(page intentionally blank) CENTRE FOR THE STUDY OF BIOLOGICAL DIVERSITY UNIVERSITY OF GUYANA

Contributions to the Study of Biological Diversity Volume 1: 1 - 46

The use of biodiversity data in developing Kaieteur National Park, Guyana for ecotourism and conservation

by Carol L. Kelloff

edited by Phillip DaSilva and V.A. Funk

Centre for the Study of Biological Diversity University of Guyana Faculty of Natural Science Turkeyen Campus Georgetown, Guyana 2003 ABSTRACT

Carol L. Kelloff. Smithsonian Institution. The use of biodiversity data in developing Kaieteur National Park, Guyana for ecotourism and conservation. Contributions to the Study of Biological Diversity, volume 1: 46 pages (including 8 plates).- Under the auspices of the National Protected Areas System (NPAS), Guyana is developing policies to incorporate conservation and management of its tropcial forest. Kaieteur National Park was selected as the first area under this program. Information on the plants (and animals) is vital in order to make informed conservation or management policy for this unique ecosystem of the Potaro Plateau. Understanding and identifying important ecosystems and the locations of endemic plant taxa will assist Guyana in formulating a comprehensive management and conservation policy that can be incorporated into the development of Kaieteur National Park. KEY WORDS: Guyana, Kaieteur, conservation, management, biodiversity

DATE OF PUBLICATION: June 2003

Cover: Photo of Kaieteur Falls by Carol L. Kelloff. Cover design courtesy of Systematic Biology: Journal of the Society of Systematic Biology published by Taylor and Frances, Inc. in April 2002. Back cover: photo of the Centre for the Study of Biological Diversity, UG by T. Hollowell.

All photographs Copyright, Carol L. Kelloff , except as noted.

Contributions from the Centre for the Study of Biological Diversity, University of Guyana, Faculty of Natural Science, Turkeyen Campus, Georgetown, Guyana, South America

POSTMASTER: Send address changes to the Editor, Centre for the Study of Biological Diversity, University of Guyana, Faculty of Natural Science, Turkeyen Campus, Georgetown, Guyana, South America CONTENTS

INTRODUCTION ...... 5 Guyana’s Forests ...... 6 Forest management in Guyana ...... 7 Establishing a National Protected Area System (NPAS) ...... 8 KAIETEUR NATIONAL PARK ...... 8 Biophysical setting of Kaieteur ...... 8 Location and size of the Park ...... 9 Legal status, administration and management ...... 9 Infrastructure ...... 10 Local communities ...... 10 Access to Kaieteur ...... 11 BIODIVERSITY OF THE PARK ...... 11 Biological diversity ...... 11 Biological exploration ...... 13 Producing maps ...... 14 PARK DEVELOPMENT ...... 17 Tourisn-for-Nature ...... 18 Involving local communities ...... 20 Infrastructure development of the area around Kaieteur Falls and Tukeit ...... 22 Using signage for interpretation ...... 23 Major concerns to consider ...... 25 CONCLUSION ...... 25 ACKNOWLEDGEMENTS ...... 27 REFERENCES ...... 28 APPENDIX A: Protected Area Categories, Purposes and Practices Permitted to Local Communities ...... 31 APPENDIX B: Lichenized fungi and brytophytes of Kaietuer National Park, Guyana ...... 32 APPENDIX C: Invertebrates of Kaieteur National Park, Guyana: Butterflies, Moths, and Termites ...... 35 APPENDIX D: Vertebrates of Kaieteur National Park, Guyana: Birds, Herpetofauna and Mammals ...... 38 PLATES ...... 45

Kaieteur National Park, Guyana: C.L. Kelloff 5

The Use of Biodiversity Data in Developing Kaieteur National Park, Guyana for Ecotourism and Conservation

Carol L. Kelloff 1

INTRODUCTION

Guyana is a small country encompassing unique geology, tropical climate and an an area of about 215,000 square km and is ecosystem that has remained relatively free of located on the Atlantic seaboard of northeastern human disturbance makes Guyana rich in South America (Figure 1). It is one of the six biological resources. countries that lie over the rich biogeographic region of the Guiana Shield. On the Shield, Underlying these biological resources, Roraima sandstone forms the Pakaraima Guyana is rich in mineral resources such as Mountains in northwestern Guyana and extends bauxite, diamonds, gold, kaolin, and manganese into the Venezuelan Highland as a series of (Oleynik et al., 1999). In the past few decades, table-top mountains (Gibbs and Barron, 1993). pressure has increased to exploit these resources Guyana is covered with a rich rain forest and a for economic growth and national development. variety of other natural habitats. Over 70% of The burden of foreign debt and local poverty has the country is still covered with forest, ranging increased the need for the Government of from montane and lowland evergreen rain Guyana (GoG) to consider releasing larger tracts forests to dry evergreen and seasonal forests (J. of land for mineral and timber mining. This has Singh, pers. comm.) Tropical dry savannas of lead to environmental concerns expressed by the Rupununi extend into Brazil while tropical both national and international parties. The loss wet of biodiversity on a global scale is being recognized by the international community. The importance of biodiversity to the environment, the economy, and to society led the United Nations Conference on Environmental and Development to hold a conference in Brazil in 1992. The purpose of the conference was to place emphasis on local and global resources and how nations can take actions to conserve and use biodiversity in a sustainable way. Since ratifying the Convention on Biological Diversity in August 1994, Guyana has agreed to develop strategies for environmental protection and natural resource conservation. Guyana recognizes that biological and other natural resources are major components of the national Figure 1: Location of Guyana on the Guiana patrimony. One of the initial steps towards this Shield, northeastern coast of South America goal was the production of the “National and semi-wet savannas are centered on the Strategy for the Conservation and Sustainable Berbice River system. This combination of Use of Guyana’s Biodiversity” (EPA-Guyana, 1: Biological Diversity of the Guianas Program, MRC 166, National Museum of Natural History, Smithsonian Institution, Washington DC 20013-7012, USA. 6 Kaieteur National Park: C.L. Kelloff

1997)to define its position on biodiversity. This portion of the earth’s biodiversity. These forests process later produced the “National constitute approximately 22 percent of the Biodiversity Action Plan” (EPA-GEF/UNDP, original forest cover of the earth (Bryant et al., 1999) to promote the responsible use of 1997). In addition to providing ecological biodiversity and biological resources and to functions of climate stabilization and watershed develop a system for their conservation. protection, the natural forests are habitats for 50 – 90 percent of all terrestrial species still on With the aid of international agencies, earth. Large areas of natural forest provide Guyana has developed a policy to incorporate habitat needed for the survival of far-ranging the conservation of biodiversity and unique animals such as the harpy eagle, the timber wolf, geological formations within the management of and the grizzly bear. They are storehouses of its tropical forests under the auspices of its genetic resources and homelands to much of the National Protected Area System (NPAS). remainder of the world’s indigenous peoples. Kaieteur National Park is the first focus of the NPAS project. However, before the If the riches and resources of the forests Government of Guyana (GoG) can initiate any are to be available for future generations, the informed conservation or management policy concept of stewardship must be incorporated for this unique ecosystem, a basic knowledge is into precepts of how we, the citizens, required of the plants and animals in the park, businessmen, and policy-makers, use the forests. their relative abundance and location, and what For the future of these and all forests, good areas may need further study. This paper stewardship may mean setting aside areas for examines Kaieteur National Park’s present complete conservation and preservation while management and infrastructure and the current other areas may need to be carefully managed knowledge of the biological diversity of the park for both timber and non-timber products. and provides some suggestions on developing the park for ecotourism and conservation. There are only eight countries in the world that have large tracts of original forest remaining Fear of global climate change, species that are not yet under some form of threat. extinction, pollution, and habitat destruction has These countries are Russia, Canada, Colombia, caused the nations of the world to realize that Venezuela, Brazil, Surinam, French Guiana and forests are no longer an unlimited resource. It Guyana (Bryant et al., 1997; Miranda et al., has been estimated that almost half of the six 1998). billion hectares of forest that once covered the earth are gone, and of the remaining forest 16 Guyana’s Forests million hectares fall to the chain saw, bulldozer, Geographically, Guyana is situated on or fire each year (Bryant et al., 1997; Miranda et the Guiana Shield, a granitic formation overlaid al., 1998). What has developed in some areas of with sandstone, quartzite, shales, conglomerates the world (such as eastern North America) is a and boulder beds (Fanshawe, 1952; Gansser, mosaic of small or highly disturbed fragments of 1954). Contiguous with the Amazon Basin, the forest that have become refuges for some of the Shield represents one of the most intact parts of world’s most endangered species, such as the the region. Approximately 70% of Guyana’s spotted owl in northwestern United States tropical forest (11.5 million hectares) remain (Arcese and Sinclair, 1997), the lemurs in intact. The forests may be classified as rain, Madagascar (Jolly, 1988; Mittermeier et al., seasonal, dry evergreen, montane, marsh, and 1992), or the black caiman (Asanza, 1992) in swamp (Bernard, 1999). Guyana also has two South America. Although these fragments are classes of savannas: tropical wet and semi-wet still important for providing economic products savannas in the northeast and tropical dry and environmental services, they may not be savannas in the southwest of the country (World able to sustain these uses in the long term. Bank, 1996). The different forest and savanna classifications are depicted on the ecoregion However, the ecologically intact, map of Guyana (Figure 2), based on the relatively pristine natural forests may survive Preliminary Vegetation Map of Guyana (Huber and continue to harbour and maintain a good et al., 1995) that was compiled from existing Kaieteur National Park: C.L. Kelloff 7

literature, LANDSAT-5 satellite images, and Mountains in the northwestern region and the existing knowledge of the natural vegetation. Rupununi savanna. However, it has lagged behind in setting aside land for conservation. Guyana’s conservation tradition was nominal under the British (Sizer and Rice, 1995), and after independence in 1966 most initiatives focused on managing the forests. In 1979, the Guyana Forestry Commission (GFC) replaced the previous forestry service in the administration of timber resources. GFC drafted the National Forest Policy in 1985 and the National Forestry Action Plan in 1989. The objective of both of these documents was to efficiently manage forest resources, develop forest industries, and protect and conserve forest ecosystems and wildlife. The lack of adequately trained staff, funding, and infrastructure has made it difficult for GFC to implement and monitor its forest policy (The Tropenbos Foundation, 1991). It wasn’t until the late 1980’s that two initiatives began looking at conservation of Guyana’s forests: The Tropenbos Programme and the Iwokrama Rain Forest Reserve.

Figure 2: Map of the ecoregions of Guyana. The Tropenbos Programme, set up by the Government of the Netherlands, was established The forests of Guyana are relatively in Guyana in 1988 as the research component of pristine in large part because of their Demerara Timbers Limited (DTL). Its mandate inaccessibility, although the Guyana Forestry was to develop an appropriate management Commission (Guyana Department of Forestry- system for all forest types within DTL’s CIDA, 1989) considers approximately 3.6 235,000-hectare concession near Mabura Hill. million hectares (approximately 31% of the The Tropenbos Programme established a 900- remaining forest) exploitable. Access into much hectare ecological reserve with an adjacent of the interior of Guyana is possible only by buffer zone in the area of Mabura Hill. The small aircraft, boat or canoe, or on foot. There main objective of the Programme was to are very few roads into the interior beyond the conserve and wisely use the tropical forest coastal region, and on these pavement is through knowledge of biological processes and replaced by laterite clay that becomes impassible to develop methods for sustainable development during the rainy season. Approximately 95% of (The Tropenbos Foundation, 1991). Although Guyana’s 750,000 people live in the fertile area several good projects have contributed to this of the coastal plain, mainly from the capital, research, culminating in a number of Georgetown, east to the Corentyne River. In the publications (i.e.: Ek, 1997) in the Tropenbos- interior, or hinderland, about 35,000 indigenous Guyana Series (1 – 8), the Government of the people, collectively known as Amerindians, live Netherlands decided to restructure its in about 65 reserved areas scattered throughout Tropenbos Programme worldwide, and its work the country (Guyana Department of Forestry- in Guyana ended in late 2001. CIDA, 1989). The Iwokrama Rain Forest Programme Forest Management in Guyana was established in 1989 when the Government Guyana is richer in biological diversity of Guyana set aside 360,000 hectares of primary than either Surinam or French Guiana, due in rainforest in the central portion of the country. part the diverse habitats of to the Pakaraima The purpose of this reserve was to develop and 8 Kaieteur National Park: C.L. Kelloff

demonstrate methods of conservation and by the IUCN classification system (Appendix sustainable management of tropical forests A). The first two areas under consideration for (Hawkes and Wall, 1993; Pennington, 1993). NPAS were the Kanuku Mountains, inhabited Since its establishment, plant and animal by the Harpy Eagle, and Kaieteur National Park. surveys including birds (Ridgely and Agro, Since Kaieteur is the only legally established 1997); herpetofauna (Lee and Arjoon, ms.); park in Guyana, it became the first priority of mammals (Lim and Engstrom, 2001); plants NPAS. (Clarke et al., 2001); and fish (Watkins et al., 2001) have been undertaken in this diverse, low- elevation rainforest by a number of KAIETEUR NATIONAL PARK organizations. These surveys are providing the framework for conservation and sustainable Biophysical Setting of Kaieteur management efforts, as well as providing actual The Pakaraima Mountains (Figure 3) data for comparing Iwokrama with other areas in extend for 73,000 square km or about 34% of Guyana and with other countries (Clarke et al., 2001).

