Edge-effects on tree regeneration in the Colombian Andes
conclude that random effects prevail and no edge principles can be uncovered. The
experimental design in future research should take into account larger sample sizes of
edges per fragment and more fragments, to uncover the general trends of edge-effects
within a landscape.
Species-specific responses: Tree species responses varied in terms of abundance,
germination, seedling growth, seed predation and seedling herbivory as a function of
distance from the forest edge. This is a common pattern found in other studies on tree
regeneration (Osunkoya 1994; Meiners et al. 2000), and seems to be a generality of edgeeffects
(Laurance et al. 1997). I found that tree species varied in their responses to edgeeffects
based on regeneration habit, seed size, cotyledon morphology and susceptibility to
herbivory. My results also suggested that seed predators and herbivores might have
species-specific responses. Future work on the effects of fragmentation on tree
regeneration should focus on examining the response of multiple plant species with
different attributes. Such responses should be linked to the species physiology and
tolerance to fragmentation and to seed predation and herbivory. In addition, it is
essential to know the composition of seed predators and seedling herbivores, as well as
their diets and habitat preferences. Only when the ecology of the study species is taken
into account, will general patterns for the basis of species-specific responses to
fragmentation emerge.
The knowledge of species regeneration ecology is critical to implement and apply
practices for forest management and recovery. For example, open edges can be
protected by sowing fast-growing trees at the boundary of the forest-pasture to buffer the
224
impact of climatic extremes (Bierregaard et al. 1997). In addition, the matrix vegetation
type and its structure can determine the severity of edge-effects; cattle pastures, as in my
system are often the least preferable matrix land use (Laurance et al. 1997; Mesquita et
al. 1999). Abandoning pastures for natural regeneration is recommended but it is a slow
process and will not necessarily result in a natural forest structure. Thus, reforestation
with native species would be a complementary practice, but it is expensive and requires
extensive knowledge of native species and their ecology (Bierregaard et al. 1997).
Edge-effects on tree regeneration of Andean remnant forest generated changes in
the abiotic environment that were linked to changes in forest structure and composition as
a function of distance from the forest edge. Some of the mechanisms behind these
responses seem to be edge-mediated effects of the abiotic environment on seed
germination and seedling growth for some species. On the other hand, seed predation
and seedling herbivory, though critical for plant survival, were not edge-mediated effects
of tree regeneration. Consequently, edge-effects impacted the community integrity of
these remnants by affecting ecological patterns, mechanisms, and dynamics of tree
regeneration. Edge-effects are very complex to interpret and predict due to multiple
processes involved. Future research with rigorous experimental design should continue
testing mechanistic hypotheses to interpret observed edge-related patterns.
225
LITERATURE CITED
Angulo-Sandoval, P. and T. M. Aide. 2000b. Effect of plant density and light availability
on leaf damage in Manilkara bidentata (Sapotaceae). Journal of Tropical Ecology
6:447-464.
Augspurger, C. J. and K. Kitajima. 1992. Experimental studies of seedling recruitment
from contrasting seed distributions. Ecology 73:1270-1284.
Benítez-Malvido, J. 1995. The ecology of seedlings in Central Amazonian forest
fragments. Downing Street, Cambridge, Department of Plant Sciences,
Cambridge University: 160.
Benítez-Malvido, J. 1998. Impact of forest fragmentation on seedling abundance in a
tropical rain forest. Conservation Biology 12:380-389.
Bierregaard Jr., R. O., Laurance, W. F. , Sites, Jr. J. W., Lynam, A. J., Didham, R. K.,
Andersen, M., Gascon, C., Tocher, M. D., Smith, A. P., Viana, V. M., Lovejoy,
T. E., Sieving, K. E., Kramer, E. A., Restrepo, C. and C. Moritz. 1997. Key
Priorities for the Study of Fragmented Tropical Ecosystems. in J. W. F. Laurance
and R. O. Bierregaard, editor. Tropical Forest Remnants. The University of
Chicago Press, Chicago.
Blundell, A. G. and D. R. Peart. 1998. Distance-dependence in herbivory and foliar
condition for juvenile Shorea trees in Bornean dipterocarp rain forest. Oecologia:
117:151-160.
Burkey, T. V. 1993. Edge effects in seed and egg predation at 2 neotropical rainforest
sites. Biological Conservation 66: 139-143.
226
Camargo, J. L. and V. Kapos. 1995. Complex edge effects on soil moisture and
microclimate in Central Amazonian forest. Journal of Tropical Ecology 11:205-
221.
Chen, J., Franklin, J.F. and T. A. Spies. 1992. Vegetation responses to edge environments
in old-growth Douglas-fir forests. Ecological Applications 2:387-396.
Chen, J., Franklin, J. F. and T. A. Spies. 1995. Growing-season microclimatic gradients
from clearcut edges into old-growth douglas-fir forests. Ecological Applications
5:74-86.
