ford, and 

W. Platt. 2001. Regeneration in fringe 

mangrove forests damaged by Hurricane 

Andrew. Plant Ecology 157:151-164. 

Baran, E., and J. Hambrey. 1998. Mangrove 

conservation and coastal management in 

southeast Asia: What impact on fishery 

resources? Marine Pollution Bulletin 

37:431-440. 

Barbosa, R. I., and P. M. Fearnside. 1999. 

IncL ndios na Amazônia brasileira: 

Estimativa da emissno de gases do efeito 

estufa pela queima de diferentes 

ecossistemas de Roraima na passagem do 

evento “El NiZo” (1997/98). Acta 

Amazonica 29:513-534. 

Barbosa, R. I., M. R. Xaud, G. N. F. Silva, and 

A. C. Cattâneo. 2003. Forest Fires in 

Roraima, Brazilian Amazonia. 

International Forest Fire News 28:51-56. 

Blasco, F., J. L. Carayon, and M. Aizpuru. 2001. 

World mangrove resources. ISME/ 

GLOMIS Electronic Journal 1:1-3. 

Bossi, R. H., and G. Cintrón 1990. Mangroves 

121 

of the Wider Caribbean: Toward 

Sustainable Management. Caribbean 

Conservation Association, St. Michael, 

Barbados. 

Breen, C. M., and B. J. Hill. 1969. A mass 

mortality of mangroves in the Kosi Estuary. 

Transactions of the Royal Society of South 

Africa 38:285-303. 

Burns, K. A., S. D. Garrity, D. Jorissen, J. 

MacPherson, and M. Stoelting. 1994. The 

Galeta oil spill. II: Unexpected persistence 

of oil trapped in mangrove sediments. 

Estuarine, Coastal, and Shelf Science 

38:349-364. 

Cardona, P., and L. Botero. 1998. Soil 

characteristics and vegetation structure in 

a heavily deteriorated mangrove forest in 

the Caribbean coast of Colombia. 


Cerulean, S. I., and R. T. Engstrom, editors. 

1995. Fire in Wetlands: A Management 

Perspective. Proceedings of the Tall 

Timbers Fire Ecology Conference, No. 19. 

Tall Timbers Research Station, Tallahassee, 

FL. 

Charles-Dominique, P., P. Blanc, D. Larpin, M.- 

P. Ledru, B. Riera, C. Sarthou, M. Servant, 

and C. Tardy. 1998. Forest perturbations 

and biodiversity during the last ten thousand 

years in French Guiana. Acta Oecologia 

19:295-302. 

Christensen, B. 1983. Mangroves - What are 

they worth? Unasylva 35:2-15. 

Cintrón-Molero, G., and Y. Schaeffer-Novelli. 

1992. Ecology and management of New 

World mangroves. Pages 233-258 in U. 

Seeliger, editor. Coastal Plant Communities 

of Latin America. Academic Press. 

Clarkson, B. R. 1997. Vegetation recovery 

following fire in two Waikato peatlands at 

Whangamarino and Moanatuatua, New 

Zealand. New Zealand Journal of Botany 

35:167-179. 

Cochrane, M. A. 2000. Forest fire, deforestation 

and landcover change in the Brazilian 

Amazon. Pages 170-176 in L. F. 

Neuenschwander, K. C. Ryan, G. E. 

Gollberg, and J. D. Greer, editors. 

Proceedings from, The Joint Fire Science 

Conference and Workshop, June 15-17, 

1999, “Crossing the Millennium:

122 


Integrating Spatial Technologies and 

Ecological Principals for a New Age in Fire 

Management”. University of Idaho and the 

International Association of Wildland Fire, 

Moscow, Idaho. 

Cochrane, M. A. 2001. Synergistic Interactions 

between Habitat Fragmentation and Fire in 

Evergreen Tropical Forests. Conservation 

Biology 15:1515-1521. 

Cochrane, M. A. 2002. Spreading Like 

Wildfire—Tropical Forest Fires in Latin 

America and the Caribbean: Prevention, 

Assessment and Early Warning. Page 96. 

United Nations Environment Programme 

(UNEP), Regional Office for Latin America 

and the Caribbean, Mexico City. 

Cochrane, M. A. 2003. Fire science for 

rainforests. Nature 421:913-919. 

Cochrane, M. A., A. Alencar, M. D. Schulze, C. 

M. Souza, Jr., P. Lefebvre, and D. Nepstad. 

2002a. Investigating positive feedbacks in 

the fire dynamic of closed canopy tropical 

forests. Pages 285-298 in C. H. Wood, and 

R. Porro, editors. Deforestation and 

Landuse in the Amazon. University of 

Florida Press, Gainesville. 

Cochrane, M. A., and W. F. Laurance. 2002. Fire 

as a large-scale edge effect in Amazonian 

forests. Journal of Tropical Ecology 18:311- 

325. 

Cochrane, M. A., D. L. Skole, E. A. T. 

Matricardi, C. Barber, and W. 

Chomentowski. 2002b. Interaction and 

Synergy between Selective Logging, Forest 

Fragmentation and Fire Disturbance in 

Tropical Forests: Case Study Mato Grosso, 

Brazil. Pages 1-20. Michigan State 

University, East Lansing, Michigan. 

Colonnello, G., and E. Medina. 1998. Vegetation 

changes induced by dam construction in a 

tropical estuary: the case of the Mánamo 

river, Orinoco Delta (Venezuela). Plant 

Ecology 139:145-154. 

Conner, W. H., J. W. Day, R. H. Bauman, and J. 

M. Randall. 1989. Influence of hurricanes 

on coastal ecosystems along the northern 

Gulf of Mexico. Wetlands Ecology and 


Cook, S., and K. C. Ewel. 1992. Regeneration 

in burned cypress swamps. Florida Scientist 

56:62-64. 

Daly, V. T. 1995. A Short History of the 

Guyanese People. Macmillan, London. 

Davis, J. H. 1940. The ecology and geologic role 

of mangroves of Florida. Papers of the 

Tortugas Laboratory 32:303-341. 

de la Cruz, A. A. 1983. Traditional practices 

applied to the management of mangrove 

areas in Southeast Asia. International 

Journal of Ecology and Environmental 

Science 9:13-19. 

DeFilipps, R. A., S. L. Maina, and J. Crepin 

2004. Medicinal Plants of the Guianas 

(Guyana, Surinam, French Guiana), web 

version. Department of Botany, 


DeWalt, B. R., P. Vergne, and M. Hardin. 1996. 

Shrimp aquaculture development and the 

environment: People, mangroves and 

fisheries on the Gulf of Fonseca, Honduras. 

World Development 24:1193-1208. 

Dugan, P. J. 1993. Distilling lessons from the 

case studies. Pages 153-156 in T. J. Davis, 

editor. Towards the wise use of wetlands: 

Report of the Ramsar Convention Wise Use 

Project. Ramsar Convention Bureau, Gland, 

Switzerland. 

Duke, N. C. 2001. Gap creation and regenerative 

processes driving diversity and structure of 

mangrove ecosystems. Wetlands Ecology 

and Management 9:257-269. 

Duke, N. C., M. C. Ball, and J. Ellison. 1998. 

Factors influencing biodiversity and 

distributional gradients in mangroves. 

Global Ecology and Biogeography Letters 

7:27-47. 

Duke, N. C., A. M. Bell, D. K. Pedersen, C. M. 

Roelfsema, L. M. Godson, K. N. Zahmel, 

J. Mackenzie, and S. Bengtson-Nash 2003. 

Mackay Mangrove Dieback. Investigations 

in 2002 with recommendations for further 

research, monitoring and management. 

Report to Queensland Department of 

Primary Industries Northern Fisheries 

Centre, and the Community of Mackay. 11 

September 2003. Marine Botany Group, 

Centre for Marine Studies, The University 

of Queensland, Brisbane. 

Duke, N. C., Z. S. Pinzón M., and M. C. Prada 

T. 1997. Large-scale damage to mangrove 

forests following two large oil spills in 

Panama. Biotropica 29:2-14.


Eden, M. J. 1964. The Savanna Ecosystem - 

Northern Rupununi, British Guiana. 

Department of Geography, McGill 

University, Montreal, Canada. 

Egler, F. E. 1948. The dispersal and 

establishment of red mangrove, 

Rhizophora, in Florida. Caribbean Forester 

9:299-310. 

Ellison, A. M., and E. J. Farnsworth. 1992. The 

ecology of Belizean mangrove-root fouling 

communities: patterns of epibiont 

distribution and abundance, and effects on 

root growth. Hydrobiologia 247:87-98. 

Ellison, A. M., and E. J. Farnsworth. 1996. 

Anthropogenic disturbance of Caribbean 

mangrove ecosystems: Past impacts, 

present trends, and future predictions. 


Elster, C., L. Perdomo, and M.-L. Schnetter. 

1999. Impact of ecological factors on the 

regeneration of mangroves in the Cienaga 

Grande de Santa Marta, Colombia. 


Elster, C., and J. Polanía. 2000. Regeneration 

of mangrove forests in the Cienaga Grande 

de Santa Marta (Colombia). Actualidades 

Biologicas 22:29-36. 

EPA Guyana. 2000. Integrated Coastal Zone 

Management Action Plan. Page 31 pp. 

Environmental Protection Agency of 


Epler, B. 1992. Social ramifications of 

mariculture on coastal Ecuadorian 

communities. Pages 24-32 in R. B. Pollnac, 

and P. Weeks, editors. Coastal Aquaculture 

in Developing Countries: Problems and 

Perspectives. International Center for 

Marine Resource Development (ICMRD) 

at the University of Rhode Island, 

Providence. 

Ewel, K. C. 1995. Fire in cypress swamps in 

the southeastern United States. Pages 111- 

116 in S. I. Cerulean, and R. T. Engstrom, 

editors. Fire in Wetlands: A Management 




FL. 

Ewel, K. C., R. R. Twilley, and J. E. Ong. 1998a. 

Different kinds of mangrove forests provide 

different goods and services. Global 

123 

Ecology and Biogeography Letters 7:83-94. 

Ewel, K. C., S. Zheng, Z. S. Pinzón, and J. A. 

Bourgeois. 1998b. Environmental effects of 

canopy gap formation in high-rainfall 

mangrove forests. Biotropica 30:510-518. 

FAO 1994. Mangrove Management Guidelines. 

U.N. Food and Agriculture Organization, 

Rome. 

Farnsworth, E. J., and A. M. Ellison. 1997. The 

global conservation status of mangroves. 

Ambio 26:328-334. 

Fearnside, P. M. 1990. Fire in the tropical rain 

forest of the Amazon basin. Pages 106-116 

in J. G. Goldammer, editor. Fire in the 

Tropical Biota: Ecosystem Processes and 

Global Challenges. Springer-Verlag, Berlin. 

Fosberg, F. R. 1947. Micronesian mangroves. 

Journal of the New York Botanical Garden 

48:128-138. 

Fuller, D. O., and M. Fulk. 2001. Burned area 

in Kalimantan, Indonesia mapped with 

NOAA-AVHRR and Landsat TM imagery. 

International Journal of Remote Sensing 

22:691-697. 

Garrity, S. D., S. D. Levings, and K. A. Burns. 

1994. The Galeta oil spill. I: Long term 

effects on the physical structure of the 

mangrove fringe. Estuarine, Coastal, and 

Shelf Science 38:327-348. 

Gentry, A. H. 1993. A Field Guide to the 

Families and Genera of Woody Plants of 

Northwest South America (Colombia, 

Ecuador, Peru), with Supplementary Notes 

on Herbaceous Taxa. Conservation 

International, Washington, DC. 

GFC, and CIDA 1989. National Forestry Action 

Plan 1990-2000. Guyana Forestry 

Commission and Canadian International 

Development Agency, Georgetown, 

Guyana. 

Glasser, J. E. 1985. Successional trends on tree 

islands in the Okeefenokee Swamp as 

determined by interspecific association 

analysis. American Midland Naturalist 

113:287-293. 

Gleason, H. A. 1927. Further views on the 

succession-concept. Ecology 8:299-326. 

Görts-van Rijn, A. R. A., editor. 1985 -. Flora 

of the Guianas. Royal Botanic Gardens, 

Kew; Richmond, Surrey, United Kingdom. 

Granville, J.-J. d. 1992. Les formations végétales

124 


actuelles des zones côtiPre et subcôtiPre des 

Guyanes. Pages 231-249 in M. T. Proust, 

editor. Évolution des Littoraux de Guyane 

et de la Zone Caríbe Méridionale Pendant 

le Quaternaire. ORSTOM Editions, Paris. 

Gujja, B., and A. Finger-Stich. 1996. What price 

prawn? Shrimp aquaculture’s impact in 

Asia. Environment 38:12-15 & 33-39. 

Guyana Agency for Health, Science, Education, 

Environment and Food Policy. 1992. 

Guyana/UNEP Country Study of Biological 

Diversity. Coordinated by the Guyana 

Agency for Health, Science, Education, 

Environment, and Food Policy, Liliandaal, 


Hackney, C. T., and A. A. de la Cruz. 1981. 

Effects of fire on brackish marsh 

communities: management implications. 

Wetlands 1:75-86. 

Hamilton, L. S., and S. C. Snedaker, editors. 

1984. Handbook for Mangrove Area 

Management. United Nations Environment 

Programme and East-West Center, 

Environment and Policy Institute, Paris. 

Hammond, D. S., and H. t. Steege. 1998. 

Propensity for fires in Guianan Rainforests. 


Hillis, T. L., and R. E. Randall, editors. 1968. 

The Ecology of the Forest/Savanna 

Boundary (Proceedings of the I.G.U. Humid 

Tropics Commission Symposium, 

Venezuela, 1964). Department of 

Geography, McGill University, Montreal, 

Canada. 

Huber, O., G. Gharbarran, and V. Funk 1995. 

Vegetation Map of Guyana (Preliminary 

Version). Centre for the Study of Biological 

Diversity, University of Guyana, 

Georgetown. 

Hungerford, R. D., W. H. Frandsen, and K. C. 

Ryan. 1995. Ignition and burning 

characteristics of organic soils. Pages 78- 

91 in S. I. Cerulean, and R. T. Engstrom, 





FL. 

IFFN. 2004. Regional South America Wildland 

Fire Network. International Forest Fire 

News 31:38-47. 

Imbert, D., P. Labbé, and A. Rousteau. 1996. 

Hurricane damage and forest structure in 

Guadeloupe, French West Indies. Journal 

of Tropical Ecology 12:663-680. 

Imbert, D., A. Rousteau, and P. Scherrer. 2000. 

Ecology of mangrove growth and recovery 

in the Lesser Antilles: State of knowledge 

and basis for restoration projects. 

Restoration Ecology 8:230-236. 

Jiménez, J. A., A. E. Lugo, and G. Cintrón. 1985. 

Tree mortality in mangrove forests. 


Kathiresan, K., and B. L. Bingham. 2001. 

Biology of mangroves and mangrove 

ecosystems. Advances in Marine Biology 

40:81-251. 

Kauffman, J. B., and C. Uhl. 1990. Interactions 

of anthropogenic activities, fire, and rain 

forests in the Amazon basin. Pages 117-134 

in J. G. Goldammer, editor. Fire in the 

Tropical Biota: Ecosystem Processes and 

Global Challenges. Springer-Verlag, Berlin. 

Kinnaird, M. F., and T. G. O’Brien. 1998. 

Ecological effects of wildfire on lowland 

rainforest in Sumatra. Conservation 

Biology 12:954-956. 

Kirby, R. E., S. J. Lewis, and T. N. Sexton. 1988. 

Fire in North American wetland ecosystems 

and fire-wildlife relations: An annotated 

bibliography. U.S. Fish and Wildlife 

Services, Biological Report 88:1-146. 

Kodandapani, N., M. A. Cochrane, and R. 

Sukumar. 2003. Conservation threat of 

increasing fire frequencies in the Western 

Ghats, India. Conservation Biology 

18:1553-1561. 

Kovacs, J. M. 1999. Assessing mangrove use at 

the local scale. Landscape and Urban 

Planning 43. 

Kozlowski, T. T., and C. E. Ahlgren, editors. 

1974. Fire and Ecosystems. Academic 

Press, New York. 

Lacerda, L. D., and R. V. Marins. 2002. River 

damming and changes in mangrove 

distribution. ISME/GLOMIS Electronic 

Journal 2. 

Lanjouw, J., editor. 1964-. Flora of Suriname. 

E. J. Brill, Leiden. 

Lasser, T., editor. 1964-1992. Flora de 

Venezuela. Instituto Botánico, Caracas. 

Laurance, W. F. 2003. Slow burn: the insidious


effects of surface fires on tropical forests. 

Trends in Ecology and Evolution 18:209- 

212. 

Lewis, R. R. 1990. Creation and restoration of 

coastal wetlands in Puerto Rico and the 

U.S. Virgin Islands. Pages 103-123 in J. A. 

Kusler, and M. E. Kentula, editors. Wetland 

Creation and Restoration: The Status of the 

Science. Island Press, Washington, DC. 

Lindeman, J. C. 1953. The Vegetation of the 

Coastal Region of Suriname. 

Rijksuniversiteit te Utrecht, Utrecht, 


Lindeman, J. C., and S. A. Mori. 1989. The 

Guianas. In D. G. Campbell, editor. Floristic 

Inventory of Tropical Countries: Plant 

Systematics, Collections, and Vegetation, 

plus Recommendations for the Future. New 

York Botanical Garden, New York. 

Lugo, A. E. 1980. Mangrove ecosystems: 

successional or steady state? Biotropica 

12:65-72. 

Lugo, A. E. 1995. Fire and wetland 

management. Pages 1-9 in S. I. Cerulean, 

and R. T. Engstrom, editors. Fire in 

Wetlands: A Management Perspective. 

Proceedings of the Tall Timbers Fire 

Ecology Conference, No. 19. Tall Timbers 

Research Station, Tallahassee, FL. 

Lugo, A. E. 1997. Old growth mangrove forests 

in the United States. Conservation Biology 

11:11-20. 

Lugo, A. E. 2002. Conserving Latin American 

and Caribbean mangroves: issues and 

challenges. Madera y Bosques Número 

especial:5-25. 

Lugo, A. E., and G. Cintrón. 1975. The 

mangroves of Puerto Rico and their 

management. Pages 825-846 in G. E. 







Lugo, A. E., and S. C. Snedaker. 1974. The 

ecology of mangroves. Annual Review of 

Ecology and Systematics 5:39-64. 

Marshall, N. 1994. Mangrove conservation in 

relation to overall environmental 

considerations. Hydrobiologia 285:303- 

125 

309. 

Means, D. B. 1995. Fire ecology of the Guayana 

region, Northeastern South America. Pages 

61-77 in S. I. Cerulean, and R. T. Engstrom, 





FL. 

Medina, E., H. Fonseca, F. Barboza, and M. 

Francisco. 2001. Natural and man-induced 

changes in a tidal channel mangrove system 

under tropical semiarid climate at the 

entrance of the Maracaibo lake (Western 

Venezuela). Wetlands Ecology and 


Meggers, B. J. 1994. Archaeological evidence 

for the impact of Mega- NiZo events on 

Amazonia during the past two millenia. 

Climatic Change 28:321-338. 

Merrill, T. L., editor. 1993. Guyana and Belize: 

country studies. Federal Research Division, 

U.S. Department of Army, Washington, DC. 

Miao, S. L., P. V. McCormick, S. Newman, and 

S. Rajagopalan. 2001. Interactive effects of 

seed availability, water depth, and 

phosphorus enrichment on cattail 

colonization in an Everglades wetland. 

Wetlands Ecology and Management 9:39- 

47. 

Middleton, B. 1999. Wetland Restoration, Flood 

Pulsing and Disturbance Dynamics. John 

Wiley & Sons, Inc., New York. 

Mitsch, W. J., and J. G. Gosselink 1993. 

Wetlands. Van Nostrand Rheinhold, New 

York. 

Motzkin, G., W. A. Patterson, III, and N. E. R. 

Drake. 1993. Fire history and vegetation 

dynamics of a Chamaecyparis thyoides 

wetland on Cape Cod. Journal of Ecology 

81:391-402. 

Mueller-Dombois, D., and J. G. Goldammer. 

1990. Fire in the tropical ecosystems and 

global environmental change: an 

introduction. Pages 1-10 in J. G. 

Goldammer, editor. Fire in the Tropical 

Biota: Ecosystem Processes and Global 

Challenges. Springer-Verlag, Berlin. 

Myers, R. L. 1990. Palm swamps. Pages 267- 

286 in A. E. Lugo, M. M. Brinson, and S. 

Brown, editors. Ecosystems of the World

126 


15, Forested Wetlands. Elsevier, 

Amsterdam. 

Newman, S., J. B. Grace, and J. W. Koebel. 

1996. Effects of nutrients and hydroperiod 

on Typha, Cladium, and Eleocharis: 

Implications for Everglades restoration. 

Ecological Applications 6:774-783. 

Nickerson, D. J. 1999. Trade-offs of mangrove 

area development in the Philippines. 

Ecological Economics 28:279-298. 

Noble, I. R., and R. O. Slatyer. 1980. The use of 

vital attributes to predict successional 

changes in plant communities subject to 

recurrent disturbances. Vegetatio 43:5-21. 

