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29

Núm. 27, pp. 29-50, ISSN 1405-2768; México, 2009

SMALL-SCALE ENVIRONMENTAL GRADIENTS IN A PINE-OAK FORESTCOMMUNITY IN NUEVA COLONIA, MEZQUITIC, JALISCO, MEXICOGregorio Nieves-Hernández

1

, J. Antonio Vázquez-García

1

, Yalma L. Vargas-Rodriguez

2

, Marcelino Vázquez-García

1

and Jesús González-Gallegos

1

1

Centro Universitario de Ciencias Biológicas y Agropecuarias, Universidad deGuadalajara, Km 15 carretera Guadalajara-Nogales, Las Agujas, Zapopan, Jalisco,45110, México. Correo electrónico: jvazquez@cucba.udg.mx

2

107 Life Sciences Building, Department of Biological Sciences, Louisiana State University, Baton Rouge, Louisiana, 70803, USA.

A

BSTRACTWe explore what environmental variablesare related to pine-oak forest communitystructure at one locality in Jalisco, Mexico.We used a NMS ordination, in connectionwith the Sørensen distance, and identi

ﬁ

edmajor small-scale community gradientsalong 25 contiguous quadrats (25 m x 25m, 400 m

2

each) of pine-oak forest in NuevaColonia, Mezquitic, Jalisco, Mexico. Mainmatrix (25 stands x 7 tree species) included basal area data and the environmental matrixconsisted of 19 quantitative environmentalvariables. Community structure, throughsociological ordination, showed a directcorrelation with the vertical altitudinal gra-dient, the apparent soil density and the slopeinclination across the horizontal gradient;It also showed an inverse correlation withcation exchange capacity, Ca+Mg, Mg, andaltitude across the horizontal gradient. Di-rect gradient analyses showed an increase of cation exchange capacity, Ca+Mg, Mg andK with decreasing altitude along the verticalgradient (from north to south); and totalnitrogen increased with decreasing altitudeacross the horizontal gradient (from West toEast). In addition, we identi

ﬁ

ed three maincommunity groups using UPGMA cluster analysis; however, groups were weakly rela-ted to the ordination results or to the physicalspace. At a 1 hectare scale, species compo-sition and basal area of pine-oak forest in Nueva Colonia can be explained by niche partitioning of altitude and soil gradients.The nitrogen relevance for this communitycould be the result of habitat specializationor disturbance history.

Key words

: calcium, magnesium, environ-mental gradients, habitat differentiation,niche partitioning.

R

ESUMEN Nosotros exploramos cuáles variablesambientales se relacionan con la estructurade comunidades de pino-encino en NuevaColonia, Mézquitic Jalisco, México. Através de una ordenación NMS y usandola distancia Sørensen se identi

ﬁ

caron en laescala pequeña los principales gradientesde comunidades entre las 25 parcelas con-tiguas (25 x 25 m, cada una de 400 m²) de bosque de pino-encino en Nueva Colonia,Mezquitic, Jalisco, México. La matriz de principal (25 rodales x 7 especies de árboles)incluyó datos de área basal y la matriz am- biental consistió de 19 variables ambientales

30

Marzo 2009Núm. 27: 29-50

cuantitativas. La estructura de la comunidad,mediante la ordenación sociológica, mostróuna correlación directa con la altitud sobreel gradiente vertical y la densidad aparentedel suelo, así como con la inclinación de pendiente sobre el gradiente horizontal;también mostró una correlación inversa conla capacidad de intercambio catiónico, Ca +Mg, Mg y con la altitud sobre el gradientehorizontal. El análisis de gradientes directosmostró un incremento de la capacidad deintercambio catiónico, Ca+Mg, Mg y K conla disminución de la altitud, en el gradientevertical (de Norte a Sur). El nitrógeno totalincrementó con la disminución de la altituden el gradiente horizontal (de Oeste a Este).Además, se identi

ﬁ

caron tres grupos prin-cipales de comunidades usando el análisisde conglomerado de UPGMA, sin embargo,los grupos mostraron una relación débilcon los resultados de la ordenación y con elespacio físico. A la escala de una hectárea, lacomposición de las especies y el área basaldel bosque de pino-encino en Nueva Colonia pueden ser explicados por la partición denicho en gradientes de altitud y de suelo. Larelevancia del nitrógeno para esta comuni-dad podría resultar de la especialización dehábitat o por la historia de disturbio.

Palabras clave

: calcio, magnesio, gradien-tes ambientales, diferenciación de hábitat, partición de nicho.

