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Nutrient Relations of Dwarf Rhizophora Mangle L. Mangroves on Peat in Eastern Puerto Rico

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Nutrient Relations of Dwarf Rhizophora Mangle L. Mangroves on Peat in Eastern Puerto Rico Plant Ecol (2010) 207:13–24 DOI 10.1007/s11258-009-9650-z Nutrient relations of dwarf Rhizophora mangle L. mangroves on peat in eastern Puerto Rico Ernesto Medina Æ Elvira Cuevas Æ Ariel E. Lugo Received: 21 June 2007 / Accepted: 30 July 2009 / Published online: 19 August 2009 Ó Springer Science+Business Media B.V. 2009 Abstract Dwarf mangroves on peat substrate grow- resorption of these nutrients. On an area basis, ing in eastern Puerto Rico (Los Machos, Ceiba State resorption was complete for P but not for N. Sulfur Forest) were analyzed for element concentration, leaf accumulated markedly with leaf age, reaching values sap osmolality, and isotopic signatures of C and N in up to 400%, compared with relatively modest accu- leaves and substrate. Mangrove communities behind mulation of Na (40%) in the same leaves. This the fringe presented poor structural development with suggests a more effective rejection of Na than sulfate maximum height below 1.5 m, lacked a main stem, at the root level. Dwarf mangrove leaves had more and produced horizontal stems from which rhizo- positive d13C values, which were not related to phores developed. This growth form departs from salinity, but possibly to drought during the dry season other dwarf mangrove sites in Belize, Panama, and due to reduced flooding, and/or reduced hydraulic Florida. The dwarf mangroves were not stressed by conductance under P limitation. Negative leaf d15N salinity but by the low P availability reflected in low values were associated with low leaf P concentra- P concentrations in adult and senescent leaves. Low P tions. Comparison with other R. mangle communities availability was associated with reduced remobiliza- showed that P concentration in adult leaves below tion of N and accumulation of K in senescent leaves, 13 mmol kg-1 is associated with negative d15N contrasting with the behavior of this cation in values, whereas leaves with P concentrations above terrestrial plants. Remobilization of N and P before 30 mmol kg-1 in non-polluted environments had leaf abscission on a weight basis indicated complete positive d15N values. Keywords Mangroves Á Nutrient resorption Á & E. Medina ( ) Salinity Á 13C Á 15N Centro de Ecologı´a, Instituto Venezolano de Investigaciones Cientı´ficas, Aptdo. 21827, Caracas 1020 A, Venezuela e-mail: [email protected] Introduction E. Medina Á A. E. Lugo International Institute of Tropical Forestry, USDA Forest Mangroves are tropical and subtropical intertidal Service, Jardin Bota´nico Sur, 1201 Calle Ceiba, San Juan, communities, which grow along protected marine PR 00926-1119, USA coastlines and in estuaries with a steady supply of nutrients and fresh water (Lugo and Snedaker 1974). E. Cuevas Department of Biology, University of Puerto Rico, The species Rhizophora mangle L. occurs along the San Juan, PR 00931-3360, USA Atlantic coast from the tropic of Cancer in Florida to 123 14 Plant Ecol (2010) 207:13–24 the tropic of Capricorn in southern Brazil (Tomlinson document the changes from P to N limitations in 1986). The environments along this large geographic mangrove ecotones (Feller et al. 2002), and the range vary from high nutrient and low salinity in the analysis of stable isotopes showed that phosphorus estuaries of many tropical rivers in Brazil and limitation was the cause of large variations in d15N Venezuela, to hypersaline, low nutrient environments along the transects submitted to fertilization (McKee in many islands and continental coastlines in the et al. 2002). The dwarf mangrove communities Caribbean. Structural variations across these envi- described so far are found behind fringes of R. ronments have been frequently attributed to avail- mangle, their habit resulting from an inhibition of the ability of fresh water and salinity (Walter and Steiner growth in length of the terminal meristem. Therefore, 1936; Lugo and Snedaker 1974; Pool et al. 1977; the plants branch and produce adventitious roots from Medina 1999). Reduction in structural development a shortened stalk resulting in two round-shaped, is well known in R. mangle communities found in the umbrella-like, superimposing structures, the canopy, Florida and Yucata´n peninsulas on calcareous sub- and the adventitious root system (or rhizophore strate (Thom 1982; Tomlinson 1986). The morpho- system, Feller 1995). logical characteristics of these communities, Dwarf R. mangle communities are also found in however, are not uniform. In south Florida, mangrove the eastern coast of Puerto Rico in areas subjected to communities on marl substrate show strongly reduced a limited tidal flooding and not exposed to any wave structural development (dwarf mangroves), with little action (Cintro´n et al. 1978; Helmer et al. 2002; accumulation of organic matter at the surface, and Gonza´lez et al. 2006). The dwarf mangrove commu- low interstitial soil salinity (Koch and Snedaker nities at this site differ substantially in growth form 