In August 1994, Guyana ratified the Convention on Biological Diversity. With this ratification and several other related international treaties signed earlier; including the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), Treaty for Amazonian Co-operation, and Convention Concerning the Preservation of World Cultural and Natural Heritage. Guyana’s commitment to environmental protection and natural resource conservation was underway (World Bank, 1996).

Establishing a National Protected Area System The first objective of the Government of Guyana was to establish the National Protected Areas System (NPAS) which would conserve globally-important biological diversity. The system was designed to work within Guyana’s strategy for sustainable development by contributing to watershed protection and productivity, ecosystem and biodiversity Figure 3 : Elevation map showing the conservation, and the maintenance of Guyana’s extension of the Roraima Formation, the cultural heritage (World Bank, 1996). There are Pakaraima Mountians in Guyana, and the a number of studies, reports, and documents that location of Kaieteur National Park. focus on the need, rationale, and criteria for the land area in Guyana (Gibbs and Barron, selecting protected areas at a countrywide level, 1993). The mountains are located in the mid- a regional level (Ramdass, 1990) or of a specific western portion of Guyana and are part of the type such as a World Heritage Site or Biosphere Roraima formation that also comprises a large Reserve (P. Hitchcock, pers. comm.). A portion of the Venezuelan Guayana. The problem with these reports is the lack of any geological formation of the Pakaraima type of procedure for selecting a proposed area. Mountains are composed mostly of a sandstone The design and implementation of NPAS was to conglomerate with rocks of quartz up to 7.5 cm encompass a full array of protected areas within in diameter. The range is comprised of a plateau Kaieteur National Park: C.L. Kelloff 9

including table-topped mountains (tepuis) and a and its left bank tributaries, the Uewang and series of steep-sloped escarpments. The average Kuribrong Rivers, as well as the Potaro River range in elevation of the Pakaraima Mountains itself on the west, to the source of the Kwitaro is from 300 to 500 meters with the highest peak, River; thence down the midstream of the Mt. Roraima, reaching (2,730 m). Kwitaro River through its mouth to the mid- stream of the Potaro River; then down the mid- The Pakaraima Mountains are the stream of the Potaro River for about six decimal source of many rivers that are renowned for their seven five (6.75) miles or (10.86) kilometers; waterfalls which drop off the edge of the thence in a southeasterly direction along the escarpment. These include the Sakaika Falls on watershed between the Echerak and Amakwa the Ekereko River and the Kamarang Great Falls rivers, right bank Potaro River; thence on the Kamarang River (Bernard, 1999), but northwards along the watershed between the certainly the most famous waterfall in Guyana is Potaro River and its right bank tributary, the Kaieteur Falls. It is the highest and most Amu River, to the source of the Waraparu beautiful of the falls on the Potaro River, located Creek, thence down the mid-stream of the where the Roraima formation gives way to the Waraparu Creek through it mouth to the mid- lowlands. Here the water from the Potaro River stream of the Potaro River; thence down the drops 226 m to the splash basin below after mid-stream of the Potaro River to a point which it travels some 32 km through a gorge opposite the mouth of the Haiabaru Creek, before leaving the escarpment behind. thence to the point of commencement; containing an area of two hundred and forty two Location and Size of the Park (242) square miles, or six hundred and twenty- Kaieteur Falls is the centerpiece of the six decimal eight (626.8) square kilometers national park. It was established in 1929 by the approximately.” British Commonwealth and encompassed 11,400 hectares. In the early 1960's, the park’s area were reduced to 1,940 hectares (19.4 sq. km), from Menzie’s Landing to the Tukeit Rest House Compound (Department of Lands and Surveys, 1961). This was done in order to take advantage of the rich mineral resources (gold and diamonds) of the area (Lechner, 1997). The radar image (JERS-1 SAR) of the Potaro Plateau (Figure 4) provides an aerial view of the topography of the area. The 1973 boundaries (Figure 4) covered nothing more than the falls and an approximately 9.5 km stretch of the Potaro River. On March 9, 1999, President Cheddi Jagan issued an Order under the Kaieteur National Park Act of 1929 extending the boundaries of the Park to 62,680 hectares. The demarcation of the new park boundaries (Figure 4) fall within the limits of the “Peoples Figure 4: Radar image of the Potaro Plateau Cooperative Unit No. 812121, Division: provides an aerial view of the topography of Waratak/Echerak” and is defined as follows the region. Areas indicate the size of the 1973 (Anon, 1999): (19.4 sq. km) and 1999 (627 sq. km) boundries of Kaieteur National Park. “Commencing at the mouth of the Haiabaru Creek at mid-stream, left bank Potaro River; extending thence up the mid-stream of the Legal Status, Administration and Haiabaru Creek to its source; thence in the Management westerly and southerly direction, along the In 1973, the Kaieteur National Park Act watershed between the Potaro River on the east was passed followed by the National Parks 10 Kaieteur National Park: C.L. Kelloff

Commission Act (NPCA) in 1977. The NPCA behind the guesthouse has little to no facilities. gave management authority to the National This is where the park wardens have lived. Two Parks Commission (NPC), an independent other buildings in the vicinity of the falls are the agency (World Wildlife Fund, 1991), radio beacon house just south of the runway and responsible for all regional and national parks. a small recently replaced house for the As an independent agency, the NPC has been attendants of the meteorological station. variously required, over the years, to report to either the Ministry of Communication and Along the edge of the gorge, about 150 Works; Guyana Agency for Health Sciences, meters from the falls, stand the cement footings Education, Environment and Food Policy of another structure that once overlooked the (GAHEF); the Ministry of Health; or the falls. In the 1920’s the British built a small Ministry of Culture, Youth and Sports. In 1995, house upon these cement footings to serve as the a Kaieteur Board was established through the original guesthouse. The outhouse stood several Office of the President. This board meets yards away near the edge of the gorge with a independently of NPC on issues dealing with large bucket serving as the base of the seat. Kaieteur. The NPC continues to collect the Although the guest- and out- houses eventually landing fees required of all aircraft using the fell, the bucket and footings of this historical site Kaietuer airstrip and is responsible for paying still remain. the rangers and park wardens stationed at the park. Along with the NPC and the Kaieteur Menzie’s Landing is located approximately Board, Kaieteur now comes under the one mile upstream from the falls. It has always administration of the Office of the President. In been a way station for gold and diamond miners the past, management of the park has been (locally known as porkknockers) heading into minimal, at best. Guyana is currently in the the bush or waiting for a flight to Georgetown. process of establishing a single centralized Although considered outside of the 19.4 square agency to oversee and manage Kaieteur. Until km boundary of the park, the Official Gazette this is done, it will be difficult for Guyana to (Department of Lands and Surveys, 1961) develop and carry out any kind of a included Menzie’s Landing as part of the park. comprehensive management plan. Today, Menzie’s Landing has several wooden buildings and an active shop where miners can Infrastructure restock supplies, sell their gold and diamonds, or Since the World Wildlife Fund’s first have a drink (mostly “High Wine,” a cheap form major attempt to develop a management plan of of rum). Although generally quiet, Menzie’s Kaieteur in 1991 at the request of the Guyana Landing can be quite rough when there is a large Ministry of Trade and Tourism, little has been influx of miners passing through. done to address the inferstructure problem at Kaieteur. Most of the current infrastructure is Approximately seven kilometers below located in the immediate vicinity of the falls. Kaieteur Falls, on the Potaro River, is Tukeit. Kaieteur has two airstrips, a small dirt airstrip Historically, Tukeit had several small structures that is no longer in use and the main 610 m which were built by the British around the airstrip. The larger airstrip was paved with 1950’s. The buildings housed a kitchen/dining concrete in 1993 and is capable of handling up area, sleeping quarters with bunks, and a to an Islander aircraft (eight-seater). One bathhouse with its row of toilets. It was a hundred ten meters from the head of the airstrip resting place for those visitors who selected to there stands a large “benab” or hut to shelter hike the steep old Amerindian trail (believed to visitors from the sun and rain and a small shed be 1000 years old) along the Korume Creek with two non-functional toilets. There are three gorge to and from the Kaieteur Falls. Today, main buildings at Kaieteur. The larger building, Tukeit stands as a mute reminder of the days of closest to the falls, was built in the 1980’s. It tourism under British rule. What remains are a has two bedrooms, inadequate kitchen and few boards, a pile of rusting equipment and sanitary facilities, a leaky roof, broken windows some broken toilets. and no lights. This building is considered the “guesthouse”. A smaller wooden house just Kaieteur National Park: C.L. Kelloff 11

Local Communities Guyana, and compares to other protected areas The only permanent settlement near little is known about Kaieteur (Table 1). Plant Kaieteur is the Amerindian village of Chenapau. and animal data need to be collected before It is approximately 25 km upstream from the important questions can be answered about taxa falls and has a population of 317 (Ministry of present in the area, their distribution, population Amerindian Affairs, no date). The people dynamics, and endemicity. Some may question belong to the Patamona tribe and have a the need for detailed collecting in the Kaieteur traditional lifestyle of hunting, fishing, area, but collections build vouchered inventories subsistence agriculture, and collection of minor that, in turn, provide accurate information for forest products. Chenapau has also become a verifiable environmental assessments, center of mining activities for gold and biodiversity conservation, and development diamonds by both the Amerindians and non- projects (Bojorquez-Tapia et al., 1994). From Amerindians. the foundation of inventories, guides and checklists can be produced (Mori et al., 1998). Access to Kaieteur The Preliminary Checklist of the Plants of There are two modes of transportation to Kaieteur National Park (Kelloff and Funk, Kaieteur, by air-charter or by boat up the Potaro 1998), the floristic comparison between the River. The most popular way to Kaieteur from forests at Kaieteur and Kwakwani, Guyana and Georgetown is by air. Kaieteur can be reached Barro Colorado Island, Panama (Kelloff, ms.), by a 1.5 hour flight in a small plane which flies and the distribution and broad floral affinities of up the Potaro River gorge for a spectacular plants of the Potaro Plateau (Kelloff and Funk, aerial view of the falls. Depending on the pilot, in review) represent the first serious attempts to the plane may circle overhead to provide photo understand the plant diversity at Kaieteur. opportunities to the guests on board. The However, the checklist of vascular plants (the drawbacks to air charters are that they tend to be best known floral group of the area), the costly and limit visitors to 1 – 2 hours at the preliminary checklists of the lichenized fungi falls. The other option is a 3 – 4 day trek and bryophytes (Appendix B) and the two overland, which requires a sturdy pair of ecological plots established at Kaieteur are only walking boots. The traveler must carry all gear the beginning of the environmental information and food needed for the trip. The overland trip necessary for any valuable protected area’s begins with a rough truck from Georgetown to management. Pamela Landing near Madia on the Potaro River. From the landing a boat takes the traveler The Potaro Plateau supports many upstream including a portage around Amatuk different habitats. The “guiana type” savanna is and Waratuk Falls (carrying boat and gear characterized by a shrub-herb plant community overland to the other side of the falls) before with only very few small trees. The pink sands arriving at Tukeit. Here they can set up camp support scattered shrubs and a dense mat of and rest before beginning the acsent up the small herbaceous plants. During the wet Korume Creek gorge on the old Amerindian trail season, on the almost bare, flat sandstone, to Kaieteur. The hike up the gorge takes an numerous species of lichens (Cladonia spp. and average of 2-3 hours, depending on a person’s Cladina spp.), the small blue flowered herb physical condition. Travel overland on foot Burmannia bicolor, two types of carnivorous from Amatuk to Kaieteur may not be feasible at plants, the tiny terrestrial bladderworts present because of the degradation of the trail by (Utricularia spp.), and the red sundew (Drosera mining activities and overgrowth. kaieteurensis), spring up out of tiny cracks and on the surface of the rock. This habitat is also the home of a species of small frog (Leptodactylus rugosus). During the brief rainy BIODIVERSITY OF THE PARK season, its tadpoles can been seen swimming through the thin sheet of water on the bare rocks. Biological Diversity Science is only beginning to understand the diversity of plants and animals found in 12 Kaieteur National Park: C.L. Kelloff

Table 1 : Current knowledge of the flora and fauna within the 1970’s boundary (19.4 square km) at Kaieteur National Park as compared to what is known for Guyana. Information in brackets indicates the theoretical percentage of species represented based on the current knowledge of the taxa in the Neotropics. Floral and Faunal Groups No. of species: Guyana No. of species: Kaieteur Lichenized fungi (lichens) 337 (90%) 1 87 (low) 2 Bryophytes (mosses) 238 (low) 1 72 (low) 3 Anthocerophyte (hornworts) 1 (?)1 Unknown Hepatics (liverworts) 259 (low) 1 4 (low) 1 Gymnosperms 5 (low) 1 Unknown Ferns 523 (90%) 4 115 5 Flowering plants (seed) 6334 (90%) 4 1176 5 Reptiles and Amphibians 254 (70%) 6 29 (low) 7 Fish (freshwater) 673 (low) 8 180 (low) 8 Butterflies and moths ca. 12,000 (50%) 9 164 (low) 10 Termites 130 (90%) 11 22 (low) 12 Mammals 227 (95%) 13 151 (65%) 14 Birds 786 (90%) 15 180 (60%) 16 1. T. Hollowell (pers. comm.) Biological Diversity of the Guianas database; 2. P. Depriest (pers. comm.) unpublished data; 3. A. Newton (pers. comm.) unpublished data; 4. Boggan et al., 1997; 5. Kelloff and Funk, 1998; 6. based on current knowledge of neotropical herpetofauna 7. based on museum specimens; 8. L. Page (pers. comm.); 9. L. Page (pers. comm.); 10. based on field observations and collections, S. Fratello (pers. comm.); 11. P. Eggleton and M. Kane (pers. comm.); 12. Eggleton et al., 1999; 13. Engstrom and Lim, 2002, Lim and Engstrom, in prep., Lehman, 2002; 14. based on Lim and Engstrom, in prep.; Emmons, 1997; 15. Braun et al., 2000; 16. based on field observations.