Noble, I. R., and R. O. Slatyer. 1981. Concepts 

and models of successional in vascular plant 

communities subject to recurrent fire. Pages 

311-335 in A. M. Gill, R. H. Groves, and I. 

R. Noble, editors. Fire and the Australian 

Biota. Australian Academy of Science, 

Canberra. 

Nyman, J. A., and R. H. Chabreck. 1995. Fire 

in coastal marshes: history and recent 

concerns. Pages 134-141 in S. I. Cerulean, 

and R. T. Engstrom, editors. Fire in 

Wetlands: A Management Perspective. 

Proceedings of the Tall Timbers Fire 

Ecology Conference, No. 19. Tall Timbers 

Research Station, Tallahassee, FL. 

Odum, W. E., C. C. McIvor, and T. J. Smith, III. 

1982. The Ecology of the Mangroves of 

South Florida: A Community Profile. U.S. 

Fish and Wildlife Service, Washington, DC. 

Ong, J.-E. 1995. The ecology of mangrove 

conservation & management. 


Pernetta, J. C. 1993. Mangrove Forests, Climate 

Change and Sea Level Rise: Hydrological 

Influences on Community Structure and 

Survival, with Examples from the Indo- 

West Pacific. IUCN, Gland, Switzerland. 

Pons, L. J., and J. L. Fislier. 1991. Sustainable 

development of mangroves. Landscape and 

Urban Planning 20:103-109. 

Pons, T. L., and L. J. Pons. 1975. Mangrove 

vegetation and soils along a more or less 

stationary part of the coast of Surinam, 

South America. Pages 548-560 in G. E. 







Pulle, A. A., editor. 1932-. Flora of Suriname. 

Koninklijke Vereeniging Indisch Instituut, 

Amsterdam. 

Quinn, W. H. 1992. A study of Southern 

Oscillation-related climatic activity for 

A.D. 622-1900 incorporating Nile River 

flood data. Pages 119-149 in H. F. Diaz, 

and V. Markgraf, editors. El NiZo Historical 

and Paleoclimatic Aspects of the Southern 

Oscillation. Cambridge University Press, 

Cambridge. 

Ramdass, I., J. Singh, M. Tamessar, and D. 

Gopaul 1997. National Activity on the use 

of Biodiversity, Guyana. UNDP, The Office 

of the President, and The University of 


Richardson, B. C. 1987. Men, water, and 

mudflats in coastal Guyana. Resource 

Management and Optimization 5:213-236. 

Rittibhonbhun, N., P. Chansanoh, S. Tongrak, 

and R. Suwannatchote. 1993. Communitybased 

mangrove rehabilitation and 

management: a case study in Sikao district, 

Trang Province, Southern Thailand. 

Regional Development Dialogue 14:111- 

122. 

Robadue, D., Jr. 1995. Eight Years in Ecuador: 

The Road to Integrated Coastal 

Management. Coastal Resources Center, 

University of Rhode Island. 

Roberts, S. J. 2000. Tropical fire ecology. 

Progress in Physical Geography 24:281- 

288. 

Rubin, J. A., C. Gordoni, and J. K. Amatekpor. 

1998. Causes and consequences of 

mangrove deforestation in the Volta estuary, 

Ghana: Some recommendations for 

ecosystem rehabilitation. Marine Pollution 


Rützler, K., and C. Feller. 1987. Mangrove 

swamp communities. Oceanus 30:16-24. 

Rützler, K., and I. C. Feller. 1996. Caribbean 

mangrove swamps. Scientific American 

274:74-79. 

Saenger, P., E. J. Hegerl, and J. D. S. Davie. 

1983. Global status of mangrove 

ecosystems. The Environmentalist, IUCN, 

Gland, Switzerland 3:1-88.


Saldarriaga, J. G., and D. C. West. 1986. 

Holocene fires in the Northern Amazon 

Basin. Quaternary Research 26:358-366. 

Schomburgk, M. R. 1922. Travels in British 

Guiana During the Years 1840-1844. 

Translated by Walter Edmund Roth. Daily 

Chronicle, Georgetown, Guyana. 

Sherman, R. E., T. J. Fahey, and J. J. Battles. 

2000. Small-scale disturbance and 

regeneration dynamics in a neotropical 

mangrove forest. Journal of Ecology 

88:165-178. 

Singhroy, V. 1996. Interpretation of SAR images 

for coastal zone mapping in Guyana. 

Canadian Journal of Remote Sensing 

22:317-328. 

Smith, N., S. A. Mori, A. Henderson, D. W. 

Stevenson, and S. V. Heald, editors. 2004. 

Flowering Plants of the Neotropics. 

Princeton University Press, Princeton. 

Smith, S. M., and S. Newman. 2001. Growth of 

southern cattail (Typha domingensis Pers.) 

seedlings in response to fire-related soil 

transformations in the northern Florida 

Everglades. Wetlands 21:363-369. 

Smith, T. J., III, M. B. Robblee, H. R. Wanless, 

and T. W. Doyle. 1994. Mangroves, 

hurricanes, and lightning strikes: 

assessment of Hurricane Andrew suggests 

an interaction across two differing scales. 

Bioscience 44:256-262. 

Snedaker, S. C. 1986. Traditional uses of South 

American mangrove resources and the 

socio-economic effect of ecosystem 

changes. Pages 104-112 in P. Kunstader, E. 

C. F. Bird, and S. Sabhasri, editors. Man in 

the Mangroves: The Socio-economic 

Situation of Human Settlements in 

Mangrove Forests. United Nations 

University Press, Tokyo. 

Spalding, M., F. Blasco, and C. Field 1997. 

World Mangrove Atlas. International 

Society for Mangrove Ecosystems, 

Okinawa, Japan. 

Stevenson, N. J. 1997. Disused shrimp ponds: 

Options for redevelopment of mangrove. 

Coastal Management 25:425-435. 

Steyermark, J. A., P. E. Berry, and B. K. Holst, 

editors. 1995 - 2005. Flora of the 

Venezuelan Guayana. Missouri Botanical 


127 

Thom, B. G. 1975. Mangrove ecology from a 

geomorphic viewpoint. Pages 469-481 in 

G. E. Walsh, S. C. Snedaker, and H. J. Teas, 






Timmins, S. M. 1992. Wetland recovery after 

fire: Eweburn Bog, Te Anau, New Zealand. 

New Zealand Journal of Botany 30:383- 

399. 

Tomlinson, P. B. 1986. The Botany of 

Mangroves. Cambridge University Press. 

Trenberth, K. E. 1999. The extreme weather 

events of 1997 and 1998. Consequences 

5:3-15. 

Twilley, R. R. 1989. Impacts of shrimp 

mariculture practices on the ecology of 

coastal ecosystems in Ecuador. Pages 91- 

120 in S. Olsen, and L. Arriaga, editors. 

Establishing a Sustainable Shrimp 

Mariculture Industry in Ecuador. 276 pp. 

Coastal Resources Center, University of 

Rhode Island, Providence. 

Uhl, C. 1998. Perspectives on wildfire in the 

humid tropics. Conservation Biology 

12:942-943. 

Uhl, C., and R. Buschbacher. 1985. A disturbing 

synergism between cattle ranch burning 

practices and selective tree harvesting in the 

Eastern Amazon. Biotropica 17:265-268. 

Uhl, C., and J. B. Kauffman. 1990. 

Deforestation, fire susceptibility, and 

potential tree responses to fire in the Eastern 

Amazon. Ecology 71:437-449. 

Valiela, I., J. L. Bowen, and J. K. York. 2001. 

Mangrove forests: One of the world’s 

threatened major tropical environments. 

BioScience 51:807-815. 

van Andel, T. R. 2000. Non-Timber Forest 


Guyana, Part II: A Field Guide. Tropenbos 

Guyana Series 8B. Tropenbos-Guyana 


Vann, J. H. 1959. The Physical Geography of 

the Lower Coastal Plain of the Guiana 

Coast. Louisiana State University, Baton 

Rouge. 

Vann, J. H. 1969. Landforms, Vegetation, and 

Sea Level Change along the Guiana Coast

128 


of South America. Office of Naval 

Research, Washington, DC. 

Vogl, R. J. 1975. Fire: A destructive menace or 

a natural process? Recovery and 

Restoration of Damaged Ecosystems. 

Proceedings of the International 

Symposium on the Recovery of Damaged 

Ecosystems, held at Virginia Polytechnic 

Institute and State University, Blacksburg, 

Virginia, on March 23-25, 1975. University 

Press of Virginia, Charlottesville. 

Wade, D., J. Ewell, and R. Hofstetter. 1980. Fire 

in South Florida Ecosystems. Page 125. 

Southeast For. Exp. Stn., U.S. Department 

of Agriculture Forest Service, Asheville, 

NC. 

Weinstock, J. A. 1994. Rhizophora mangrove 

agroforestry. Economic Botany 48:210- 

213. 

West, R. C. 1977. Tidal salt-marsh and mangal 

formations of Middle and South America. 

Pages 193-213 in V. J. Chapman, editor. 

Ecosystems of the World 1, Wet Coastal 

Ecosystems. Elsevier, Amsterdam. 

Whelan, R. J. 1995. The Ecology of Fire. 

Cambridge University Press. 

WMO. 1999. The 1997-1998 El NiZo Event: A 

Scientific and Technical Retrospective. 

Page 96. World Meteorological 

Organization, Geneva. 

Wolanski, E. 1995. Transport of sediment in 

mangrove swamps. Hydrobiologia 295:31- 

42. 

Wolanski, E., S. Spagnol, S. Thomas, K. Moore, 

D. M. Alongi, L. Trott, and A. Davidson. 

2000. Modelling and visualizing the fate of 

shrimp pond effluent in a mangrove-fringed 

tidal creek. Estuarine, Coastal, and Shelf 

Science 50:85-97. 

Woodroffe, C. 1992. Mangrove sediments and 

geomorphology. In A. I. Robertson, and D. 

M. Alongi, editors. Tropical Mangrove 

Ecosystems. American Geophysical Union, 

Washington, DC.


APPENDIX 2. 

PLANT SPECIES LISTED FOR THE WAINI PENINSULA, 

WITH SYNONYMY 

CHLOROPHYTES (Green Algae) 

Cladophoraceae 

Rhizoclonium africanum Kützing 

PTERIDOPHYTES 

Blechnaceae 

Blechnum serrulatum Rich. 

Lygodiaceae 

Lygodium venustum Sw. 

Oleandraceae 

Nephrolepis biserrata (Sw.) Schott 

Nephrolepis rivularis (Vahl) Mett. ex Krug 

Parkeriaceae 

Ceratopteris thalictroides (L.) Brongn. 

Polypodiaceae 

Campyloneurum phyllitidis (L.) C. Presl 

Pteridaceae 

Acrostichum aureum L. 

Acrostichum danaeifolium Langsd. & Fisch. 

Pteris pungens Willd. 

DICOTILEDONEAE 

Aizoaceae 

Sesuvium portulacastrum (L.) L. 

Amaranthaceae 

Alternanthera sessilis (L.) R. Br. ex DC. 

Amaranthus australis (A. Gray) J.D. Sauer 

Amaranthus dubius Mart. ex Thell. 

Blutaparon vermiculare (L.) Mears 

Anacardiaceae 

Spondias mombin L. 

Annonaceae 

Annona glabra L. 

Apocynaceae 

Allamanda cathartica L. 

Malouetia tamaquarina (Aubl.) A. DC. 

Rhabdadenia biflora (Jacq.) Müll. Arg. 

Aristolochiaceae 

Aristolochia trilobata L. 

Asclepiadaceae 

Sarcostemma clausum (Jacq.) Schult. 

Asteraceae 

Bidens alba (L.) DC. 

Cyanthillium cinereum (L.) H. Rob. 

Eclipta prostrata (L.) L. 

Mikania micrantha Kunth 

Pluchea odorata (L.) Cass. 

Avicenniaceae 

Avicennia germinans (L.) Stearn 

Bignoniaceae 

Cydista aequinoctialis (L.) Miers 

Cactaceae 

Epiphyllum phyllanthus (L.) Haw. 

Rhipsalis baccifera (J.S. Muell.) Stearn 

Caricaceae 

Carica papaya L. 

Cecropiaceae 

Coussapoa asperifolia Trécul 

Ceratophyllaceae 

Ceratophyllum muricatum Cham. 

Clusiaceae 

Clusia palmicida Rich. ex Planch. & Triana 

Combretaceae 

Conocarpus erectus L. 

Laguncularia racemosa (L.) C.F. Gaertn. 

Terminalia catappa L. 


Ipomoea pes-caprae (L.) R. Br. 

Ipomoea tiliacea (Willd.) Choisy 

129

130 

Ipomoea violacea L. 

Merremia cissoides (Lam.) Hallier f. 

Merremia umbellata (L.) Hallier f. 

Cucurbitaceae 

Melothria pendula L. 

Cuscutaceae 

Cuscuta umbellata Kunth 

Euphorbiaceae 

Euphorbia hirta L. 

Manihot esculenta Crantz (cultivated) 

Fabaceae-Caesal. 

Caesalpinia bonduc (L.) Roxb. 

Fabaceae-Mimos. 

Entada polystachya (L.) DC. 

Inga ingoides (Rich.) Willd. 

Zygia latifolia (L.) Fawc. & Rendle 

Fabaceae-Papil. 

Aeschynomene sensitiva Sw. 

Canavalia rosea (Sw.) DC. 

Machaerium lunatum (L. f.) Ducke 

Pterocarpus officinalis Jacq. 

Sesbania sericea (Willd.) DC. 

Vigna luteola (Jacq.) Benth. 

Hippocrateaceae 

Hippocratea volubilis L. 

Lauraceae 

Cassytha filiformis L. 

Malpighiaceae 

Stigmaphyllon bannisterioides (L.) C.E. 

Anderson 

Malvaceae 

Hibiscus bifurcatus Cav. 

Talipariti tiliaceum (L.) Fryxell 

Thespesia populnea (L.) Sol. ex CorrLa 

Moraceae 

Ficus amazonica (Miq.) Miq. 

Ficus eximia Schott 

Ficus maxima Mill. 

Myrtaceae 

Calyptranthes sp. 


Psidium guajava L. 

Nyctaginaceae 

Boerhavia diffusa L. 

Onagraceae 

Ludwigia affinis (DC.) H. Hara 

Ludwigia leptocarpa (Nutt.) H. Hara 

Passifloraceae 

Passiflora foetida L. 

Piperaceae 

Peperomia glabella (Sw.) A. Dietr. 

Polygalaceae 

Securidaca diversifolia (L.) S.F. Blake 


Cassipourea guianensis Aubl. 

Rhizophora mangle L. 

Rhizophora racemosa G. Mey. 

Rubiaceae 

Morinda citrifolia L. 

Sapindaceae 

Paullinia pinnata L. 

Scrophulariaceae 

Capraria biflora L. 

Solanaceae 

Physalis angulata L. 

Solanum stramoniifolium Jacq. 

Vitaceae 

Cissus verticillata (L.) Nicolson & C.E. Jarvis 

MONOCOTILEDONEAE 

Araceae 

Anthurium gracile (Rudge) Schott 

Monstera adansonii Schott 

Montrichardia linifera (Arruda) Schott 

Philodendron acutatum Schott 

Syngonium podophyllum Schott 

Arecaceae 

Cocos nucifera L. (cultivated) 

Desmoncus orthacanthos Mart.

Euterpe oleracea Mart. 

Nypa fruticans Wurmb. 

Roystonea oleracea (Jacq.) O.F. Cook 


Bromeliaceae 

Aechmea nudicaulis (L.) Griseb. 

Bromelia plumieri (E. Morren) L.B. Sm. 

Guzmania lingulata (L.) Mez 

Costaceae 

Costus arabicus L. 

Cyperaceae 

Cyperus ligularis L. 

Cyperus odoratus L. 

Cyperus polystachyos Rottb. 

Eleocharis mutata (L.) Roem. & Schult. 

Fimbristylis cymosa R. Br. 

Dioscoreaceae 

Dioscorea polygonoides Humb. & Bonpl. ex 

Willd. 

Heliconiaceae 

Heliconia psittacorum L.f. 

Hydrocharitaceae 

Limnobium laevigatum (Humb. & Bonpl. ex 

Willd.) Heine 

Lemnaceae 

Lemna aequinoctialis Welw. 

Liliaceae 

Crinum erubescens L. f. ex Sol. 

Orchidaceae 

Dimerandra elegans ? (H. Focke) Siegerist 

Epidendrum ciliare L. 

Prosthechea aemula (Lindl.) W.E. Higgins 

Trichocentrum lanceanum (Lindl.) M.W. 

Chase & N.H. Williams 

Poaceae 

Echinochloa polystachya (Kunth) Hitchc. 

Leptochloa scabra Nees 

Paspalum distichum L. 

Sporobolus virginicus (L.) Kunth 

Smilacaceae 

Smilax cumanensis Humb. & Bonpl. ex Willd. 

Typhaceae 

Typha domingensis Pers. 

Zingiberaceae 

Renealmia alpinia (Rottb.) Maas 

SYNONYMY 

131 

Achyranthes sessilis (L.) Desf. ex Steud. = 

Alternanthera sessilis 

Acnida australis A. Gray = Amaranthus 

australis 

Acnida cuspidata Bertero ex Spreng. = 

Amaranthus australis 

Acrostichum guineense Gaudich. = Acrostichum 

aureum 

Acrostichum lomarioides (Jenman) Jenman 

[nom. illeg.] = Acrostichum danaeifolium 

Acrostichum thalictroides L. = Ceratopteris 

thalictroides 

Agrostis littoralis Lam. = Sporobolus virginicus 

Agrostis virginica L. = Sporobolus virginicus 

Alpinia tubulata Ker Gawl. = Renealmia alpinia 

Annona palustris L. = Annona glabra 

Anthurium scolopendrinum (Ham.) Kunth = 

Anthurium gracile 

Arum liniferum Arruda = Montrichardia linifera 

Asclepias clausa Jacq. = Sarcostemma clausum 

Aspidium biserratum Sw. = Nephrolepis 

biserrata 

Aulizia ciliaris (L.) Salisb. = Epidendrum ciliare 

Avicennia nitida Jacq. = Avicennia germinans 

Avicennia tomentosa Jacq. = Avicennia 

germinans 

Banisteria ovata Cav. = Stigmaphyllon 

bannisterioides 

Bidens pilosa forma radiata Sch. Bip. = Bidens 

alba 

Bignonia aequinoctialis L. = Cydista 

aequinoctialis 

Bignonia picta Kunth = Cydista aequinoctialis 

Bihai silvestris Gleason = Heliconia psittacorum 

Blechnum indicum auct. non Burm. f. = 

Blechnum serrulatum 

Boerhavia paniculata Rich. = Boerhavia diffusa 

Boerhavia surinamensis Miq. = Boerhavia 

diffusa 

Brachypterys borealis A. Juss. = Stigmaphyllon 


Bromelia karatas L. = Bromelia plumieri

132 


Bryonia guadalupensis Spreng. = Melothria 

pendula 

Cacalia cinerea (L.) Kuntze = Cyanthillium 

cinereum 

Cactus caripensis Kunth = Rhipsalis baccifera 

Cactus phyllanthus L. = Epiphyllum phyllanthus 

Calliandra latifolia (L.) Griseb. = Zygia latifolia 

Calonyction tuba (Schltdl.) Colla = Ipomoea 

violacea 

Cameraria tamaquarina Aubl. = Malouetia 

tamaquarina 

Campyloneurum costatum (Kunze) C. Presl = 

Campyloneurum phyllitidis 

Canavalia maritima Thouars = Canavalia rosea 

Canavalia obtusifolia DC. = Canavalia rosea 

Capraria hirsuta Kunth = Capraria biflora 

Caraguata lingulata (L.) Lindl. = Guzmania 

lingulata 

Caraguata splendens Planch. = Guzmania 

lingulata 

Caraxeron vermicularis (L.) Raf. = Blutaparon 

vermiculare 

Carica sativa Tussac = Carica papaya 

Cassia paramariboensis Miq. = Aeschynomene 

sensitiva 

Cassipourea belizensis Lundell = Cassipourea 

guianensis 

Cassipourea elliptica (Sw.) Poir. = Cassipourea 

guianensis 

Cassipourea macrodonta Standl. = Cassipourea 

guianensis 

Cassipourea podantha Standl. = Cassipourea 

guianensis 

Cassytha americana Nees = Cassytha filiformis 

Cassytha baccifera Sol. ex J.S. Muell. = 

Rhipsalis baccifera 

Ceratophyllum demersum L. var. cristatum 

(Spruce) K. Schum. = Ceratophyllum 

muricatum 

Ceratophyllum lleranae Fassett = 

Ceratophyllum muricatum 

Cereus caripensis (Kunth) DC. = Rhipsalis 

baccifera 

Chamaesyce hirta (L.) Millsp. = Euphorbia 

hirta 

Chrysodium aureum (L.) Mett. = Acrostichum 

aureum 

Chrysodium lomarioides Jenman = Acrostichum 

danaeifolium 

Cissus cordifolia L. = Cissus verticillata 

Cissus sicyoides L. = Cissus verticillata 

Conocarpus racemosa L. = Laguncularia 

racemosa 

Convolvulus caracassanus Roem. & Schult. = 

Merremia umbellata 

Convolvulus cissoides Lam. = Merremia 

cissoides 

Convolvulus pes-caprae L. = Ipomoea pescaprae 

Convolvulus riparius Kunth = Merremia 

cissoides 

Convolvulus tiliaceus Willd. = Ipomoea tiliacea 

Convolvulus umbellatus L. = Merremia 

umbellata 

Conyza cinerea L. = Cyanthillium cinereum 

Conyza cortesii Kunth = Pluchea odorata 

Conyza odorata L. = Pluchea odorata 

Coreopsis alba L. = Bidens alba 

Coronilla sericea Willd. = Sesbania sericea 

Costus discolor Roscoe = Costus arabicus 

Costus niveus G. Mey. = Costus arabicus 

Costus ramosus Woodson = Costus arabicus 

Crinum commelyni Jacq. = Crinum erubescens 

Crinum guianense M. Roem. = Crinum 

erubescens 

Crinum lancei Herbert ex Sweet = Crinum 

erubescens 

Crinum lindleyanum Schult. f. ex Seub. = 

Crinum erubescens 

Cynanchum clausum (Jacq.) Jacq. = 

Sarcostemma clausum 

Cyperus eggersii Boeck. = Cyperus odoratus 

Cyperus macrocephalus Liebm. = Cyperus 

odoratus 

Desmoncus apureanus L.H. Bailey = 

Desmoncus orthacanthos 

Desmoncus horridus Splitg. ex Mart. = 

Desmoncus orthacanthos 

Desmoncus multijugus Steyerm. = Desmoncus 

orthacanthos 

Desmoncus palustris Trail = Desmoncus 

orthacanthos 

Desmoncus velezii L.H. Bailey & H.E. Moore 

= Desmoncus orthacanthos 

Digitaria disticha (L.) Fiori & Paol. = Paspalum 

distichum 

Digitaria paspalodes Michx. = Paspalum 

distichum 

Dimerandra isthmii Schltr. = Dimerandra 

elegans ? 

