Aquatic Botany 79 (2004) 137143

Structure and litterfall of an arid mangrove stand on the Gulf of California, Mexico
Jos Alfredo Arreola-Lizrraga a, , Francisco Javier Flores-Verdugo b,1 , Alfredo Ortega-Rubio c,2
a Centro de Investigaciones Biolgicas del Noroeste, S.C., Unidad Guaymas, Carr. a Las Tinajas Perdio El Tular s/n, A.P. 349, Guaymas, Sonora 85454, Mexico b UNAM Instituto de Ciencias del Mar y Limnolog a, A.P. 811, Mazatln, Sinaloa 82000, Mexico c Centro de Investigaciones Biolgicas del Noroeste, S.C., A.P. 128, La Paz, BCS 23000, Mexico

Received 22 October 2002; received in revised form 27 January 2004; accepted 27 January 2004

Abstract Structure and litterfall in an Avicennia germinans mangrove stand in an arid zone on the Gulf of California, Mxico were studied. The density (4700 ha1 ) and basal area (13.5 m2 ha1 ) were more characteristic of a fringe-type mangrove stand, while average height (2.7 m), number of species in 0.1 ha (1), and complexity index (1.7) were closer to those of a dwarf-type mangrove. Structure is jointly inuenced by a high rate of evaporation (2982 mm per year), scarce rainfall (290 mm per year, mainly in summer) and a brief period of tidal ooding (also only in the summer). Litterfall averaged 1.75 t ha1 per year, placing it even lower than the lowest value reported so far for mangrove litterfall (216 t ha1 per year). Litterfall showed a unimodal seasonal pattern, with higher values in summer. Rainfall, sea level and the ratio rainfall/evaporation explained 86% of the seasonal litterfall variability. 2004 Elsevier B.V. All rights reserved.
Keywords: Litterfall; Mangroves; Avicennia germinans; Gulf of California; Arid coast

1. Introduction Mangroves have been very successful in colonizing protected coasts throughout the tropics (Tomlinson, 1986) under conditions ranging from very humid with rainfall >2000 mm per
Corresponding author. Tel.: +52-622-12237; fax: +52-622-12238. E-mail addresses: aarreola04@cibnor.mx (J.A. Arreola-Liz arraga), verduz@mar.icmyl.unam.mx (F.J. Flores-Verdugo), aortega@cibnor.mx (A. Ortega-Rubio). 1 Tel.: +52-69-852845; fax: +52-69-826133. 2 Tel.: +52-112-53633; fax: +52-112-53625.

0304-3770/$ see front matter 2004 Elsevier B.V. All rights reserved. doi:10.1016/j.aquabot.2004.01.012

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year to arid with rainfall <300 mm per year (Shaefer-Novelli and Cintron-Molero, 1993). The total world-wide mangrove area is estimated at no less than 170,000 km2 (Field, 1999) and among tropical marine ecosystems, mangrove are estimated rank second in importance only to coral reef in terms of gross productivity and sustained tertiary yield (Valiela, 1995, p. 29; Duarte and Cebrin, 1996). Mangroves increase in structural complexity and productivity as size progresses from the dwarf to the riverine type (Lugo and Snedaker, 1974; Pool et al., 1975; Pool et al., 1977; Brown and Lugo, 1982). Recently, Twilley (1998) has proposed an ecogeomorphological classication of mangroves based on the hypothesis that a gradient in the forcing functions (i.e. geophysical processes) of a coastal region will result in a variation of energy ow and material cycling that in turn will be reected in different patterns of forest structure, biomass, productivity, biogeochemistry, and an exchange of nutrients and organic matter with coastal waters. Our study contributes information about the structure and productivity of mangroves in an arid region. Its objectives were (1) to determine the structural aspects of mangrove, (2) to estimate the production of litterfall, and (3) to assess the inuence of environmental factors affecting the seasonal variability of mangrove productivity.

