Checklist of Venezuelan Bromeliaceae with Notes on Species Distribution by State and Levels of Endemism

By Bruce K. Holst, Marie Selby Botanical Gardens, 811 South Palm Ave., Sarasota, FL 34236, USA

Note (12 Dec. 1996): This is a slightly modified version of a paper published in Volume 15, part 1, 1994 of the journal Selbyana, and reflects the generic circumscription that I accepted at that time. For updated generic combinations see: Luther & Sieff, An Alphabetical List of Bromeliad Binomials, Fifth Edition, April 1996, Bromeliad Society, Incorporated. Since the original publication of this paper in 1994, 5 species have been added to the Venezuelan bromeliad flora and are reflected in the numbers given in this version: Connellia nahoumii Leme (Bolívar), Lindmania candelabriformis B. Holst (Amazonas), Lindmania oliva-estevae B. Holst (Bolívar), Navia glandulifera B. Holst (Bolívar), and Navia tentaculata B. Holst (Bolívar).

ABSTRACT.

A checklist of the 24 genera and 365 native species of Bromeliaceae known from Venezuela is presented, including their occurrence by state and indications of which are endemic to the country. A comparison of the number of genera and species known from Mesoamerica (southern Mexico to Panama), Colombia, Venezuela, the Guianas (Guyana, Suriname, French Guiana), Ecuador, and Peru is presented, as well as a summary of the number of species and endemic species in each Venezuelan state.

RESUMEN.

Se presenta un listado de los 24 generos y 365 especies nativas de Bromeliaceae que se conocen de Venezuela, junto con sus distribuciones por estado y una indicación cuales son endémicas a Venezuela. Se presenta tambien una comparación del número de los géneros y especies de Mesoamerica (sur de Mexico a Panama), Colombia, Venezuela, las Guayanas (Guyana, Suriname, Guyana Francesa), Ecuador, y Perú, y un resumen del número de especies y número de especies endémicas de cada estado de Venezuela.

INTRODUCTION

The checklist of Venezuelan Bromeliaceae presented below (Appendix 1) adds three genera (Brewcaria, Neoregelia, and Steyerbromelia) and 72 species to the totals for the country since the last summary of Venezuelan bromeliads in the Flora de Venezuela series which contained 293 species (Smith 1971). The checklist, with 24 genera and 365 species, was originally prepared for use in citing species totals for Venezuela in the forthcoming Flora of the Venezuelan Guayana (Steyermark, Berry, & Holst in preparation, covering the southern half of Venezuela). Checklists such as this one, are also useful for comparing species totals from other countries and for assessing species diversity across geographical gradients, which help point out centers of diversity and directions of radiation.

METHODS

The checklist was prepared from both herbaria and literature sources. Herbaria surveyed were: Field Museum of Natural History (F), Missouri Botanical Garden (MO), Facultad de Agronomia-Universidad Central de Venezuela (MY), Herbario Ovalles, Universidad Central (MYF), New York Botanical Garden (NY), Herbario Universitario, Universidad de los Llanos (PORT), United States National Herbarium (US), and the Herbario Nacional de Venezuela (VEN). Principal literature references used were the Flora Neotropica Bromeliaceae treatments (Smith & Downs 1974, 1977, 1979), Flora de Venezuela Bromeliaceae (Smith 1971), and Revision of the Guayana Highland Bromeliaceae (Smith 1986). Several additional country records were reported in works by Smith & Read (1982), Luther (1984), Morillo (1986), and Oliva-Esteva & Steyermark (1987). Author abbreviations used in the checklist follow Brummit & Powell (1992). Abbreviation of literature citations follow Stafleu & Cowan (1976--1988) for books, and Lawrence et al. (1968) for journals.

States in Venezuela (Figure 1) where the species occur are listed after each species, and the word "Endemic" is added if the species is wholly confined within Venezuela. Species that occur very near the border with Venezuela, and selected excluded taxa are listed at the end of the checklist. Selected synonyms from the appendix are cross-referenced with accepted taxa at the end of the checklist.

Subspecies, varieties, and forms are either not included in the list, or in a few instances in Tillandsia, are merely listed after the species where widely recognized species have been recently reduced to infraspecific levels. Some of the infraspecific taxa that have been recognized in Venezuela are likely valid biological entities, but many also appear to be artifacts of incomplete collecting or poor understanding of variation within a species. Thorough monographs involving field studies are needed before most infraspecific taxa can attain greater taxonomic or biological meaning.

RESULTS AND DISCUSSION

Venezuela ranks among the top Neotropical countries in regard to diversity and number of Bromeliaceae genera and species. Species totals from recent checklists of the Guianas (Guyana, Suriname, French Guiana; Boggan et al., 1992), Ecuador (Luther 1989), and Peru (Brako & Zarucchi 1993), and floras from Mesoamerica (including southern Mexico to Panama; Utley & Burt-Utley, in press) and Colombia (Smith 1977, updated through 1988, Smith, pers. com.) are compared in Table 1 with a list of genera and species known from Venezuela. Brazil is not included, but when a list of species from that country is assembled, it will have the highest number of genera and species of Bromeliaceae of any country. It is also much larger in area (8,506,663 km²) than any other neotropical country and is the center of diversity of many genera, especially those of the Bromelioideae. An example of the high bromeliad diversity in Brazil was illustrated by Fontoura et al. (1991) where they found 20 genera and 245 species in the southeastern state of Rio de Janeiro alone (6321 km²).