Dioscorea kegeliana Griseb. = Dioscorea 

polygonoides


Diplachne scabra (Nees) Nicora = Leptochloa 

scabra 

Dolichos luteolus Jacq. = Vigna luteola 

Dolichos maritimus Aubl. = Canavalia rosea 

Dolichos roseus Sw. = Canavalia rosea 

Drepanocarpus lunatus (L. f.) G. Mey. = 

Machaerium lunatum 

Echinochloa polystachya var. spectabilis (Nees 

ex Trin.) Mart. Crov. = Echinochloa 

polystachya 

Echinochloa spectabilis (Nees ex Trin.) Link = 

Echinochloa polystachya 

Echites biflora Jacq. = Rhabdadenia biflora 

Eclipta alba (L.) Hassk. = Eclipta prostrata 

Elsota diversifolia (L.) S.F. Blake = Securidaca 

diversifolia 

Encyclia aemula (Lindl.) Carnevali & I. 

Ramírez = Prosthechea aemula 

Encyclia ciliaris (L.) Lemée = Epidendrum 

ciliare 

Encyclia fragrans (Sw.) Lemée var. aemula 

(Lindl.) Dressler & G.E. Pollard = 

Prosthechea aemula 

Entadopsis polystachya (L.) Britton = Entada 

polystachya 

Epidendrum aemulum Lindl. = Prosthechea 

aemula 

Epidendrum fragans auct. non Sw. 1788 = 

Prosthechea aemula 

Epidendrum fragans Sw. var. aemulum (Lindl.) 

Barb. Rodr. = Prosthechea aemula 

Epiphyllum hookeri Haw. = Epiphyllum 

phyllanthus 

Epiphyllum phyllanthus (L.) Haw. var. hookeri 

(Haw.) Kimnach = Epiphyllum phyllanthus 

Eupatorium denticulatum Vahl = Mikania 

micrantha 

Euterpe badiocarpa Barb. Rodr. = Euterpe 

oleracea 

Euterpe beardii Bailey = Euterpe oleracea 

Euterpe edulis auct. = Euterpe oleracea 

Feuilleea ingoides (Rich.) Kuntze = Inga 

ingoides 

Ficus angustifolia (Miq.) Miq. = Ficus 

amazonica 

Ficus expansa Pittier = Ficus eximia 

Ficus foveata Pittier = Ficus eximia 

Ficus foveolata Pittier = Ficus eximia 

Ficus glandulosa Pittier = Ficus eximia 

Ficus glaucescens (Liebm.) Miq. = Ficus 

maxima 

133 

Ficus guanarensis Pittier = Ficus eximia 

Ficus parkeri Miq. = Ficus maxima 

Ficus radula Humb. & Bonpl. ex Willd. = Ficus 

maxima 

Ficus surinamensis Miq. = Ficus amazonica 

Ficus turbinata Pittier = Ficus eximia 

Fimbristylis cymosa R. Br. subsp. spathacea 

(Roth) T. Koyama = Fimbristylis cymosa 

Fimbristylis glomerata (Retz.) Urb. = 

Fimbristylis cymosa 

Fimbristylis obtusifolia (Vahl) Kunth = 

Fimbristylis cymosa 

Fimbristylis spathacea Roth = Fimbristylis 

cymosa 

Funastrum clausum (Jacq.) Schltr. = 

Sarcostemma clausum 

Gomphrena sessilis L. = Alternanthera sessilis 

Gomphrena vermicularis L. = Blutaparon 

vermiculare 

Guilandina bonduc L. = Caesalpinia bonduc 

Guzmania lingulata (L.) Mez var. minor (Mez) 

L.B. Sm. & Pittendr. = Guzmania lingulata 

Guzmania minor Mez = Guzmania lingulata 

Heliconia ballia Rich. = Heliconia psittacorum 

Heliconia cannoidea Rich. = Heliconia 

psittacorum 

Heliconia humilis (Aubl.) Jacq. = Heliconia 

psittacorum 

Heliconia schomburgkiana Klotzsch = 

Heliconia psittacorum 

Heliconia silvestris (Gleason) L.B. Sm. = 

Heliconia psittacorum 

Hibiscus elatus Sw. = Talipariti tiliaceum 

Hibiscus pernambucensis = Talipariti tiliaceum 

Hibiscus populneus L. = Thespesia populnea 

Hibiscus tiliaceus L. = Talipariti tiliaceum 

Hippocratea laevigata Rich. = Hippocratea 

volubilis 

Hydromystria laevigata (Humb. & Bonpl. ex 

Willd.) Díaz-Mir. & Philcox = Limnobium 

laevigatum 

Hydromystria laevigata (Willd.) Hanzeker = 

Limnobium laevigatum 

Ilex acuminata Willd. = Ilex guianensis 

Ilex celastroides Klotzsch ex Garcke = Ilex 

guianensis 

Ilex cumanensis Turcz. = Ilex guianensis 

Ilex macoucoua Pers. = Ilex guianensis 

Inga latifolia (L.) Willd. = Zygia latifolia 

Ipomoea cissoides (Lam.) Griseb. fma. viscidula 

Meisn. = Merremia cissoides

134 


Ipomoea macrantha Roem. & Schult. = Ipomoea 

violacea 

Ipomoea mollicoma Miq. = Merremia umbellata 

Ipomoea polyanthes Roem. & Schult. = 

Merremia umbellata 

Ipomoea tuba (Schltdl.) G. Don = Ipomoea 

violacea 

Iresine vermicularis (L.) Moq. = Blutaparon 

vermiculare 

Isochilus elegans H. Focke = Dimerandra 

elegans ? 

Jatropha manihot L. = Manihot esculenta 

Jussiaea affinis DC. = Ludwigia affinis 

Jussiaea aluligera Miq. = Ludwigia leptocarpa 

Jussiaea hexamera Miq. = Ludwigia affinis 

Jussiaea leptocarpa Nutt. = Ludwigia 

leptocarpa 

Jussiaea leptocarpa Nutt. var. aluligera (Miq.) 

Jonker = Ludwigia leptocarpa 

Jussiaea leptocarpa Nutt. var. genuina Munz = 

Ludwigia leptocarpa 

Jussiaea leptocarpa Nutt. var. meyeriana 

(Kuntze) Munz = Ludwigia leptocarpa 

Jussiaea surinamensis Miq. = Ludwigia 

leptocarpa 

Jussiaea variabilis G. Mey. var. affinis (DC.) 

Kuntze = Ludwigia affinis 

Jussiaea variabilis G. Mey. var. meyeriana 

Kuntze = Ludwigia leptocarpa 

Karatas plumieri E. Morren = Bromelia plumieri 

Lemna paucicostata Hegelm. = Lemna 

aequinoctialis 

Leptochloa langloisii Vasey = Leptochloa 

scabra 

Leptochloa liebmannii E. Fourn = Leptochloa 

scabra 

Limnobium spongia (Bosc) Steud. subsp. 

laevigatum (Humb. & Bonpl. ex Willd.) 

Lowden = Limnobium laevigatum 

Limnobium stoloniferum (G. Mey.) Griseb. = 


Lophiaris fragrans Raf. = Trichocentrum 

lanceanum 

Lophiaris lanceana (Lindl.) Braem = 

Trichocentrum lanceanum 

Lygodium mexicanum C. Presl = Lygodium 

venustum 

Lygodium polymorphum auct. non (Cav.) Kunth 

= Lygodium venustum 

Macoucoua guianensis Aubl. = Ilex guianensis 

Malouetia furfuracea Spruce ex Müll. Arg. = 

Malouetia tamaquarina 

Malouetia guianensis (Aubl.) Miers = Malouetia 


Malouetia obtusiloba A. DC. = Malouetia 


Malouetia odorata DC. = Malouetia 


Malpighia bannisterioides L. = Stigmaphyllon 


Manihot diffusa Pohl = Manihot esculenta 

Manihot dulcis Pax = Manihot esculenta 

Manihot utilissima Pohl = Manihot esculenta 

Mariscus ligularis (L.) Urb. = Cyperus ligularis 

Melothria fluminensis Gardner = Melothria 

pendula 

Melothria guadalupensis (Spreng.) Cogn. = 

Melothria pendula 

Mikania denticulata (Vahl) Willd. = Mikania 

micrantha 

Mikania orinocensis Kunth = Mikania 

micrantha 

Milium distichum (L.) Muhl. = Paspalum 

distichum 

Mimosa bipinnata Aubl. = Entada polystachya 

Mimosa ingoides Rich. = Inga ingoides 

Mimosa latifolia L. = Zygia latifolia 

Mimosa polystachia L. = Entada polystachya 

Moutouchi suberosa Aubl. = Pterocarpus 

officinalis 

Musa humilis Aubl. = Heliconia psittacorum 

Oncidium lanceanum Lindl. = Trichocentrum 

lanceanum 

Oplismenus polystachyus Kunth = Echinochloa 

polystachya 

Orelia grandiflora Aubl. = Allamanda 

cathartica 

Panicum bonplandianum Steud. = Echinochloa 

polystachya 

Panicum spectabile Nees ex Trin. = Echinochloa 

polystachya 

Paullinia hostmannii Steud. = Paullinia pinnata 

Pharmacosycea guyanensis Miq. = Ficus 

maxima 

Phaseolus luteolus (Jacq.) Gagnep. = Vigna 

luteola 

Philodendron cyclops A.D. Hawkes = 

Philodendron acutatum 

Philoxerus vermicularis (L.) R. Br. = Blutaparon 

vermiculare 

Phyllocactus phyllanthus (DC.) Link = 

Epiphyllum phyllanthus


Physalis capsicifolia Dunal = Physalis angulata 

Physalis lanceifolia Nees = Physalis angulata 

Piper glabellum Sw. = Peperomia glabella 

Pithecellobium latifolium (L.) Benth. = Zygia 

latifolia 

Pluchea cortesii (Kunth) DC. = Pluchea 

odorata 

Polygala diversifolia L. = Securidaca 

diversifolia 

Polypodium phyllitidis L. = Campyloneurum 

phyllitidis 

Polypodium rivulare Vahl = Nephrolepis 

rivularis 

Portulaca portulacastrum L. = Sesuvium 

portulacastrum 

Pothos gracilis Rudge = Anthurium gracile 

Pothos scolopendrinus Ham. = Anthurium 

gracile 

Pteris biaurita var. pungens (Willd.) H. Christ 

= Pteris pungens 

Pteris longicauda H. Christ = Pteris pungens 

Pterocarpus draco L. = Pterocarpus officinalis 

Pterocarpus lunatus L. f. = Machaerium 

lunatum 

Pterocarpus suberosa (Aubl.) Pers. = 

Pterocarpus officinalis 

Pycreus polystachyos (Rottb.) P. Beauv. = 

Cyperus polystachyos 

Renealmia exaltata L. f. = Renealmia alpinia 

Rhipsalis cassutha Gaertn. = Rhipsalis 

baccifera 

Rhipsalis minutiflora K. Schum. = Rhipsalis 

baccifera 

Rhizophora americana Nutt. = Rhizophora 

mangle 

Rhizophora mangle var. racemosa (G. Mey.) 

Engl. = Rhizophora racemosa 

Rhizophora mangle var. samoensis Hochr. = 

Rhizophora mangle 

Rhizophora samoensis (Hochr.) Salvoza = 

Rhizophora mangle 

Roystonea venezuelana L.H. Bailey = 

Roystonea oleracea 

Salvinia laevigata Humb. & Bonpl. ex Willd. = 


Sarcostemma cumanense Kunth = Sarcostemma 

clausum 

Sarcostemma pubescens Kunth = Sarcostemma 

clausum 

135 

Scirpus glomeratus Retz. = Fimbristylis cymosa 

Scirpus mutatus L. = Eleocharis mutata 

Scirpus obtusifolius Vahl = Fimbristylis cymosa 

Sesuvium acutifolium Miq. = Sesuvium 

portulacastrum 

Smilax globifera G. Mey. = Smilax cumanensis 

Smilax hostmanniana Kunth = Smilax 

cumanensis 

Smilax pirarensis Kunth & M.R. Schomb. = 

Smilax cumanensis 

Smilax surinamensis Miq. = Smilax cumanensis 

Solanum demerarense Dunal = Solanum 

stramoniifolium 

Solanum toxicarium Rich. = Solanum 


Solanum trichocarpum Miq. = Solanum 


Spondias lutea L. = Spondias mombin 

Sporobolus littoralis (Lam.) Kunth = 

Sporobolus virginicus 

Stigmaphyllon ovatum (Cav.) Nied. = 

Stigmaphyllon bannisterioides 

Tillandsia lingulata L. = Guzmania lingulata 

Torulinium ferax (Rich.) Urb. = Cyperus 

odoratus 

Torulinium odoratum (L.) S.S. Hooper = 

Cyperus odoratus 

Typha angustifolia var. domingensis (Pers.) 

Hemsl. = Typha domingensis 

Typha tenuifolia Kunth = Typha domingensis 

Typha truxillensis Kunth = Typha domingensis 

Urostigma amazonicum Miq. = Ficus 

amazonica 

Urostigma angustifolium Miq. = Ficus 

amazonica 

Verbesina alba L. = Eclipta prostrata 

Verbesina prostrata L. = Eclipta prostrata 

Vernonia cinerea (L.) Less. = Cyanthillium 

cinereum 

Vigna repens (L.) Kuntze = Vigna luteola 

Vilfa virginica (L.) P. Beauv. = Sporobolus 

virginicus 

Vitis sicyoides (L.) Baker = Cissus verticillata 

Wedelia psammophila Poepp. = Eclipta 

prostrata 

Willoughbya micrantha (Kunth) Rusby = 

Mikania micrantha 

Willoughbya scandens Kuntze var. orinocensis 

(Kunth) Kuntze = Mikania micrantha

136 


APPENDIX 3. 

PRELIMINARY SPECIES LIST OF THE VASCULAR PLANTS 

OF THE NORTHWEST DISTRICT OF GUYANA 

(* denotes taxon added by collections for this volume.) 

LYCOPHYTES 

Lycopodiaceae 

Lycopodiella cernua (L.) Pic. Serm. 

Selaginellaceae 

Selaginella epirrhizos Spring 

Selaginella parkeri (Hook. & Grev.) Spring 

Selaginella porelloides (Lam.) Spring 

Selaginella producta Baker 

PTERIDOPHYTES 

Adiantaceae 

Adiantum latifolium Lam. 

Adiantum phyllitidis J. Sm. 

Pityrogramma calomelanos (L.) Link 

Aspleniaceae 

Asplenium cristatum Lam. * 

Asplenium juglandifolium Lam. 

Asplenium salicifolium L. 

Asplenium serratum L. 

Blechnaceae 

Blechnum serrulatum Rich. 

Cyatheaceae 

Cnemidaria spectabilis (Kunze) R.M. Tryon 

Cyathea cyatheoides (Desv.) K.U. Kramer 

Cyathea microdonta (Desv.) Domin 

Cyathea pungens (Willd.) Domin 

Cyathea surinamensis (Miq.) Domin 

Dennstaedtiaceae 

Lindsaea lancea (L.) Bedd. 

Lindsaea portoricensis Desv. * 

Saccoloma elegans Kaulf. 

Dryopteridaceae 

Cyclodium meniscioides (Willd.) C. Presl 

Didymochlaena truncatula (Sw.) J. Sm. 

Polybotrya caudata Kunze 

Gleicheniaceae 

Dicranopteris pectinata (Willd.) Underw. 

[moss] 

Grammitidaceae 

Cochlidium serrulatum (Sw.) L.E. Bishop 

Hymenophyllaceae 

Trichomanes diversifrons (Bory) Mett. ex 

Sadeb. 

Trichomanes martiusii C. Presl 

Trichomanes polypodioides L. 

Trichomanes radicans Sw. 

Lomariopsidaceae 

Elaphoglossum flaccidum (Fée) T. Moore 

Elaphoglossum glabellum J. Sm. 

Lomariopsis japurensis (Mart.) J. Sm. 

Lomariopsis latiuscula (Maxon) Holttum 

Lygodiaceae 

Lygodium venustum Sw. * 

Lygodium volubile Sw. 

Metaxyaceae 

Metaxya rostrata (Kunth) C. Presl 

Oleandraceae 

Nephrolepis biserrata (Sw.) Schott 

Nephrolepis rivularis (Vahl) Mett. ex Krug 

Parkeriaceae 

Ceratopteris thalictroides (L.) Brongn. * 

Polypodiaceae 

Campyloneurum phyllitidis (L.) C. Presl 

Campyloneurum repens (Aubl.) C. Presl 

Dicranoglossum desvauxii (Klotzsch) Proctor


Microgramma fuscopunctata (Hook.) Vareschi 

Microgramma lycopodioides (L.) Copel. 

Microgramma persicariifolia (Schrad.) C. Presl 

Microgramma reptans (Cav.) A.R. Sm. 

Phlebodium aureum (L.) J. Sm. * 

Phlebodium pseudoaureum (Cav.) Lellinger 

Polypodium adnatum Kunze ex Klotzsch 

Polypodium attenuatum Humb. & Bonpl. ex 

Willd. 

Pteridaceae 

Acrostichum aureum L. * 

Acrostichum danaeifolium Langsd. & Fisch. * 

Pteris altissima Poir. 

Pteris pungens Willd. * 

Schizaeaceae 

Schizaea fluminensis Miers ex J.W. Sturm 

Schizaea incurvata Schkuhr 

Tectariaceae 

Tectaria incisa Cav. 

Tectaria trifoliata (L.) Cav. 

Triplophyllum funestum (Kunze) Holttum 

Thelypteridaceae 

Thelypteris abrupta (Desv.) Proctor 

Thelypteris interrupta (Willd.) K. Iwats. 

Thelypteris leprieurii (Hook.) R.M. Tryon 

Thelypteris opulenta (Kaulf.) Fosberg 

[naturalized] 

Thelypteris serrata (Cav.) Alston 

Vittariaceae 

Anetium citrifolium (L.) Splitg. 

Antrophyum cajenense (Desv.) Spreng. 

Woodsiaceae 

Diplazium celtidifolium Kunze 

GYMNOSPERMS 

Gnetaceae 

Gnetum nodiflorum Brongn. 

DICOTILEDONEAE 

Acanthaceae 

Aphelandra pulcherrima (Jacq.) Kunth 

Asystasia gangetica (L.) T. Anderson 

[cultivated] 

Blechum pyramidatum (Lam.) Urb. * 

Justicia calycina (Nees) V.A.W. Graham 

137 

Justicia pectoralis Jacq. 

Justicia secunda Vahl 

Ruellia tuberosa L. [naturalized] 

Thunbergia alata Bojer ex Sims [cultivated] 

Thunbergia grandiflora (Roxb. ex Rottl.) Roxb. 

[cultivated] 

Trichanthera gigantea (Bonpl.) Nees 

Aizoaceae 

Sesuvium portulacastrum (L.) L. * 

Amaranthaceae 

Alternanthera philoxeroides (Mart.) Griseb. * 

Alternanthera sessilis (L.) R. Br. ex DC. 

Alternanthera tenella Colla 

Amaranthus australis (A. Gray) J.D. Sauer * 

Amaranthus blitum L. * 

Amaranthus dubius Mart. ex Thell. 

Amaranthus viridis L. 

Blutaparon vermiculare (L.) Mears * 

Celosia argentea L. [cultivated] 

Cyathula achyranthoides (Kunth) Moq. 

Cyathula prostrata (L.) Blume 

Gomphrena globosa L. [cultivated] 

Iresine diffusa Humb. & Bonpl. ex Willd. 

