Long-Term Changes in Structure and Composition Following Hurricanes in a Primary Lower Montane Rain Forest in Puerto Rico
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BOIS ET FORÊTS DES TROPIQUES, 2013, N° 317 (3) OURAGANS, DÉGÂTS ET STRUCTURE / LE POINT SUR… 7 Long-term changes in structure and composition following hurricanes in a primary lower montane rain forest in Puerto Rico Peter L. Weaver U.S. Department of Agriculture Forest Service International Institute of Tropical Forestry San Juan Puerto Rico Dacryodes excelsa Vahl (Burseraceae) in lower montane rain forest: The tree, called tabonuco in Puerto Rico and gommier in the Lesser Antilles, was used by indigenous peoples for making and caulking boats, for torches, as incense in religious ceremonies, and for medicinal purposes. During the early 20th century, the timber was used for making furniture and general construction. The scene shows the impacts of Hurricane Hugo on TR-1 in 1989. Photograph P. L. Weaver. BOIS ET FORÊTS DES TROPIQUES, 2013, N° 317 (3) 8 FOCUS / HURRICANES, DAMAGES & STRUCTURE P. L. Weaver RÉSUMÉ ABSTRACT RESUMEN CHANGEMENTS A LONG TERME DE LONG-TERM CHANGES IN STRUCTURE CAMBIOS A LARGO PLAZO DE ESTRUC- STRUCTURE ET DE COMPOSITION D’UNE AND COMPOSITION FOLLOWING TURA Y COMPOSICIÓN DE UN BOSQUE FORET PRIMAIRE HUMIDE DE BASSE HURRICANES IN A PRIMARY LOWER PRIMARIO PLUVIAL MONTANO BAJO MONTAGNE SUITE AU PASSAGE MONTANE RAIN FOREST IN PUERTO RICO TRAS EL PASO DE HURACANES EN D’OURAGANS A PUERTO RICO PUERTO RICO Dacryodes excelsa, espèce arborée Ridges within the lower montane rain Dacryodes excelsa, especie de árbol connue sous les noms de tabonuco à forests (sensu Beard) of the Caribbean conocido como tabonuco en Puerto Rico Puerto Rico et de gommier aux Petites Basin are dominated by Dacryodes y gommier en las Antillas Menores, Antilles, domine les sommets de la forêt excelsa, a tree species known as domina en las cumbres del bosque plu- tropicale humide de basse montagne tabonuco in Puerto Rico and gommier in vial montano bajo (según Beard) del (selon Beard) de l’archipel des Caraïbes ; the Lesser Antilles. Periodically, hurri- archipiélago del Caribe. Periódicamente, où périodiquement les ouragans passent canes traverse the islands causing los huracanes atraviesan las islas cau- à travers les îles modifiant la structure des changes in structure, species composi- sando cambios en la estructura, compo- massifs forestiers, leur composition en tion, and dynamics of forests. The sición de especies y dinámica de los bos- essences et leur dynamique. Au nord-est chronology of post-hurricane vegetation ques. La cronología de cambios post- de Puerto Rico, l’étude chronologique change from 1946 to 2010 on a 0.4 ha huracanes en la vegetación entre 1946 y post-ouragans menée entre 1946 et 2010 ridge plot ~380 m in elevation in the 2010 en una parcela de 0.40 hectárea a a permis de suivre les modifications de la Luquillo Experimental Forest of northeast- unos 380 metros de altitud en el Bosque végétation sur une parcelle de 0,40 hec- ern Puerto Rico showed: (1) defoliation, Experimental de Luquillo, en el noreste tare à 380 metres d’altitude dans la forêt breakage, and uprooting by wind accom- de Puerto Rico, evidenció: (1) defolia- expérimentale de Luquillo. Celle-ci a mon- panied by immediate mortality resulting ción, rotura y caída de árboles causadas tré : (1) la défoliation, la casse et les cha- in a loss in forest structure, including por el viento; todo ello acompañado por blis provoqués par le vent induisant une stem numbers, tree height, and biomass; la mortalidad inmediata ocasionando un mortalité immédiate entraînent une dés- (2) delayed mortality of some impacted destructuración forestal, con pérdidas de tructuration de forêt en perte du nombre trees, along with prompt, abundant densidad y altura de árboles , así como de tiges, de hauteur des arbres et de bio- ingrowth of Cecropia schreberiana, and de biomasa; (2) una mortalidad diferida masse ; (2) la mortalité différée de d’au- other secondary species, increasing the de ciertos árboles dañados, con un tres arbres blessés s’accompagne d’un number of stems within the forest; (3) a rápido y abundante reclutamiento de rapide et abondant recrutement de Cecro- post-hurricane increase in diameter Cecropia schreberiana y otras especies pia schreberiana et d’espèces cicatri- growth and biomass accumulation for secundarias, lo que aumenta el número cielles, augmentant le nombre de tiges ~20 years, followed by a decline in both de tallos del bosque; (3) un aumento dans la forêt ; (3) une accélération de la rates; (4) a high post-hurricane survival post-huracán del incremento de creci- croissance en diamètre et l’accumulation and increase in dbh increment of Dacryo- miento de diámetros y la acumulación de de la biomasse pendant une vingtaine des excelsa; (5) a post-hurricane increase biomasa durante unos veinte años, d’années après l’ouragan, suivie du déclin in species richness for ~20 years when seguido de un descenso de ambas tasas; de leur ratio ; (4) un très fort taux de survie primary and secondary species grow (4) una alta supervivencia