• Embed Doc
  • Copy Link
  • Readcast
  • Collections
  • CommentGo Back
Download
 
-1
Environmental correlates of tree and seedling–sapling distributionsin a Mexican tropical dry forest
Yalma Luisa Vargas-Rodriguez
1,2,
*, J. Antonio Va ´zquez-Garcı ´a
3
and G. Bruce Williamson
1
1
Department of Biological Sciences, 107 Life Sciences Building, Louisiana State University, Baton Rouge, LA70803, USA;
2
Current address: Carlos Fuero 543, Colonia Universitaria, Guadalajara, 44840, Jalisco,Me´ xico;
3
Centro Universitario de Ciencias Biolo´  gicas y Agropecuarias, Departamento de Bota´ nica y Zoo-logı´ a, Universidad de Guadalajara, Km 15 carretera Guadalajara-Nogales, Las Agujas, Zapopan, 45110,Jalisco, Me´ xico; *Author for correspondence (e-mail: yvarga1@lsu.edu)
Received 28 June 2004; accepted in revised form 28 February 2005
Key words:
Bray and Curtis ordination, Disturbance, Importance value, Regeneration, Specialization, Treediversity, Tropical deciduous forest
Abstract
Bray and Curtis ordination was used to explore which environmental variables explained importance valuesand the presence–absence of tropical tree seedlings, saplings and adults in La Escondida-La Caban ˜a, Sierrade Manantla ´n, Jalisco, Mexico. The diameters of trees
‡
2.5 cm DBH and the presence and height of seedlings and saplings were measured in nine 0.1 ha sites. Four matrices including presence–absence dataand importance value indices for trees and seedlings and saplings were analyzed through Bray and Curtisordination. The matrices were based on density, frequency, and dominance of adult trees as well asseedlings and saplings. The environmental matrix consisted of 18 variables, including elevation, slope,canopy gaps, disturbance, and soil variables. We recorded 63 tree species and 38 seedling and saplingspecies in the nine sites. The ordination explained 70.9% of the variation in importance value data for treesand 62.6% for seedlings and saplings. The variation explained in presence–absence data for trees was 67.1and 77.4% for seedlings and saplings. The variance in the ordination axes of seedlings and sapling pres-ence–absence data was poorly explained by the number of gaps in the tree, shrub, or herb layer, suggestinglittle light specialization by seedlings and saplings. Habitat specialization for soil nutrients appears to beimportant in explaining the presence–absence of seedlings and saplings. Seedling and sapling specializationalong different soil microsites could promote species coexistence in this forest, while heterogeneity in lightconditions may instead determine differences in growth and, thus, importance value of trees. Wehypothesize that in tropical dry forest in Jalisco, Mexico, a habitat specialization for soil resources is likelymore important at early stages in tree life histories than in later life history.
Introduction
In Mexico, 60% of the tropical vegetation istropical dry forest (TDF). Mexican TDF is widelydistributed across the Pacific lowlands on thewestern side of the country, covering most of Jalisco state and confined to small patches ineastern Mexico (Rzedowski 1978; Trejo and Dirzo
Plant Ecology (2005) 180:117–134
Ó
Springer 2005DOI 10.1007/s11258-005-3026-9
 
