Forest Structure and Tree Species Composition of the Grand Etang Forest on Grenada, West Indies, Pre-Hurricane Ivan
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Caribbean Journal of Science, Vol. 44, No. 3, 395-401, 2008 Copyright 2008 College of Arts and Sciences University of Puerto Rico, Mayagu¨ez Forest structure and tree species composition of the Grand Etang Forest on Grenada, West Indies, pre-Hurricane Ivan MARY E. GLENN1,2 AND KEITH J. BENSEN2 1Department of Anthropology, Humboldt State University, Arcata, CA 95521 USA 2Windward Islands Research and Education Foundation, Grenada, West Indies. Contact information for corresponding author: Tel: (707) 826-3126 Fax: (707) 826-4418 Email: [email protected] ABSTRACT.—The last comprehensive native forest structure survey made in the Grand Etang National Park and Forest Reserve of Grenada was in the 1940s, before Hurricane Janet hit the island in 1955. The data from this study were collected in the mid-1990s, previous to the devastation of Hurricane Ivan in 2004, the first hurricane to strike Grenada since 1955. A total of 584 trees representing 41 species were identified in twenty 2m × 50m plots. Mean tree height, basal area per hectare, tree density per hectare, and species richness were determined for the park and forest reserve. Licania ternatensis was the dominant tree in terms of density per hectare (18%), while Dacryodes excelsa was dominant in terms of basal area per hectare (30.3 m2). Trichillia septentrionalis had the largest mean height (23.6 m) even though the tallest tree in the sample was a Dac- ryodes excelsa (51.9 m). The composition of the Grand Etang forest in the 1990s consisted of a mosaic of stands of different size classes and species composition, most likely indicating varying damage and recovery from Hurricane Janet. Additional surveys are recommended to reassess Grenada’s Grand Etang native rainforest structure in order to determine which tree species are recovering and at what rate after the devastation of Hurricane Ivan. These results could then be compared to the significant body of work done on ecologically similar hurricane affected wet/rain and lower montane wet/rain forests found on Puerto Rico. KEYWORDS.—Rainforest, Caribbean, Licania ternatensis, Dacryodes excelsa, Trichillia septentrionalis INTRODUCTION order to better understand the forest’s cur- rent condition and species survivorship In the Grand Etang National Park and and succession rates, future surveys de- Forest Reserve, only one comprehensive scribing Grand Etang’s forest structure and forest structure survey was made previous species composition should be conducted to this study (Beard 1944, 1949). In 1955, and then compared to the material pre- Hurricane Janet passed directly over sented here. Grenada and caused damage to the Grand Etang forest, altering much of its previous structure (Caribbean Conservation Asso- MATERIALS AND METHODS ciation 1991). Our forest structure and com- Study Site position surveys were conducted in the mid-1990s as part of a larger study concen- Grenada, which is located 150 km north trating on the natural history and ecology of South America, is the southernmost is- of the introduced mona monkey (Cercopi- land in the Lesser Antilles’ archipelago thecus mona) (e.g., Glenn 1996, 1997, 1998, (Figure 1). The island is 33.5 km long and Horsburgh et al. 2002). In 2004, Hurricane 19 km wide, and has a total area of approxi- Ivan devastated most of the Grand Etang mately 312 km2. The topography of Gre- forest. Our surveys were completed after nada consists of a series of highly eroded Hurricane Janet and before Hurricane Ivan, volcanic peaks (the highest being Mt. St. and provide a picture in time of a Lesser Catherine at 850 m) that run along the cen- Antillean rainforest almost a half century ter of the island. The lower elevations con- after a major disturbance and less than ten tain cultivated lands consisting mainly of years before an even larger disturbance. In tree crops (primarily nutmeg, cocoa, and 395 396 M. E. GLENN AND K. J. BENSEN FIG. 1. Location of the study site on the island of Grenada. bananas), dry scrub woodlands, and man- streams on the bottom of deep ravines and grove swamps that occur along shallow valleys. The elevation within the Grand bays and inlets. Most towns and villages Etang forest ranges between 340 and 710 m. are located at lower elevations and along The Grand Etang forest receives an aver- the coast. Upper elevations receive more age of 4,060 mm of rain per year (Grenada rain and have a thicker forest cover, with Government and Organization of Ameri- rain forest generally occupying elevations can States 1988). A total of 3,422 mm of rain above 350 m (Caribbean Conservation As- fell in the one continuous year (1993) dur- sociation 1991). Ridge tops and peaks are ing which climactic data were collected at covered in short, thick forest (termed “elfin the Grand Etang field station. Mean woodland”) that does not exceed 5 m in monthly rainfall at the field station was height (Beard 1949). 