1997). Sites located nearer to the sea, characterized from what has been described for Florida, Belize, and by much higher interstitial soil salinity, also con- Panama´ (Feller 1995; Koch and Snedaker 1997; Chen tained short mangroves (Lin and Sternberg 1992a, b, and Twilley 1999; Cheeseman and Lovelock 2004; c). The former appeared to be mainly nutrient limited Lovelock et al. 2004). They grow on 1–6 m deep while the latter appeared to be affected by saline peat, deposited on top of calcareous material of concentrations above seawater. biological origin. The plants do not develop a main Nutrients have been identified as determinant of stem, but grow almost horizontally following the mangrove structural development. Boto and Welling- direction of the predominant winds and produce ton (1984) concluded that N and P availability rhizophores that sustain the horizontal stems. regulate mangrove development, correlating soil Windward-oriented stem-ends dry out and die pro- parameters with community structure in transects gressively. In addition, it appears that plants seldom through mangroves around Sydney, Australia. The flower, with the recruitment of new plants occurring effect of nutrients is somewhat hidden by lower through the eventual supply of propagules from the salinity in estuarine environments (Lugo and Sneda- fringe belts surrounding the dwarf communities. ker 1974; Medina and Francisco 1997). High fresh- The peculiar growth form of R. mangle in Los water runoff is naturally associated with higher Machos lagoon, and the probable association of nutrient supply, and the positive effects of both dwarfing of this species with availability of nutrients ecophysiological factors result in development of tall or hypersaline conditions prompted us to perform an and dense estuarine mangrove communities (e.g., analysis of the nutritional and osmotic conditions of Rivera-Monroy et al. 2004). Feller (1995) established soil and vegetation in this area. We hypothesized that a long-term fertilization experiment in the Twin Cays the extreme reduction in tree size and scarcity of in Belize and showed elegantly that for those propagules of these communities could result from communities on calcareous soils P availability was nutrient limitation and possibly high salinity because the ecological factor determining plant growth. of restricted tidal flushing determined by the topog- Subsequent articles revealed complex relationships raphy of the site. The combination of these factors between P availability and N utilization in coastal probably generated extreme low nutrient availability mangroves throughout the Caribbean and the Gulf of and physiologically detrimental K and Na concentra- Mexico (Feller et al. 1999, 2002; Koch and Snedaker tions in leaf sap, resulting in diminished capacity for 1997; Lovelock et al. 2006). Recent publications organic matter production and canopy development. 123 Plant Ecol (2010) 207:13–24 15 Site description The area is under the influence of shallow tides typical of the Caribbean (around 30 cm, http:// The mangrove site selected belongs to the Ceiba coops.nos.noaa.gov/tides06/), and is isolated from State Forest of the Commonwealth of Puerto Rico, significant transport of allochthonous sediments. and is located south of the Nuevo Mundo Bay, Sampling area was selected after observation of connected to the sea through tidal channels, some of aerial photographs, looking for accessible, undis- which were historically blocked by road construc- turbed sectors showing a pattern of tall–dwarf tions or hurricanes (Fig. 1). The tidal channels are communities from the berm along the main water lined by 3–6-m tall R. mangle mangrove communi- channel to the area landwards. The area selected was ties, followed by dwarf mangroves of the same near the center of the Los Machos lagoon and species, \1.5 m tall. The substrate is mangrove- conformed a nearly peninsular shape, located derived peat that began to accumulate since around between 18°1404200N; 65°3605600W and 18°1403700N; 4,500 years ago (E. Cuevas, personal communication). 65°3605600W (Fig. 1). A total of five points along a Fig. 1 Los Machos mangroves study site in the Ceiba State Forest, Humacao County, Puerto Rico. A and B showed the main water channels allowing the sea-water flushing of the wetland. The numbers indicate location of the points sampled in December 1995 for leaf area and nutrient analyses. A similar transect was sampled in March 1999 for estimation of leaf osmolality and measurement of carbon and nitrogen concentration, and isotope ratios in peat and leaves 123 16 Plant Ecol (2010) 207:13–24 transect from land to berm were sampled in Methods December 1995 for the analysis of leaf morphology and nutrient concentrations. Point 1 was located at the Leaves were sampled without petioles in a sequence of inland-most border, where R. mangle grew together relative ages from young (1st and 2nd pair), adult (3rd with Avicennia germinans and a few trees of and 4th pair), old (5th pair and beyond), senescent Laguncularia racemosa. Substrate was composed of (yellow-colored leaves) (two replicates of each leaf very dark-colored peat.
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