2. Methods Las Gusimas is a coastal lagoon located on the eastern coast of the Gulf of California (27 49 55 N, 110 29 45 W), inuenced by a dry desert climate (Garc a, 1973). Coastal vegetation surrounding the lagoon is composed of mangroves, halophytes (saltworts), and coastal dune vegetation. Mangroves cover approximately 5.4 km2 of the surface area. The dominant species is Avicennia germinans (Linnaeus 1764), although Rhizophora mangle (Linnaeus 1753), Laguncularia racemosa (Linnaeus 1807), and Conocarpus erectus (Linnaeus 1753), are also present. Structure of the mangrove was estimated by means of four transects perpendicular to the coastline and equidistant 10 m2 quadrants were established along the transects in accordance with the criteria proposed by Cintron et al. (1978). The average height, density, basal area, and complexity index were assessed (Holdridge et al., 1971). A topographical survey by means of a microtopographic technique (Garc a-Marquz, 1984) was made along each transect in order to develop the lagoons prole with reference to tidal levels. The tide level reference was obtained by using Version 0.32 of the Mar tide prediction program (Gonzlez, 2001). Litterfall was collected in 1 m2 litter baskets constructed of berglass screening (1 mm mesh). Litterfall was estimated monthly from March 1998 to February 2000 using the method described by Heald (1971). Leaves, owers fruits and branches were separated from the material collected monthly, dried at 70 C over a 24 h period and subsequently weighed. The data on climatic variables such as air temperature, winds, rainfall, and evaporation were obtained from a weather station of the National Meteorological Service located 15 km northwest of the lagoon. In addition, mean sea level for each month of the year was estimated. Multiple regression analysis was used to relate litterfall production values to average records

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of rainfall, sea level, and ratio rainfall/evaporation. The Statistica 6.0 package was used (Stat Soft, Tulsa, OK, USA).

3. Results 3.1. Environmental variables Summer was the season with the greatest rainfall in both of the annual cycles studied, with 19981999 having more (342 mm) rainfall than 19992000 (175 mm). Evaporation was highest during May and June and lowest during December, with annual values of 2693 mm in 19981999 and 2732 mm in 19992000. Air temperature was highest in July and August (3031 C) and lowest in December and January (1617 C; Fig. 1b). The mean sea level interval was 0.30.9 m, with the maximum value occurring in August and the minimum in March. Information of the topographic proles and mean sea level reference showed that the forest is inundated by high tides only during June, July, August and September.

Fig. 1. (a) Litterfall and (b) climatic variables during two annual cycles (19981999, 19992000) in Las Gu asimas, M exico.

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Table 1 Components of mangrove litterfall during two annual cycles in Las Gu asimas, M exico Components litterfall Leaves Flowers and fruits Branches Total 19981999 (g dry wt. m2 per year) 141 26 37 204 19992000 (g dry wt. m2 per year) 133 5 11 149 Average (g dry wt. m2 per year) 137 15 24 175

3.2. Structure The mangrove forest was almost monospecically A. germinans. R. mangle, L. racemosa and C. erectus specimens were scarce and none were observed in the transects. Quantied structural characteristics were canopy height (2.7 0.7 m), stem density (4700 1100 ha1 ), basal area (13.5 3.5 m2 ha1 ), number of species in 0.1 ha (1), and complexity index (1.7). 3.3. Litterfall Annual production of litterfall appeared higher in the 19981999 cycle than in that of 19992000 (Table 1). However, the values of leaves (paired t = 0.41, P = 0.687), owers and fruits (t = 1.6, P = 0.137), and branches (t = 1.25, P = 0.236), were not signicantly different between two annual cycles. Litterfall was observed in all months of the period studied. Specically, leaves were observed throughout the year, while branches, owers and fruits were observed only during the summers of the two cycles studied (Fig. 1a). Litterfall showed unimodal seasonal pattern and yielded the highest values during summer (Fig. 1a). Multiple regression analyses selected average rainfall, sea level and the ratio rainfall/evaporation as independent variables which were highly signicant in explaining temporal litterfall variability over the period of study. The equation that best describes litterfall variability is: Y = 0.498 3.524(X1 ) + 0.927(X2 ) + 3.619(X3 ) r 2 = 0.86

where X1 is rainfall (P < 0.001), X2 is sea level (P < 0.001), and X3 is ratio rainfall/evaporation (P < 0.001).

4. Discussion 4.1. Structural aspects Location of the mangrove in terms of both coastal geomorphology, and topographic setting is an important descriptor in classifying mangrove forests (Twilley, 1998). At our study sitea lagoon with a sand bar dominated by high wave energygeomorphological conditions corresponded to Type III suggested by Thom (1982). Microtopography, seasonal

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Table 2 Structural characteristics and litterfall for different types of mangroves compared to the Las Gu asimas stand, investigated here Parameter Mean canopy height Stem density (no. ha1 )a Basal area (m2 ha1 )a No. Species in 0.1 haa Complexity indexb Litterfall (g m2 per year)c
a b