252.8 mm (SD = 143), and temperatures av- The Grand Etang National Park and For- eraged 21.3° C (SD = 1.3) over 28 months est Reserve on Grenada (12°6’ N, 61°42’ S) between September 1992 and April 1995. occupies approximately 1,540 ha of moun- Trade winds blow across the Grand Etang tainous, tropical rain forest (Figure 1). The forest year-round. forest reserve was originally set aside in 1902 in order to protect vital watersheds for Forest Structure Surveys lower elevation croplands. No tree harvest has been conducted within the reserve over Forest structure surveys were conducted the past century except within small areas between 28 October 1994 and 5 April 1995. that were replanted with exotic hardwoods Twenty, 2m × 50m plots were installed after the 1955 hurricane damaged parts of throughout the study area. Plots were the forest (Caribbean Conservation Asso- placed at random perpendicular distances ciation 1991). The terrain is characterized (0-500m, generated from a random num- by volcanic craters (one of which contains bers table) from monkey survey trails every the Grand Etang Lake), steep slopes, nar- 500m with the direction determined by the row ridges, and numerous small perennial flip of a coin. Several small plots were used GRAND ETANG FOREST STRUCTURE, GRENADA 397 instead of a few large plots because pre- primary and secondary, were identified by liminary surveys indicated that the Grand successional stages; the third forest type, Etang forest was highly variable in struc- palm brake (Beard 1949), was a specific ture and species composition over small ar- vegetation category. Primary forest was eas and had low overall species richness. characterized by emergent species such as Each tree within each plot was identified Dacryodes excelsa and Sloanea spp., while sec- to species and measured with a diameter- ondary forest was characterized by species at-breast-height (DBH) tape. Only trees of woody shrubs such as Psychotria berteri- with a DBH of 3 cm or greater were in- ana, Miconia spp., and Citharexylum spino- cluded in this analysis. Tree density was sum. Palm brake consisted of a minimum of calculated as the number of trees per spe- 33% and up to 60% of palm trees (Prestoea cies per hectare or the number of trees per montana and Euterpe dominicana). forest type per hectare. The basal area of Primary forest was the prevalent forest each tree was calculated from DBH mea- type and occurred in 11 of the 20 plots. Sec- surements using the following formula: ondary forest and palm brake occurred in (DBH/2)2 (). Basal area per species per six and three of the 20 plots, respectively. hectare and basal area per forest type per Primary forest had significantly lower spe- hectare were then derived from these cal- cies richness than secondary forest (t = 4.79, culations. Each tree within each plot was n = 17, p < 0.0001). In addition, primary measured for height with a clinometer. forest had a significantly lower tree density Canopy cover for each plot was visually per hectare (t = 4.79, n = 17, p < 0.0002) but estimated with a spherical densiometer. Fi- a significantly higher mean tree height (t = nal tree identification and taxonomic clas- 4.76, n = 533, p < 0.0005) than did second- sification were based on Howard’s Flora of ary forest. No differences were found be- the Lesser Antilles, Leeward and Windward Is- tween primary and secondary forest for lands (1989). Mean tree height, basal area basal area per hectare (t = 0.97, n = 17, ns). per hectare, tree density per hectare and Palm brake had the lowest mean tree den- species richness between primary and sec- sity per hectare, basal area per hectare, tree ondary forest types were compared using height, and species diversity of all three t-tests (one-tailed) with a 95% confidence forest types. No differences were found for level. percentage of canopy cover among forest types. Mean canopy coverage for all forest types combined was 97.8% (SD = 5.6, n = RESULTS 20) with a range between 75% and 100%. A total of 584 trees were measured within the 20 plots. A summary of tree spe- DISCUSSION cies composition and structure is provided in Table 1. Forty-one tree species were rep- The forest of Grenada’s Grand Etang Na- resented. Six of the 41 species were not tional Park and Forest Reserve during the identified. Unidentified trees, however, 1990s was similar to the subtropical wet/ made up only 2.3% (n = 12) of the total rain forest and lower montane wet/rain number of trees within the plots. Licania ter- forest types of Puerto Rico. These Puerto natensis was the dominant tree in terms of Rican forests receive similar amounts of density per hectare (18%, n = 525), whereas rainfall, are found at similar elevations and Dacryodes excelsa was dominant in terms of have been disturbed periodically by hurri- basal area per hectare (30.3 m2).