INTRODUCTION
The factors that influence the ecology of popula-
tions and communities have been traditionally stud-
ied at the local level (Dunning et al., 1992), howev-
er the development of analytical tools such as
Geographic Information Systems (GIS) makes it
possible to evaluate these factors across much
broader scales. The concurrent analysis of habitat
characteristics (i.e., the structure and composition of
vegetation) and landscape characteristics (the com-
position and configuration of habitat patches) are
important for evaluating the use of habitat remnants
by native wildlife in fragmented landscapes (Gor-
resen and Willig, 2004; Pinto and Keitt, 2008;
Klingbeil and Willig, 2009).
Phytophagous bats provide crucial environmen-
tal services by pollinating (nectarivorous species)
and dispersing the seeds (frugivorous species) of na-
tive woody plants across anthropogenic landscapes,
and they do this over larger areas than other fauna,
including birds, owing to their greater vagility in
these landscapes (Galindo-Gonzalez, 2004). The
ability of some bat species to fly long distances and
cross open areas has made these volant mammals
a useful group for evaluating the response of native
fauna to habitat fragmentation (Gorresen and Willig,
2004; Klingbeil and Willig, 2009). In environments
that have been modified by humans, it has been
shown that bat species respond differently to changes
in the habitat; some bat species that depend directly
on plants as their main source of food benefit from
Acta Chiropterologica, 15(2): 399–409, 2013
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
doi: 10.3161/150811013X679026
Response of phytophagous bats to patch quality and landscape attributes
in fragmented tropical semi-deciduous forest
BEATRIZ BOLÍVAR-CIMÉ
1, 5, 6
, JAVIER LABORDE
1
, M. CRISTINA MACSWINEY G.
2
, CARLOS MUÑOZ-ROBLES
3
,
and JUAN TUN-GARRIDO
4
1
Red de Ecología Funcional, Instituto de Ecología, A.C., Carretera antigua a Coatepec No. 351, El Haya, C.P. 91070, Xalapa,
Veracruz, México
2
Centro de Investigaciones Tropicales, Universidad Veracruzana, Casco de la ExHacienda Lucas Martín,
Privada de Araucarias S/N, Col. Periodistas, C.P. 91019, Xalapa, Veracruz, México
3
Instituto de Investigación de Zonas Desérticas y Coordinación de Ciencias Sociales y Humanidades, UASLP,
Altair # 200 Col. Del Llano, C.P. 78377, SLP, México
4
Campus de Ciencias Biológicas y Agropecuarias, Universidad Autónoma de Yucatán, Carretera a Xmatkuil Km 15.5,
C.P. 97315, Mérida, Yucatán, México
6
Corresponding author: E-mail: bolivar_cime@yahoo.com
Forest fragmentation reduces the amount of forest cover and negatively affects the habitat quality of forest remnants. Landscape
attributes and habitat quality should therefore be evaluated together to improve our understanding of how fauna respond to
fragmentation. We evaluated how patch quality (vegetation structure) and landscape characteristics influence the abundance of
phytophagous bats in two contrasting types of landscape that differ in percent forest cover and matrix type: landscapes dominated
by man-made pastures and landscapes with large tracts of continuous forest (tropical semi-deciduous forest). Bats were sampled in
forest patches in both types of matrices, and for each matrix two sites with a cenote (water filled sink-holes, typical of the Yucatan)
and two with no cenotes were sampled. Sites with cenotes offer better habitat quality than sites without cenotes: the richness and
basal area of plants eaten (flower or fruit) by bats are higher in the forest vegetation surrounding them. At the landscape level,
phytophagous bat abundance was negatively correlated with the amount of forest cover and proximity to other forest fragments, but
positively correlated with forest edge density, patch density and landscape heterogeneity. At the patch level, bat abundance was
positively correlated with plant richness and the basal area of edible tree species. In the Yucatan’s agricultural landscapes the area
and spatial distribution of forest remnants are not the only variables affecting bats. Habitat patch quality and high heterogeneity of
land cover types are also important, and have a positive effect on phytophagous bat abundance and movement.
Key words: frugivorous bats, landscape complementation, landscape heterogeneity, nectarivorous bats, vegetation structure