Riverine (m)a 17.7 1760 41.3 3 36.4 1770

Basin 9.0 3580 18.5 2 15.4 730

Fringe 8.2 5930 17.9 2 14.6 906

Dwarf 1.0 25302 0.6 1 1.5 120

Las Gu asimas 2.7 4700 13.5 1 1.7 175

From Pool et al. (1977) and Brown and Lugo (1982). Canopy basal area stem density no. of species in 0.1 ha 105 . c From Brown and Lugo (1982) and Twilley et al. (1986).

variation in sea level and amplitude of the tide (0.8 m) are specic factors that cause the mangrove to be subject to ooding only during the period of summer high tides when the sea level is highest (0.9 m); these conditions, plus the high evaporation rate and the scarcity of fresh water (which comes only from rain), result in this mangrove sharing structural characteristics with both fringe and dwarf-type forests. The values for density and basal area were closer to those of a fringe-type forest, while the values for average height, number of species and complexity index were closer to those of a dwarf-type (Table 2). Basal area of the mangrove in Las Guasimas (13.5 m2 ha1 ) was similar and average height (2.7 m) was lower than those found in other arid and semi-arid localities of the Pacic coast of Mexico, where mangrove forests range from 11.416.9 m2 ha1 in basal area and 4.57 m in average height. In sub-humid localities the ranges are 12.529.6 m2 ha1 basal area and 7.58.8 m average height (Flores-Verdugo et al., 1992). The hypersalinity in the mangrove soils has been attributed to a high evaporation rate (Shaefer-Novelli and Cintron-Molero, 1993), and the species best adapted to this high soil salinity environment is reportedly A. germinans (Painner and Painner, 1976). The mangroves in Las Guasimas are subject to high evaporation rate (2982 mm per year) and the dominant species is A. germinans, suggesting hypersalinity to be important here as well. 4.2. Aspects of litter production Litterfall values for mangrove forests worldwide range from 2 to 16 t ha1 per year (Twilley and Day, 1999); thus the values of litterfall from the Las Gusimas mangrove forest (avg. annual production: 1.7 t ha1 per year) actually fall below the minimum of this range and are similar to those of a dwarf-type mangrove wetland (Table 2). The low rate of mangrove litterfall in Las Gusimas is related both to minimal tidal ooding that only occurs during summer high tide conditions and to minimal availability of fresh water that occurring only during the summer rainy season in this arid region where the evaporation rate (2982 mm per year) is ten times greater than the rainfall rate (290 mm per year). The average annual rainfall in Las Gusimas is 290 mm per year; thus the 342 mm of the 19981999 cycle was above average and the 175 mm of the 19992000 cycle below average. Our results show that litterfall increases during years of greater rainfall, but that

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this increase is due to the greater quantities of fallen branches, owers and fruits (Table 1). It is also worth noting that this region has both a summer and winter rainy season, the former with more abundant rainfall (Garc a, 1973). Beginning in 1996, winter rains have been scarce or non-existent (as was observed during the period of our study) and predictions are that this pattern could continue during the present decade (Brito-Castillo et al., 2002). In Las Gusimas, A. germinans produces most litterfall (53%) during the summer months, when the hydrological characteristics of the forest are inuenced both by seasonal runoff (summer being the rainiest season of the year) and by tidal ooding, thus allowing the totality of its litter to be contributed to the lagoon. In autumn, litterfall decreases and subsequently falls to its lowest level in winter when the forest has already contributed almost the entire quantity of litter available. Spring is marked by litterfall likewise increasing until it again reaches its maximum during the summer rainy season. Eighthy six percent of seasonal litterfall variability was explained by rainfall, sea level and ratio rainfall/evaporation. Litterfall may occur throughout the year with little (Shunula and Whittick, 1999) or marked (Day et al., 1996; Wafar et al., 1997) seasonal variation, depending on the forcing functions and their intensity in each region. In our site (dry region) and in sub-humid and humid regions along Mexicos Pacic coasts (Flores-Verdugo et al., 1987, 1992) a marked litterfall variability has been observed with a seasonal unimodal pattern. In our site maximum litterfall occurred during the rainy season (JulySeptember), while in sub-humid and humid regions this takes place during the dry season (MayJune).

Acknowledgements We are grateful for the technical support of personnel attached to coastal management of the Northwest Center of Biological Researchs (CIBNOR) Guaymas Unit. The principal author would also like to thank CONACYT for the grant that has allowed him to pursue his doctoral studies. Thanks to Albert Brandt for editing the English.

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