Anacardiaceae 

Anacardium giganteum W. Hancock ex Engl. 

Anacardium occidentale L. 

Astronium lecointei Ducke 

Mangifera indica L. [cultivated] 

Spondias dulcis Parkinson [cultivated] 

Spondias mombin L. 

Tapirira guianensis Aubl. 

Tapirira obtusa (Benth.) J.D. Mitch. 

Thyrsodium guianense Sagot ex Marchand 

Annonaceae 

Anaxagorea dolichocarpa Sprague & Sandwith 

Annona glabra L. * 

Annona montana Macfad. 

Annona muricata L. [cultivated] 

Annona sericea Dunal 

Annona symphyocarpa Sandwith 

Bocageopsis multiflora (Mart.) R.E. Fr. 

Duguetia calycina Benoist 

Duguetia megalophylla R.E. Fr. 

Duguetia pauciflora Rusby 

Duguetia pycnastera Sandwith 

Duguetia yeshidan Sandwith 

Fusaea longifolia (Aubl.) Saff.

138 

Guatteria flexilis R.E. Fr. 

Guatteria schomburgkiana Mart. 

Rollinia exsucca (DC. ex Dunal) A. DC. 

Rollinia mucosa (Jacq.) Baill. 

Unonopsis glaucopetala R.E. Fr. 

Xylopia benthamii R.E. Fr. 

Xylopia cayennensis Maas 

Xylopia surinamensis R.E. Fr. 

Apiaceae 

Coriandrum sativum L. [cultivated] 

Eryngium foetidum L. 


Apocynaceae 

Allamanda cathartica L. 

Ambelania acida Aubl. 

Aspidosperma cruentum Woodson 

Aspidosperma excelsum Benth. 

Aspidosperma marcgravianum Woodson 

Catharanthus roseus (L.) G. Don [cultivated] 

Condylocarpon intermedium Müll. Arg. 

Forsteronia gracilis (Benth.) Müll. Arg. 

Forsteronia guyanensis Müll. Arg. 

Himatanthus articulatus (Vahl) Woodson 

Himatanthus bracteatus (A. DC.) Woodson 

Macoubea guianensis Aubl. 

Malouetia flavescens (Willd. ex Roem. & 

Schult.) Müll. Arg. 

Malouetia tamaquarina (Aubl.) A. DC. * 

Mandevilla hirsuta (Rich.) K. Schum. 

Mandevilla symphitocarpa (G. Mey.) Woodson 

Mesechites trifida (Jacq.) Müll. Arg. 

Odontadenia geminata (Hoffmanns. ex Roem. 

& Schult.) Müll. Arg. 

Odontadenia macrantha (Roem. & Schult.) 

Markgr. 

Odontadenia puncticulosa (Rich.) Pulle 

Odontadenia sandwithiana Woodson 

Plumeria inodora Jacq. 

Prestonia tomentosa R. Br. 

Rhabdadenia biflora (Jacq.) Müll. Arg. 

Tabernaemontana disticha A. DC. 

Tabernaemontana divaricata (L.) R. Br. ex 

Roem. & Schult. 

Tabernaemontana heterophylla Vahl 

Tabernaemontana lorifera (Miers) Leeuwenb. 

Tabernaemontana undulata Vahl 

Aquifoliaceae 

Ilex guianensis (Aubl.) Kuntze 

Ilex martiniana D. Don 

Araliaceae 

Oreopanax capitatus (Jacq.) Decne. & Planch. 

Schefflera decaphylla (Seem.) Harms 

Schefflera morototoni (Aubl.) Maguire, 

Steyerm. & Frodin 


Aristolochia daemoninoxia Mast. 

Aristolochia hians Willd. 

Aristolochia rugosa Lam. 

Aristolochia trilobata L. * 


Asclepias curassavica L. 

Blepharodon nitidus (Vell.) J.F. Macbr. 

Matelea badilloi Morillo 

Matelea stenopetala Sandwith 

Sarcostemma clausum (Jacq.) Schult. 

Stenomeria decalepis Turcz. 

Tassadia propinqua Decne. 

Asteraceae 

Ageratum conyzoides L. 

Bidens alba (L.) DC. * 

Bidens cynapiifolia Kunth 

Bidens pilosa L. 

Chromolaena odorata (L.) R.M. King & H. Rob. 

Clibadium surinamense L. 

Clibadium sylvestre (Aubl.) Baill. 

Conyza bonariensis (L.) Cronquist 

Cosmos caudatus Kunth [cultivated] 

Cosmos sulphureus Cav. [cultivated] 

Cyanthillium cinereum (L.) H. Rob. 

[naturalized] 

Cyrtocymura scorpioides (Lam.) H. Rob. 

Eclipta prostrata (L.) L. * 

Elephantopus mollis Kunth 

Emilia sonchifolia (L.) DC. ex Wight 

Erechtites hieracifolia (L.) Raf. ex DC. 

Hebeclinium macrophyllum (L.) DC. 

Mikania banisteriae DC. 

Mikania congesta DC. 

Mikania cordifolia (L. f.) Willd. 

Mikania guaco Bonpl. 

Mikania hookeriana DC. 

Mikania micrantha Kunth 

Mikania microptera DC. 

Mikania parviflora (Aubl.) H. Karst. 

Mikania psilostachya DC. 

Mikania trinitaria DC. 

Pluchea odorata (L.) Cass. *


Rolandra fruticosa (L.) Kuntze 

Sonchus asper (L.) Hill * 

Sphagneticola trilobata (L.) Pruski 

Struchium sparganophorum (L.) Kuntze 

Synedrella nodiflora (L.) Gaertn. 

Tagetes erecta L. [cultivated] 

Unxia camphorata L. f. 

Wulffia baccata (L.) Kuntze 

Zinnia elegans Jacq. [cultivated] 

Avicenniaceae 

Avicennia germinans (L.) Stearn 

Balanophoraceae 

Helosis cayennensis (Sw.) Spreng. 

Basellaceae 

Basella alba L. [cultivated] 

Begoniaceae 

Begonia humilis Dryand. 

Bignoniaceae 

Anemopaegma chrysoleucum (Kunth) Sandwith 

Anemopaegma karstenii Bureau & K. Schum. 

Anemopaegma oligoneuron (Sprague & 

Sandwith) A.H. Gentry 

Arrabidaea candicans (Rich.) DC. 

Callichlamys latifolia (Rich.) K. Schum. 

Ceratophytum tetragonolobum (Jacq.) Sprague 

& Sandwith 

Clytostoma binatum (Thunb.) Sandwith 

Crescentia amazonica Ducke 

Crescentia cujete L. [cultivated] 

Cydista aequinoctialis (L.) Miers 

Distictella elongata (Vahl) Urb. 

Jacaranda copaia (Aubl.) D. Don 

Jacaranda obtusifolia Bonpl. 

Lundia densiflora DC. 

Macfadyena uncata (T.F. Andrews) Sprague & 

Sandwith 

Macfadyena unguis-cati (L.) A.H. Gentry 

Mansoa kerere (Aubl.) A.H. Gentry 

Martinella obovata (Kunth) Bureau & K. 

Schum. 

Parabignonia steyermarkii Sandwith 

Pleonotoma albiflora (Salzm. ex DC.) A.H. 

Gentry 

Pleonotoma echitidea Sprague & Sandwith 

Schlegelia spruceana K. Schum. 

Schlegelia violacea (Aubl.) Griseb. 

Tabebuia fluviatilis (Aubl.) DC. 

Tabebuia insignis (Miq.) Sandwith 

Tabebuia serratifolia (Vahl) G. Nicholson 

Bixaceae 

Bixa orellana L. 

Bombacaceae 

Catostemma commune Sandwith 

Catostemma fragrans Benth. 

Ceiba pentandra (L.) Gaertn. 

Pachira aquatica Aubl. 

Pachira insignis (Sw.) Sw. ex Savigny 

139 

Boraginaceae 

Cordia curassavica (Jacq.) Roem. & Schult. 

Cordia exaltata Lam. 

Cordia fallax I.M. Johnst. 

Cordia nodosa Lam. 

Cordia schomburgkii DC. 

Cordia sericicalyx DC. 

Cordia tetrandra Aubl. 

Heliotropium indicum L. [naturalized] 

Tournefortia bicolor Sw. 

Tournefortia cuspidata Kunth 

Burseraceae 

Protium decandrum (Aubl.) Marchand 

Protium guianense (Aubl.) Marchand 

Protium heptaphyllum (Aubl.) Marchand 

Protium tenuifolium (Engl.) Engl. 

Protium unifoliolatum Spruce ex Engl. 

Tetragastris altissima (Aubl.) Swart 

Trattinnickia boliviana (Swart) Daly 

Trattinnickia burserifolia Mart. 

Trattinnickia rhoifolia Willd. 

Cabombaceae 

Cabomba aquatica Aubl. 

Cactaceae 

Epiphyllum phyllanthus (L.) Haw. 

Opuntia cochenillifera (L.) Mill. [cultivated] 

Pereskia aculeata Mill. [naturalized] 

Rhipsalis baccifera (J.S. Muell.) Stearn 


Bauhinia guianensis Aubl. 

Bauhinia scala-simiae Sandwith 

Bauhinia siqueiraei Ducke 

Brownea coccinea Jacq.

140 


Brownea grandiceps Jacq. * 

Caesalpinia bonduc (L.) Roxb. * 

Caesalpinia pulcherrima (L.) Sw. [cultivated] 

Chamaecrista ramosa (Vogel) H.S. Irwin & 

Barneby 

Crudia glaberrima (Steud.) J.F. Macbr. 

Dicorynia guianensis Amshoff 

Eperua falcata Aubl. 

Eperua rubiginosa Miq. 

Hymenaea courbaril L. 

Macrolobium acaciifolium (Benth.) Benth. 

Macrolobium angustifolium (Benth.) R.S. 

Cowan 

Macrolobium bifolium (Aubl.) Pers. 

Mora excelsa Benth. 

Peltogyne venosa (Vahl) Benth. 

Senna alata (L.) Roxb. 

Senna bacillaris (L. f.) H.S. Irwin & Barneby 

Senna multijuga (Rich.) H.S. Irwin & Barneby 

Senna occidentalis (L.) Link 

Senna quinquangulata (Rich.) H.S. Irwin & 

Barneby 

Senna reticulata (Willd.) H.S. Irwin & Barneby 

Senna sandwithiana H.S. Irwin & Barneby 

Tachigali micropetala (Ducke) Zarucchi & 

Pipoly 

Tachigali paniculata Aubl. 

Campanulaceae 

Centropogon cornutus (L.) Druce 

Capparaceae 

Cleome parviflora Kunth 

Cleome serrata Jacq. 

Cleome speciosa Raf. 

Caricaceae 

Carica papaya L. [naturalized] 

Caryocaraceae 

Caryocar microcarpum Ducke 

Caryocar nuciferum L. 

Casuarinaceae 

Casuarina equisetifolia J.R. Forst. & G. Forst. 

[cultivated] 

Cecropiaceae 

Cecropia angulata I.W. Bailey 

Cecropia obtusa Trécul 

Cecropia peltata L. 

Cecropia sciadophylla Mart. 

Coussapoa asperifolia Trécul * 

Coussapoa microcephala Trécul 

Pourouma guianensis Aubl. 

Celastraceae 

Goupia glabra Aubl. 

Maytenus guyanensis Klotzsch ex Reissek 

Ceratophyllaceae 

Ceratophyllum muricatum Cham. * 


Chrysobalanus icaco L. 

Couepia parillo DC. 

Hirtella paniculata Sw. 

Hirtella racemosa Lam. 

Hirtella silicea Griseb. 

Hirtella triandra Sw. 

Licania alba (Bernoulli) Cuatrec. 

Licania apetala (E. Mey.) Fritsch 

Licania boyanii Tutin 

Licania densiflora Kleinhoonte 

Licania divaricata Benth. 

Licania guianensis (Aubl.) Griseb. 

Licania heteromorpha Benth. 

Licania incana Aubl. 

Licania kunthiana Hook. f. 

Licania laxiflora Fritsch 

Licania majuscula Sagot 

Licania membranacea Sagot ex Laness. 

Licania micrantha Miq. 

Licania octandra (Hoffmanns. ex Roem. & 

Schult.) Kuntze 

Licania persaudii Fanshawe & Maguire 

Licania rufescens Klotzsch ex Fritsch 

Parinari campestris Aubl. 

Parinari rodolphii Huber 

Clusiaceae 

Calophyllum brasiliense Cambess. 

Caraipa richardiana Cambess. 

Clusia cuneata Benth. 

Clusia flavida (Benth.) Pipoly 

Clusia gaudichaudii Choisy 

Clusia grandiflora Splitg. 

Clusia myriandra (Benth.) Planch. & Triana 

Clusia nemorosa G. Mey. 

Clusia palmicida Rich. ex Planch. & Triana 

Clusia panapanari (Aubl.) Choisy 

Mammea americana L. [cultivated]


Rheedia macrophylla (Mart.) Planch. & Triana 

Rheedia virens Planch. & Triana 

Symphonia globulifera L. f. 

Tovomita brevistaminea Engl. 

Tovomita calodictyos Sandwith 

Tovomita choisyana Planch. & Triana 

Tovomita obscura Sandwith 

Tovomita schomburgkii Planch. & Triana 

Vismia cayennensis (Jacq.) Pers. 

Vismia gracilis Hieron. 

Vismia guianensis (Aubl.) Choisy 

Vismia japurensis Reichardt 

Vismia laxiflora Reichardt 

Vismia macrophylla Kunth 

Vismia sessilifolia (Aubl.) Choisy 

Combretaceae 

Buchenavia grandis Ducke 

Combretum cacoucia Exell ex Sandwith 

Combretum fruticosum (Loefl.) Stuntz 

Combretum laxum Jacq. 

Conocarpus erectus L. * 

Laguncularia racemosa (L.) C.F. Gaertn. 

Terminalia amazonia (J.F. Gmel.) Exell 

Terminalia catappa L. [naturalized] 

Terminalia dichotoma G. Mey. 

Connaraceae 

Cnestidium guianense (G. Schellenb.) G. 

Schellenb. 

Connarus coriaceus G. Schellenb. 

Pseudoconnarus macrophyllus (Poepp.) Radlk. 


Aniseia martinicensis (Jacq.) Choisy 

Dicranostyles sp. 

Ipomoea asarifolia (Desr.) Roem. & Schult. 

Ipomoea batatas (L.) Lam. [cultivated] 

Ipomoea carnea Jacq. 

Ipomoea indica (Burm.) Merr. 

Ipomoea mauritiana Jacq. 

Ipomoea pes-caprae (L.) R. Br. * 

Ipomoea phillomega (Vell.) House 

Ipomoea quamoclit L. 

Ipomoea tiliacea (Willd.) Choisy 

Ipomoea violacea L. * 

Jacquemontia guyanensis (Aubl.) Meisn. 

Jacquemontia tamnifolia (L.) Griseb. 

Maripa scandens Aubl. 

Merremia cissoides (Lam.) Hallier f. * 

Merremia macrocalyx (Ruiz & Pav.) O’Donell 

141 

Merremia umbellata (L.) Hallier f. * 

Crassulaceae 

Kalanchoe integra (Medic.) O. Kuntze 

[cultivated] 

Kalanchoe pinnata (Lam.) Pers. [cultivated] 

Cucurbitaceae 

Cayaponia jenmanii C. Jeffrey 

Cayaponia simplicifolia ? (Naudin) Cogn. 

Citrullus lanatus (Thunb.) Matsum. & Nakai 

[cultivated] 

Cucumis melo L. [cultivated] 

Cucumis sativus L. [cultivated] 

Cucurbita moschata Duchesne [cultivated] 

Gurania lobata (L.) Pruski 

Gurania subumbellata (Miq.) Cogn. 

Helmontia leptantha (Schltdl.) Cogn. 

Lagenaria siceraria (Molina) Standl. 

[cultivated] 

Luffa cylindrica (L.) M. Roem. [cultivated] 

Melothria pendula L. 

Momordica charantia L. [naturalized] 

Posadaea sphaerocarpa Cogn. 

Psiguria triphylla (Miq.) C. Jeffrey 

Cuscutaceae 

Cuscuta umbellata Kunth * 

Cyrillaceae 

Cyrilla racemiflora L. 

Dichapetalaceae 

Dichapetalum pedunculatum (DC.) Baill. 

Tapura guianensis Aubl. 

Dilleniaceae 

Davilla kunthii A. St.-Hil. 

Davilla nitida (Vahl) Kubitzki * 

Doliocarpus dentatus (Aubl.) Standl. 

Pinzona coriacea Mart. & Zucc. 

Tetracera asperula Miq. 

Tetracera tigarea DC. 

Tetracera volubilis L. 

Droseraceae 

Drosera intermedia Hayne 

Ebenaceae 

Diospyros cayennensis A. DC. 

Diospyros discolor Willd. [cultivated] 

Diospyros guianensis (Aubl.) Gürke

142 

Diospyros tetrandra Hiern 

Elaeocarpaceae 

Sloanea eichleri K. Schum. 

Sloanea grandiflora Sm. 

Sloanea guianensis (Aubl.) Benth. 

Sloanea latifolia (Rich.) K. Schum. 

Sloanea obtusifolia (Moric.) K. Schum. 

Erythroxylaceae 

Erythroxylum citrifolium A. St.-Hil. 

Erythroxylum macrophyllum Cav. 


Euphorbiaceae 

Acalypha amentacea Roxb. [cultivated] * 

Acalypha macrostachya Jacq. 

Acalypha scandens Benth. 

Alchornea discolor Poepp. 

Alchornea triplinervia (Spreng.) Müll. Arg. 

Alchorneopsis floribunda (Benth.) Müll. Arg. 

Amanoa guianensis Aubl. 

Chaetocarpus schomburgkianus (Kuntze) Pax 

& K. Hoffm. 

Codiaeum variegatum (L.) A. Juss. [cultivated] 

Conceveiba guianensis Aubl. 

Croton cuneatus Klotzsch 

Croton trinitatis Millsp. 

Dalechampia brownsbergensis G.L. Webster & 

Armbr. 

Drypetes fanshawei Sandwith 

Euphorbia cotinifolia L. 

Euphorbia hirta L. * 

Euphorbia neriifolia L. [naturalized] 

Euphorbia oerstediana (Klotzsch & Garcke) 

Boiss. 

Euphorbia thymifolia L. 

Hevea brasiliensis (Willd. ex A. Juss.) Müll. 

Arg. [cultivated] 

Hieronyma alchorneoides Allemao 

Jatropha curcas L. [cultivated] 

Jatropha gossypiifolia L. [cultivated] * 

Mabea piriri Aubl. 

Manihot esculenta Crantz [cultivated] 

Maprounea guianensis Aubl. 

Microstachys corniculata (Vahl) Griseb. 

Omphalea diandra L. 

Pausandra martinii Baill. 

Pedilanthus tithymaloides (L.) Poit. 

[cultivated]* 

Pera glabrata (Schott) Poepp. ex Baill. 

Phyllanthus brasiliensis (Aubl.) Poir. 

Phyllanthus caribaeus Urb. 

Phyllanthus stipulatus (Raf.) G.L. Webster 

Phyllanthus urinaria L. [naturalized] 

Plukenetia polyadenia Müll. Arg. 

Ricinus communis L. [cultivated] 

Sandwithia guyanensis Lanj. 

Sapium jenmanii Hemsl. 

Sapium paucinervium Hemsl. 

Senefeldera sp. 

Fabaceae 

Aeschynomene sensitiva Sw. * 

Alexa confusa Pittier 

Alexa imperatricis (R.H. Schomb.) Baill. 

Alexa surinamensis Yakovlev 

Andira surinamensis (Bondt) Splitg. ex Amshoff 

Cajanus cajan (L.) Millsp. [cultivated] 

Calopogonium caeruleum (Benth.) C. Wright 

Calopogonium mucunoides Desv. 

Canavalia rosea (Sw.) DC. * 

Centrosema brasilianum (L.) Benth. 

Centrosema capitatum (Rich.) Amshoff 

Centrosema molle Mart. ex Benth. 

Centrosema plumieri (Turpin ex Pers.) Benth. 

Clathrotropis brachypetala (Tul.) Kleinhoonte 

Clitoria arborescens R. Br. 

Crotalaria incana L. 

Crotalaria nitens Kunth 

Crotalaria stipularia Desv. 

Dalbergia monetaria L. f. 

Derris amazonica Killip 

Derris pterocarpus (DC.) Killip 

Desmodium adscendens (Sw.) DC. 

Desmodium axillare (Sw.) DC. 

Desmodium barbatum (L.) Benth. 

Desmodium incanum DC. 

Dioclea reflexa Hook. f. 

Dioclea scabra (Rich.) R.H. Maxwell 

Dioclea wilsonii Standl. 

Diplotropis purpurea (Rich.) Amshoff 

Dipteryx odorata (Aubl.) Willd. 

Hymenolobium flavum Kleinhoonte 

Indigofera suffruticosa Mill. [naturalized] 

Lablab purpureus (L.) Sweet [cultivated] 

Lonchocarpus chrysophyllus Kleinhoonte 

Lonchocarpus heptaphyllus (Poir.) DC. 

Lonchocarpus martynii A.C. Sm. 

Lonchocarpus rufescens Benth. 

