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Ecophysiology of a Mangrove Forest in Jobos Bay, Puerto Rico

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Ecophysiology of a Mangrove Forest in Jobos Bay, Puerto Rico Caribbean Journal of Science, Vol. 43, No. 2, 200-219, 2007 Copyright 2007 College of Arts and Sciences University of Puerto Rico, Mayagu¨ez Ecophysiology of a Mangrove Forest in Jobos Bay, Puerto Rico ARIEL E. LUGO1,ERNESTO MEDINA2,ELVIRA CUEVAS1,2,3,GILBERTO CINTRÓN4, EDDIE N. LABOY NIEVES5 AND YARA SCHÄEFFER NOVELLI6 1USDA Forest Service International Institute of Tropical Forestry, Ceiba 1201, Jardín Botánico Sur Río Piedras, PR 00926-1119. 2Instituto Venezolano de Investigaciones Científicas, Caracas, Venezuela 3Department of Biology, University of Puerto Rico, Río Piedras, PR. 4Department of Natural Resources, P.O. Box 5887, Commonwealth of Puerto Rico, San Juan, PR. Current address: US Department of Interior Fish and Wildlife Service, Washington, D.C. 5School of Environmental Affairs, Metropolitan University, Río Piedras, PR. 6Instituto Oceanográfico da Universidade de São Paulo, São Paulo, Brazil. [email protected] ABSTRACT.—We studied gas exchange, leaf dimensions, litter production, leaf and litterfall chemistry, nutrient flux to the forest floor, retranslocation rates, and nutrient use efficiency of mangroves in Jobos Bay, Puerto Rico. The fringe forest had a salinity gradient from the ocean (35‰) to a salt flat (100‰) and a basin (about 80‰). Red (Rhizophora mangle), white (Laguncularia racemosa), and black (Avicennia germinans) mangroves were zoned along this gradient. Photosynthetic rates, stomatal conductance, leaf area and weight, leaf specific area, and nutrient use efficiency decreased with increasing salinity, while xylem tension, nu- trient retranslocation, and leaf respiration increased with increasing salinity. The concentration of some leaf elements increased with salinity (N, P, Mg, Na) while others decreased (Ca). Leaf specific area was less −2 −1 variable than leaf area or weight. Maximum photosynthesis (12.7 µmol CO2 m s ), leaf conductance (283 −2 −1 −1 −1 mmol CO2 m s ), and litterfall (16.9 Mg ha yr ) in the fringe were high in comparison with those of other world mangroves. Nutrient flux to the forest floor was also high and nutrient use efficiency was low in this forest. Part of the reason for these high values and low use-efficiencies was the high proportion of nutrient-rich flowers and propagules in litterfall. Nevertheless, access to freshwater in the form of ground- water discharge and about1mofannual rainfall play a role in the high productivity of the Jobos mangroves. INTRODUCTION nated by Rhizophora mangle (there are many) and studies of environmental gradi- The literature on mangrove forests is ents and vegetation response (very few). abundant (Rollet 1981) and only recently Yet, progress in understanding the re- have there been quantitative studies on the sponses of mangrove species to variable ecophysiological responses of trees to envi- conditions requires precise paired mea- ronmental gradients. For example, Feller et surements of biotic responses and gradi- al. (2002) studied phosphorus limitation ents of abiotic conditions. Without such across an ecotonal gradient that included paired measurements, very little generali- several edaphic variables, Lovelock and zation will be possible because short dis- Feller (2003) focused on a salinity gradient, tances in a fringe mangrove may involve a and McKee et al. (2002) studied a nutrient large range of environmental conditions gradient in Belize, Florida, and Panamá, re- (Cintrón et al. 1978, Feller et al. 2002). These spectively. A review of fringe mangrove broad ranges are not usually included in forest’s literature (Lugo 1990) uncovered a experiments on the ecophysiology of man- wide gap between the description of veg- groves. For example, Björkman et al. (1988) etation zonation in fringe forests domi- and Ball and Pidsley (1995) studied gas ex- change and tree growth (respectively) in mangrove trees and seedlings but restricted their salinity experiment to values between ms. received July 4, 2004; accepted Jan. 28, 2007 seawater strength (35‰) and 90% dilution 200 ECOPHYSIOLOGY OF MANGROVES IN PUERTO RICO 201 (or less than 5‰). However, salinity ranges trees to species, mapped their location, and of0to100‰ are common in mangroves, measured tree height. Land elevation rela- particularly in arid coastlines or on steep tive to sea level was measured along the topographic gradients (Cintrón et al. 1978). transect using a level and readings to a pre- Lovelock and Feller (2003) studied photo- cision of 1 cm. We used a refractometer synthetic performance of Florida man- calibrated for seawater to measure pore groves at salinities as high as 55‰ as did and surface water salinity. Feller et al. (2003), but most other ecophysi- We measured photosynthetically active ological research is at lower salinities radiation (PAR) with a light integrator (Feller et al. 1999, Cheeseman and Lovelock (Li190 quantum sensor, Li-Cor, Lincoln, 2004, Lovelock et al. 2004). NE) and air temperature and humidity We expand previous analyses