Brocchinia micrantha, one of the largest Small trees such as Andira grandistipula of all bromeliads growing to the height of 3-3.5 and shrubs like Clusia and Erythroxylum can meters, takes advantage of humus caught in develop into “bush islands” which support an larger cracks and crevices. The water that entire community of plants and often differ from collects in the base, ‘tank’, of B. micrantha island to island. Orchids, ferns and bromeliads leaves provides a home for an assortment of are frequently observed both on the ground and animals (e.g., the golden frog, Colostethus on other plants. The cabbage head bromeliad, beebei, endemic to Guyana) and plants. One Aechmea brassicoides, is a common plant of the such plant is the largest of the bladderworts, bush islands. This bromeliad is so named for the Utricularia humboldtii. Tiny bladders on its habit of its inner leaves being folded into a aquatic roots capture insects that live in the cabbage-like head. It can be found on the stagnant waters and digests them to obtain ground and up on the trees that make up the nutrients. From the bromeliad tank, the small bush islands on the savanna, and it is inflorescence of U. humboldtii, with its light endemic to Kaieteur. purple flowers, can reach the height of 1.8 meters. The bright yellow flowers of the "yellow- eyed grass" (Xyridaceae) are seen everywhere Another species of Brocchinia, B. reducta, on the savanna. If one looks closely upon some with tall, narrow, yellowish leaves, grows on the of these flowering heads, a yellow fungus can be grass savanna and on the lower limbs of trees. found. It is believe that the fungus mimics the First collected and described from Kaieteur flower and may be dispersed by flying insects Falls, this species was thought to be endemic to landing on the flowers. Other small herbs found the area until it was later collected in Venezuela in the savanna on moist sand are members of (Soderstrom, 1963). Eriocaulaceae family, “Bog Buttons”. These Kaieteur National Park: C.L. Kelloff 13

small white flowers resemble a tuft of cotton on 1898), E.F. Im Thurn (1879-99), T.G. Tutin a pin. (1933), N.Y. Sandwith (1937), D.B. Fanshawe (1944), B. Maguire and D.B. Fanshawe (1955), The white sand forests along the rivers are and R. Cowan and T. Soderstrom (1962). composed of numerous tree species, including Recent collectors from the Smithsonian's wallaba (Eperua), aromata (Clathrotropis Biological Diversity of the Guianas (BDG) macrocarpa), kakaralli (Eschweilera spp., Program and the University of Guyana (UG) Lecythidaceae) and members of the coffee include L. Gillespie (BDG), D. Gopaul (UG), T. family (Rubiaceae). The understory of these Henkel (BDG), C.L. Kelloff (BDG), L.P. Kvist forests support Heliconia, Marantaceae, and (BDG), H. Lall Persaud (UG), J.J. Pipoly many species of Melastomataceae. (BDG), and S. Tiwari (UG).

Because of the cool mist that rises up from There have been numerous species of the gorge, a cloud forest habitat is found along ferns and flowering plants discovered in Guyana the top of the gorge just downstream from the that are new to science. George S. Jenman, falls. This riparian forest supports more Government Botanist from 1879 to 1901 (Ek, epiphytes than a typical rain forest, with tree 1990) described many new species. The branches covered with mosses, orchids, ferns Cambridge University Expedition of 1933 and aroids. Down in the gorge near the splash collected many new taxa (Tutin, 1934a; Tutin, basin, the vegetation is bathed in a continuous 1934b; Lindeman and Mori, 1989; Ek, 1990). fine mist from the falls. The cool, ambient air Maguire and Fanshawe’s expedition (Maguire creates a unique habitat that scientists are just and collaborators, 1948) to Kaieteur as well as beginning to understand. Collecting in this area Cowan and Soderstrom’s expedition produced a has been difficult. The steep cliff face and the number of new taxa from the Potaro Plateau. slippery rocks make it almost impossible to get Kaieteur is still yielding new species. While near the falls when the volume of the Potaro inventorying the trees of the Korume Creek River is high. A small fern, Grammitis gorge, a tiny fern less than 2 cm tall, mollissima, grows on moss-laden rocks and tree Hecistopteris kaieteurensis (Vittariaceae), was trunks, and bromeliads line the steep walls of the discovered growing in moss on a trunk of a tree gorge. A primitive species of bromeliad, Navia (Kelloff and McKee, 1998). sandwithii, is found in the moist, shaded areas between boulders. Found growing only in the Even less is know about the animal splash basin habitat are two species of species of the Potaro Plateau. A preliminary Rhoogeton (Gesneriaceae). This red-flowered checklist of the birds of Kaieteur was compiled member of the African violet family has been using observation data from several reported near Mt. Roraima in a similar habitat ornithologists who visited the park between the (Toogood, 1983). years 1991 – 2002. Although thorough faunal inventories of the park are yet to be initiated, There are 22 endemic species currently lists of the butterflies, moths, and termites recorded for Kaieteur National Park (Table 2). (Appendix C) and birds, mammals, and In Rapateaceae, a family centered on the Guiana herpetofauna (Appendix D) are being compiled Shield, nine species can be found near the falls from the literature and collection data. These including Potarophytum riparium, a preliminary faunal lists provide a basis from monospecific genus endemic to Kaieteur which future collections and inventories can be (Maguire, 1970). planned. Preliminary studies from recent visits by specialists including G. Bourne (University Biological Exploration of Missouri – St. Louis), S. Lehman The first European to see Kaieteur Falls was (Smithsonian Institution), R. Reynolds (U.S. a British surveyor, C. Barrington Brown, in Geological Survey), M. Tamessar (University of 1870. Since that time, the Kaieteur area has Guyana), and D. Wilson (Smithsonian been explored by a number of botanical Institution) have indicated that this area is collectors including G.S. Jenman (1881 and particularly rich in animal life, and that the 14 Kaieteur National Park: C.L. Kelloff

Table 2: Plants species endemic the original a number of animals considered under CITES to 19.4 square km of Kaieteur National Park, be extremely rare (in a broad geographic range). Guyana. Important species known to be in the area are the bird Cock-of-the-Rock (Rupicola rupicola), PLANT ENDEMICS as well as bush dogs (Speothos venaticus, Aquifoliaceae CITES-extremely rare). A pair of hyacinth Ilex soderstromii var. soderstromii macaws (Anodorhynchus hyacinthinus, CITES- Ilex soderstromii var. ovata extremely rare) were seen flying over Kaieteur Bromeliaceae (author’s field notes, Kelloff, 1993-1995) in Aechmea brassicoides July1994. Although not officially reported for Chrysobalanaceae Guyana, it would be interesting to investigate Couepia cognata the possibility of a pair nesting in the area. Fabaceae – Caesalpinioideae Dicymbe jenmanii The cave behind the falls is home to three Macrolobium huberianum var. species of swifts: the white-chinned swift, pubirachis Cypseloides cryptus (globally rare), which has a Swartzia eriocarpa colony of several thousand birds; the white- Swartzia lamellata var. kaieteurensis tipped swift, Aeronautes montivagus (found only Tachigali pubiflora at Kaieteur and the Kanuku Mountains); and the Fabaceae – Papilionoideae large white-collared swift, Streptoprogne Clitoria kaieteurensis zonaris (Michael Braun, pers. comm.). These Melastomataceae birds can be seen circling above the falls in the Graffenrieda irwinii early morning and at dusk. Miconia maguirei Myricaceae Producing maps Myrcia extranea There are no published studies of the Myrtaceae ecological systems or natural habitats of Marlierea buxifolia Kaieteur. These types of studies are needed in Poaceae order to base decisions on where critical areas Thrasya achlysophila for preservation might be located. What has Paspalum bifidifolium been published from various expeditions and Podostemaceae inventories are primarily in the form of Jenmaniella guianensis checklists with relatively no other information. Jenmaniella isoetifolia However, when these lists are combined with Rhyncholacis jenmanii f. dolichophylla specimen label data, they can provide valuable Rapateaceae information concerning the distribution and Potarophytum riparium ranges of species (Yahnke et al., 1998). At Rubiaceae present, the plants are the best-known group of Psychotria kaieteurensis organisms of Kaieteur. Since the area most Sapotaceae affected by ecotourism is centered in the 19.4 Pouteria kaieteurensis square km portion of the park, this area will be used to demonstrate species distribution maps. presence of previously unidentified species is probable. Historically, agouti (Dasyprocta), A database of the plants of Kaieteur has paca (Agouti paca), tapir (Tapirus), red brocket been compiled using current and historical plant deer (Mazama americana), collared peccary collections from the U.S. National Museum (Tayassu tajacu), jaguarundi (Herpailurs), (US), New York Botanical Garden (NY), British raccoon (Procyon), bushmaster (Lachesis Museum (BM), Royal Botanic Gardens – Kew stenophrys), labaria (Bothrops spp.), and tegu or (K), the University of Utrecht (U), and from the salempenter (Tupinabis nigropunctatus) have literature. The database is specimen based and been recorded for this area. Although the fauna contains information on collector(s), collection inventories for the area are incomplete, there are number, taxa, locality, and other important data. Each record has been verified and, if the latitude Kaieteur National Park: C.L. Kelloff 15

and longitude (georeference) are missing, they information on the location where important have been added using either a reading from a species, i.e.: rare or endemic, have been found. global positioning system (GPS) taken at This information can be useful when developing collection locations at Kaieteur or from 1:50,000 a park for ecotourism and conservation. Since it topographic maps. Large tables and lists of raw is impossible in the context of this paper to data can be overwhelming to the user, presenting produce a range map of every species found in a hindrance to meaningful interpretation of the the park, this section will focus on the 22 taxa information. Maps are useful tools for endemic to Kaieteur. Five area maps have been consolidating large amounts of environmental produced of Kaieteur National Park between the information into a visual format that can be area of Menzie’s Landing and just below Tukeit. understood by most users. They provide a quick Each map displays the general area where plants reference for collection localities of particular endemic to the park have been collected (Table species. Mapping the patterns of species 3). occurrences can help in the analysis of species richness and diversity (Stoms, 1994) and Table 3: Known collection points for the indicate areas that require further study. endemic plants of Kaieteur National Park Although this database is far from a illustrated on maps with reference to figure comprehensive catalogue of the distribution of numbers. plants at Kaieteur, it is a beginning. COLLECTION LOCALITIES OF PLANT Compilation and digitization of spatial ENDEMICS features of Kaieteur National Park has been Figure 6 performed in Guyana through collaborations - Thrasya achlysophila with the Natural Resources Management Project - Tachigalia pubiflora (NRMP) and the Centre for the Study of - Psychotria kaieteurensis Biological Diversity (CSBD) at the University - Pouteria kaieteurensis of Guyana. The expanded Kaieteur National - Potarophytum riparium Park is covered by four topographic maps Figure 7 (Survey Department-Guyana, 1970, 1:50,000 - Swartzia eriocarpa scale; UTM projection, zone 21.): the NE and - Myrcia extranea SE of sheets of the Ayanganna quadrangle and - Miconia maguirei NW and SW sheets of the Kaieteur quadrangle. - Macrolobium huberianum var. pubirachis The topographic contours were digitized in Figure 8 increments of 50 feet. - Marlierea buxifolia - Jenmaniella isoetifolia ArcView was used to integrate - Swartzia lamellata var. kaieteurensis information from the database with the digitized - Ilex soderstromii var. soderstromii map of the Kaieteur area to produce a Figure 9 distribution or dot map (Figure 5) of collection - Graffenrieda irwinii localities. From the mapped collection data, it is - Couepia cognata apparent that plant collecting on the Potaro - Ilex soderstromii var. ovata Plateau has generally focused within the vicinity - Aechmea brassicoides of the falls and along the banks of the Potaro - Rhyncholacis jenmanii f. dolichophylla River. This type of map is useful in providing a Figure 10 graphic representation of the general area where - Licania microphylla plants have been collected and suggesting where - Dicymbe jenmanii more collecting and study needs to be done. - Clitoria kaieteurensis What this simple map does not indicate is what - Jenmaniella guianensis was collected or the intensity of collection for a particular locality. Distribution maps of individual plant species can provide some 16 Kaieteur National Park: C.L. Kelloff

Figure 5: Plant collection data, georeferenced and represented by dots (●), have been plotted on the topographic map of Potaro Plateau. Most plants collected in Kaeiteur National Park have been in the proximity of the falls and along the Potaro River. When developing the park for ecotourism, Aechmea brassicoides, as well as other plants, the plants that must be considered first and including orchids, that occupy this habitat. This foremost are the endemics. These plants have a damage could have been avoided, or at least range that is restricted to the park within the minimized, if a maintenance plan had been in immediate vicinity of the falls and are the most place for this area. vulnerable. From collection localities and site surveys (i.e., ground-truthing), the habitats can The area at the north end of the airstrip is then be assessed to determine whether protection one of the main trails visited by tourists. from foot traffic or park infrastructure Slashing the plants of the bush islands and development should be considered in the leaving the vegetation to rot also detracts from management plan. An example of a well- the natural beauty that attracts so many tourists meaning decision leading to the destruction of to Kaieteur. The bush islands are an important some of vulnerable habitat can be seen with the component of habitat in this area, and they could clearing of the bush from the north end of the be included as a scenic point along a self guided airstrip. Whether this was done to avoid any tour of the park (to be discussed later). The perceived interference with air traffic is cabbage head bromeliad, A. brassicoides, easily unknown. It is known that the slashing of these distinguished by the habit of its inner leaves, is bush islands damaged not only the general another “point of interest” that can bring habitat, but also the endemic bromeliad, awareness to the importance of preserving these Kaieteur National Park: C.L. Kelloff 17