Lonchocarpus sericeus (Poir.) Kunth ex DC. 

Lonchocarpus spruceanus Benth. 

Lonchocarpus utilis A.C. Sm.


Machaerium floribundum Benth. 

Machaerium inundatum (Mart. ex Benth.) 

Ducke 

Machaerium kegelii Meisn. 

Machaerium leiophyllum (DC.) Benth. 

Machaerium lunatum (L. f.) Ducke * 

Machaerium quinatum (Aubl.) Sandwith 

Mucuna urens (L.) Medik. 

Muellera frutescens (Aubl.) Standl. 

Ormosia coccinea (Aubl.) Jacks. 

Ormosia coutinhoi Ducke 

Ormosia nobilis Tul. 

Phaseolus lunatus L. [cultivated] 

Platymiscium pinnatum (Jacq.) Dugand 

Pterocarpus officinalis Jacq. 

Pterocarpus santalinoides L’Hér. ex DC. 

Pueraria phaseoloides (Roxb.) Benth. 

[cultivated] * 

Rhynchosia phaseoloides (Sw.) DC. 

Sesbania sericea (Willd.) DC. * 

Swartzia conferta Spruce ex Benth. 

Swartzia guianensis (Aubl.) Urb. 

Swartzia schomburgkii Benth. 

Swartzia steyermarkii R.S. Cowan 

Tephrosia sinapou (Buc’hoz) A. Chev. 

Vatairea guianensis Aubl. 

Vigna luteola (Jacq.) Benth. 

Vigna sinensis (L.) Savi ex Hassk. [cultivated] 

Vigna unguiculata (L.) Walp. [cultivated] 

Zornia diphylla (L.) Pers. 

Flacourtiaceae 

Banara guianensis Aubl. 

Casearia acuminata DC. 

Casearia commersoniana Cambess. 

Casearia guianensis (Aubl.) Urb. 

Casearia javitensis Kunth 

Casearia rusbyana Briq. 

Flacourtia jangomas (Lour.) Raeusch. 

[cultivated] 

Homalium guianense (Aubl.) Oken 

Homalium racemosum Jacq. 

Laetia procera (Poepp.) Eichler 

Ryania speciosa Vahl 

Xylosma benthamii (Tul.) Triana & Planch. 

Gentianaceae 

Chelonanthus alatus (Aubl.) Pulle 

Chelonanthus purpurascens (Aubl.) Struwe, S. 

Nilsson & V.A. 

Coutoubea ramosa Aubl. 

143 

Voyria aphylla (Jacq.) Pers. 

Voyria aurantiaca Splitg. 

Voyria pittieri (Standl.) L.O. Williams 

Gesneriaceae 

Besleria flavovirens Nees & Mart. 

Chrysothemis pulchella (Donn ex Sims) Decne. 

Chrysothemis villosa (Benth.) Leeuwenb.* 

Codonanthe calcarata (Miq.) Hanst. 

Codonanthe crassifolia (H. Focke) C.V. Morton 

Drymonia serrulata (Jacq.) Mart. * 

Nautilocalyx coccineus Feuillet & L.E. Skog 

Nautilocalyx cordatus (Gleason) L.E. Skog 

Nautilocalyx mimuloides (Benth.) C.V. Morton 

Paradrymonia maculata (Hook. f.) Wiehler 


Cheiloclinium cognatum (Miers) A.C. Sm. 

Hippocratea volubilis L. 

Peritassa pruinosa (Seem.) A.C. Sm. 

Tontelea glabra A.C. Sm. 

Humiriaceae 

Humiria balsamifera Aubl. 

Humiriastrum obovatum (Benth.) Cuatrec. 

Sacoglottis cydonioides Cuatrec. 

Hydrophyllaceae 

Hydrolea spinosa L. 

Icacinaceae 

Discophora guianensis Miers 

Emmotum fagifolium Ham. 

Leretia cordata Vell. 

Poraqueiba guianensis Aubl. 

Lacistemataceae 

Lacistema aggregatum (P.J. Bergius) Rusby 

Lamiaceae 

Coleus amboinicus Lour. [cultivated] 

Coleus blumei Benth. [cultivated] 

Coleus hybridus Hort. ex Cobeau [cultivated] 

Hyptis lanceolata Poir. 

Hyptis parkeri Benth. 

Hyptis pectinata (L.) Poit. 

Leonotis nepetifolia (L.) W.T. Aiton 

Ocimum campechianum Mill. 

Lauraceae 

Aiouea guianensis Aubl. 

Aniba guianensis Aubl.

144 

Aniba hostmanniana (Nees) Mez 

Aniba hypoglauca Sandwith 

Aniba jenmanii Mez 

Aniba kappleri Mez 

Aniba terminalis Ducke 

Cassytha filiformis L. * 

Licaria debilis (Mez) Kosterm. 

Licaria oppositifolia (Nees) Kosterm. 

Nectandra amazonum Nees 

Nectandra canescens Nees 

Nectandra cuspidata Nees 

Nectandra globosa (Aubl.) Mez 

Ocotea cernua (Nees) Mez 

Ocotea puberula (Rich.) Nees 

Ocotea schomburgkiana (Nees) Mez 

Ocotea splendens (Meisn.) Baill. 

Ocotea tomentella Sandwith 

Persea americana Mill. [cultivated] 

Persea nivea Mez 


Lecythidaceae 

Couratari guianensis Aubl. 

Couratari multiflora (Sm.) Eyma 

Eschweilera alata A.C. Sm. 

Eschweilera coriacea (DC.) S.A. Mori 

Eschweilera decolorans Sandwith 

Eschweilera micrantha (O. Berg) Miers 

Eschweilera parviflora (Aubl.) Miers 

Eschweilera sagotiana Miers 

Eschweilera wachenheimii (Benoist) Sandwith 

Gustavia augusta L. 

Gustavia poeppigiana O. Berg 

Lecythis chartacea O. Berg 

Lecythis corrugata Poit. 

Lecythis zabucajo Aubl. 

Lentibulariaceae 

Utricularia benjaminiana Oliv. 

Utricularia foliosa L. 

Utricularia myriocista A. St.-Hil. & Girard 

Loganiaceae 

Strychnos erichsonii M.R. Schomb. ex Progel 

Strychnos mitscherlichii M.R. Schomb. 

Strychnos toxifera R.H. Schomb. ex Benth. 

Loranthaceae 

Oryctanthus florulentus (Rich.) Tiegh. 

Phthirusa pyrifolia (Kunth) Eichler 

Phthirusa stelis (L.) Kuijt 

Lythraceae 

Cuphea melvilla Lindl. 

Malpighiaceae 

Banisteriopsis caapi (Griseb.) C.V. Morton 

Banisteriopsis lucida (Rich.) Small 

Banisteriopsis martiniana (A. Juss.) Cuatrec. 

Burdachia sphaerocarpa A. Juss. 

Byrsonima aerugo Sagot 

Byrsonima crassifolia (L.) Kunth 

Byrsonima spicata (Cav.) DC. 

Byrsonima stipulacea A. Juss. 

Diplopterys pauciflora (G. Mey.) Nied. 

Heteropterys leona (Cav.) Exell 

Heteropterys macrostachya A. Juss. 

Hiraea fagifolia (DC.) A. Juss. 

Hiraea faginea (Sw.) Nied. 

Lophopterys euryptera Sandwith 

Malpighia emarginata DC. 

Mascagnia macrodisca (Triana & Planch.) Nied. 

Mezia includens (Benth.) Cuatrec. 

Spachea elegans (G. Mey.) A. Juss. 

Stigmaphyllon bannisterioides (L.) C.E. 

Anderson 

Stigmaphyllon convolvulifolium A. Juss. 

Stigmaphyllon puberum (Rich.) A. Juss. 

Stigmaphyllon sinuatum (DC.) A. Juss. 

Tetrapterys discolor (G. Mey.) DC. 

Tetrapterys fimbripetala A. Juss. 

Malvaceae 

Abelmoschus moschatus Medik. 

Gossypium barbadense L. [cultivated] 

Gossypium hirsutum L. [cultivated] 

Hibiscus bifurcatus Cav. 

Hibiscus furcellatus Desr. 

Hibiscus rosa-sinensis L. [cultivated] 

Hibiscus sabdariffa L. [cultivated] 

Malachra alceifolia Jacq. 

Pavonia fruticosa (Mill.) Fawc. & Rendle 

Sida acuta Burm. f. 

Sida glomerata Cav. 

Sida linifolia Juss. ex Cav. 

Sida rhombifolia L. 

Sida setosa Mart. ex Colla 

Talipariti tiliaceum (L.) Fryxell 

Thespesia populnea (L.) Sol. ex Corr?a * 

[naturalized] * 

Urena lobata L.


Marcgraviaceae 

Marcgravia coriacea Vahl 

Marcgravia magnibracteata Lanj. & Heerdt 

Marcgravia pedunculosa Triana & Planch. 

Marcgravia purpurea I.W. Bailey 

Norantea guianensis Aubl. 

Souroubea guianensis Aubl. 


Aciotis annua (Mart. ex DC.) Triana 

Aciotis indecora (Bonpl.) Triana 

Aciotis ornata (Miq.) Gleason 

Aciotis purpurascens (Aubl.) Triana 

Bellucia grossularioides (L.) Triana 

Clidemia capitellata (Bonpl.) D. Don 

Clidemia conglomerata DC. 

Clidemia dentata D. Don 

Clidemia hirta (L.) D. Don 

Clidemia japurensis DC. 

Clidemia microthyrsa R.O. Williams 

Clidemia pustulata DC. 

Clidemia rubra (Aubl.) Mart. 

Clidemia venosa (Gleason) Wurdack 

Comolia villosa (Aubl.) Triana 

Desmoscelis villosa (Aubl.) Naudin 

Henriettea multiflora Naudin 

Henriettea succosa (Aubl.) DC. 

Henriettella caudata Gleason 

Leandra divaricata (Naudin) Cogn. 

Leandra rufescens (DC.) Cogn. 

Loreya mespiloides Miq. 

Miconia acinodendron (L.) Sweet 

Miconia affinis DC. 

Miconia bubalina (D. Don) Naudin 

Miconia ceramicarpa (DC.) Cogn. 

Miconia chrysophylla (Rich.) Urb. 

Miconia ciliata (Rich.) DC. 

Miconia egensis Cogn. 

Miconia fragilis Naudin 

Miconia gratissima Benth. ex Triana 

Miconia hypoleuca (Benth.) Triana 

Miconia ibaguensis (Bonpl.) Triana 

Miconia lateriflora Cogn. 

Miconia lepidota DC. 

Miconia matthaei Naudin 

Miconia minutiflora (Bonpl.) DC. * 

Miconia mirabilis (Aubl.) L.O. Williams 

Miconia myriantha Benth. 

Miconia nervosa (Sm.) Triana 

Miconia plukenetii Naudin 

Miconia prasina (Sw.) DC. 

Miconia pubipetala Miq. 

Miconia racemosa (Aubl.) DC. 

Miconia rubiginosa (Bonpl.) DC. 

Miconia ruficalyx Gleason 

Miconia serrulata (DC.) Naudin 

Myriaspora egensis DC. 

Nepsera aquatica (Aubl.) Naudin 

Pterolepis glomerata (Rottb.) Miq. 

Rhynchanthera dichotoma (Desr.) DC. 

Tococa aristata Benth. 

Meliaceae 

Azadirachta indica A. Juss. [cultivated] 

Carapa guianensis Aubl. 

Cedrela odorata L. 

Guarea guidonia (L.) Sleumer 

Guarea pubescens (Rich.) A. Juss. 

Trichilia rubra C. DC. 

Trichilia schomburgkii C. DC. 

Mendonciaceae 

Mendoncia bivalvis (L. f.) Merr. 

Mendoncia glabra (Poepp. & Endl.) Nees 

Mendoncia hoffmannseggiana Nees 

145 

Menispermaceae 

Cissampelos andromorpha DC. 

Curarea candicans (Rich. ex DC.) Barneby & 

Krukoff 

Orthomene schomburgkii (Miers) Barneby & 

Krukoff 

Telitoxicum sp. 

Menyanthaceae 

Nymphoides indica (L.) Kuntze 

Mimosaceae 

Abarema jupunba (Willd.) Britton & Killip 

Abarema laeta (Benth.) Barneby & J.W. Grimes 

Abarema mataybifolia (Sandwith) Barneby & 

J.W. Grimes 

Calliandra surinamensis Benth. 

Entada polystachya (L.) DC. * 

Hydrochorea corymbosa (Rich.) Barneby & 

J.W. Grimes 

Hydrochorea gonggrijpii (Kleinhoonte) 

Barneby & J.W. Grimes 

Inga acreana Harms 

Inga acrocephala Steud. 

Inga alba (Sw.) Willd. 

Inga bourgonii (Aubl.) DC.

146 


Inga edulis Mart. 

Inga graciliflora Benth. 

Inga gracilifolia Ducke 

Inga huberi Ducke 

Inga ingoides (Rich.) Willd. * 

Inga java Pittier 

Inga jenmanii Sandwith 

Inga lateriflora Miq. 

Inga leiocalycina Benth. 

Inga marginata Willd. 

Inga melinonis Sagot 

Inga nobilis Willd. 

Inga pezizifera Benth. 

Inga pilosula (Rich.) J.F. Macbr. 

Inga rubiginosa (Rich.) DC. 

Inga sarmentosa Glaz. ex Harms 

Inga sertulifera DC. 

Inga splendens Willd. 

Inga thibaudiana DC. 

Inga umbellifera (Vahl) Steud. ex DC. 

Macrosamanea pubiramea (Steud.) Barneby & 

J.W. Grimes 

Mimosa polydactyla Humb. & Bonpl. ex Willd. 

Mimosa myriadenia (Benth.) Benth. * 

Pentaclethra macroloba (Willd.) Kuntze 

Pithecellobium longiflorum Benth. 

Zygia cataractae (Kunth) L. Rico 

Zygia latifolia (L.) Fawc. & Rendle 

Moraceae 

Artocarpus altilis (Parkinson) Fosberg 

[cultivated] 

Bagassa guianensis Aubl. 

Brosimum guianense (Aubl.) Huber 

Ficus amazonica (Miq.) Miq. 

Ficus caballina Standl. 

Ficus eximia Schott 

Ficus gomelleira Kunth & Bouché 

Ficus guianensis Desv. ex Ham. 

Ficus malacocarpa Standl. 

Ficus mathewsii (Miq.) Miq. 

Ficus maxima Mill. 

Ficus nymphaeifolia Mill. 

Ficus paludica Standl. 

Ficus paraensis (Miq.) Miq. 

Ficus roraimensis C.C. Berg 

Ficus trigona L. f. 

Pseudolmedia laevis (Ruiz & Pav.) J.F. Macbr. 

Moringaceae 

Moringa oleifera Lam. [cultivated] * 

Myristicaceae 

Iryanthera juruensis Warb. 

Iryanthera lancifolia Ducke * 

Iryanthera macrophylla (Benth.) Warb. 

Virola calophylla (Spruce) Warb. 

Virola elongata (Benth.) Warb. 

Virola sebifera Aubl. 

Virola surinamensis (Rol. ex Rottb.) Warb. 

Myrsinaceae 

Ardisia guianensis (Aubl.) Mez 

Cybianthus surinamensis (Spreng.) G. Agostini 

Stylogyne orinocensis (Kunth) Mez 

Myrtaceae 

Calycolpus goetheanus (DC.) O. Berg 

Calyptranthes sp. 

Eugenia cucullata Amshoff 

Eugenia florida DC. 

Eugenia patrisii Vahl 

Eugenia punicifolia (Kunth) DC. 

Eugenia uniflora L. [cultivated] 

Marlierea montana (Aubl.) Amshoff 

Marlierea schomburgkiana O. Berg 

Myrcia fallax (Rich.) DC. 

Myrcia graciliflora Sagot 

Myrcia guianensis (Aubl.) DC. 

Myrcia servata McVaugh 

Myrcia sylvatica (G. Mey.) DC. 

Psidium cattleianum Sabine [cultivated] 

Psidium guajava L. [naturalized] 

Syzygium cumini (L.) Skeels [cultivated] 

Syzygium jambos (L.) Alston [cultivated] 

Syzygium malaccense (L.) Merr. & Perry 

[cultivated] 


Boerhavia diffusa L. * 

Guapira eggersiana (Heimerl) Lundell 

Guapira salicifolia (Heimerl) Lundell 

Neea constricta Spruce ex J.A. Schmidt 

Neea floribunda Poepp. & Endl. 

Nymphaeaceae 

Nymphaea odorata Aiton 

Nymphaea pulchella DC. 

Nymphaea rudgeana G. Mey. 

Ochnaceae 

Ouratea candollei (Planch.) Tiegh. 

Ouratea leblondii (Tiegh.) Lemée

Ouratea macrocarpa Sastre 

Ouratea rorida Sastre 

Sauvagesia elata Benth. 

Sauvagesia erecta L. 

Sauvagesia sprengelii A. St.-Hil. 

Olacaceae 

Heisteria maguirei Sleumer 


Onagraceae 

Ludwigia affinis (DC.) H. Hara 

Ludwigia foliobracteolata (Munz) H. Hara 

Ludwigia hyssopifolia (G. Don) Exell 

Ludwigia latifolia (Benth.) H. Hara 

Ludwigia leptocarpa (Nutt.) H. Hara * 

Ludwigia nervosa (Poir.) H. Hara 

Ludwigia octovalvis (Jacq.) P.H. Raven 

Ludwigia torulosa (Arn.) H. Hara 

Oxalidaceae 

Averrhoa carambola L. [cultivated] 

Oxalis barrelieri L. 

Oxalis debilis Kunth 


Passiflora amicorum Wurdack * 

Passiflora auriculata Kunth 

Passiflora cirrhiflora Juss. 

Passiflora coccinea Aubl. 

Passiflora foetida L. 

Passiflora garckei Mast. 

Passiflora glandulosa Cav. 

Passiflora laurifolia L. 

Passiflora nitida Kunth 

Passiflora quadrangularis L. [cultivated] 

Passiflora quadriglandulosa Rodschied 

Phytolaccaceae 

Microtea debilis Sw. 

Petiveria alliacea L. 

Phytolacca rivinoides Kunth & Bouché 

Piperaceae 

Peperomia duidana Trel. ex Gleason 

Peperomia elongata Kunth 

Peperomia glabella (Sw.) A. Dietr. 

Peperomia macrostachya (Vahl) A. Dietr. 

Peperomia magnoliifolia (Jacq.) A. Dietr. 

Peperomia obtusifolia (L.) A. Dietr. 

Peperomia pernambucensis Miq. 

Peperomia rotundifolia (L.) Kunth 

Peperomia serpens (Sw.) Loudon 

Piper aduncum L. 

Piper aequale Vahl 

Piper anonifolium (Kunth) C. DC. 

Piper arboreum Aubl. 

Piper avellanum (Miq.) C. DC. 

Piper coruscans Kunth 

Piper dilatatum Rich. 

Piper divaricatum G. Mey. 

Piper glabrescens (Miq.) C. DC. 

Piper hispidum Sw. 

Piper hostmannianum (Miq.) C. DC. 

Piper nigrispicum C. DC. 

Piper peltatum L. * 

Piper pseudoglabrescens Trel. & Yunck. 

Piper pulleanum Yunck. 

Polygalaceae 

Bredemeyera densiflora A.W. Benn. 

Moutabea guianensis Aubl. 

Securidaca diversifolia (L.) S.F. Blake 

Securidaca paniculata Rich. 

147 

Polygonaceae 

Antigonon leptopus Hook. & Arn. [cultivated] 

* 

Coccoloba ascendens Duss ex Lindau 

Coccoloba densifrons Mart. ex Meisn. 

Coccoloba marginata Benth. 

Coccoloba parimensis Benth. 

Polygonum punctatum Elliott 

Triplaris weigeltiana (Rchb.) Kuntze 

Portulacaceae 

Portulaca oleracea L. 

Portulaca pilosa L. 

Portulaca sedifolia N.E. Br. 

Quiinaceae 

Quiina guianensis Aubl. 

Quiina indigofera Sandwith 

Rhamnaceae 

Gouania lupuloides (L.) Urb. s.l. 

Gouania polygama (Jacq.) Urb. 


Cassipourea guianensis Aubl. 

Cassipourea lasiocalyx Alston 

Rhizophora x harrisonii Leechm. *? 

Rhizophora mangle L.

148 

Rhizophora racemosa G. Mey. 


Rubiaceae 

Amaioua corymbosa Kunth 

Amaioua guianensis Aubl. 

Bertiera guianensis Aubl. 

Borreria assurgens (Ruiz & Pav.) Griseb. 

Borreria capitata (Ruiz & Pav.) DC. 

Borreria densiflora DC. 

Borreria latifolia (Aubl.) K. Schum. 

Borreria prostrata (Aubl.) Miq. 

Borreria suaveolens G. Mey. 

Borreria verticillata (L.) G. Mey. 

Chimarrhis microcarpa Standl. 

Cinchona sp. [cultivated] 

Coccocypselum guianense (Aubl.) K. Schum. 

Coccocypselum tontanea Kunth 

Coffea arabica L. [cultivated] 

Coffea liberica W. Bull ex Hiern [cultivated] 

Cosmibuena grandiflora (Ruiz & Pav.) Rusby 

Coussarea leptoloba (Spreng. ex Benth. & 

Hook. f.) Müll. Arg. 