On a per-area basis, Ecuador has the highest bromeliad species diversity of countries compared in Table 1, though its small size automatically gives it a relatively high concentration of species per area. Pitcairnioideae species are especially well represented in Venezuela due mostly to the presence of three genera that have centers of diversity there (Navia, Lindmania, and Brocchinia), and the Andes are species-rich in Tillandsioideae. The high number of genera in Venezuela is not only due to the Guayanan pitcairnioid genera (Ayensua, Brewcaria, Brocchinia, Connellia, Lindmania, Navia, and Steyerbromelia), but also because parts of Venezuela belong to several other important neotropical phytogeographical regions: Caribbean, Andean, and Amazonian.

Table 1. Species diversity of Bromeliaceae in some neotropical countries and regions.

1. Total species listed for the subfamilies and genera include all species known, not just those presented in the table. Numbers listed here are mostly from Luther (1991), with some minor modifications to the genera that are largely centered in Venezuela.
2. Only genera from countries and regions compared in the table are listed here.
3. Including Pepinia.

Table 2 summarizes information presented in the checklist of Venezuelan Bromeliaceae (Appendix 1) with the total number of species and number of endemic species per state. In addition, the number of species per 1000 km² is calculated for each state and overall species numbers are compared for southern and northern Venezuela.

Table 2. Bromeliad species diversity and number of endemic species per state in Venezuela.

1. Includes Amazonas, Bolívar, and Delta Amacuro.
2. Includes all other states not listed under Southern Venezuela.

The southern Venezuelan states of Amazonas and Bolívar are by far the richest in the country for bromeliad species, though they are also considerably larger than any other state. These states contain numerous, topographically varied (50--3,000 m elevation) habitats including mostly oligotrophic soils and large expanses of igneous and sandstone rock outcrops where many pitcairnioid species grow. The Llanos states of Anzoátegui, Apure, Barinas, Cojedes, and Guárico are relatively species-poor, they contain large expanses of mostly open-savannas with small amounts of rock outcrops, relatively few forested areas, and they are only slightly varied topographically (mostly below 500 m elevation). The richest states on a per-area basis are those of the coastal range (Aragua, Carabobo, Distrito Federal, Miranda, Yaracuy, Sucre) and the Andes (Mérida, Táchira, Trujillo). These states vary greatly in elevation (sea level to 5,000 m elevation) and contain numerous forest types; epiphytic Tillandsioideae are numerous in the montane habitats that occur there. The islands that form the state of Nueva Esparta are also species-rich for their size, though nearly all of the species that occur there are relatively widespread.

Summarizing the number of endemic species in political regions can be misleading, especially in the tropics where intensive exploration frequently follows country borders. The numbers presented in Table 2, however, point out the two Venezuelan states where there is a high amount of local endemism: Amazonas and Bolívar. This high number of endemic species can partially be explained by the fact that some of the most speciose genera that occur there have non- or only scarcely appendaged seeds (Navia, Lindmania, most Pitcairnia), and therefore poor dispersal abilities via wind. This implies that once a population becomes established away from its parents, it is likely to remain genetically isolated. The numerous, mostly steep- or vertical-walled mountains in these states provide a further barrier to dispersal. In addition, and for reasons that are not entirely clear, habitats with oligotrophic soils, which are frequent in Amazonas and Bolívar states, tend to contain higher numbers of local endemic species. The Llanos states do not have any endemic species of Bromeliaceae. Of the 395 species of Bromeliaceae that occur in Venezuela, 176 are endemic to the country.

There are an additional 8--10 unidentified taxa known from Venezuela that are either new species for the country or new for science, however there also some species included in the checklist presented here that are not well-defined species and may eventually need to be synonymized. Future collecting, though, will likely turn up even more species and new country records, and may push the total number of species for Venezuela closer to 400.

Several Guayana Shield bromeliad genera, particularly Lindmania and Navia, are in need of monographs that include extensive fieldwork. Many of the species in these genera are known only from their type collections and have proved difficult to recollect and study because of their remote localities. Recent collecting efforts in southern Venezuela, mainly by botanists from the herbarium of the University of the Llanos in Guanare (PORT) and Otto Huber, associated with the Herbario Ovalles in Caracas (MYF), have provided critical collections that will necessitate further refinements of generic limits and additions of species to the appended list of Venezuelan bromeliads. Some of these will be presented in the Flora of the Venezuelan Guayana.

ACKNOWLEDGMENTS.

I thank Paul Berry, Harry Luther, and Susan Murphy for providing useful comments on the manuscript, Gerrit Davidse for providing the Bromeliaceae genus and species totals for the Mesoamerican flora, Ivón Ramírez for making her thesis on Neoregelia subgenus Hylaeaicum available to me and for helpful comments, and Germán Carnevali for helping to add several additional species to the checklist.

LITERATURE CITED

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