post-huracán y après l’ouragan et une augmentation de together, followed by a decline due to un aumento en el incremento del diáme- la vitesse de croissance en diamètre de competition; (6) asymptotic conditions tro de parte de Dacryodes excelsa; (5) un Dacryodes excelsa ; (5) toujours suite à for stems, biomass, and number of aumento post-huracán de la riqueza de l’ouragan, une augmentation de la species after more than 40 years. In addi- especies durante los primeros 20 años richesse en espèces pendant une période tion, related studies in the same forest en los que las especies primarias y de 20 ans au cours de laquelle les type showed that hurricanes and thinning secundarias crecen juntas, seguido por espèces primaires et secondaires grandis- evoke comparable growth responses. un consecuente ocaso debido a la com- sent ensemble, suivie dune chute d’abon- This study should be of regional interest petencia; (6) y por fin, condiciones de dance liée la concurrence ; (6) enfin, des because Puerto Rico’s lower montane evolución asintóticas para los tallos, la conditions d’évolution asymptotiques rain forest shares similar environmental biomasa y el número de especies des- aussi bien pour les tiges et la biomasse conditions as well as tree species with pués de más de 40 años. Además, algu- que pour le nombre d’espèces, après plus the mountainous Antilles. nos estudios relacionados con el mismo de 40 ans. En outre, d’autres études tipo de bosque demostraron que los connexes menées dans des types de Keywords: lower montane rain forest, huracanes y las entresacas de árboles forêts semblables ont montré que les hurricanes, forest monitoring, structure, provocan respuestas de crecimiento ouragans et les coupes intensives indui- forest dynamics, Puerto Rico. comparables. Este estudio resulta ser de sent des réactions de croissance compa- interés regional porque el bosque pluvial rables. Cette étude s’avère être d’intérêt montano bajo de Puerto Rico comparte régional puisque la forêt humide de basse similares condiciones ambientales y las montagne de Puerto Rico partage des mismas especies de árboles con las Anti- conditions environnementales sembla- llas montañosas. bles ainsi que les mêmes essences fores- tières avec les montagnes des Antilles. Palabras clave: bosque pluvial montano bajo, huracanes, monitoreo forestal, Mots-clés: forêt humide de basse mon- estructura, dinámica forestal, Puerto Rico. tagne, ouragans, suivi des forêts, struc- ture, dynamique forestière, Puerto Rico. BOIS ET FORÊTS DES TROPIQUES, 2013, N° 317 (3) OURAGANS, DÉGÂTS ET STRUCTURE / LE POINT SUR… 9 Introduction Hurricanes, also called typhoons, are strong tropical cyclones that form in the South Atlantic, the Caribbean Forest monitoring provides resource information Basin, the Gulf of Mexico, and in the eastern Pacific Ocean. needed to develop forest management policies and prac- Their trajectories pass over the Antilles, the Caribbean and tices, planning activities, and programs. The U.S. Forest Gulf coasts of Central America and Mexico, the southern and Service and collaborators have carried out studies in the eastern United States, the East Indies, Southeast Asia, Luquillo Mountains for nearly a century, including at least Australia, and Madagascar, where they recurrently affect five separate timber surveys of stocking and timber volumes vegetation and human settlements. Hurricanes that traverse between 1905 and 1948 (WADSWORTH, 1951, 1970). In the Caribbean basin occur from June through November. 1939, the International Institute of Tropical Forestry, first Those that pass near or over Puerto Rico typically originate called the Tropical Experiment Station, was established to near the Cape Verde Islands off western Africa and enter the conduct research on forest management, protection, and island from the southeast. Since 1700, apparent and con- utilization, along with soil and water conservation in Puerto firmed trajectories of 31 hurricanes of varying intensity Rico and the U.S. Virgin Islands. Since 1943, the Institute’s passed within 50 km of the forest, including four over the staff has set up ~620 plots of different sizes totaling 67 ha mountains (figure 1; SALIVIA, 1972; WEAVER, 2012). Based in primary and secondary stands within the Luquillo on these data, hurricanes may be expected to pass directly Experimental Forest (hereafter called El Yunque). The plots, over the forest once every 78 years and within 50 km once established for a variety of reasons, are located between every 10 years. Five hurricanes heavily impacted the moun- 200 and ~1000 m in elevation and contained ~134 000 tains during the 20th century: San Felipe (1928), the island’s trees at first measurement (WEAVER, 2012). Nearly 80% of legendary storm; San Nicolas (1931), passed along the the plots were permanently marked, and some of the earli- coast north of the forest; San Cipriano (1932), with a trajec- est plots were re-measured. In 1980, the Institute initiated tory directly over El Yunque; Hugo (1989), the first major an inventory program to monitor the island’s secondary hurricane after a hiatus of 57 years; and Georges (1998).