Coussarea violacea Aubl. 

Diodia ocymifolia (Willd. ex Roem. & Schult.) 

Bremek. 

Diodia sarmentosa Sw. 

Duroia eriopila L. f. 

Faramea guianensis (Aubl.) Bremek. 

Faramea multiflora A. Rich. ex DC. 

Genipa americana L. 

Genipa spruceana Steyerm. 

Geophila repens (L.) I.M. Johnst. 

Geophila tenuis (Müll. Arg.) Standl. 

Gonzalagunia bunchosioides Standl. 

Gonzalagunia cornifolia (Kunth) Standl. 

Gonzalagunia dicocca Cham. & Schltdl. 

Gonzalagunia spicata (Lamb.) M. Gómez 

Hamelia patens Jacq. 

Hillia illustris (Vell.) K. Schum. 

Ixora coccinea L. [cultivated] 

Ixora schomburgkiana Benth. 

Malanea hypoleuca Steyerm. 

Malanea macrophylla Bartl. ex Griseb. 

Morinda citrifolia L. [naturalized] * 

Notopleura uliginosa (Sw.) Bremek. 

Oldenlandia lancifolia (Schumach.) DC. 

[naturalized] 

Palicourea crocea (Sw.) Roem. & Schult. 

Palicourea croceoides Ham. 

Palicourea guianensis Aubl. 

Palicourea triphylla DC. 

Posoqueria coriacea M. Martens & Galeotti 

Posoqueria longiflora Aubl. 

Posoqueria trinitatis DC. 

Psychotria acuminata Benth. 

Psychotria anceps Kunth 

Psychotria apoda Steyerm. 

Psychotria bahiensis DC. 

Psychotria barbiflora DC. 

Psychotria callithrix (Miq.) Steyerm. 

Psychotria capitata Ruiz & Pav. 

Psychotria cupularis (Müll. Arg.) Standl. 

Psychotria deflexa DC. 

Psychotria erecta (Aubl.) Standl. & Steyerm. 

Psychotria gracilenta Müll. Arg. 

Psychotria hoffmannseggiana (Willd. ex Roem. 

& Schult.) Müll. 

Psychotria horizontalis Sw. 

Psychotria iodotricha Müll. Arg. 

Psychotria irwinii Steyerm. 

Psychotria mapourioides DC. 

Psychotria platypoda DC. 

Psychotria poeppigiana Müll. Arg. 

Psychotria racemosa Rich. 

Psychotria tillettii Steyerm. 

Psychotria ulviformis Steyerm. 

Psychotria wessels-boeri Steyerm. 

Randia armata (Sw.) DC. 

Rudgea hostmanniana Benth. 

Rudgea standleyana Steyerm. 

Rudgea stipulacea (DC.) Steyerm. 

Sabicea aspera Aubl. 

Sabicea glabrescens Benth. 

Sabicea oblongifolia (Miq.) Steyerm. 

Sabicea velutina Benth. 

Schradera polycephala DC. 

Sipanea biflora (L. f.) Cham. & Schltdl. 

Sipanea pratensis Aubl. 

Uncaria guianensis (Aubl.) J.F. Gmel. 

Uncaria tomentosa (Willd. ex Roem. & Schult.) 

DC. 

Rutaceae 

Citrus aurantifolia (Christm.) Swingle 

[cultivated] 

Citrus aurantium L. [cultivated] 

Citrus deliciosa Terr. [cultivated] 

Citrus medica L. [cultivated] 

Citrus paradisi Macfad. [cultivated] 

Citrus reticulata Blanco [cultivated] 

Citrus sinensis (L.) Osbeck [cultivated] 

Ertela trifolia (L.) Kuntze


Murraya paniculata (L.) Jack [cultivated] 

Zanthoxylum rhoifolium Lam. 

Sabiaceae 

Meliosma herbertii Rolfe [cultivated] 


Allophylus racemosus Sw. 

Cardiospermum halicacabum L. 

Cupania hirsuta Radlk. 

Cupania scrobiculata Rich. 

Matayba camptoneura Radlk. 

Matayba opaca Radlk. 

Paullinia caloptera Radlk. 

Paullinia capreolata (Aubl.) Radlk. 

Paullinia hitchcockii Gleason 

Paullinia pinnata L. 

Paullinia rufescens Rich. ex Juss. 

Paullinia xestophylla Radlk. 

Pseudima frutescens (Aubl.) Radlk. 

Serjania membranacea Splitg. 

Serjania paucidentata DC. 

Serjania pyramidata Radlk. 

Talisia guianensis Aubl. 

Talisia hemidasya Radlk. 

Talisia hexaphylla Vahl 

Sapotaceae 

Chrysophyllum argenteum Jacq. 

Chrysophyllum cainito L. [naturalized] 

Chrysophyllum pomiferum (Eyma) T.D. Penn. 

Chrysophyllum sanguinolentum (Pierre) Baehni 

Manilkara bidentata (A. DC.) A. Chev. 

Manilkara zapota (L.) P. Royen [cultivated] 

Micropholis venulosa (Mart. & Eichler) Pierre 

Pouteria ambelaniifolia (Sandwith) T.D. Penn. 

Pouteria bilocularis (H. Winkl.) Baehni 

Pouteria caimito (Ruiz & Pav.) Radlk. 

Pouteria coriacea (Pierre) Pierre 

Pouteria cuspidata (A. DC.) Baehni 

Pouteria durlandii (Standl.) Baehni 

Pouteria guianensis Aubl. 

Pouteria hispida Eyma 

Pouteria venosa (Mart.) Baehni 

Pradosia schomburgkiana (A. DC.) Cronquist 


Achetaria guianensis Pennell 

Angelonia biflora Benth. 

Asarina erubescens (D. Don) Pennell 

[cultivated] 

Bacopa aquatica Aubl. 

Capraria biflora L. 

Lindernia crustacea (L.) F. Muell. 

Lindernia diffusa (L.) Wettst. 

Scoparia dulcis L. 

Simaroubaceae 

Picramnia guianensis (Aubl.) Jans.-Jac. 

Picramnia latifolia Tul. 

Quassia amara L. [naturalized] 

Simarouba amara Aubl. 

Siparunaceae 

Siparuna decipiens (Tul.) A. DC. 

Siparuna guianensis Aubl. 

Solanaceae 

Capsicum annuum L. [naturalized] 

Cestrum latifolium Lam. 

Markea camponoti Ducke 

Markea longiflora Miers * 

Nicotiana tabacum L. [cultivated] 

Physalis angulata L. 

Physalis pubescens L. 

Solanum adhaerens Roem. & Schult. 

Solanum asperum Rich. 

Solanum leucocarpon Dunal 

Solanum lycopersicum L. [cultivated] 

Solanum pensile Sendtn. 

Solanum rugosum Dunal 

Solanum schlechtendalianum Walp. 

Solanum stramoniifolium Jacq. 

Solanum subinerme Jacq. 

149 

Sterculiaceae 

Herrania kanukuensis R.E. Schult. 

Herrania lemniscata (M.R. Schomb.) R.E. 

Schult. 

Sterculia pruriens (Aubl.) K. Schum. 

Sterculia rugosa R. Br. 

Theobroma cacao L. [cultivated] 

Waltheria indica L. 

Theophrastaceae 

Clavija lancifolia Desf. 

Tiliaceae 

Apeiba petoumo Aubl. 

Corchorus aestuans L. 

Triumfetta althaeoides Lam.

150 

Turneraceae 

Turnera scabra Millsp. 

Turnera ulmifolia L. 

Ulmaceae 

Trema micrantha (L.) Blume 

Urticaceae 

Laportea aestuans (L.) Chew 

Pilea pubescens Liebm. 

Urera baccifera (L.) Gaudich. ex Wedd. 


Verbenaceae 

Aegiphila racemosa Vell. 

Citharexylum macrophyllum Poir. 

Clerodendrum fragrans (Vent.) Willd. 

[naturalized] 

Lantana camara L. 

Lippia alba (Mill.) N.E. Br. 

Lippia micromera Schauer 

Petrea volubilis L. 

Priva lappulacea (L.) Pers. 

Stachytarpheta cayennensis (Rich.) Vahl 

Stachytarpheta jamaicensis (L.) Vahl 

Vitex compressa Turcz. 

Vitex stahelii Moldenke 

Vitex triflora Vahl 

Violaceae 

Amphirrhox longifolia (St.-Hil.) Spreng. 

Paypayrola grandiflora Tul. 

Paypayrola longifolia Tul. 

Rinorea flavescens (Aubl.) Kuntze 

Rinorea pubiflora (Benth.) Sprague & Sandwith 

Rinorea riana (DC.) Kuntze 

Viscaceae 

Phoradendron bathyoryctum Eichler 

Phoradendron perrottetii (DC.) Eichler 

Phoradendron piperoides (Kunth) Trel. 

Phoradendron racemosum (Aubl.) Krug & Urb. 

Vitaceae 

Cissus erosa Rich. 

Cissus verticillata (L.) Nicolson & C.E. Jarvis 

Vochysiaceae 

Vochysia guianensis Aubl. 

MONOCOTILEDONEAE 

Agavaceae 

Agave americana L. [cultivated] 

Furcraea foetida (L.) Haw. 

Alismataceae 

Sagittaria lancifolia L. 

Araceae 

Anthurium gracile (Rudge) Schott 

Anthurium obtusum (Engl.) Grayum 

Caladium bicolor (Aiton) Vent. 

Caladium humboldtii Schott [cultivated] 

Caladium schomburgkii Schott 

Colocasia esculenta (L.) Schott [cultivated] 

Dieffenbachia humilis Poepp. 

Dieffenbachia paludicola N.E. Br. ex Gleason 

Heteropsis flexuosa (Kunth) G.S. Bunting 

Monstera adansonii Schott 

Monstera obliqua Miq. 

Montrichardia arborescens (L.) Schott 

Montrichardia linifera (Arruda) Schott * 

Philodendron acutatum Schott * 

Philodendron brevispathum Schott 

Philodendron deflexum Poepp. ex Schott 

Philodendron fragrantissimum (Hook.) G. Don 

Philodendron grandifolium (Jacq.) Schott 

Philodendron hederaceum (Jacq.) Schott 

Philodendron linnaei Kunth 

Philodendron melinonii Brongn. ex Regel 

Philodendron ornatum Schott 

Philodendron pedatum (Hook.) Kunth 

Philodendron rudgeanum Schott 

Philodendron scandens K. Koch & Sello 

Philodendron surinamense (Miq.) Engl. 

Rhodospatha oblongata Poepp. 

Spathiphyllum cannifolium (Dryand. ex Sims) 

Schott 

Spathiphyllum cuspidatum Schott 

Spathiphyllum maguirei G.S. Bunting 

Stenospermation maguirei A.M.E. Jonker & 

Jonker * 

Syngonium podophyllum Schott 

Urospatha sagittifolia (Rudge) Schott 

Xanthosoma belophyllum (Willd.) Schott 

[naturalized] 

Xanthosoma brasiliense (Desf.) Engl. 

[cultivated] 

Xanthosoma sagittifolium (L.) Schott 

[cultivated] 

Xanthosoma undipes (K. Koch & C.D. Bouché) 

K. Koch 

Arecaceae 

Astrocaryum aculeatum G. Mey.


Astrocaryum gynacanthum Mart. 

Astrocaryum vulgare Mart. 

Attalea maripa (Aubl.) Mart. 

Bactris acanthocarpa Mart. 

Bactris brongniartii Mart. 

Bactris campestris Poepp. ex Mart. 

Bactris gasipaes Kunth [naturalized] 

Bactris major Jacq. 

Bactris maraja Mart. 

Bactris oligoclada Burret 

Bactris simplicifrons Mart. 

Cocos nucifera L. [cultivated] 

Desmoncus orthacanthos Mart. 

Desmoncus polyacanthos Mart. 

Elaeis guineensis Jacq. [cultivated] 

Euterpe oleracea Mart. 

Euterpe precatoria Mart. 

Geonoma baculifera (Poit.) Kunth 

Geonoma macrostachys Mart. 

Geonoma maxima (Poit.) Kunth 

Manicaria saccifera Gaertn. 

Mauritia flexuosa L. f. 

Nypa fruticans Wurmb. [naturalized] * 

Oenocarpus bataua Mart. 

Roystonea oleracea (Jacq.) O.F. Cook * 

Socratea exorrhiza (Mart.) H. Wendl. 

Bromeliaceae 

Aechmea angustifolia Poepp. & Endl. 

Aechmea bromeliifolia (Rudge) Baker 

Aechmea lingulata (L.) Baker 

Aechmea mertensii (G. Mey.) Schult. & Schult. 

f. 

Aechmea nudicaulis (L.) Griseb. 

Ananas comosus (L.) Merr. [naturalized] 

Araeococcus micranthus Brongn. 

Bromelia plumieri (E. Morren) L.B. Sm. 

Disteganthus lateralis (L.B. Sm.) Gouda 

Guzmania lingulata (L.) Mez 

Guzmania monostachia (L.) Rusby ex Mez 

Guzmania roezlii (E. Morren) Mez 

Mezobromelia pleiosticha (Griseb.) Utley & H. 

Luther 

Tillandsia bulbosa Hook. 

Tillandsia monadelpha (E. Morren) Baker 

Vriesea heliconioides (Kunth) Hook. ex Walp. 

Vriesea procera (Mart. ex Schult. f.) Wittm. * 

Werauhia gigantea (Mart. ex Schult. f.) J.R. 

Grant * 

Burmanniaceae 

Burmannia tenella Benth. 

Cannaceae 

Canna indica L. [cultivated] 

Canna x generalis L.H. Bailey [cultivated] 

Commelinaceae 

Commelina diffusa Burm. f. 

Commelina erecta L. * 

Commelina rufipes Seub. 

Dichorisandra hexandra (Aubl.) Standl. 

Gibasis geniculata (Jacq.) Rohweder 

Tripogandra serrulata (Vahl) Handlos 

151 

Costaceae 

Costus amazonicus (Loes.) J.F. Macbr. 

[cultivated] 

Costus arabicus L. 

Costus congestiflorus Rich. ex Gagnep. 

Costus erythrothyrsus Loes. 

Costus guanaiensis Rusby 

Costus scaber Ruiz & Pav. 

Costus spiralis (Jacq.) Roscoe 

Cyclanthaceae 

Asplundia brachyphylla Harling 

Asplundia glandulosa (Gleason) Harling 

Asplundia gleasonii Harling 

Asplundia guianensis Harling 

Cyclanthus bipartitus Poit. 

Evodianthus funifer (Poit.) Lindm. 

Thoracocarpus bissectus (Vell.) Harling 

Cyperaceae 

Becquerelia cymosa Brongn. 

Calyptrocarya bicolor (H. Pfeiff.) T. Koyama 

Cyperus aggregatus (Willd.) Endl. 

Cyperus articulatus L. 

Cyperus comosus Poir. 

Cyperus digitatus Roxb. 

Cyperus haspan L. 

Cyperus laxus Lam. 

Cyperus ligularis L. 

Cyperus luzulae (L.) Rottb. ex Retz. 

Cyperus odoratus L. 

Cyperus polystachyos Rottb. * 

Cyperus simplex Kunth 

Cyperus sphacelatus Rottb.

152 


Cyperus surinamensis Rottb. 

Diplasia karatifolia Rich. 

Eleocharis interstincta (Vahl) Roem. & Schult. 

Eleocharis mitrata (Griseb.) C.B. Clarke 

Eleocharis mutata (L.) Roem. & Schult. 

Eleocharis plicarhachis (Griseb.) Svenson 

Eleocharis subfoliata C.B. Clarke 

Fimbristylis cymosa R. Br. * 

Fimbristylis dichotoma (L.) Vahl 

Fimbristylis ferruginea (L.) Vahl 

Fimbristylis miliacea (L.) Vahl 

Fuirena umbellata Rottb. 

Hypolytrum longifolium (Rich.) Nees 

Kyllinga brevifolia Rottb. 

Lagenocarpus guianensis Lindl. & Nees ex 

Nees 

Rhynchospora cephalotes (L.) Vahl 

Rhynchospora ciliata (Vahl) Kük. 

Rhynchospora corymbosa (L.) Britton 

Rhynchospora hassleri C.B. Clarke 

Rhynchospora holoschoenoides (Rich.) Herter 

Rhynchospora pubera (Vahl) Boeck. 

Scleria latifolia Sw. 

Scleria macrophylla J. Presl & C. Presl 

Scleria melaleuca Rchb. ex Schltdl. & Cham. 

Scleria microcarpa Nees ex Kunth 

Scleria secans (L.) Urb. 

Dioscoreaceae 

Dioscorea alata L. 

Dioscorea cayenensis Lam. 

Dioscorea esculenta (Lour.) Prain [cultivated] 

Dioscorea oblonga Gleason 

Dioscorea pilosiuscula Bertero ex Spreng. 

Dioscorea polygonoides Humb. & Bonpl. ex 

Willd. 

Dioscorea riparia Kunth & R.H. Schomb. 

Dioscorea samydea Griseb. 

Dioscorea trichanthera Gleason 

Dioscorea trifida L. f. 

Eriocaulaceae 

Paepalanthus bifidus (Schrad.) Kunth 

Syngonanthus longipes Gleason 

Syngonanthus umbellatus (Lam.) Ruhland 

Tonina fluviatilis Aubl. 

Haemodoraceae 

Xiphidium caeruleum Aubl. 

Heliconiaceae 

Heliconia acuminata Rich. 

Heliconia bihai (L.) L. 

Heliconia chartacea Lane ex Barreiros 

Heliconia hirsuta L. f. 

Heliconia psittacorum L. f. 

Heliconia richardiana Miq. 

Heliconia spathocircinata Aristeg. 

Musa x paradisiaca L. [cultivated] 

Hydrocharitaceae 

Limnobium laevigatum (Humb. & Bonpl. ex 

Willd.) Heine * 

Iridaceae 

Eleutherine bulbosa (Mill.) Urb. 

Lemnaceae 

Lemna aequinoctialis Welw. * 

Liliaceae 

Aloe vera (L.) Burm. f. [cultivated] 

Cordyline fructicosa (L.) A. Chev. [cultivated] 

Crinum erubescens L. f. ex Sol. 

Hippeastrum puniceum (Lam.) Kuntze 

Hymenocallis littoralis (Jacq.) Salisb. 

[naturalized] 

Hymenocallis tubiflora Salisb. 

Marantaceae 

Calathea cyclophora Baker 

Calathea elliptica (Roscoe) K. Schum. 

Calathea legrelleana (Linden) Regel 

Calathea micans (Mathieu) Körn. 

Calathea variegata Linden ex Körn. 

Ischnosiphon arouma (Aubl.) Körn. 

Ischnosiphon enigmaticus L. Andersson 

Ischnosiphon foliosus Gleason 

Ischnosiphon obliquus (Rudge) Körn. 

Ischnosiphon puberulus Loes. 

Maranta arundinacea L. 

Monotagma spicatum (Aubl.) J.F. Macbr. 

Orchidaceae 

Brassia neglecta Rchb. f. 

Brassia verrucosa Lindl. 

Catasetum barbatum (Lindl.) Lindl. 

Cyclopogon olivaceus (Rolfe) Schltr. 

Dichaea picta Rchb. f.


Dichaea rendlei Gleason 

Dimerandra elegans (H. Focke) Siegerist 

Encyclia diurna (Jacq.) Schltr. 

Encyclia granitica (Bateman ex Lindl.) Schltr. 

Epidendrum anceps Jacq. 

Epidendrum ciliare L. * 

Epidendrum flexuosum G. Mey. 

Epidendrum ibaguense Kunth 

Epidendrum longicolle Lindl. 

Epidendrum macrocarpum Rich. 

Epidendrum nocturnum Jacq. 

Epidendrum purpurascens H. Focke 

Epidendrum rigidum Jacq. 

Epidendrum strobiliferum Rchb. f. 

Erycina pusilla (L.) N.H. Williams & M.W. 

Chase 

Eulophia alta (L.) Fawc. & Rendle 

Habenaria longicauda Hook. 

Ionopsis utricularioides (Sw.) Lindl. 

Jacquiniella sp. 

Koellensteinia graminea (Lindl.) Rchb. f. 

Lockhartia imbricata (Lam.) Hoehne 

Maxillaria camaridii Rchb. f. 

Maxillaria parviflora (Poepp. & Endl.) Garay 

Maxillaria rufescens Lindl. 

Maxillaria uncata Lindl. 

Maxillaria villosa (Barb. Rodr.) Cogn. 

Oncidium baueri Lindl. 

Pleurothallis corniculata Lindl. 

Pleurothallis exigua Cogn. 

Pleurothallis glandulosa Ames 

Pleurothallis lanceana Lodd. 

Pleurothallis pruinosa Lindl. 

Pleurothallis sclerophylla Lindl. 

Pleurothallis yauaperyensis Barb. Rodr. 

Polystachya concreta (Jacq.) Garay & H.R. 

Sweet 

Prosthechea aemula (Lindl.) W.E. Higgins 

Prosthechea vespa (Vell.) W.E. Higgins 

Rodriguezia lanceolata Ruiz & Pav. 

Scaphyglottis graminifolia (Ruiz & Pav.) Poepp. 

& Endl. 