Figure 6: Area map of Kaieteur National Park between Menzie’s Landing and Tukeit showing collection points for the following plants endemic to the park: Thrasya achlysophila, Poaceae (O); Tachigalia pubiflora, Fabaceae-Caesalpinoideae (■); Psychotria kaieteurensis, Rubiaceae ( V ); Pouteria kaieteurensis, Sapotaceae ( 7 ) and Potarophytum riparium, Rapateaceae (  ). fragile habitats. Understanding the plant There have been several papers written on community at Kaieteur and the species the development of a management strategy for distribution is an important first step in Kaieteur (MacKnight, 1991; Schuerholz, 1992; developing a comprehensive management plan Lechner, 1997; World Bank, 1998). Most of for the park. these papers focus on a large-scale development project for the park and involves a huge capital investment. Although these papers provide PARK DEVELOPMENT excellent suggestions on how Kaieteur National Park can be developed, they would require a In most countries, the nation’s protected tremendous amount of person hours to area system is the responsibility of the implement, and they assume that a management government. The land set aside for parks and plan or set of guiding principles are already in reserves represents a significant portion of the place. Guyana is just beginning to articulate its national territory. However, funding and position on biodiversity. The National administration for protected areas tends to rank Biodiversity Action Plan (EPA-GEF/UNDP, low in priority within a government’s budget 1999) and the National Protected Area System and as a result the efficiency of any management (World Bank, 1996) recognize biodiversity as an policy is reduced (Barzetti, 1993). 18 Kaieteur National Park: C.L. Kelloff

Figure 7: Area map of Kaieteur National Park between Menzie’s Landing and Tukeit showing collection points for the following plants endemic to the park: Swartzia eriocarpa, Fabaceae- Caesalpinoideae ( O ); Myrcia extranea, Myrtaceae ( V ); Miconia maguirei, Melastomataceae (7) and Macrolobium huberianum var. pubirachis, Fabaceae-Caesalpinoideae ( ). important national asset that can offer Guyana jumped from 25 million to 664 million, and the economic options, but the manpower and numbers are still rising (World Tourism funding will have to come from a variety of Organization, 2000). In 2000, the number of sources. tourists traveling rose to a record high of 698 million with an estimated US$476 billion spent There are various options that can be (World Tourism Organization, 2001). Traveling pursued for the development of Kaieteur abroad is not limited to special events such as National Park based on the current use of the the Olympics or the Soccer World Cup, many park, local community involvement, biological people are traveling for the sole purpose of diversity, and habitat. Beginning with the area enjoying and studying nature. Nature tourism, around the falls, the trail to and including Tukeit or ecotourism, is the fastest-growing sector of can be developed for tourism-for-nature. the tourism industry. Humans derive satisfaction and inspiration from observing the Tourism-for-Nature wonders of nature (Iltis, 1997), whether it is a Tourism is becoming one of the fastest geological formation, a landscape, or wildlife. growing industries in the world. The world population is taking part in more tourism Protecting land in a pristine state in order activities than a decade ago. Between 1950 and to promote ecotourism has the potential to 1999, the number of international arrivals generate income. It also provides an additional Kaieteur National Park: C.L. Kelloff 19

Figure 8: Marlierea buxifolia, Myrtaceae ( O ); Jenmaniella isoetifolia, Podostemaceae ( ▪ ); Swartzia lamellate var. kaieteurensis, Fabaceae-Caesalpinoideae ( + ) and Ilex soderstromii var. soderstromii, Aquifoliaceae (  ). species and all other plants and animals justification for setting aside land for associated with this bird. It also preserves future conservation. Observing birds in their income from birders for the local and national natural habitats is one of the leading sources of economies (Mori et al., 1998). ecotourism and prime example of how tourism can lead to conservation. For instance, in Costa Kaieteur is no exception to these Rica the quetzal (Pharomachrus mocinno) is the principles. In addition to the waterfall, the most sought after bird by birdwatchers. Its spectacular bird life attracts some visitors but breeding grounds are located in the Monteverde has the potential to bring in many more. Scarlet Cloud Forest and the International Children’s macaws frequent this area. Gliding through the Forest, where they are protected. However, gorge, they are frequently seen flying close to during certain times of the year the quetzal the edge of the escarpment. The Cock-of-the- migrates to the lowlands to feed on fruits of the Rock can be found in the area considered its Lauraceae (Wille, 1993; Powell and Bjork, home range near Johnson’s view. It is here, on 1994; Powell and Bjork, 1995). In order to the forest floor, that bright orange males perform protect this bird species, lowland forests, and their courtship display to attract potential female cloud forests must both be conserved. The partners. At the waterfall there are large profits from ecotourism have been used to set numbers of swifts that roost on the canyon wall aside and preserve the habitats used by the behind the falls. It is quite a sight to witness quetzel, thus contributing to the survival of the hundreds, if not thousands, of birds darting in 20 Kaieteur National Park: C.L. Kelloff

Figure 9: Area map of Kaieteur National Park between Menzie’s Landing and Tukeit showing collection points for the following plants endemic to the park: Graffenrieda irwinii, Melastomataceae ( O ); Couepia cognata, Chrysobalanaceae ( ▲ ); Ilex soderstromii var. ovata, Aquifoliaceaea (▄); Aechmea brassicoides, Bromeliaceae (  ) and Rhyncholacis jenmanii f. dolichophylla, Podostemaceae (P). management process, with an increased and out of the thin edges of the waterfall at dawn emphasis on socio-economic and cultural and dusk each day. In addition to these birds, factors. In a draft decision on tourism submitted excellent birdwatching can be realized in the to the United Nation’s Commission on forest on the way to Tukeit. Sustainable Development, Hanif (1999) urged governments to consult local communities and Involving local communities major stakeholders in the tourism development Many early studies of development and process “including policy formulation, planning, management strategies for protected areas have management and sharing of benefits”. He proposed a “top-down” approach. This cautioned that without involving local approach advocates high levels of technical and communities and major stakeholders in the capital inputs with an external control over development process, ecotourism would be resources. Local communities are generally left detrimental to the harmonious relationship out of such development plans or have very little among the people, the community and the input. environment.

In recent years, it has been acknowledged An example of where the establishment of that more emphasis is needed on contributions a park caused hardship in surrounding of local communities in the development and communities can been seen in the case of the Kaieteur National Park: C.L. Kelloff 21

Figure 10: Area map of Kaieteur National Park between Menzie’s Landing and Tukeit showing collection points for the following plants endemic to the park: Dicymbe jenmanii, Fabaceae – Caesalpinoideae ( ● ); Clitoria kaieteurensis, Fabaceae – Papilionoideae ( X ); Jenmaniella guianensis, Podostemataceae ( ▲ ) and Licania microphylla, Chrysobalanaceae (no georeference, not illustrated on map). 2001). The park also suffers from poachers and Keoladeo National Park in India. In 1982, India other encroachments. declared Keoladeo a national park. The local communities were not brought into the decision- At Kaieteur, the indigenous community making process, and as a result officials were closest to the park is Chenapau. The Patamona not aware of potential problems that could arise Amerindians at Chenapau and other surrounding out of limiting access to the park. The communities have had long-term occupation of establishment of the park deprived one local this area on the Potaro Plateau. The residents community of the rights to graze their cattle or practice traditional subsistence activities are to gather grass for thatching. Another slash and burn agriculture, hunting, and fishing community was denied its main route to market. as well as small-scale mining. When the This brought on civil unrest and several people Chenapau community learned of the March died. Today, the local communities are still 1999 order expanding Kaieteur National Park to paying the price in lost resources because their its present 627 square km without any needs were not taken into consideration when meaningful consultation with the Patamona the area was converted into a conservation area people of the Potaro Plateau, they immediately (Department for International Development, protested to the Government. The general reluctance of the government to deal with 22 Kaieteur National Park: C.L. Kelloff

indigenous peoples, has set the first stage in the guides, operators, and eco-tourism managers. development of the NPAS project into abeyance As tour guides, the people of the local (Richards, 1999; Colchester, 2001). communities are provided with employment opportunities. Their intimate knowledge of the In many countries indigenous people view area, the local plants and animals, and their tourism as an activity coming from outside with culture will give the visitor a richer no benefits to the community at large, or when understanding of the cultural and environmental their communities are included in the tours, they history of the area. Research has shown that may feel they are being exploited. This was true tourists enjoy “informative interactions” with in the Quichua community of Napo Province, local persons and consider their guide their Ecuador until the village of Capirona decided to “main source of learning” (Department for investigate how they could benefit from tourism. International Development, 2001). The idea project was conceived of a communal ecotourism that allowed local community Infrastructure development of the area management. The people of Capirona built around Kaieteur Falls and Tukeit several tourist cabins in the traditional bamboo There are three catagories of visitors that and thatch style, designed guided walks through come to Kaieteur each year. The first category the forest, prepared traditional meals, of tourist is the “nature” tourist who comes to demonstrated hunting weapons, and presented a view the falls and the Potaro River Gorge. It is cultural program. The income came directly estimated that 2,500 – 5,000 visitors come to from the visitors and was invested into a Kaieteur each year (OAS, 1997). The majority community fund. From there, the community of these visitors spend 1-2 hours on the plateau decided how the money was to be spent; e.g., as part of a packet tour including other areas; towards health emergencies, education i.e., Orinduk Falls on the Ireng River between instruction, or transportation improvements Brazil and Guyana or Baganara Island in the (Schaller, 1996). Essequibo River. The second category of visitors is comprised of researchers engaged in The development and expansion of various scientific projects, students, and Kaieteur should not be viewed as a threat by educational groups. These visitors spend local communities. A mechanism for the anywhere from two to twenty-one days at the development of a communal ecotourism project falls. The third category includes miners, may provide an avenue for Chenapau to feel that Amerindians, and others that pass though the they have a stake in the development of that area. portion of the park that is near their communities. Creating economic incentives Kaieteur Falls has been touted by may encourage local stewardship to biological Guyana’s nature tourist industry as having resources (Harlan, 2000). potential as a world-class attraction, but there is the need to improve the facilities. Extensive Recently, the Patamona at Chenapau have development has been proposed but this would signed an agreement with the World Wildlife take time, resources, and large amounts of Fund (WWF) and the Guyana Forest and money and the result may not be appropriate for Environmental Conservation Project (GFECP) Kaieteur National Park. The attractions of to develop a sustainable eco-tourism programme Kaieteur are, of course, the falls, but also the at Chenapau. The grant will help construct a wildness and beauty of the park. For this type of guesthouse and provide transportation to and experience, most nature tourists don’t mind from the airstrip at Kaieteur with the intention “roughing it” for a while. They feel that it is that Chenapau become a tourist destination. As part of the “jungle” experience. What they do a purely community-based venture, the require are two basic standards: security and Patamona at Chenapau hope the project will hygiene. preserve both their traditional land and culture, and in turn, benefit the community (Anon., 2001). This venture will provide members of the community with training to become tour Kaieteur National Park: C.L. Kelloff 23

As funding allows, infrastructure and recharge a bank of batteries for back-up. development of the park can proceed by stages. Electricity supplied to the guesthouse could be The initial stage should focus on the area limited to a few hours each evening (i.e., sunset immediately around the falls. This is the area – 10 pm) to provide lights for visitors. Visitors that is most widely visited by tourists. Kaieteur wishing to stay at the guesthouse would have to currently has a large thatched “benab” and a make arrangements in advance, and would be shed that houses two toilets. The benab is required to provide their own hammocks and generally in good condition but may need some mosquito netting, food and toiletries. work on the thatching. The toilets are non- functional, with no water supply for flushing. Waste disposal is another issue that needs to There are several toilet designs that do not be addressed. Proper containers for trash can be depend on water that work well. One that is the provided in several locations throughout the most environmentally friendly and maintenance park (i.e., airstrip, guesthouse, and along trails). free is the composting toilet suggested by the A planned system of trash disposal should be National Plan for Ecotourism Development developed that is environmentally friendly. In (Organization of Amercian States, 1997). the past problems have arisen when visitors as well as wardens have created a trash dump or The main guesthouse near the falls should burned trash in open, uncontrolled areas near the be redesigned to accommodate overnight falls. visitors. The design can be rustic, providing shelter, areas for hanging hammocks, sanitation The condition of the smaller house used facilities, and an area for cooking. The current by the park wardens is uncertain. A house has two bedrooms, two bathrooms with maintenance schedule for this facility needs to showers, a living/dining area and a small take into consideration sanitation, cooking, kitchen. The basic structure of the house is lights, roof, etc. The wardens should be trained sound but with some modifications it can be in emergency medicine, snake bite treatment, developed for low maintenance. The roof needs etc. They will need to be equipped with a radio- repair since it leaks in several areas. A transmitter to communicate with Georgetown, a catchment system for rainwater off the roof medical kit, and a toolbox with a few basic could provide water to the kitchen and bathroom tools. Other supplies can be provided or areas. Experience shows that toilets and replaced as needed, such as generator parts or showers can stress the water supply at Kaieteur fluorescent light bulbs for the houses. when there is a large group visiting. Compost style toilets, a design that does not entail Development of Tukeit for tourism will flushing, can help conserve water during the dry depend on the demand. This area could be set season and avoid wastewater disposal problems. up as a rustic campsite. A simple hut or benab The two bedrooms could be opened into a bay placed on high ground to prevent flooding can with pairs of opposing hammock hooks evenly provide visitors a place to hang a hammock. A spaced so that several visitors can hang small shelter where food can be prepared along hammocks without “bumping”. with a designated cooking area to prevent fires should be placed separate from but close to the The kitchen could be equipped with a LP sleeping quarters. Finally, an area for waste gas stove and a sink with running water for disposal and sanitation facilities needs to be washing dishes. Central lighting in each room designed to keep the area clean. would be needed in the house. Electricity could be provided by several means. A generator is Using Signage for Interpretation one method but this requires a supply of Interpretation is a way of conveying gasoline to be shipped and stored at the complex information to people and could be a key and creates noise which negatively impacts component of communicating the importance of tourist experience. Other methods include a Kaieteur National Park to visitors. The three wind turbine or photovoltaic panels. Energy most popular forms of interpretation generated would supply lights to both the implemented by many parks worldwide are: guesthouse and the smaller house in the back guided walks, trail brochures, and signs. 24 Kaieteur National Park: C.L. Kelloff