I

NTRODUCTIONMany studies of large-scale plant communi-ty patterns have supported the individualistichypothesis of plant community organization(Gleason, 1926) in both, temperate areas(Whittaker, 1956, 1960; Curtis and McIn-tosh, 1951; Peet, 1980) and subtropicalto tropical areas (Vázquez-García, 1995,1998; Cuevas, 2002), some other authorsdocument vegetational discontinuities(Beals, 1969; Kitayama, 1992). However,studies of small-scale community patternsto assess whether or not the individualistichypothesis is supported, have largely beenneglected.Understanding patterns of species distribu-tion and abundance across landscapes have been central goals in Ecology. Ecologicalstudies have focused on determining whe-ther species specialize to partition envi-ronments or whether distributions are theresult of stochastic processes (Brokaw andBusing, 2000; Hubbell, 2001). A number of mechanisms have been proposed to explainhow diversity and abundance are regulatedin tropical and temperate systems (Wills

et al.

, 1997; Wright, 2002). Niche differentia-tion among species, arising from resource partitioning across resource gradients, has been hypothesized to be a major mechanismmaintaining tree diversity in both temperateand tropical forests. Many studies (Dens-low, 1980; Kabakof and Chazdon, 1996;Chazdon

et al.

, 1999; Capers

et al.

, 2005;Iriarte and Chazdon, 2005; Gouvenain

et al.

,2007) have focused only on light gradientsas driving force or mechanism in resource partitioning among tree species, however,and little or no attention has been given toother resources such as soil variables.Climatic variables might determine plantcommunity distribution at a landscape level.Latitude, longitude, precipitation, tempera-ture, and moisture variables have an effectin plant composition and distribution (Rze-dowski, 1978, Gentry, 1982). However, atsmall-scale gradients (< 200 m in altitude),topography is considered a more relevantvariable that affects tree community spatial

31

Nieves

et al.

: Small-scale environmental gradients in a pine-oak forest community in Nva. Colonia, Mezquitic, Méx.

variation than it is altitude or any other climatic variable (Basnet, 1992; Vázquez-García, 1995). Topography is also relatedto patterns in moisture regimes and soilchemical properties (Oliveira-Filho

et al.

,1999, 1994; Ratter, 1980; Bourgeron, 1983;Johnston, 1992). For instance, soil leaching patterns along an elevational gradient areaffected by topography (Smith, 1990).Soil characteristics and biotic interactions,at a small scale (within a hectare), might bemore important determining plant communi-ty patterns (Fowler, 1981). The soil organichorizon is related to chemical composition, biological activity, fertility, and soil development. Soil nutrients distributions vary at asmall-scale, affecting plant development andspecies distributions. For instance, nitrogenavailability commonly limits plant growth inforested ecosystems (Aber

et al.

, 1989). Inupland forests, N mineralization rates have been related to tree species composition(Pastor

et al.

1982; Zak

et al.

, 1986; Garten

et al.

, 1994) and productivity (Pastor

et al.

,1984; Liu and Muller, 1993).Habitat specialization for soil resources in atropical dry forest is likely to be more impor-tant at early stages in tree life histories thanit is in later life history (Vargas-Rodríguez

et al.

, 2005). This fact is very relevant for understanding tree establishment and rege-neration and could help guide land managersin implementing restoration policies. Howe-ver, environmental variables explainingmain small-scale gradients in communitycomposition of pine-oak forests in northernJalisco have not been explored yet. Treecommunity variation needs to be evaluatedat a much larger scale, however communityvariation of early stages in tree life histories(e.g. saplings) may be studied at smaller scales. Here we use 25 quadrats (0.04 ha)within 1 ha plot to focus on tree communities patterns at early stages in life tree histories(woody species > 2.5 cm in diam.) within asmall area, with the advantage of low habitatand climatic heterogeneity.In the present study, we aim to generatehypotheses, rather than to test them, aboutwhat environmental gradients are evidentat a hectare plot and which environmen-tal variables possibly explain small-scalecommunity gradients at early stages of treelife histories of a pine-oak forest in NuevaColonia, Mezquitic, Jalisco, Mexico.

M

ETHODS

Study area

The study area was located at 6 km northeastof Nueva Colonia, southeast of Mezquitic,Mezquitic municipality, Jalisco, Mexico. Welocated a hectare plot within the tributarywatershed Los Coyotes, along an 1800-2400altitudinal gradient, at the southern portionof Sierra Madre Occidental (Fig. 1). Theclimate at Nueva Colonia was temperatewith cold and moist winters (Vázquez-Gar-cía

et al.

, 2004). Mean annual rainfall was600 mm with the wet season from July toAugust. Relief ranged from 5-20% slope, of southern aspect. Volcanic acidic extrusiverocks and igneous Toba rocks were prevalentin the area. Silt soils and ferric Luvisol soilswere common in the area (INEGI 1970a,1970b).Floristic checklists for the municipalityincluded over 373 vascular plant species(Vázquez-García et al. 2004). The dominantvegetation was pine and pine-oak forests. Inaccordance with vegetation class, the most

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