Scaphyglottis sickii Pabst 

Selenipedium palmifolium (Lindl.) Rchb. f. 

Sobralia sessilis Lindl. 

Stanhopea grandiflora (Lodd.) Lindl. 

Stelis argentata Lindl. 

Trichocentrum lanceanum (Lindl.) M.W. Chase 

& N.H. Williams * 

Trigonidium acuminatum Bateman ex Lindl. 

Vanilla fimbriata Rolfe 

Vanilla grandiflora Lindl. 

Vanilla latisegmenta Ames & C. Schweinf. 

Vanilla mexicana Mill. 

Wullschlaegelia calcarata Benth. 

Zygosepalum labiosum (Rich.) Garay 

153 

Poaceae 

Acroceras zizanioides (Kunth) Dandy 

[naturalized] 

Andropogon bicornis L. 

Andropogon leucostachyus Kunth 

Andropogon virgatus Desv. * 

Axonopus compressus (Sw.) P. Beauv. 

Bambusa vulgaris Schrad. ex J.C. Wendl. 

[cultivated] 

Coix lacryma-jobi L. [naturalized] 

Cymbopogon citratus (DC.) Stapf [naturalized] 

Digitaria horizontalis Willd. 

Echinochloa polystachya (Kunth) Hitchc. * 

Eleusine indica (L.) Gaertn. [naturalized] 

Eragrostis ciliaris (L.) R. Br. [naturalized] 

Eragrostis tephrosanthos Schult. 

Eragrostis unioloides (Retz.) Nees [naturalized] 

Gynerium saccharoides ? Bonpl. 

Gynerium sagittatum (Aubl.) P. Beauv. 

Homolepis isocalycia (G. Mey.) Chase 

Hymenachne amplexicaulis (Rudge) Nees 

Hymenachne donacifolia (Raddi) Chase 

Ichnanthus pallens (Sw.) Munro ex Benth. 

Ichnanthus panicoides P. Beauv. 

Ichnanthus ruprechtii Döll 

Imperata contracta (Kunth) Hitchc. 

Ischaemum timorense Kunth [naturalized] 

Lasiacis ligulata Hitchc. & Chase 

Leersia hexandra Sw. 

Leptochloa scabra Nees 

Leptochloa virgata (L.) P. Beauv. 

Olyra latifolia L. 

Olyra longifolia Kunth 

Orthoclada laxa (Rich.) P. Beauv. 

Oryza rufipogon Griff. [naturalized] 

Panicum elephantipes Nees ex Trin. 

Panicum laxum Sw. 

Panicum millegrana Poir. 

Panicum parvifolium Lam. 

Panicum pilosum Sw. 

Panicum polygonatum Schrad.

154 


Panicum rudgei Roem. & Schult. 

Panicum stoloniferum Poir. 

Panicum trichoides Sw. 

Paspalum conjugatum P.J. Bergius 

Paspalum distichum L. * 

Paspalum melanospermum Desv. ex Poir. 

Paspalum millegrana Schrad. 

Paspalum repens P.J. Bergius 

Pharus latifolius L. 

Piresia sympodica (Döll) Swallen 

Saccharum officinarum L. [cultivated] 

Sacciolepis striata (L.) Nash 

Setaria parviflora (Poir.) Kerguélen 

Setaria poiretiana (Schult.) Kunth 

Sporobolus jacquemontii Kunth 

Sporobolus virginicus (L.) Kunth * 

Vetiveria zizanioides (L.) Nash [cultivated] 

Zea mays L. [cultivated] 

Pontederiaceae 

Eichhornia azurea (Sw.) Kunth 

Eichhornia diversifolia (Vahl) Urb. 

Rapateaceae 

Rapatea linearis Gleason 

Rapatea paludosa Aubl. 

Smilacaceae 

Smilax cumanensis Humb. & Bonpl. ex Willd. 

Smilax latipes Gleason 

Smilax schomburgkiana Kunth 

Taccaceae 

Tacca parkeri Seem. 

Triuridaceae 

Sciaphila albescens Benth. 

Typhaceae 

Typha domingensis Pers. * 

Xyridaceae 

Xyris jupicai Rich. 

Xyris laxifolia Mart. 

Zingiberaceae 

Aframomum melegueta K. Schum. [cultivated] 

Curcuma zanthorrhiza Roxb. [cultivated] 

Hedychium coronarium J. Koenig [naturalized] 

Renealmia alpinia (Rottb.) Maas 

Renealmia guianensis Maas 

Renealmia orinocensis Rusby 

Zingiber officinale Roscoe [cultivated] 

Zingiber zerumbet (L.) Roscoe ex Sm. 

[cultivated]


APPENDIX 4. 

SPECIES DISPARITIES BETWEEN THE NORTHWEST 

DISTRICT, 

GUYANA AND DELTA AMACURO, VENEZUELA 

A: Taxa Found in the Northwest District not Recorded for Delta Amacuro 

Including all Pteridophytes, and Spermatophytes (seed plants) 

PTERIDOPHYTES 

Lycopodiaceae 

Lycopodiella cernua (L.) Pic. Serm. 


Selaginella parkeri (Hook. & Grev.) Spring 

Selaginella porelloides (Lam.) Spring 

Selaginella producta Baker 

Aspleniaceae 

Asplenium juglandifolium Lam. 

Dennstaedtiaceae 

Lindsaea lancea (L.) Bedd. 

Lindsaea portoricensis Desv. 

Saccoloma elegans Kaulf. 

Grammitidaceae 

Cochlidium serrulatum (Sw.) L.E. Bishop 

Hymenophyllaceae 

Trichomanes diversifrons (Bory) Mett. ex 

Sadeb. 

Trichomanes martiusii C. Presl 

Trichomanes polypodioides L. 

Trichomanes radicans Sw. 

Lomariopsidaceae 

Elaphoglossum flaccidum (Fée) T. Moore 

Elaphoglossum glabellum J. Sm. 

Metaxyaceae 

Metaxya rostrata (Kunth) C. Presl 

Polypodiaceae 

Dicranoglossum desvauxii (Klotzsch) Proctor 

Microgramma fuscopunctata (Hook.) Vareschi 

Phlebodium pseudoaureum (Cav.) Lellinger 

155 

Polypodium adnatum Kunze ex Klotzsch 

Polypodium attenuatum Humb. & Bonpl. ex 

Willd. 

Pteridaceae 

Pteris altissima Poir. 

Schizaeaceae 

Schizaea fluminensis Miers ex J.W. Sturm 

Schizaea incurvata Schkuhr 

Tectariaceae 

Triplophyllum funestum (Kunze) Holttum 


Thelypteris interrupta (Willd.) K. Iwats. 

Thelypteris leprieurii (Hook.) R.M. Tryon 

Woodsiaceae 

Diplazium celtidifolium Kunze 

GYMNOSPERMS 

Gnetaceae 

Gnetum nodiflorum Brongn. 

ANGIOSPERMS 

Acanthaceae 

Justicia pectoralis Jacq. 

Agavaceae 

Furcraea foetida (L.) Haw. 

Alismataceae 

Sagittaria lancifolia L. 

Amaranthaceae 

Alternanthera tenella Colla

156 


Anacardiaceae 

Anacardium giganteum W. Hancock ex Engl. 

Anacardium occidentale L. 

Astronium lecointei Ducke 

Thyrsodium guianense Sagot ex Marchand 

Annonaceae 

Annona sericea Dunal 

Annona symphyocarpa Sandwith 

Bocageopsis multiflora (Mart.) R.E. Fr. 

Duguetia calycina Benoist 

Fusaea longifolia (Aubl.) Saff. 

Guatteria schomburgkiana Mart. 

Rollinia mucosa (Jacq.) Baill. 

Xylopia benthamii R.E. Fr. 

Xylopia cayennensis Maas 

Xylopia surinamensis R.E. Fr. 

Apocynaceae 

Ambelania acida Aubl. 

Himatanthus bracteatus (A. DC.) Woodson 

Macoubea guianensis Aubl. 

Odontadenia geminata (Hoffmanns. ex Roem. 

& Schult.) Müll. Arg. 

Odontadenia puncticulosa (Rich.) Pulle 

Odontadenia sandwithiana Woodson 

Plumeria inodora Jacq. 

Tabernaemontana disticha A. DC. 

Tabernaemontana lorifera (Miers) Leeuwenb. 

Araceae 

Caladium bicolor (Aiton) Vent. 

Caladium schomburgkii Schott 

Dieffenbachia humilis Poepp. 

Dieffenbachia paludicola N.E. Br. ex Gleason 

Heteropsis flexuosa (Kunth) G.S. Bunting 

Philodendron brevispathum Schott 

Philodendron fragrantissimum (Hook.) G. Don 

Philodendron hederaceum (Jacq.) Schott 

Philodendron surinamense (Miq.) Engl. 

Rhodospatha oblongata Poepp. 

Spathiphyllum cuspidatum Schott 

Spathiphyllum maguirei G.S. Bunting 

Stenospermation maguirei A.M.E. Jonker & 

Jonker 

Araliaceae 

Schefflera decaphylla (Seem.) Harms 

Arecaceae 

Astrocaryum aculeatum G. Mey. 

Astrocaryum vulgare Mart. 

Bactris acanthocarpa Mart. 

Bactris campestris Poepp. ex Mart. 

Bactris oligoclada Burret 

Bactris simplicifrons Mart. 

Geonoma baculifera (Poit.) Kunth 

Geonoma macrostachys Mart. 


Aristolochia daemoninoxia Mast. 

Aristolochia hians Willd. 


Blepharodon nitidus (Vell.) J.F. Macbr. 

Matelea badilloi Morillo 

Matelea stenopetala Sandwith 

Stenomeria decalepis Turcz. 

Asteraceae 

Clibadium surinamense L. 

Clibadium sylvestre (Aubl.) Baill. 

Conyza bonariensis (L.) Cronquist 

Cyrtocymura scorpioides (Lam.) H. Rob. 

Elephantopus mollis Kunth 

Erechtites hieracifolia (L.) Raf. ex DC. 

Mikania banisteriae DC. 

Mikania cordifolia (L. f.) Willd. 

Mikania hookeriana DC. 

Mikania parviflora (Aubl.) H. Karst. 

Mikania psilostachya DC. 

Sonchus asper (L.) Hill 

Sphagneticola trilobata (L.) Pruski 

Unxia camphorata L. f. 

Begoniaceae 

Begonia humilis Dryand. 

Bignoniaceae 

Anemopaegma oligoneuron (Sprague & 

Sandwith) A.H. Gentry 

Arrabidaea candicans (Rich.) DC. 

Ceratophytum tetragonolobum (Jacq.) Sprague 

& Sandwith 

Crescentia amazonica Ducke 

Distictella elongata (Vahl) Urb. 

Lundia densiflora DC. 

Martinella obovata (Kunth) Bureau & K. 

Schum. 

Pleonotoma echitidea Sprague & Sandwith 

Schlegelia spruceana K. Schum. 

Tabebuia serratifolia (Vahl) G. Nicholson

Bixaceae 

Bixa orellana L. 

Bombacaceae 

Catostemma fragrans Benth. 

Pachira insignis (Sw.) Sw. ex Savigny 


Cordia schomburgkii DC. 


Bromeliaceae 

Aechmea angustifolia Poepp. & Endl. 

Disteganthus lateralis (L.B. Sm.) Gouda 

Vriesea heliconioides (Kunth) Hook. ex Walp. 

Burmanniaceae 

Burmannia tenella Benth. 

Burseraceae 

Protium unifoliolatum Spruce ex Engl. 

Trattinnickia boliviana (Swart) Daly 

Trattinnickia burserifolia Mart. 


Brownea grandiceps Jacq. 

Chamaecrista ramosa (Vogel) H.S. Irwin & 

Barneby 

Dicorynia guianensis Amshoff 

Eperua falcata Aubl. 

Eperua rubiginosa Miq. 

Hymenaea courbaril L. 

Macrolobium angustifolium (Benth.) R.S. 

Cowan 

Peltogyne venosa (Vahl) Benth. 

Senna alata (L.) Roxb. 

Senna bacillaris (L. f.) H.S. Irwin & Barneby 

Senna quinquangulata (Rich.) H.S. Irwin & 

Barneby 

Senna reticulata (Willd.) H.S. Irwin & Barneby 

Senna sandwithiana H.S. Irwin & Barneby 

Tachigali micropetala (Ducke) Zarucchi & 

Pipoly 

Capparaceae 

Cleome serrata Jacq. 

Cleome speciosa Raf. 

Cecropiaceae 

Cecropia obtusa Trécul 

Coussapoa microcephala Trécul 


Couepia parillo DC. 

Licania boyanii Tutin 

Licania incana Aubl. 

Licania kunthiana Hook. f. 

Licania laxiflora Fritsch 

Licania majuscula Sagot 

Licania membranacea Sagot ex Laness. 

Licania micrantha Miq. 

Licania persaudii Fanshawe & Maguire 

157 

Clusiaceae 

Clusia cuneata Benth. 

Clusia gaudichaudii Choisy 

Clusia myriandra (Benth.) Planch. & Triana 

Rheedia virens Planch. & Triana 

Tovomita calodictyos Sandwith 

Tovomita choisyana Planch. & Triana 

Tovomita obscura Sandwith 

Tovomita schomburgkii Planch. & Triana 

Vismia gracilis Hieron. 

Vismia japurensis Reichardt 

Vismia sessilifolia (Aubl.) Choisy 

Combretaceae 

Buchenavia grandis Ducke 

Commelinaceae 

Commelina erecta L. 

Gibasis geniculata (Jacq.) Rohweder 

Connaraceae 

Connarus coriaceus G. Schellenb. 

Pseudoconnarus macrophyllus (Poepp.) Radlk. 


Ipomoea asarifolia (Desr.) Roem. & Schult. 

Ipomoea quamoclit L. 

Costaceae 

Costus erythrothyrsus Loes. 

Costus guanaiensis Rusby 

Costus spiralis (Jacq.) Roscoe 

Cucurbitaceae 

Cayaponia jenmanii C. Jeffrey 

Helmontia leptantha (Schltdl.) Cogn. 

Cyclanthaceae 

Asplundia brachyphylla Harling

158 

Asplundia glandulosa (Gleason) Harling 

Asplundia gleasonii Harling 

Asplundia guianensis Harling 


Cyperaceae 

Calyptrocarya bicolor (H. Pfeiff.) T. Koyama 

Cyperus comosus Poir. 

Cyperus simplex Kunth 

Diplasia karatifolia Rich. 

Eleocharis interstincta (Vahl) Roem. & Schult. 

Eleocharis mitrata (Griseb.) C.B. Clarke 

Eleocharis plicarhachis (Griseb.) Svenson 

Eleocharis subfoliata C.B. Clarke 

Fimbristylis ferruginea (L.) Vahl 

Kyllinga brevifolia Rottb. 

Rhynchospora cephalotes (L.) Vahl 

Rhynchospora ciliata (Vahl) Kük. 

Rhynchospora hassleri C.B. Clarke 

Rhynchospora holoschoenoides (Rich.) Herter 

Rhynchospora pubera (Vahl) Boeck. 

Scleria macrophylla J. Presl & C. Presl 

Cyrillaceae 

Cyrilla racemiflora L. 

Dichapetalaceae 

Dichapetalum pedunculatum (DC.) Baill. 

Dilleniaceae 

Davilla nitida (Vahl) Kubitzki 

Tetracera asperula Miq. 

Dioscoreaceae 

Dioscorea cayenensis Lam. 

Dioscorea oblonga Gleason 

Dioscorea pilosiuscula Bertero ex Spreng. 

Dioscorea riparia Kunth & R.H. Schomb. 

Dioscorea samydea Griseb. 

Dioscorea trichanthera Gleason 

Dioscorea trifida L. f. 

Droseraceae 

Drosera intermedia Hayne 

Ebenaceae 

Diospyros tetrandra Hiern 


Sloanea eichleri K. Schum. 

Sloanea latifolia (Rich.) K. Schum. 

Eriocaulaceae 

Paepalanthus bifidus (Schrad.) Kunth 

Syngonanthus longipes Gleason 

Syngonanthus umbellatus (Lam.) Ruhland 

Erythroxylaceae 

Erythroxylum citrifolium A. St.-Hil. 

Erythroxylum macrophyllum Cav. 

Euphorbiaceae 

Alchornea discolor Poepp. 

Alchornea triplinervia (Spreng.) Müll. Arg. 

Alchorneopsis floribunda (Benth.) Müll. Arg. 

Croton cuneatus Klotzsch 

Euphorbia cotinifolia L. 

Euphorbia oerstediana (Klotzsch & Garcke) 

Boiss. 

Euphorbia thymifolia L. 

Microstachys corniculata (Vahl) Griseb. 

Phyllanthus brasiliensis (Aubl.) Poir. 

Fabaceae 

Alexa confusa Pittier 

Alexa surinamensis Yakovlev 

Centrosema capitatum (Rich.) Amshoff 

Crotalaria nitens Kunth 

Crotalaria stipularia Desv. 

Derris amazonica Killip 

Derris pterocarpus (DC.) Killip 

Desmodium axillare (Sw.) DC. 

Dioclea scabra (Rich.) R.H. Maxwell 

Dioclea wilsonii Standl. 

Hymenolobium flavum Kleinhoonte 

Lonchocarpus chrysophyllus Kleinhoonte 

Lonchocarpus martynii A.C. Sm. 

Lonchocarpus rufescens Benth. 

Lonchocarpus spruceanus Benth. 

Machaerium leiophyllum (DC.) Benth. 

Machaerium quinatum (Aubl.) Sandwith 

Ormosia coccinea (Aubl.) Jacks. 

Ormosia coutinhoi Ducke 

Ormosia nobilis Tul. 

Platymiscium pinnatum (Jacq.) Dugand 

Swartzia conferta Spruce ex Benth. 

Swartzia guianensis (Aubl.) Urb. 

Swartzia schomburgkii Benth. 

Swartzia steyermarkii R.S. Cowan 

Tephrosia sinapou (Buc’hoz) A. Chev. 

Zornia diphylla (L.) Pers.



Banara guianensis Aubl. 

Casearia acuminata DC. 

Casearia javitensis Kunth 

Ryania speciosa Vahl 

Xylosma benthamii (Tul.) Triana & Planch. 

Gentianaceae 

Chelonanthus purpurascens (Aubl.) Struwe, S. 

Nilsson & V.A. Albert 

Gesneriaceae 

Besleria flavovirens Nees & Mart. 

Chrysothemis villosa (Benth.) Leeuwenb. 

Codonanthe calcarata (Miq.) Hanst. 

Nautilocalyx coccineus Feuillet & L.E. Skog 

Nautilocalyx cordatus (Gleason) L.E. Skog 

Nautilocalyx mimuloides (Benth.) C.V. Morton 

Paradrymonia maculata (Hook. f.) Wiehler 

Heliconiaceae 

Heliconia acuminata Rich. 

Heliconia chartacea Lane ex Barreiros 


Peritassa pruinosa (Seem.) A.C. Sm. 

Tontelea glabra A.C. Sm. 

Humiriaceae 

Humiria balsamifera Aubl. 

Humiriastrum obovatum (Benth.) Cuatrec. 

Icacinaceae 

Discophora guianensis Miers 

Emmotum fagifolium Ham. 

Poraqueiba guianensis Aubl. 

Iridaceae 

Eleutherine bulbosa (Mill.) Urb. 

Lamiaceae 

Hyptis parkeri Benth. 

Lauraceae 

Aiouea guianensis Aubl. 

Aniba guianensis Aubl. 

Aniba hostmanniana (Nees) Mez 

Aniba hypoglauca Sandwith 

Aniba jenmanii Mez 

Aniba terminalis Ducke 

Licaria oppositifolia (Nees) Kosterm. 

Nectandra amazonum Nees 

Ocotea puberula (Rich.) Nees 

Ocotea splendens (Meisn.) Baill. 

Persea nivea Mez 

159 

Lecythidaceae 

Eschweilera alata A.C. Sm. 

Eschweilera micrantha (O. Berg) Miers 

Eschweilera sagotiana Miers 

Eschweilera wachenheimii (Benoist) Sandwith 

Lentibulariaceae 

Utricularia benjaminiana Oliv. 

Utricularia myriocista A. St.-Hil. & Girard 

Liliaceae 

Hippeastrum puniceum (Lam.) Kuntze 

Loganiaceae 

Strychnos erichsonii M.R. Schomb. ex Progel 

Strychnos mitscherlichii M.R. Schomb. 

Malpighiaceae 

Banisteriopsis caapi (Griseb.) C.V. Morton 

Burdachia sphaerocarpa A. Juss. 

Diplopterys pauciflora (G. Mey.) Nied. 

Heteropterys macrostachya A. Juss. 

Hiraea fagifolia (DC.) A. Juss. 

Mascagnia macrodisca (Triana & Planch.) Nied. 

Mezia includens (Benth.) Cuatrec. 

Stigmaphyllon convolvulifolium A. Juss. 

Tetrapterys fimbripetala A. Juss. 

Malvaceae 

Sida glomerata Cav. 

Urena lobata L. 

Marantaceae 

Calathea legrelleana (Linden) Regel 

Calathea variegata Linden ex Körn. 

Ischnosiphon enigmaticus L. Andersson 

Ischnosiphon foliosus Gleason 

Maranta arundinacea L. 