Interpretation is educational and can be used to which is inhabited by sundews, lichens, provide historical background on some of the Utricularia and small frogs and tadpoles of landmarks or areas of interest, to emphasize Leptodactylus rugosus. At both sides of the important habitats or ecosystems, to promote forest entrance are large bromeliads (Brocchinia conservation of the park, and to communicate micrantha) with Utricularia humboldtii, the rules and their function (Honig, 2000). tallest and largest bladderwort in the world, growing out of their tanks. The conspicuous Kaieteur National Park is visited by bird, Cock-of-the-rock, has his lek in this area, thousand of people each year. Most of these and tarantulas, lizards, snakes, and other animals visitors arrived by air-charters that are scheduled have been seen or have left their mark on this based on availability of passagers. This trail. The path also has three sets of cement irregularly may make signage the best (or a very steps and two bridges that were built during important) method for communication and British rule. Visitors may be interested in the interpretation in park. Signs are useful in that historical ruins at Kaieteur. Signs at these sites they are available to the visitor at all times. can provide the visitor with information on They can be set into place without a high cost, when and why these structures were built. reducing the requirement for tour guides and staff or excessive maintenance. There are many On the “Airstrip – Falls Trail” the publications that discuss the development and vegetation of the grass- and shrub-savanna is use of signs in park development (Roff, 1995; found, including many showy flowers of Ryan, 1995; Honig, 2000; Ham, 1992) and these Bignoniaceae, Xyridaceae, Gentianaceae, and can be reviewed to determine the best options other plants. A sign can point out the trail of the for Kaieteur. leaf cutter ants that utilize the forest to cut sections of leaves to bring back to their nest. At Signage can be used to designate a the top of the falls are several plants endemic to direction or location on a trail and provide a Kaieteur. A sign near these plants can point out general orientation on a map of the area. the importance of preserving the habitat in order Smaller signs can be used as trail guides and to that these rare plants may be enjoyed by future provide information on an area of interest. At visitors. present, Kaieteur has three paths that are used by tourists. From the northern end of the airstrip, Signs placed along the “Johnson’s View – visitors may walk directly to the falls or head Falls Trail” can be used to bring the visitors northward towards Johnson’s View. The other attention to the many different plant species that path runs from Johnson’s View to the falls. grow near the edge of the gorge. Historical Although these paths are generally well defined features along this trail include the remains of by the trampling of many feet over the years, the “footing” of the old British guest house and visitors have been known to become lost, the bucket that once served as the base to the requiring searches by rangers (M. Phang, pers. outhouse. Can you imagine the surprise when comm.). Signs at Kaieteur should therefore the visitor realizes how close to the edge of the serve two purposes, enrichment and safety. gorge the outhouse once stood?

Currently, visitors are escorted or directed Besides information on the flora, fauna to the falls or Johnson’s View and are returned and historical landmarks, signage can convey a to the planes often without detailed information strong message of conservation. Messages such on the habitats that they have walked through or as “take only pictures and leave only footprints” the history or cultural aspects of the area. are used in a number of parks in the United Signage along these trails can be used to States. The visitor understands that defacing highlight various points along the way and landmarks, littering, and actions that may lead to enrich the experience. the destruction of habitat are not acceptable.

Points of interest along the “Airstrip – The messages that are passed on to the Johnson’s View Trail” are the bush island visitor by signage can be very positive ones. habitats and the seemingly bare sandstone rock Signs can point out features that may otherwise Kaieteur National Park: C.L. Kelloff 25 be overlooked such as unique plant CONCLUSION communities, areas that animals may frequently visit, and historical events or landmarks. Guyana stands at the threshold of Signage can convey the importance of staying developing its park and reserves system under on designated paths to prevent damage to fragile the National Protected Area System (NPAS). ecosystems and to keep visitors safe. And The first area selected under the NPAS project finally, signage can enrich the experience and was Kaieteur National Park. Although the give visitors the sense that it is not just the falls Government of Guyana recognized the need to that make Kaieteur unique but the splendor of its conserve a larger portion of the ecosystem and forests, its wildlife and the indigenous people watershed and expanded the park to 627 square who live there. km, it failed to identify the needs of the local communities and the importance of the flora and Major Concerns to Consider fauna. There are several major concerns about the welfare and condition of the park as it stands There are several ways in which local today. The Potaro River and smaller tributaries communities can be incorporated into the above and below the falls have been under siege development and management of the park. As by gold mining dredges. These dredges dig into with many park projects of the past, the main the river banks and silt the waters (Anon., problems confronted by governments have been 1998a; Anon., 1998b). Diamond mining the lack of understanding of the socio-economic operations, though smaller in scale, also leave and cultural realities of the local communities. their marks on the landscape. Damage from Including the local communities in the decision trails and pits cut into the earth leave a legacy of making process and providing opportunities for environmental degradation. Although Guyana local communities to develop and manage a has placed a moratorium on any mining activity communal ecotourism project that would benefit within the park (Ministry of Health, pers. the people directly, may be the answer to some comm.) and has enacted laws restricting this problems with park selection and development type of activity, lack of staff and financial in Guyana. The Patamona people have lived in resources has limited Guyana’s ability to enforce the area longer than most can remember. The these laws. legend of Kaieteur is passed down from their oral history. The Amerindians can enrich the Over 2,500 tourists visit Kaieteur each year. visiting tourist through their local knowledge, The park is raw and rugged, which gives the customs and traditions. visitor the thrill of being in the rainforest. But without proper signs, boardwalks, and guides, Kaieteur National Park is not just a single the potential for injury or losing a visitor along a feature park. The area has a rich geological, trail is great. The unwary visitor wandering off biological, and cultural history. Although the the paths can also threaten delicate ecosystems. falls is a spectacular geological feature (single Crushing plants underfoot and breaking drop falls of 226 m), as noted in the branches can lead to habitat degradation and phytogeography study of Kaieteur Falls (Kelloff erosion. and Funk, in review), the plants represent a unique phytogeographic region of South Mining and tourism are not the only America. There are several rare and endemic threats to the environment. Other potential animals found at Kaieteur in the immediate pressures on the environment are the capture of vicinity of the falls, but more inventories are animals for the wildlife trade and commercial needed in both plants and animals to fully logging within the park. understand the biological diversity of the Potaro Plateau.

Kaieteur needs to be developed in such a way that protection of the ecosystems generates income. The expansion of ecotourism within 26 Kaieteur National Park: C.L. Kelloff

Kaieteur National Park is one option. Kaieteur the diversity and complexity of the ecosystems can be easily and cost effectively developed by will be gained. simply renovating the existing buildings and sanitation facilitates. Signage can be used to And finally, management and guide and educate the visitor at a low cost and administration of national parks generally fall provide an increased level of safety as the visitor under the responsibility of the Government. If moves around the park. It can also provide Kaieteur is to have a comprehensive markers to prevent damage to fragile habitats management plan, there needs to be a single and keep tourist safe. Signage can convey a centralized agency to oversee and manage the message that visitors to Kaieteur can take with park and all its activity. Table 4 is a short list of them - “It is not just the falls that make Kaieteur suggestions to help provide an initial framework unique but the splendor of its forests, its wildlife towards developing and managing Kaieteur and the indigenous people who live there”. This National Park. It is understood that there may strong conservation message needs to be be actions already taken by the Guyana conveyed in order to protect Kaieteur National Government toward this goal and unknown at Park. It is Guyana’s natural heritage and it the time of this writing. It should be noted that needs to be protected and preserved for future neither the list nor this paper constitute an generations. Gold and diamond mining, infrastructure design plan for Kaieteur. Proper unchecked and unplanned development, and the planning and development can only occur after a capture of animals for the wildlife trade present management plan has been formulated and major threats to the ecosystems and can be adopted. detrimental to ecotourism as these activities can spoil the intrinsic beauty of the park and can Perhaps, for Guyana, the legend of El reduce the safety of the tourists. Dorado will come true. However, the lure of gold and riches will not be from those that are With the recent expansion of the park and dug out of the ground but come from the beauty the desire to make Kaieteur a World Hertitage of its natural heritage. Site, more information is needed on the flora and fauna of this region. As more inventories are completed for both plants and animals in the park, a better knowledge and understanding of Kaieteur National Park: C.L. Kelloff 27

Table 4: Some suggestion to better meet the objectives towards developing conservation and ecotourism at Kaieteur National Park.

Topic Suggested Action Benefit Forestry Legislation Develop clear objectives in the Reduces conflict with National Forest Policy to prevent Forestry. conflicts with the development of the NPAS. Concession Moratorium Stop all future mining and timber Prevents further damage to concession permits in Kaieteur this fragile ecosystem. National Park, including exploratory permits. Current Mining and Do not renew permits. Reduces concessions as Timber Permit in the permits run out. Park Land Issues Demarcate boundaries with Reduces border conflicts and Chenapau & establish zoning in provides tradition and the park near the village. subsistence use for local communities. Chenapau Village Establish a dialog to resolve Establishes communications to conflicts. Help Chenapau develop help reduce conflicts with a project to help village benefit local communities. from ecotourism. Land Zoning Establish areas within the park Clearly defines the area for boundaries and around the use. perimeter of the park. Administration Establish one Government agency Centralizes management of the to be responsible for management park. of the park. Infrastructure Facilitate the overall management Protection of the park and Development: Phase I of the protected and improvement increased satisfaction of of lodging and signage. tourists and provide a safe and educational experience for visitors. Infrastructure Develop a modest building plan for Repair existing structure to Development: Phase II core area keep initial cost low. Scientific Research Provides data and analysis on the Provides policy makers with plants and animals in the park biodiversity information for zoning, developing, conservation; petition for World Heritage Site

ACKNOWLEDGEMENTS the Department of Botany, National Museum of Natural History, Smithsonian Institution, that There were many people that helped has been operating since 1983. during the course of this research in one way or I thank Dr. Lee Talbot for his advice another. I thank the Smithsonian Institution’s and suggestions on the writing of a policy paper, Biological Diversity of the Guianas (BDG) Major General (retired) Joseph Singh for his Program for its financial support during this comments on the content, Dr. Ben J.H. ter Welle project. The BDG is a field-oriented program of and his staff at the Natural Resource 28 Kaieteur National Park: C.L. Kelloff

Management Project – Guyana and Naseem Bojorquez-Tapia, L., et al. (1994). “Biological Nasir, GIS Specialist, Centre for the Study of inventories and computer data bases: their role Biological Diversity, UG for their assistance in in environmental assessments.” producing the digital topographic map of Environmental Management 18(5): 775-785. Kaieteur. Special thanks are given to Tom Bryant, D., et al. (1997). The last frontier Hollowell, Data Manager, BDG for his forests: Ecosystems and Economies on the assistance with the distribution maps and finally, edge. Washington, DC, World Resources Dr. Ted Bradley, Dr. V.A. Funk, Dr. James Institute. Lawrey, Dr. Larry Rockwood, Dr. Berry Haack Clarke, D., et al. (2001). Plant Diversity of the for their patience and guidance throughout my Iwokrama Forest, Guyana. Fort Worth, Texas. research. Colchester, M. (2001). Proposed listing of the And finally, one cannot forget all the Kaieteur National Park, Guyana, as a World park wardens: M. Phang, L. Gibson, P. Heritage Site (letter dated: 31 January 2001). Benjamin and others; the porkknockers that Department-for-International-Development went under such names as “god” and “red- (2001). Changing the nature of tourism: dread”, and the shopkeepers at Menzie’s Developing an agenda for action. London, Landing that have made the experience at Environmental Policy Department, Dept for Kaieteur interesting and colorful. I thank you International Development: 14. all. Department of Lands and Surveys (1961). This is number 77 in the Smithsonian’s Boundaries of Kaieteur National Park. Biological Diversity of the Guianas Program Georgetown, Guyana. publication series. Ek, R. C. (1990). Index of Guyana Plant Collectors. Koenigstein, Koeltz Scientific REFERENCES Books. Ek, R. C. (1997). Botanical Diversity in the Anon. (1998a). Top team in emergency check of Tropical Rain Forest of Guyana. Tropenbos - Kaieteur Mining signs seen 11 miles Guyana Series 4. The Netherlands, upstream. Stabroek News. Georgetown, Tropenbos-Guyana Programme: 237. Guyana. EPA-GEF/UNDP (1999). Guyana: National Anon. (1998b). Kaieteur Warnings. Stabroek Biodiversity Action Plan. Georgetown, News. Georgetown, Guyana. Guyana, Environmental Protection Agency - Anon. 1999. Kaieteur park demarcation Guyana: 135 pp. Anon. (2001). $8.9M Chenapowu eco-tourism EPA-Guyana (1997). National Strategy for the deal signed. conservation and sustainable use of Guyana's Asanza, E. (1992). Population dynamics, Biological Diversity. Georgetown, Guyana: ecology and conservation of the black caiman, 44. Melanosuchus niger in Ecuadorian Amazonia. Fanshawe, D. B. (1952). The vegetation of Crocodiles. Preceedings of the 11th Working British Guiana: A preliminary review, Meeting of the Crocodile Specialist Group. Imperial Forestry Institute: 1-96. Gland, Switzerland, IUCN: 22-30. Gansser, A. (1954). “The Guiana Shield (South Arcese, P. and A. R. E. Sinclair (1997). “The America).” Ecolog. Geol. Helv. 47: 77-117. role of protected areas as ecological Gibbs, A. K. and C. N. Barron (1993). The baselines.” Journal of Wildlife Management Geology of the Guiana Shield. New York, 61: 587-602. Oxford University Press. Barzetti, V. (1993). Parks and progress. Guyana Department of Forestry – CIDA (1989). Cambridge, England, IUCN-the World National Forestry Action Plan: 1990-2000. Conservation Union in collaboration with the Georgetown, Guyana Forestry Commission: Inter-American Development Bank. 77 pp. Bernard, D. M. (1999). A New Geography of Ham, S. (1992). Environmental Interpretation - a Guyana. London, England, Macmillan practical guide for people with big ideas and Caribbean. small budgets, North American Press. Kaieteur National Park: C.L. Kelloff 29