of man- with an aspirated psychrometer. Xylem grove vegetation response to pore salinity tension was measured with a Scholander (Cintrón et al. 1978) to studies of ecophysi- pressure chamber (Scholander et al. 1965). ological responses to salinity gradients. By Leaf samples used for measurement of leaf relating structural and ecophysiological re- sap osmotic pressure were cleaned in the sponses of mangrove species to specific en- field by washing the leaf surface with dis- vironmental conditions, we contribute to tilled water (to eliminate external sea salt the understanding of robust relationships deposition), dried with paper towels, and between mangroves and their environ- enclosed inside tight glass vials. The vials ment. were frozen in dry ice, transported to the laboratory, and boiled for 2 h. We extracted METHODS leaf sap from the samples with a hand press providing a pressure of about 100 psi, and We conducted measurements in two ad- measured the osmolality of cell sap with a jacent fringe forests. One was an inland water vapor osmometer (Wescor, Logan fringe forest lining a dead-end canal that UT) calibrated against NaCl solutions of connects the forest to the Mar Negro in Jo- known concentration. We transported sub- bos Bay National Estuarine Research Re- samples of pore water taken at 30 cm depth serve in Puerto Rico. We will refer to this to the laboratory for the measurement of site as the inland fringe forest (IFF). Lugo osmolality with a water vapor osmometer. (1987), Montero and Soler (1987), Dávila Salinity (‰) measured with a hand refrac- (1987) and Ruiz Bernard and Lugo (1999) tometer and osmolality (mol kg−1) mea- have described the site. Between October surements of soil water samples were 11, 1985 and January 11, 1987, we collected highly correlated as expected: litterfall monthly in this forest. A second mangrove stand fringed the ocean and was Osmolality (mmol kg−1) = 33.9098 × located just southwest of the IFF, across a salinity (‰) − 231.8243, r2 = 0.99 dirt road. We will refer to this site as the ocean fringe forest (OFF). On December 6, 1986 we established a 100 m × 2 m transect For natural waters, this equation has a in the OFF and conducted the following negative independent term due to the pres- measurements: vegetation profiles, topo- ence of organic compounds that influence graphic relief, environmental conditions, the refraction index without affecting salin- leaf photosynthesis, stomatal conductance, ity. water potential, leaf specific area, and nu- Photosynthetic rate, stomatal conduc- tritional status of leaves of various sizes tance, leaf temperature, and incident pho- and appearance. These measurements oc- ton flux density on leaves were measured curred at the end of the rainy season. along the transect every 3 h from dawn to All the work in the OFF was along the dusk using a portable gas exchange system 100-m transect that originated at the ocean (ADC-LCA-2, Analytical Development edge and ended inland in a basin after Corp., Hoddesdon, UK), operating in the crossing a salt flat (salina). We identified open mode (Long and Hålgren 1985). Every 202 A. E. LUGO, ET AL. 3 h we conducted triplicate measurements branch sampled. Nutrient concentrations on mature leaves of red mangroves (Rhizo- were expressed on an area basis using spe- phora mangle) on the water’s edge, white cific leaf areas, to avoid errors estimating mangroves (Laguncularia racemosa) on the absolute amounts of a given nutrient in the water’s edge, and those two species plus leaf tissue derived from increasing accumu- black mangroves (Avicennia germinans)in lation of minerals in older leaves. the basin behind the salina (basin). Litterfall was collected in the IFF using two 0.5 m × 4.0 m baskets constructed of Leaf Characteristics, Litterfall, and plastic screen mesh located in the field on Nutrient Cycling September 11, 1985. Collections from these We collected leaves still attached to the baskets were monthly. Litterfall was ini- trees to study their area/weight character- tially separated into leaves, and miscella- istics and nutritional status by species and neous between October 11,1985 and Janu- location along the OFF transect. We classi- ary 11,1986. After February 11, 1986 we fied leaves by their appearance and loca- separated leaves by species and between tion on the branch. The leaf categories were May 11, 1986 and January 11, 1987, we mature (fully expanded), old mature, yel- separated flowers and propagules by spe- low, and senescent. Senescent leaves were cies. Samples were oven dried to constant still attached to the stem but were already weight at 70°C. We used these samples for brown in color. We ignored young leaves chemical analyses (N, P, K, Ca, Mg, and that had yet to expand to full size. Leaf col- Na). Litterfall material from each basket lections included 10 leaves each of mature, was analyzed separately for each date of old mature, yellow, and senescent leaves. collection and after sorting into leaves, We measured leaf area in the laboratory flowers, and propagules by species plus with a leaf area meter (Li-cor model 3100, fine wood and miscellaneous.
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