2000). The forest is characterized by marked sea-sonality in rain fall and its occurrence on moderatesteep slopes and rocky outcrops.Tropical dry forests in western Mexico, includ-ing those in the studied area, are typically decid-uous, with short stature trees and high density of small size trees (Trejo 1998). Trees ca. 4 m tallconstitute 65%, those with heights between 4 and8 m are 31% and only 3% are 8–12 m height,some exceptional individuals reach 15 m. Treemean density is 3610 (800) individuals/ha and ba-sal area is ca. 56.8 m
2
/ha. Trees with diameter
‡
10 cm represent only 20% of individuals and lessthan 5% are
‡
30 cm, thus, the majority of treeshave diameters
£
2.5 cm (Rzedowski 1978; Trejo1998). Trees have extended crowns with bright andpeeling barks, as well as compound leaves. Most of the tree species (ca. 85%) are deciduous and few(ca. 15%) are evergreen, such as
Ziziphus mexicana
Rose and
Prosopis laevigata
(Willd.) M.C. Johnst.In addition, columnar and Opuntioideae cacti arecommon in this community (Rzedowski 1978;Trejo 1998).TDF, throughout Mexico, is severely affected bylivestock, slash and burn agricultural practices,forest fire, and selective logging. In 1990, only 27%of the original cover of tropical dry forest re-mained intact and 1.4% continues to be lostannually as a result of deforestation (Trejo andDirzo 2000). Forest fires and livestock are alsocommon in Sierra de Manantla ´n Biosphere Re-serve (SMBR). Fire in the area has been mainlyassociated with agricultural burning, occurring inthe dry season (December–May). Livestock activ-ity occurs during the rainy season (June–October).However, forest protection is enforced in corezones through various mechanisms negotiatedwith agrarian communities (68% of reserve’s area)and private landowners (32% of reserve’s area). Inaddition, there are management goals in the bufferzone directed to implement sustainable practices inforestry, agriculture, livestock, and other naturalresource management activities (INE 2000).TDF in Mexico is characterized by high
a
and
b
diversity (Balvanera et al. 2002; Trejo and Dirzo2002). While species diversity has been shown tobe positively correlated with increasing precipita-tion across wet and dry forest in Puerto Rico(Murphy and Lugo 1986), a comprehensiveanalysis of Mexican TDF does not show such arelationship (Trejo and Dirzo 2002). Instead, it hasbeen suggested that Mexican TDF diversity ispositively related to potential evapotranspiration(Trejo and Dirzo 2002).In addition, local site factors such as thepresence and size of canopy gaps, soil properties,anthropogenic disturbance, and total annualprecipitation may have important effects on TDFtree species richness, composition, abundance,and structure (Gonzalez and Zak 1996; Gentry1988; Oliveira-Filho et al. 1998; Gillespie et al.2000; Segura et al. 2003). For instance, variationin species diversity in the Neotropics could beexplained by total annual precipitation (Gentry1982, 1988). Associations of species with thosefactors and specialization to particular micro-habitats are hypothesized to contribute to speciesdiversity (Connell 1978; Gentry 1982; Denslow1987; Welden et al. 1991; Clark et al. 1993).Different tree species are best suited to differenthabitats, which may lead to habitat specializa-tion. Differential resource utilization might ex-press itself as microhabitat specializationbetween the species or as differences in geo-graphical distribution. On the other hand, theavailability of different resources may be sepa-rate in time and may become available duringdifferent seasons. For example, canopy gapsprovide different soil, light, and moisture condi-tions than the forest understory and could beexploited by species with specialized competitiveabilities (Denslow 1987; Oliveira-Filho et al.1998). Differences in species’ growth and survivalmay also occur in the absence of gaps alongsmaller light gradients, such as between 0.2 and6.5% available diffuse light (Lieberman et al.1989; Montgomery and Chazdon 2002). Inaddition, natural disturbances can cause spatialand temporal variability in the availability of resources leading to habitat differentiation.Consequently, variation in growth and survival of tree species under differing conditions of resourceavailability (for example micro-topography orlight) could result in habitat specialization (Kobe1999; Pearson et al. 2003).Previous studies have shown that microhabitatspecialization with regard to topography and soilcharacteristics affect the distribution of severalplant groups, including tropical trees, melasto-mate shrubs, herbs, pteridophytes, and palms(Kahn and De Castro 1985; Liberman et al.1985; Poulsen and Baslev 1991; Basnet 1992;118
 