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Appendix A: PROTECTED AREA CATEGORIES, PURPOSES AND PRACTICES PERMITTED TO LOCAL COMMUNITIES

1A. Strict Nature Reserve Practices Permitted to Local Communities: Purpose: Strict protection and scientific - Subsistence fishing research - Limited commercial hunting and fishing Practices Permitted to Local Communities*: - Sport hunting and fishing Strict protection; no extraction - Usually, no human settlement inside permitted 5. Protected Landscape or Seascape 1B. Wilderness Area Purpose: Landscape/seascape conservation Purpose: Wilderness protection and recreation Practices Permitted to Local Communities: Practices Permitted to Local Communities: - Controlled, very limited extraction of - Sustainable traditional uses, such as - certain natural products permitted: nuts, hunting, fishing, logging, fruits, seeds, oils, medicinal plants - agriculture, but not at industrial levels - Limited subsistence hunting and fishing - Limited traditional mining - Usually no human settlements inside - Some communities, usually within the area 2. National Park Purpose: Ecosystem protection and 6. Managed Resource Protected Area recreation Purpose: Sustainable use of national Practices Permitted to Local Communities: ecosystems - Controlled, very limited extraction of Practices Permitted to Local Communities: certain natural products permitted: nuts, - Sustainable logging, grazing, agriculture fruits, seeds, oils, medicinal plants - Small land dredges for mining - Limited subsistence hunting and fishing - Commercial fishing - Very limited traditional mining - Some communities usually with the area - Sometimes limited human settlements inside the area 7. Biosphere Reserve Purpose: Combination of strict protection, 3. Natural Monument ecosystem protection and recreation, Purpose: Conservation of specific natural and sustainable use of national features, such as a waterfall, mountain resources, through zoning (Very often Practices Permitted to Local Communities: made up of other protected areas at - Controlled, very limited extraction of national level) certain natural products permitted: nuts, Practices Permitted to Local Communities: fruits, seeds, oils, medicinal plants - Combination of 1, 2 and 6 practices - Occasional tree removal for subsistence use (canoes, tools, construction) - Usually, no human settlements inside * Local Communities defined as indigenous peoples, ethnic groups and other local people. 4. Habitat or Species Management Area Purpose: Conservation through specific management interventions for species and habitats 32 Kaieteur National Park: C.L. Kelloff

APPENDIX B: LICHENIZED FUNGI AND BRYTOPHYTES OF KAIETEUR NATIONAL PARK, GUYANA (compiled using collection data records of U.S.National Herbarium)

LICHENIZED FUNGI Cladonia rugulosa Ahti Cladonia secundana Nylander Cladonia signata (Eschweiler) Vainio Arthoniaceae Cladonia sipmanii Ahti Cryptothecia sp. Cladonia spinea Ahti Bacidiaceae Cladonia subradiata (Vainio) Sandstede Biatora sp. Cladonia substellata Vainio Crocynia pyxinoides Nylander Cladonia vareschii Ahti Crocynia pyxnoides Nylander Cladonia variegata Ahti Lecidea leucophyllina Nylander Coccocarpiaceae Phyllopsora buettneri (Mueller-Argoviensis) Coccocarpia domingensis Vainio Zahlbruckner Coccocarpia imbricascens Nylander Phyllopsora sp. Coccocarpia sp. Physcidia squamulosa Tuckerman Ectolechiaceae Physcidia wrightii Tuckerman Calopadia sp. Cladoniaceae Cladia aggregata (Swartz) Nylander Graphidaceae Cladina argentea Ahti Cyclographina sp. Cladina confusa (R. Santesson) Follmann & Graphina malmeana Ahti Graphina sp. Cladina confusa (R. Santesson) Follmann & Gyalectaceae Ahti f. confusa Coenogonium linkii Ehrenb. Cladina dendroides (Abbayes) Ahti Coenogonium sp. Cladina densissima Ahti Lichen – unknown families Cladina rangiferina (Linnaeus) Nylander Lichenothrix sp. Cladina sprucei (Ahti) Ahti Trentopholia? sp. Cladonia aggregata (Swartz) Nylander Lichen Incert. sed. Cladonia ahtii S. Stenroos Siphula carassana Mueller-Argoviensis Cladonia chimantae Ahti ined. Siphula carassana Vainio Cladonia coniocraea (Fl”rke) Sprengel f. Siphula decumbens Nylander Cladonia corallifera (Kunze) Nylander Cladonia crassiuscula Ahti Lichenized basidio. Cladonia delicatula Ahti ined. Dictyonema glabratum (Sprengel) D. Cladonia didyma (F‚e) Vainio Hawksworth Cladonia furcata (Hudson) Schrader Dictyonema sericeum (Swartz) Berkeley Cladonia furfuracea Vainio Lobariaceae Cladonia hians Ahti Sticta sp. Cladonia huberi Ahti Megalosporaceae Cladonia meridionalis Vainio Megalospora tuberculosa (Fée) Sipman Cladonia mollis Ahti Opegraphaceae Cladonia peltastica (Nylander) Mueller- Dichosporidium nigrocinctum (Ehrenberg) Argoviensis Thor Cladonia pityrophylla Nylander Parmeliaceae Cladonia polita Ahti Bulbothrix goebelii (Zenker) Hale Cladonia polyscypha Ahti & Xavier Filho Bulbothrix tabacina (Montagne & Bosch) Cladonia prancei Ahti Hale Cladonia pulviniformis Ahti Canoparmelia caroliniana Kaieteur National Park: C.L. Kelloff 33

Parmeliopsis minarum (Vainio) Elix & Hale Syrrhopodon elongatus Sullivant var. Parmeliopsis subfatiscens (Kurokowa) Elix glaziovii (Hampe) Reese & Hale Syrrhopodon hornschuchii Martius Parmotrema latissumum (F‚e) Hale Syrrhopodon leprieurii Montin Parmotrema mellissii (Dodge) Hale Syrrhopodon rigidus Hooker & Greville Parmotrema subochraceum Hale Syrrhopodon simmondsii Steere Parmotrema sulphuratum (Nees & Flotow) Syrrhopodon tortilis Hampe Hale Dicranaceae Pseudoparmelia sphaerospora (Nylander) Bryohumbertia filifolia (Hornschuch) Frahm Hale Campylopus cubensis Sullivant Relicina abstrusa (Vainio) Hale Campylopus richardii Bridel Usnea sp. Campylopus savannarum (Müller Halle) Pertusariaceae Mitten Pertusaria velata (Turner) Nylander Campylopus surinamensis Müller Halle Ramalinaceae Campylopus trachyblepharon (Müller Halle) Ramalina anceps Nylander Mitten Thelotremataceae Eucamptodontopsis pilifera (Mitten) Myriotrema neofrondosum Sipman Brotherus Myriotrema wrightii (Tucker) Hale Leucoloma cruegerianum (Müller Halle) A. Ocellularia auberiana (Montagne) Hale Jaeger Thelotrema sp. Leucoloma mariei Bescherelle Trichotheliaceae Leucoloma serrulatum Bridel Clathroporina sp. Fissidentaceae Trypetheliaceae Fissidens diplodus Mitten var. diplodus Polymeridium catapastum (Nylander) R.C. Fissidens inaequalis Mitten Harris Fissidens radicans Montagne Polymeridium simulans R.C. Harris Hookeriaceae Trypethelium aeneum (Eschweiler) Brymela parkeriana (Hooker & Greville) Zahlbruckner Buck Trypethelium sp. Callicosta bipinnata (Schwaegrichen) Müller Halle Callicostella pallida (Hornschuch) Ångström Hookeriopsis cruegeriana (Müller Halle.) Jacquin BRYTOPHYTES Hookeriopsis parkeriana (Hooker & Greville) Jacquin Bartramiaceae Hypnella pallescens (Hooker) A. Jaeger Philonotis uncinata (Schwaegrichen) Bridel Lepidopilidium portoricense (Müller Halle) Philonotis uncinata (Schwaegrichen) Bridel Crum & Steere var. glaucescens (Hornschuch) J. Lepidopilum radicale Mitten Florschütz Lepidopilum scabrisetum (Schwaegrichen) Bryaceae Steere Bryum apiculatum Schwaegrichen Lepidopilum surinamense Müller Halle Bryum billardieri Schwaegrichen Thamniopsis killippii (Williams) E.B. Bryum ellipsifolium Müller Halle Bartram Calymperaceae Hypnaceae Calymperes lonchophyllum Schwaegrichen Ectropothecium leptochaeton Calymperes nicaraguense Renauld & Cardot (Schwaegrichen) W.R. Buck Calymperes platyloma Mitten Isopterygium tenerum (Swartz) Mitten Calymperes venezuelanum (Mitten) Brotherus Rhacopilopsis trinitensis (Müller Halle.) ex Pittier Britton & Dixon 34 Kaieteur National Park: C.L. Kelloff

Leucobryaceae Leucobryum crispum Müller Halle Leucobryum martianum (Hornschuch) Hampe Octoblepharum albidum Hedwig Octoblepharum cocuiense Mitten Octoblepharum stramineum Mitten Leucomiaceae Leucomium strumosum (Hornsch.) Mitten Meteoriaceae Squamidium leucotrichum (Taylor) Brotherus Zelometeorium patulum (Hedwig) Manuel Neckeraceae Isodrepanium lentulum (Wilson) E. Britton Orthotrichaceae Groutiella obtusa (Mitten) Florsch tz Macromitrium cirrosum (Hedwig) Bridel Macromitrium podocarpi C. Mueller Macromitrium portoricense Williams Macromitrium punctatum (Hooker & Greville) Bridel Macromitrium sp. Macromitrium trinitense Williams Phyllodrepaniaceae Phyllodrepanium falcifolium (Schwaegrichen) Crosby Pottiaceae Trichostomum brachydontium Bruch Trichostomum duidense E.B. Bartram Pterobryaceae Orthostichopsis praetermissa W.R. Buck Orthostichopsis tetragona (Swartz ex Hedwig) Brotherus Renauldia paradoxica B.H. Allen Rhizogoniaceae Pyrrhobryum spiniforme (Hedwig) Mitten Sematophyllaceae Acroporium pungens (Hedwig) Brotherus Potamium vulpinum (Montagne) Mitten Sematophyllum subsimplex (Hedwig) Mitten Taxithelium concavum (Hooker) Spruce Taxithelium planum (Bridel) Mitten Trichosteleum bolivarense H. Robinson Trichosteleum hornschuchii (Hampe) A. Jaeger var. subglabratum J. Florschütz Kaieteur National Park: C.L. Kelloff 35