Marcgraviaceae 

Marcgravia magnibracteata Lanj. & Heerdt 

Marcgravia purpurea I.W. Bailey 


Aciotis annua (Mart. ex DC.) Triana 

Aciotis indecora (Bonpl.) Triana

160 

Bellucia grossularioides (L.) Triana 

Clidemia capitellata (Bonpl.) D. Don 

Clidemia dentata D. Don 

Clidemia microthyrsa R.O. Williams 

Clidemia rubra (Aubl.) Mart. 

Clidemia venosa (Gleason) Wurdack 

Comolia villosa (Aubl.) Triana 

Desmoscelis villosa (Aubl.) Naudin 

Henriettea succosa (Aubl.) DC. 

Henriettella caudata Gleason 

Loreya mespiloides Miq. 

Miconia acinodendron (L.) Sweet 

Miconia bubalina (D. Don) Naudin 

Miconia ciliata (Rich.) DC. 

Miconia egensis Cogn. 

Miconia gratissima Benth. ex Triana 

Miconia hypoleuca (Benth.) Triana 

Miconia ibaguensis (Bonpl.) Triana 

Miconia lepidota DC. 

Miconia matthaei Naudin 

Miconia minutiflora (Bonpl.) DC. 

Miconia rubiginosa (Bonpl.) DC. 

Myriaspora egensis DC. 

Pterolepis glomerata (Rottb.) Miq. 

Tococa aristata Benth. 


Mendonciaceae 

Mendoncia bivalvis (L. f.) Merr. 

Mendoncia glabra (Poepp. & Endl.) Nees 


Curarea candicans (Rich. ex DC.) Barneby & 

Krukoff 

Menyanthaceae 

Nymphoides indica (L.) Kuntze 

Mimosaceae 

Abarema jupunba (Willd.) Britton & Killip 

Abarema laeta (Benth.) Barneby & J.W. Grimes 

Abarema mataybifolia (Sandwith) Barneby & 

J.W. Grimes 

Calliandra surinamensis Benth. 

Hydrochorea corymbosa (Rich.) Barneby & 

J.W. Grimes 

Hydrochorea gonggrijpii (Kleinhoonte) 

Barneby & J.W. Grimes 

Inga acreana Harms 

Inga acrocephala Steud. 

Inga graciliflora Benth. 

Inga java Pittier 

Inga jenmanii Sandwith 

Inga marginata Willd. 

Inga pilosula (Rich.) J.F. Macbr. 

Inga sarmentosa Glaz. ex Harms 

Inga sertulifera DC. 

Inga thibaudiana DC. 

Macrosamanea pubiramea (Steud.) Barneby & 

J.W. Grimes 

Moraceae 

Bagassa guianensis Aubl. 

Ficus mathewsii (Miq.) Miq. 

Ficus roraimensis C.C. Berg 

Myristicaceae 

Iryanthera juruensis Warb. 

Iryanthera macrophylla (Benth.) Warb. 

Virola calophylla (Spruce) Warb. 

Myrsinaceae 

Stylogyne orinocensis (Kunth) Mez 

Myrtaceae 

Eugenia cucullata Amshoff 

Eugenia punicifolia (Kunth) DC. 

Myrcia graciliflora Sagot 

Myrcia sylvatica (G. Mey.) DC. 


Guapira salicifolia (Heimerl) Lundell 

Neea constricta Spruce ex J.A. Schmidt 

Neea floribunda Poepp. & Endl. 

Nymphaeaceae 

Nymphaea odorata Aiton 

Ochnaceae 

Ouratea candollei (Planch.) Tiegh. 

Ouratea macrocarpa Sastre 

Ouratea rorida Sastre 

Sauvagesia sprengelii A. St.-Hil. 

Onagraceae 

Ludwigia nervosa (Poir.) H. Hara 

Orchidaceae 

Cyclopogon olivaceus (Rolfe) Schltr. 

Dichaea rendlei Gleason 

Encyclia diurna (Jacq.) Schltr. 

Epidendrum anceps Jacq. 

Epidendrum ibaguense Kunth 

Epidendrum macrocarpum Rich.


Epidendrum purpurascens H. Focke 

Maxillaria parviflora (Poepp. & Endl.) Garay 

Maxillaria villosa (Barb. Rodr.) Cogn. 

Pleurothallis exigua Cogn. 

Pleurothallis lanceana Lodd. 

Pleurothallis sclerophylla Lindl. 

Pleurothallis yauaperyensis Barb. Rodr. 

Selenipedium palmifolium (Lindl.) Rchb. f. 

Vanilla fimbriata Rolfe 

Vanilla grandiflora Lindl. 

Vanilla latisegmenta Ames & C. Schweinf. 

Wullschlaegelia calcarata Benth. 

Oxalidaceae 

Oxalis barrelieri L. 

Oxalis debilis Kunth 


Passiflora amicorum Wurdack 

Passiflora cirrhiflora Juss. 

Passiflora garckei Mast. 

Passiflora glandulosa Cav. 

Passiflora nitida Kunth 

Piperaceae 

Peperomia duidana Trel. ex Gleason 

Peperomia pernambucensis Miq. 

Peperomia rotundifolia (L.) Kunth 

Piper aduncum L. 

Piper avellanum (Miq.) C. DC. 

Piper glabrescens (Miq.) C. DC. 

Piper nigrispicum C. DC. 

Piper pulleanum Yunck. 

Poaceae 

Andropogon leucostachyus Kunth 

Eragrostis tephrosanthos Schult. 

Gynerium saccharoides ? Bonpl. 

Homolepis isocalycia (G. Mey.) Chase 

Ichnanthus ruprechtii Döll 

Imperata contracta (Kunth) Hitchc. 

Panicum laxum Sw. 

Panicum millegrana Poir. 

Panicum polygonatum Schrad. 

Panicum rudgei Roem. & Schult. 

Panicum trichoides Sw. 

Paspalum distichum L. 

Paspalum repens P.J. Bergius 

Piresia sympodica (Döll) Swallen 

Sporobolus jacquemontii Kunth 

Sporobolus virginicus (L.) Kunth 

Polygalaceae 

Bredemeyera densiflora A.W. Benn. 

Moutabea guianensis Aubl. 

Polygonaceae 

Coccoloba densifrons Mart. ex Meisn. 

Coccoloba parimensis Benth. 

Rapateaceae 

Rapatea linearis Gleason 

Rhamnaceae 

Gouania lupuloides (L.) Urb. s.l. 


Cassipourea lasiocalyx Alston 

161 

Rubiaceae 

Amaioua corymbosa Kunth 

Borreria assurgens (Ruiz & Pav.) Griseb. 

Borreria capitata (Ruiz & Pav.) DC. 

Borreria densiflora DC. 

Coccocypselum tontanea Kunth 

Coussarea leptoloba (Spreng. ex Benth. & 

Hook. f.) Müll. Arg. 

Coussarea violacea Aubl. 

Faramea guianensis (Aubl.) Bremek. 

Faramea multiflora A. Rich. ex DC. 

Genipa spruceana Steyerm. 

Geophila tenuis (Müll. Arg.) Standl. 

Gonzalagunia bunchosioides Standl. 

Gonzalagunia cornifolia (Kunth) Standl. 

Gonzalagunia dicocca Cham. & Schltdl. 

Gonzalagunia spicata (Lamb.) M. Gómez 

Ixora schomburgkiana Benth. 

Malanea hypoleuca Steyerm. 

Palicourea guianensis Aubl. 

Palicourea triphylla DC. 

Posoqueria coriacea M. Martens & Galeotti 

Posoqueria trinitatis DC. 

Psychotria anceps Kunth 

Psychotria bahiensis DC. 

Psychotria barbiflora DC. 

Psychotria callithrix (Miq.) Steyerm. 

Psychotria capitata Ruiz & Pav. 

Psychotria erecta (Aubl.) Standl. & Steyerm. 

Psychotria horizontalis Sw. 

Psychotria mapourioides DC. 

Psychotria platypoda DC. 

Psychotria tillettii Steyerm. 

Psychotria ulviformis Steyerm.

162 

Psychotria wessels-boeri Steyerm. 

Rudgea standleyana Steyerm. 

Rudgea stipulacea (DC.) Steyerm. 

Sabicea aspera Aubl. 

Sabicea velutina Benth. 

Schradera polycephala DC. 

Rutaceae 

Ertela trifolia (L.) Kuntze 

Zanthoxylum rhoifolium Lam. 


Allophylus racemosus Sw. 

Cupania scrobiculata Rich. 

Matayba camptoneura Radlk. 

Matayba opaca Radlk. 

Paullinia caloptera Radlk. 

Paullinia hitchcockii Gleason 

Paullinia xestophylla Radlk. 

Pseudima frutescens (Aubl.) Radlk. 

Talisia guianensis Aubl. 


Sapotaceae 

Chrysophyllum pomiferum (Eyma) T.D. Penn. 

Chrysophyllum sanguinolentum (Pierre) Baehni 

Pouteria bilocularis (H. Winkl.) Baehni 

Pouteria coriacea (Pierre) Pierre 

Pouteria durlandii (Standl.) Baehni 

Pouteria guianensis Aubl. 

Pradosia schomburgkiana (A. DC.) Cronquist 


Achetaria guianensis Pennell 

Angelonia biflora Benth. 

Lindernia diffusa (L.) Wettst. 

Simaroubaceae 

Picramnia guianensis (Aubl.) Jans.-Jac. 

Smilacaceae 

Smilax latipes Gleason 

Solanaceae 

Markea camponoti Ducke (possibly =M. 

longiflora) 

Markea longiflora Miers 

Solanum leucocarpon Dunal 

Solanum rugosum Dunal 

Solanum schlechtendalianum Walp. 

Sterculiaceae 

Herrania kanukuensis R.E. Schult. 

Sterculia rugosa R. Br. 

Taccaceae 

Tacca parkeri Seem. 

Theophrastaceae 

Clavija lancifolia Desf. 

Tiliaceae 

Corchorus aestuans L. 

Triumfetta althaeoides Lam. 

Triuridaceae 

Sciaphila albescens Benth. 

Turneraceae 

Turnera scabra Millsp. 

Turnera ulmifolia L. 

Urticaceae 

Pilea pubescens Liebm. 

Urera baccifera (L.) Gaudich. ex Wedd. 

Verbenaceae 

Aegiphila racemosa Vell. 

Vitex compressa Turcz. 

Vitex triflora Vahl 

Violaceae 

Paypayrola grandiflora Tul. 

Rinorea flavescens (Aubl.) Kuntze 

Viscaceae 

Phoradendron bathyoryctum Eichler 

Phoradendron perrottetii (DC.) Eichler 

Vochysiaceae 

Vochysia guianensis Aubl.


B: Taxa Found in Delta Amacuro but not Recorded for Guyana 

Including all Pteridophytes, and Spermatophytes (seed plants) 

LYCOPHYTES 


Selaginella cladorrhizans A. Braun 

PTERIDOPHYTES 

Adiantaceae 

Cheilanthes bonariensis (Willd.) Proctor 

Pteridaceae 

Pteris tripartita Sw. 


Thelypteris angustifolia (Willd.) Proctor 

SPERMATOPHYTES 

Acanthaceae 

Barleria cristata L. 

Barleria lupulina Lindl. 

Justicia laevilinguis (Nees) Lindau 

Justicia moritziana Wassh. 

Sanchezia pennellii Leonard 

Thunbergia erecta (Benth.) T. Anderson 

Alismataceae 

Echinodorus horizontalis Rataj 

Sagittaria sprucei Micheli 

Amaranthaceae 

Celosia virgata Jacq. 

Anacardiaceae 

Astronium graveolens Jacq. 

Aquifoliaceae 

Ilex parvifructa Edwin 

Araceae 

Anthurium digitatum (Jacq.) G. Don 

Philodendron delascioi G.S. Bunting 

Philodendron fendleri K. Krause 

Xanthosoma sagittifolium (L.) Schott 


Aristolochia sprucei Mast. 


Cynanchum strictum R.W. Holm 

163 

Asteraceae 

Ambrosia peruviana Willd. 

Ayapana trinitensis (Kuntze) R.M. King & H. 

Rob. 

Oyedaea alba Pruski 

Pseudogynoxys chenopodioides (Kunth) 

Cabrera 

Spilanthes urens Jacq. 

Tessaria integrifolia Ruiz & Pav. 

Bignoniaceae 

Arrabidaea chica (Bonpl.) B. Verl. 

Memora patula Miers 


Cordia dentata Poir. 

Cordia panamensis Riley 

Bromeliaceae 

Hohenbergia stellata Schult. f. 

Tillandsia balbisiana Schult. f. 

Tillandsia elongata Kunth 

Tillandsia gardneri Lindl. 

Tillandsia polystachia (L.) L. 


Campsiandra implexicaulis Stergios 

Chamaecrista pilosa (L.) Greene 

Crudia aequalis Ducke 

Senna spectabilis (DC.) H.S. Irwin & Barneby 

Senna spinescens (Vogel) H.S. Irwin & Barneby 

Capparaceae 

Capparis osmantha Diels 

Caprifoliaceae 

Sambucus canadensis L. 

Caryophyllaceae 

Polycarpon apurense Kunth

164 

Celastraceae 

Elaeodendron xylocarpum (Vent.) DC. 

Zinowiewia aymardii Steyerm. 


Licania latistipula Prance 

Licania leucosepala Griseb. 

Licania pyrifolia Griseb. 

Clusiaceae 

Clusia candelabrum Planch. & Triana 

Clusia comans (Meisn.) Pipoly 

Combretaceae 

Combretum spinosum Bonpl. 

Connaraceae 

Connarus venezuelanus Baill. 


Ipomoea fimbriosepala Choisy 

Maripa repens Rusby 

Cucurbitaceae 

Cayaponia metensis Cuatrec. 

Fevillea cordifolia L. 


Cyclanthaceae 

Asplundia moritziana (Klotzsch) Harling 

Cyperaceae 

Bulbostylis svensoniana Steyerm. 

Cyperus cornelii-ostenii Kük. 

Cyperus felipponei Kük. 

Cyperus gardneri Nees 

Cyperus meyenianus Kunth 


Sloanea megacarpa Steyerm. & Marc.-Berti 

Sloanea obtusifolia (Moric.) K. Schum. 

Sloanea subpsilocarpa Steyerm. 

Euphorbiaceae 

Alchornea castaneifolia (Willd.) A. Juss. 

Alchornea grandiflora Müll. Arg. 

Croton bolivarensis Croizat 

Dalechampia brownsbergensis G.L. Webster & 

Armbr. 

Phyllanthus fluitans Benth. ex Müll. Arg. 

Piranhea longepedunculata Jabl. 

Plukenetia penninervia Müll. Arg. 

Fabaceae 

Cymbosema roseum Benth. 

Dalbergia hygrophila (Mart. ex Benth.) Hoehne 

Dalbergia subcymosa Ducke 

Derris negrensis Benth. 

Desmodium affine Schltdl. 

Desmodium tortuosum (Sw.) DC. 

Lonchocarpus fendleri Benth. 

Lonchocarpus imatacensis Poppend. 

Lonchocarpus sericeus (Poir.) Kunth ex DC. 

Lonchocarpus tubicalyx Pittier ex Poppend. 

Mucuna rostrata Benth. 

Vatairea paraensis Ducke 

Vigna longifolia (Benth.) Verdc. 


Banara nitida Spruce ex Benth. 

Banara orinocensis (Cuatrec.) Sleumer 


Pristimera tenuiflora (Mart. ex Peyr.) A.C. Sm. 

Lamiaceae 

Hyptis brevipes Poit. 

Lauraceae 

Cinnamomum triplinerve (Ruiz & Pav.) 

Kosterm. 

Nectandra pichurim (Kunth) Mez 

Lemnaceae 

Spirodela intermedia W. Koch 

Loganiaceae 

Strychnos mattogrossensis S. Moore 

Loranthaceae 

Psittacanthus acinarius (Mart.) Mart. 

Lythraceae 

Cuphea elliptica Koehne 

Rotala ramosior (L.) Koehne 

Malpighiaceae 

Banisteriopsis lyrata B. Gates 

Bunchosia armeniaca (Cav.) DC. 

Clonodia complicata (Kunth) W.R. Anderson 

Malpighia emarginata DC. 

Mascagnia divaricata (Kunth) Nied. 

Stigmaphyllon adenodon A. Juss.


Malvaceae 

Abelmoschus moschatus Medik. 

Cienfuegosia heterophylla (Vent.) Garcke 

Urena sinuata L. 

Wissadula hernandioides (L’Hér.) Garcke 

Marantaceae 

Calathea inocephala (Kuntze) H. Kenn. & 

Nicolson 

Ctenanthe compressa (A. Dietr.) Eichler 


Adelobotrys spruceana Cogn. 

Clidemia acurensis Wurdack 

Miconia borjensis Wurdack 


Borismene japurensis (Mart.) Barneby 

Mimosaceae 

Enterolobium barinense L. Cardenas & H. Rodr. 

Inga sapindoides Willd. 

Macrosamanea spruceana (Benth.) Killip 

Mimosa casta L. 

Mimosa orinocoënsis Barneby 

Moraceae 

Clarisia racemosa Ruiz & Pav. 

Ficus dendrocida Kunth 

Myrtaceae 

Calycorectes enormis McVaugh 

Eugenia baileyi Britton 

Syzygium malaccense (L.) Merr. & Perry 


Guapira ferruginea (Klotsch ex Choisy) 

Lundell 

Guapira marcano-bertii Steyerm. 

Guapira rusbyana (Heimerl ex Standl.) Lundell 

Neea davidsei Steyerm. 

Olacaceae 

Heisteria acuminata (Bonpl.) Engl. 

Onagraceae 

Ludwigia densiflora (Micheli) H. Hara 

Orchidaceae 

Campylocentrum lansbergii (Rchb. f.) Schltr. 

Chelyorchis ampliata (Lindl.) Dressler & N.H. 

165 

Williams 

Dichaea panamensis Lindl. 

Dichaea robusta Schltr. 

Epidendrum congestoides Ames & C. Schweinf. 

Lockhartia acuta (Lindl.) Rchb. f. 

Macradenia rubescens Barb. Rodr. 

Mormodes carnevaliana Salazar & G.A. 

Romero 

Stelis santiagoensis Mansf. 


Dilkea acuminata Mast. 

Passiflora adenopoda DC. 

Passiflora biflora Lam. 

Passiflora cuneata Willd. 

Passiflora cyanea Mast. 

Passiflora suberosa L. 

Passiflora subpeltata Ortega 

Passiflora variolata Poepp. & Endl. 

Piperaceae 

Piper nigrum L. 

Piper tenue Kunth 

Poaceae 

Brachiaria plantaginea (Link) Hitchc. 

Guadua venezuelae Munro 

Paspalum fasciculatum Willd. ex Flüggé 

Polygalaceae 

Moutabea sp. A 

Securidaca bialata Benth. 

Polygonaceae 

Coccoloba declinata (Vell.) Mart. 

Coccoloba fallax Lindau 

Coccoloba spruceana Lindau 

Polygonum ferrugineum Wedd. 

Pontederiaceae 

Heteranthera multiflora (Griseb.) C.N. Horn 

Portulacaceae 

Portulaca pusilla Kunth 

Portulaca teretifolia Kunth 

Rubiaceae 

Chiococca pubescens Humb. & Bonpl. ex 

Roem. & Schult. 

Diodia multiflora DC. 

Psychotria occidentalis Steyerm.

166 


Rutaceae 

Angostura trifoliata (Willd.) T.S. Elias 

Conchocarpus longifolius (A. St.-Hil.) Kallunki 

& Pirani 

Neoraputia paraensis (Ducke) Emmerich 

Zanthoxylum amapaense (Albuq.) P.G. 

Waterman 

Zanthoxylum juniperinum Poepp. 

Zanthoxylum martinicense (Lam.) DC. 


Houssayanthus macrolophus (Radlk.) Hunz. 

Sapotaceae 

Diploon cuspidatum (Hoehne) Cronquist 

Pouteria reticulata (Engl.) Eyma subsp. 

reticulata 

Pradosia grisebachii (Pierre) T.D. Penn. 


Bacopa innominata (M. Gómez) Alain 

Mecardonia procumbens (Mill.) Small 

Simaroubaceae 

Simaba sp. A (sensu Steyermark et al.) 

Solanaceae 

Physalis cordata Mill. 

Schwenckia micrantha Benth. 

Solanum arboreum Dunal 

Solanum lanceifolium Jacq. 

Solanum seaforthianum Andrews 

Sterculiaceae 

Melochia manducata C. Wright 

Melochia villosa (Mill.) Fawc. & Rendle 

Sterculia abbreviata E.L. Taylor ex Mondragón 

Sterculia apetala (Jacq.) H. Karst 

Sterculia kayae P.E. Berry 

Urticaceae 

Boehmeria cylindrica (L.) Sw. 

Phenax sonneratii (Poir.) Wedd. 

Pilea fendleri Killip 

Pilea involucrata (Sims) Urb. 

Verbenaceae 

Aegiphila perplexa Moldenke 

Citharexylum poeppigii Walp. 

Lantana glutinosa Poepp. 

Viscaceae 

Phoradendron berryi Rizzini 

Vitaceae 

Cissus alata Jacq.

Plant Community Structure, Fire Disturbance, and Recovery in ...

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