Toumisto and Roukolainen 1993; Clark et al.1998; Svenning 1999). For instance, distributionand abundance of some deciduous tree families(e.g., Leguminosae), are related to ecologicalgradients such as edaphic conditions and shadetolerance (Givnish 1999). Cation exchange levelaffects the distribution of 51 tree species inGhana (Swaine 1996). Soil organic matter, soilnutrients, texture, and moisture are stronglycorrelated with main axes of a PCA ordinationof tropical trees in a Bornean tropical rain forest(Webb and Peart 2000). In addition, adult treesand seedlings had different strengths of associa-tion with those variables (Webb and Peart 2000).Other species can be specialized along moisturegradients, reflected by the greater numbers of species that persist on protected side-slope areasdue to their higher dry season moisture levels(Hubbell 1995).Even though previous studies used a quantita-tive multivariate approach to analyze how treespecies relate to environmental variables, only afew studies have considered the relationship be-tween the regeneration of tree species in seasonaltropical dry forest and environmental conditions(Lieberman and Li 1992; Oliveira-Filho et al.1998), and none have explored the possible con-tribution of environmental variables at differentstages in the life cycle of TDF trees.Here we report the results of a study of treeforest composition and regeneration along a shortaltitudinal gradient in a tropical dry forest at LaEscondida-La Caban ˜a, Sierra de Manantla ´n Bio-sphere Reserve, in the Ayuquila watershed, inwestern Mexico, to generate hypotheses abouttropical trees and regeneration patterns along soiland canopy gaps gradients. We addressed thefollowing question: What environmental variablescould explain major community gradients for bothtrees and seedlings and saplings? To answer thisquestion, we employed a multivariate gradientanalysis, Bray and Curtis ordination technique(also known as sociological ordination) (Curtisand McIntosh 1951; Beals 1984; McCune andGrace 2002), which produces pure communitygradients that can be correlated with measuredenvironmental variables. This approach has pro-vided many insights into the nature, organizationand dynamics of ecological communities (Whit-taker 1956, 1960; Terborgh 1973; Ludwig andReynolds 1988).
Methods
Study area
La Escondida-La Caban ˜a gradient is located in thenorthern boundary of the SMBR, roughly 50 kmfrom the Pacific Ocean, in Jalisco state, WesternMexico. The study area lies between El Aguacate(
municipio
El Grullo) and Zenzontla (
municipio
Tuxcacuesco), northwest of 
ejido
Zenzontla, alongthe Ayuquila-Armerı ´a river, within the tributarywatershed La Pasio ´n-Cerro Blanco. The eleva-tional gradient extends from Arroyo La Escondidaat, 880 m to the top of Cerro la Caban˜a at, 1090 ma.s.l. (19
°
42
¢
N, 104
°
07
¢
W) (Figure 1).La Escondida-La Caban ˜a occupies the steepfoothills of the northern Sierra de Manantla ´n, inthe Sierra Madre del Sur, an area of roughtopography, within a hilly and dissected land-scape. Relief ranges from 10 to 66% slope incli-nation, along La Escondida and La Caban ˜a hills.Microclimate at La Escondida-La Caban˜a ismoist, warm, and highly seasonal with a wet sea-son from June to October and a dry season fromDecember to May (Martı ´nez et al. 1991). Meanannual rainfall is 900 mm (range 600–1000 mm)(Martı ´nez et al. 1991). Mean annual temperature is22
°
C (range 22–28
°
C), free of frost. The tem-perature lapse rate is 4
°
C per 1000 m a.s.l.(Martı ´nez et al. 1991). TDF at La Escondida-LaCaban ˜a corresponds to the Lower Montane Sub-humid Tropical Dry Forests of Holdridge (1967)and occupies 12,700 ha (9%) of the SMBR(Cuevas-G. et al. 1998).Tertiary volcanic rocks are prevalent in thestudy area (Cruz 1989). Acidic intrusive rocks(granites) were found on sites 1, 2, 3, 4, 8 and 9,while intermediate extrusive rocks (rhyolites,andesites, and trachytes) prevailed on sites 5, 6,and 7. Land form is convex with boulders androcky outcrops.Shallow, well-drained lime soils (Regosol eut-rico) were common in sites 5, 6 and 7. Some limesecondary soils such as Feozem ha ´plico and someLitosols occured. Litosols (with rocks greater than7.5 cm in diameter) were frequent at sites 1, 2, 3, 4,8 and 9. Fluvial sandy soils (Fluvisol eutrico) werefound on river banks and at the foothills of LaEscondida (CETENAL 1975, 1976).Floristic checklists and descriptions of tropicaldry forest species in the SMBR (Va ´zquez-Garcı ´a119
of 00

Leave a Comment

You must be to leave a comment.
Submit
Characters: ...
You must be to leave a comment.
Submit
Characters: ...