Appendix C: INVERTEBRATES OF KAIETEUR NATIONAL PARK, GUYANA; BUTTERFLIES, MOTHS, AND TERMITES

Butterflies and Moths Eveides species This list represents a fairly complete summary Heliconius aoede of the true butterflies species encountered in the Heliconius xanthocles upper gorge between Tukeit and Kaieteur Falls Heliconius numata / ethilla and on the plateau near the falls. This list of Heliconiius melpomene / erato approximately 170 species is no where near Heliconius sara complete. A few hundred more species should Heliconius antiochus be found there and hesperide (skippers) which are not represented on the list should add at least Nymphalidae: Nymphalinae a couple of hundred more species. Prepared by Eresia eunice Steven Fratello, 2001 and reviewed by Don Tigridia acesta Harvey and Robert Robbins (SI). Ectima thecla Nessaea batesii Eunica viola Papilionidae (Swallowtails) Callicore cynosura Battus belus Dynamine tithia Eurytides dolicaon Adelpha sp. Eurytides protesilaus – complex Metamorpha stelenes Eurytides ariarathes Junonia lavinia Parides chabrias mithras Junonia sp. Parides sesostris Anartia jatrophae Parides aeneas Parides vertumnus Nymphalidae: Charaxinae Parides lysonder Agrias pericles Parides echemon Agrias claudia Papilio thoas Archaeoprepona sp. Prepona eugenes Pieridae (Sulphurs and Whites) Fountainea sp. Moschoneura pinthaeus Memphis sp. Appias drusilla Hypna clytemnestra Aphrissa statira Siderone marthesia Phoebis argante Phoebris philea Nymphalidae: Morphinae (Morphos) Phoebis trite Morpho iphidamus Leucidia brephos Morpho hecuba Morpho adonis Nymphalidae: Ithomiinae (Glass-Wings) Morpho rhetenor Melinaea “mneme” Morpho menelaus Melinaea satevis Morpho deidamia Mechanitis mazaeus Morpho achilles / helenor Mechanitis polymnia Napeogenes cyrianassa Nymphalidae: Satyrinae Hypothynis ninonia (Satyus, Wood Nymphs) Clearwing species – undetermined Pierella rhea Pierella astyoche Nymphalidae: Heliconiinae (Long-Wings) Pierella lena Philaethria dido Pierella hyalinus Dryas julia Haetera piera 36 Kaieteur National Park: C.L. Kelloff

Cithaerias andromeda Synargis abaris Bia actorion Synargis sp. Euptychia hesione Nymphidium caricae Euptychia calpurnia Nymphidium acherois Euptychia terrestris Nymphidium cachrus Euptychia batesii Nymphidium menalcus Chloreuptychia arnaea Nymphidium lisimon Euptychia tricolor Nymphidium derufuta Euptychia picea Nymphidium baeotia complex Erichthodes erichtho Stalachtis calliope Stalachtis phaedusa Riodinidae (Metalmarks) Stalachtis euterpe Euselasia eumedia Lemonias egaensis Euselasia issoria Lemonias zygia Euselasia evodias Theope sp. Euselasia melaphaea Theope nycteis Euselasia nr. inconspicua Sarota chrysus Euselasia gelanor Emesis lucinda Euselasia midas Menander hebrus Euselasia lycaeus Thisbe irenea Methone cecilia Echenais thelephus Helicupis cupido Calospila rhodope Perophthalma tullius Calospila emylius Leucochimona hyphea Adelotypa huebneri Mesosemia nina Mesosemia cippus Lycaenidae (Hairstreaks) Mesosemia ibycus Calycopis sp. Mesosemia methion Calycopis cerata Mesosemia maera Panthiades bitias Mesosemia coea Panthiades aeolus Mesosemia melaene Lamprospilus collucia Mesosemia philocles Paiwarria telemus Mesosemia machaera Evenus sp. Mesosemia magete Theritas mavors Mesosemia phace Theritas hemon Mesosemia ulrica Theritas lisus Hyphilaria anthias Ocaria thales Hyphilaria nicia Arawacus aetolus Eurybia nicaeus Chalybs janias Eurybia nr. nicaeus Thestius pholeus Pheles heliconides Janthecla sista Ithomiola floralis Theclopsis lydus Alesa amesis Thecla aruma Napaea sp. nov. Thecla aff. adela Napaea nepos Thecla ligurina Cremna actoris Thecla tarena Cremna sp. Thecla athymbra Eunogyra satyrus Thecla munditia Ancyluris meliboeus Celmia celmus Rhetus periander Charis cleonus Charis anius Juditha azan Kaieteur National Park: C.L. Kelloff 37

TERMITES (Isoptera)

Rhinotermitidae Dolichorhinotermes longilabius Dolichorhinotermes nr. tenebrosus Heterotermes tenuis Coptotermes testaceus

Termitidae Apicotermitinae Anoplotermes banksi Anoplotermes sp. A Anoplotermes sp. B Anoplotermes sp. C Termitinae Cylindrotermes parvignathus Neocapritermes sp. A Termes fatalis Nasutitermitinae Arimitermes minutus Embiratermes sp. A Araujotermes parvellus Coatitermes kartaboensis Nasutitermes banksi Nasutitermes intermedius Nasutitermes gaigei Nasutitermes sp. A Nasutitermes sp. B Nasutitermes sp. C Nasutitermes sp. D 38 Kaieteur National Park: C.L. Kelloff

Appendix D: VERTEBRATES OF KAIETEUR NATIONAL PARK, GUYANA: BIRDS, HERPETOFAUNA, AND MAMMALS

BIRDS Compiles from field observations of M. Braun, EURYPYGIDAE C. Milensky, B. O’Shea, and D.W. Finch. >>> Sunbittern (Eurypyga helias) Based on >>>>>> SCOLOPACIDAE TINAMIDAE Solitary Sandpiper (Tringa solitaria) Great Tinamou (Tinamus major) Spotted Sandpiper (Actitis macularia) Variegated Tinamou (Crypturellus variegatus) Least Sandpiper (Calidris minutilla) White-Rumped Sandpiper (Calidris ARDEIDAE fuscicollis) Zigzag Heron (Zebrilus undulatus) Striated Heron (Butorides striatus) COLUMBIDAE Rufescent Tiger-Heron (Tigrisoma lineatum) Scaled Pigeon (Columba speciosa) Great Egret (Ardea alba) Ruddy Pigeon (Columba subvinacea) Plumbeous Pigeon (Columba plumbea) THRESKIORNITHIDAE Common Ground-Dove (Columbina passerina) Green Ibis (Mesembrinibis cayennensis) Gray-fronted Dove (Leptotila rufaxilla) White-tipped Dove (Leptotila verreauxi) CATHARTIDAE Ruddy Quail-Dove (Geotrygon montana) Turkey Vulture (Cathartes aura) Greater Yellow-headed Vulture (Cathartes PSITTACIDAE melambrotus) Red-and-Green Macaw (Ara chloroptera) King Vulture (Sarcoramphus papa) Painted Parakeet (Pyrrhura picta) Parrolet species (Forpus sp.) ACCIPITRIDAE Black-headed Parrot (Pionites melanocephala) Pearl Kite (Gampsonyx swainsonii) Dusky Parrot (Pionus fuscus) White Hawk (Leucopternis albicollis) Blue-headed Parrot (Pionus menstruus) Great Black Hawk (Buteogallus urubitinga) Orange-winged Parrot (Amazona amazonica) Roadside Hawk (Buteo magnirostris) Mealy Parrot (Amazona farinosa) White-tailed Hawk (Buteo albicaudatus) Red-fan Parrot (Deroptyus accipitrinus)

FALCONIDAE CUCULIDAE Barred Forest-Falcon (Micrastur ruficollis) Black-bellied Cuckoo (Piaya melanogaster) Red-throated Caracara (Daptrius americanus) Smooth-billed Ani (Crotophaga ani) Yellow-headed Caracara (Milvago chimachima) STRIGIDAE Merlin (Falco columbarius) Tawny-bellied Screech-Owl (Otus watsoni) Laughing Falcon (Herpetotheres cachinnans) Bat Falcon (Falco rufigularis) CAPRIMULGIDAE Orange-breasted Falcon (Falco deiroleucus) Paurague (Nyctidromus albicollis)

CRACIDAE APODIDAE Little Chachalaca (Ortalis motmot) White-Collared Swift (Streptoprocne zonaris) Blue-throated Piping-Guan (Pipile White-Chinned Swift (Cypseloides cryptus) cumanensis) Black Curassow (Crax alector) White-tipped Swift (Aeronautes montivagus) Band-rumped Swift (Chaetura spinicauda) Kaieteur National Park: C.L. Kelloff 39

DENDROCOLAPTIDAE TROCHILIDAE Wedge-billed Woodcreeper (Glyphorynchus Long-tailed Hermit (Phaethornis spirurus) superciliosus) Chestnut-rumped Woodcreeper Straight-billed Hermit (Phaethornis bourcieri) (Xiphorhynchus pardalotus) Reddish Hermit (Phaethornis ruber) Plain-brown Woodcreeper (Dendrocincla White-necked Jacobin (Florisuga mellivora) fuliginosa) Tufted Coquette (Lophornis ornatus) Lineated Woodcreeper (Lepidocolaptes Fork-tailed Woodnymph (Thalurania furcata) albolineatus) Green-tailed Goldenthroat (Polytmus theresiae) THAMNOPHILIDAE Glittering-throated Emerald (Amazilia Mouse-colored Antshrike (Thamnophilus fimbriata) murinus) Black-eared Fairy (Heliothryx aurita) Dusky-throated Antshrike (Thamnomanes ardesiacus) TROGONIDAE Cinereous Antshrike (Thamnomanes caesius) White-tailed Trogon (Trogon viridis) Spot-Tailed Antwren (Herpsilochmus Black-throated Trogon (Trogon rufus) sticturus) Black-tailed Trogon (Trogon melanurus) Todd's Antwren (Herpsilochmus Violaceous Trogon (Trogon violaceus) stictocephalus) Pygmy Antwren (Myrmotherula brachyura) BUCCONIDAE Rufous-bellied Antwren (Myrmotherula Swallow-wing (Chelidoptera tenebrosa) guttata) White-flanked Antwren (Myrmotherula ALCEDINIDAE axillaris) Ringed Kingfisher (Ceryle torquata) Long-winged Antwren (Myrmotherula Amazon Kingfisher (Chloroceryle amazona) longipennis) Green Kingfisher (Ghloroceryle americana) Gray Antwren (Myrmotherula menetriesii) Ash-winged Antwren (Terenura spodioptila) GALBULIDAE Gray Antbird (Cerocmacra cinarescens) Paradise Jacamar (Galbula dea) Scale-backed Antbird (Hylophylax Yellow-Billed Jacamar (Galbula albirostris) poecilinota) Warbling Antbird (Hypocnemis cantator) RAMPHASTIDAE Black-chinned Antbird (Hypocnemoides Channel-billed Toucan (Ramphastos melanopogon) vitellinus) Ferruginous-backed Antbird (Myrmeciza Red-billed Toucan (Ramphastos tucanus) ferruginea) Black-spotted Barbet (Capito niger) Spot-winged Antbird (Schistocichla Black-necked Aracari (Pteroglossus aracari) leucostigma) White-plumed Antbird (Pithys albifrons) PICIDAE Yellow-throated Woodpecker (Piculus FORMICARIIDAE flavigula) Black-faced Antthrush (Formicarius analis) Lineated Woodpecker (Dryocopus lineatus) Thrush-like Antpitta (Myrmothera Waved Woodpecker (Celeus undatus) campanisona)

Red-necked Woodpecker (Campephilus TYRANNIDAE rubricollis) White-lored Tryannulet (Ornithion inerme) Forest Elaenia (Myiopagis gaimardii) FURNARIIDAE Tropical Kingbird (Tyrannus melancholicus) Foliage-gleaner sp. (Philydor sp.) White-ringed Flycatcher (Conopias parva) Plain Xenops (Xenops minutus) Yellow-throated Flycatcher (Conopias parva) Rusty-margined Flycatcher (Myiozetetes 40 Kaieteur National Park: C.L. Kelloff

cayanensis) HIRUNDINIDAE Bright-rumped Attila (Attila spadiceus) White-winged Swallow (Tachycineta Grayish Mourner (Rhytipterna simplex) albiventer) Dusky-capped Flycatcher (Myiarchus White-banded Swallow (Atticora fasciata) tuberculifer) Barn Swallow (Hirundo rustica) Cliff Flycatcher (Hirundinea ferruginea) Helmeted Pygmy Tyrant (Lophotriccus TROGLODYTIDAE galeatus) Coraya Wren (Thryothorus coraya) Yellow-margined Flycatcher (Tolmomyias Southern House-Wren (Troglodytes assimilis) musculus) Common Tody-Flycatcher (Todirostrum White-breasted Wood-Wren (Henicorhina cinereum) leucosticta) Ringed Antpipit (Corythopis torquata) Wing-banded Wren (Microcerculus bambla) Plain-crested Elaenia (Elaenia cristata) Musican Wren (Cyphorhinus arada) Rufous-crowned Elaenia (Elaenia ruficeps) Long-billed Gnatwren (Ramphocaenus Yellow-bellied Elaenia (Elaenia flavogaster) melanurus) Lesser Elaenia (Elaenia chiriquiensis) Ochre-bellied Flycatcher (Mionectes TURDIDAE oleagineus) Cocoa Thrush (Turdus fumigatus) McConnell's Flycatcher (Mionectes macconnelli) PARULIDAE Slender-footed Tyrannulet (Zimmerius River Warbler (Phaeothlypos flaveolus) gracilipes) Ruddy-tailed Flycatcher (Terenotriccus COEREBIDAE erythrurus) Bananaquit (Coereba flaveola) Screaming Piha (Lipaugus vociferans) THRAUPIDAE COTINGIDAE Purple Honeycreeper (Cyanerpes caeruleus) Purple-breasted Cotinga (Cotinga cotinga) Red-legged Honeycreeper (Cyanerpes Pompadour Cotinga (Xipholena punicea) cyaneus) Guianan Cock-of-the-Rock (Rupicola Green Honeycreeper (Chlorophanes spiza) rupicola) Purple Honeycreeper (Cyanerpes caeruleus) Capuchinbird (Perissocephalus tricolor) Blue Dacnis (Dacnis cayana) White Bellbird (Procnias alba) Spotted Tanager (Tangara punctata) Bay-headed Tanager (Tangara gyrola) PIPRIDAE Burnished-buff Tanager (Tangara cayana) White-throated Manakin (Corapipo gutturalis) Purple-throated Euphonia (Euphonia Golden-headed Manakin (Pipra chlorotica) erythrocephala) Orange-bellied Euphonia (Euphonia White-crowned Manakin (Pipra pipra) xanthogaster) Tepui Manakin (Lepidothrix suavissima) Plumbeous Euphonia (Euphonia plumbea) Tiny Tyrant-Manakin (Tyranneutes virescens) Blue-gray Tanager (Thraupis episcopus) Thrush-like Manakin (Schiffornis turdinus) Palm Tanager (Thraupis palmarum) Silver-beaked Tanager (Ramphocelus carbo) VIREONIDAE Fulvous Shrike-Tanager (Lanio fulvus) Buff-cheeked Greenlet (Hylophilus Flame-crested Tanager (Tachyphonus muscicapinus) cristatus) Tawny-crowned Greenlet (Hylophilus White-lined Tanager (Tachyphonus rufus) ochraceiceps) Fulvous-crested Tanager (Tachyphonus surinamus) CORVIDAE Red-shouldered Tanager (Tachyphonus Cayenne Jay (Cyanocorax cayanus) phoeniceus) Kaieteur National Park: C.L. Kelloff 41

EMBERIZIDAE Slate-colored Grosbeak (Pitylus grossus) Dendrobatidae (poison arrow frogs) Yellow-green Grosbeak (Caryothraustes Colostethus beebei canadensis) Colostethus brunneus Blue-black Grassquit (Volatinia jacarina) Hylidae Gastrotheca sp. nov. ICTERIDAE Hyla boans Green Oropendola (Psarocolius viridis) Phrynohyas venulosa Red-rumped Cacique (Cacicus haemorrhous) Stefania woodleyi Moriche Oriole (Icterus chrysocephalus) Leptodactylidae Leptodactylus fuscus Leptodactylus longirostris Leptodactylus rugosus HERPETOFAUNA Microhylidae Prepared by C.J. Cole and C. R. Townsend, Synapturanu mirandaribeiro American Museum of Natural History; R. Reynolds, U.S. Department of Agriculture MAMMALS Snakes This list of the mammals has been compiled Colubrida using Lim and Engstrom (ms.) and Engstrom Liophis purpurans and Lim (2001) and common names, where Liophis typhlus available, from Emmons (1997). Notation has Viperidae been made on whether the species has been Bothrops atrox (Laborea) collected (V), observed (OBS) or expected (E) based on distribution. Lizards Amphisbaenidae Amphisbaena fuliginosa ORDER DIDELPHIMORPHIA (opossums) Gekkonidae Family Didelphidae (opossums) Gonatodes annularis Subfamily Caluromyinae (woolly opossums) Gymnophthalmidae Caluromys philander (Bare-tailed woolly Neusticusus rudis oppossum) - E Iguanidae (Iguana family) Plica umbra Subfamily Didelphinae (opossums) Polychrotidae Chironectes minimus (Water opossum) Anolis fuscoauratus Didelphis imperfectus E Anolis sp. Didelphis marsupialis (Common opossum)E Scincidae Marmosa murina (Murine mouse opossum)- Mabuya sp. E Teiidae Marmosops pinheiroi - E Ameiva ameiva Metachirus nudicaudatus E Bachia monodactylus Micoureus demerarare E Cnemidophorus sp. Monodelphis brevicaudata (red-legged Dracaena guianensis opossum) E Kentripyx calcaratus Philander opossum (Common gray four-eyed opossum) E Frogs Bufonidae Bufo guttatus ORDER XENARTHRA (xenarthrans) Bufo marinus Family Bradypodidae (three-toed sloths) Bufo nasicus Bradypus tridactylus (pale-throated sloth) E Bufo typhonius Family Megalonychidae (two-toes sloths) 42 Kaieteur National Park: C.L. Kelloff

Choloepus didactylus (southern sloth) E Micronycteris megalotis (little big-eared bat) E Family Dasypodidae (armadillos) Micronycteris microtis (little big-eared bat) E Dasypus novemcinctus (nine-banded long Micronycteris minuta (little big-eared bat) E nose armadillo) E Micronycteris nicefori (little big-eared bat)E Priodontes maximus (giant armadillo) E Micronycteris sylvestris (little big-eared bat) E Family Myrmecophagidae (anteaters) Mimon crenulatum (hairy-nosed bat) E Cyclopes didactylus (silky or pigmy anteater) Phylloderma stenops (pale-face bat) E E Phyllostomus discolor (spear-nosed bat) E Myrmecophaga tridactyla (giant anteater) E Phyllostomus elongatus (spear-nosed bat) E Tamandua tetradactyla (collared or southern Phyllostomus hastatus (spear-nosed bat) E tamandua) E Tonatia brasiliense (round-eared bat) E Tonatia carrikeri (round-eared bat) E Tonatia saurophila (round-eared bat) E ORDER CHIROPTERA (bats) Tonatia silvicola (round-eared bat) E Family Emballonuridae (sheath-tailed bats) Trachops cirrhosus (fringe-lipped bat) E Cormura brevirostris (chestnut sac-winged Vampyrum spectrum (false vampire bat) E bat) V Diclidurus scutatus (ghost bat) E Subfamily Glossophaginae (nectar-feeding Peropteryx kappleri (doglike sac-winged bat) bats) E Anoura caudifera (hairy-legged bat) V Peropteryx macrotis (doglike sac-winged bat) Anoura geoffroyi (hairy-legged bat) V E Choeroniscus minor (long-nosed bat) E Rhynchonycteris naso (long-nosed bat) E Glossophaga longirostris (common nectar- Saccopteryx bilineata (white-lined sac-winged feeding bat) E bat) E Glossophaga soricina (common nectar- Saccopteryx canescens (white-lined sac- feeding bat) E winged bat) E Lionycteris spurrelli (dark nectar-feeding bat) Saccopteryx leptura (white-lined sac-winged V bat) E Lonchophylla thomasi (spear-nosed nectar- feeding bat) V Family Noctilionidae (bulldog bats) Noctilio albiventris (Lesser Bulldog bat) E Subfamily Carolliinae (New World fruit bats) Noctilio leporinus (Bulldog bat) E Carollia brevicauda (short-tailed bat) E Carollia perspicillata (short-tailed bat) V Family Mormoopidae (leaf-chinned bats) Rhinophylla pumilio (little fruit bat) V Pteronotus gymnonotus E Pteronotus parnellii V Subfamily Stenodermatinae (New World Pteronotus personatus E fruit-eating bats) Family Phyllostomidae Ametrida centurio (little white shouldered (New World leaf-nosed bats) bat) E Subfamily Phyllostominae (New World leaf Artibeus amplus (large fruit-eating bat) V nosed bats) Artibeus cinereus (large fruit-eating bat) E Chrotoperus auritus (woolly false vampire Artibeus concolor (large fruit-eating bat) E bat) E Artibeus gnomus (large fruit-eating bat) E Lonchorhina aurita (common sword-nosed Artibeus lituratus (large fruit-eating bat) E bat) E Artibeus obscurus (large fruit-eating bat) E Macrophyllum macrophyllum (long-legged Artibeus planirostris (large fruit-eating bat) E bat) E Chiroderma trinitatum (big-eyed bat) E Micronycteris brachyotis (little big-eared bat) Chiroderma villosum (big-eyed bat) E E Mesophylla macconnelli (Macconnell's bat) Micronycteris hirsuta (little big-eared bat) E E Kaieteur National Park: C.L. Kelloff 43

Platyrrhinus aurarius E Subfamily Cebinae (New World monkeys) Platyrrhinus brachycephalus E Cebus olivaceus (wedge-capped capuchin) Platyrrhinus helleri E OBS Sturnira lilium (yellow-shouldered bat) E Saimiri sciureus (common squirrel monkey) Sturnira tildae (yellow-shouldered bat) E E Uroderma bilobatum (tent-making bat) E Vampyressa bidens (yellowed-eared bat) E Subfamily Pitheciinae (sakis and uakaris) Vampyrodes caraccioli (yellowed-eared bat) Chiropotes satanas (brown bearded saki) E E Pithecia pithecia (guianan saki) E

Subfamily Desmodontinae (vampire bats) Desmodus rotundus (common vampire bat) E ORDER CARNIVORA (carnivores) Diaemus youngi (white-wing vampire bat) E Family Canidae (dogs) Cerdocyon thous E Family Furipteridae (thumbless bats) Speothos venaticus (bush dog) OBS Furipterus horrens (thumbless bat) E Family Felidae (cats) Family Thyropteridae (disc-winged bats) Subfamily Felinae (cats) Thyroptera tricolor (sucker-footed bat) E Herpailurus yaguarondi (jaguarundi) OBS Leopardus pardalis (ocelot) E Family Vespertilionidae (vesper bats) Leopardus tigrinus (oncilla) E Eptesicus furinalis (big brown bat) E Leopardus wiedii (margay) E Lasiurus atratus (hairy-tailed bat) E Puma concolor (puma) E Lasiurus blossevillii (hairy-tailed bat) E Lasiurus ega (hairy-tailed bat) E Subfamily Pantherinae (large cats) Myotis albescens (little brown bat) E Panthera onca (jaguar) E Myotis nigricans (little brown bat) V Myotis riparius (little brown bat) E Family Mustelidae (weasels) Subfamily Lutrinae (otters) Family Molossidae (free-tailed bats) Lontra longicaudis (southern river otter) E Cynomops abrasus (doglike bat) E Pteronura brasiliensis (giant river otter) OBS Cynomops paranus (doglike bat) E ? Cynomops planirostris (doglike bat) E Eumops auripendulus (bonneted bat) E Subfamily Mustelinae (weasels) Eumops glaucinus (bonneted bat) E Eira barbara (tayra) E Eumops hansae (bonneted bat) E Eumops trumbulli (bonneted bat) E Family Procyonidae (raccoons and allies) Molossus ater (dog-faced bat) E Subfamily Potosinae (kinkajous and olingos) Molossus molossus (dog-faced bat) E Potos flavus (kinkajou) E Nyctinomops laticaudatus (broad-eared bat) E Nyctinomops macrotis (broad-eared bat) E Subfamily Procyoninae (raccoons and coatis) Promops centralis (crested bat) E Nasua nasua (South American coatis) E Promops nasutus (crested bat) E Procyon cancrivorus (crab dog) OBS

ORDER PRIMATES (primates) ORDER PERISSODACTYLA (odd-toed Family Callitrichidae (marmosets) ungulates) Saguinus midas (Golden-handed tamerin) E Family Tapiridae (tapirs) Tapirus terrestris (Brazilian tapir) E Family Cebidae (New World monkeys) Subfamily Alouattinae (howler monkeys) Alouatta seniculus (red howler monkey) OBS 44 Kaieteur National Park: C.L. Kelloff

ORDER ARTIODACTYLA (even-toed unugulates) Family Tayassuidae (peccaries) Pecari tajacu (collared peccary) OBS Tayassu pecari (whited-lipped peccary) E

Family Cervidae (deer) Mazama americana (red brocket deer) OBS Odocoileus cariacou E

ORDER RODENTIA (rodents) Family Sciuridae (squirrels) Sciurus aestuans (Guianan squirrel) E

Subfamily Sigmodontinae (New World rats and mice) Holochilus sciureus (marsh rat) E Neacomys guianae (spiny rat) E Neacomys paracou (spiny rat) E Nectomys melanius (spiny rat) E Oecomys bicolor (arboreal rice rat) E Oligoryzomys fulvescens (pygmy rice rat) E Oryzomys macconnelli (rice rat) E Oryzomys megacephalus (rice rat) E Oryzomys yunganus (rice rat) E Rhipidomys leucodactylus (climbing rat) E Rhipidomys nitela (climbing rat) E Sigmodon alstoni E

Family Erethizontidae (porcupines) Coendou prehensilis (Brazilian porcupine) E

Family Caviidae (cavies) Cavia aperea (cavy) E

Family Hydrochaeridae (capybaras) Hydrochaeris hydrochaeris (capybara) E

Family Dasyproctidae (agoutis and acouchies) Dasyprocta leporina (red-rumped agouti) E

Family Cuniculidae (pacas) Cuniculus paca (paca) OBS

Subfamily Echimyinae (spiny rats) Makalata didelphoides (tree rat) E

Subfamily Eumysopinae (spiny rats) Mesomys hispidus (spiny tree rat) E Proechimys cuvieri (spiny rat) E Proechimys guyannensis (spiny rat) E Kaieteur National Park: C.L. Kelloff 45

Plate 1: Aerial view of Kaieteur Falls. Plate 3: The giant bromeliad, Brocchina micrantha (Baker) Mez (Bromeliaceae) in flower.

Plate 2: Vanilla bicolor Lindley (Orchidaceae) found in the savannas in the vicinity of Kaieteur Falls. Photo by M. Yoshpa.

Plate 4: The golden frog, Colostethus beebei is endemic to Guyana and can be found living in the tanks of the Brocchina micrantha (Baker) Mez at Kaieteur. 46 Kaieteur National Park: C.L. Kelloff

Plate 5: Rapatea paludosa Aublet (Rapateaceae) grows in the wet sandy soils of the bush islands at Kaieteur.

Plate 7: One of the cement steps built during British rule along the path to Johnson’s View.

Plate 6: The cabbage head bromeliad, Plate 8: Aerial view of the savanna of the Aechmea brassicoides Baker is endemic to Potaro Plateau just south of Kaieteur Falls. Kaieteur.