Cintra, R. 1997. A test of the Janzen-Connell model with two common tree species in
Amazonian forest. Journal of Tropical Ecology 13:641-658.
Dalling, J. W., Swaine, M. D. and N. C. Garwood. 1998. Dispersal patterns and seed
bank dynamics of pioneer trees in moist tropical forest. Ecolgy 79:564-578.
Davies, K. F., Gascon, C. and C. R. Margules. 2001. Habitat fragmentation.
Consequences, management, and future research priorities. in M. E. S. G. H.
Orians, editor. Conservation Biology. Island Press, Washington, D. C.
Diaz, I., Papic, C. and J. J. Armesto. 1999. An assesment of post-dispersal seed predation
in temperate rain forest fragments in Chiloe Island, Chile. Oikos 87: 228-238.
Didham, R. K. 1997. The Influence of Edge Effects and Forest Fragmentation on Leaf
Litter Invertebrates in Central Amazonia. in J. W. F. Laurance and R. O.
Bierregaard, editor. Tropical Forest Remnants. The University of Chicago Press,
Chicago.
Dudt, J. F. and D. J. Shure. 1994. The influence of light and nutrients on foliar phenolics
and insect herbivory. Ecology 75:86-98.
227
Fagan, W. F., Cantrell, R. S. and C. Consner. 1999. How habitat edges change species
interactions. American Naturalist 153:165-182.
Forman, R. T. T. 1995. Land Mosaics. The Ecology of landscapes and regions.
Cambridge University Press.
Gehlhausen, S. M., Schwartz, M. W. and C. K. Augspurger. 2000. Vegetation and
microclimatic edge effects in two mixed-mesophytic forest fragments. Plant
Ecology 147:21-35.
Holl, D. K. and M. E. Lullow. 1997. Effects of species, habitat, and distance from edge
on post-dispersal seed predation in a tropical rainforest. Biotropica 29:459-468.
Hansson, L. 1998. Local hot spots and their edge effects: Small mammals in oak-hazel
woodland. Oikos 81:55-62.
Jordano, P. C. and M. Herrera. 1995. Shuffling the offspring: Uncoupling and spatial
discordance of multiple stages in vertebrate seed dispersal. Ecoscience 2:230-237.
Jose, S., Gillespie, A. R., George, S. J. and B. M. Human. 1996. Vegetation responses
along edge-to-interior gradients in a high altitude tropical forest in peninsular
India. Forest Ecology and Management 87:51-62.
Jules, E. S. and B. J. Rathcke. 1999. Mechanisms of reduced Trillium recruitment along
edges of old-growth forest fragments. Conservation Biology 13:784-793.
Kapos, V. 1989. Effects of isolation on the water status of forest patches in the Brazilian
Amazon. Journal of Tropical Ecology 5:173-185.
Kyllo, D. 2001. Effects of a common mycorrhizal network and light on growth and
community structure of understory shrubs, Piper and Psychotria, in a moist
neotropical forest. Ph. D. University of Missouri-St. Louis, St. Louis, MO.
228
Laurance, W. F., Bierregaard Jr., R. O., Gascon, C., Didham, R. K., Smith, A. P.,
Lynam, A. J., Viana, V. M., Lovejoy, T. E., Sieving, K. E., Sites, J. W., Andersen,
M., tocher, M. D., Kramer, E. A., Restrepo, C. and C. Moritz. 1997. Tropical
Forest Fragmentation: Synthesis of a Diverse and Dynamic discipline. in Tropical
Forest Remnants. The University of Chicago Press, Chicago.
Laurance, W. F., Gascon, C. and J. M. Rankin-de Merona. 1999. Predicting effects of
habitat destruction on plant communities: A test of a model using Amazonian
trees. Ecological Applications 9:584-554.
Levey, D. J. and M. M. Byrne. 1993. Complex ant-plant interactions: rain forest ants as
secondary dispersers and post-dispersal seed predators. Ecology 74:1802-1812.
Matlack, G. R. 1993. Microenvironment variation within and among forest edge sites in
the Eastern United States. Biological Conservation 66:185-194.
Meiners, S. J., Handel, S. N. and S. T. A. Pickett. 2002. Tree seedling establishment
under insect herbivory: Edge effects and inter-annual variation. Plant Ecology
151:161-170.
Mesquita, R. C. G., Delamonica, P. and W. F. Laurance. 1999. Effect of surrounding
vegetation on edge-related tree mortality in Amazonian forest fragments.
Biological Conservation 91:129-134.
Murcia, C. 1993. Edge effects on the pollination of tropical cloud forest plants. Ph.D.
University of Florida, Gainesville.
Murcia, C. 1995. Edge effects in fragmented forests: implications for conservation. Tree
10:58-62.
Osunkoya, O. O. 1994. Postdispersal survivorship of north Queensland rainforest seeds
229
and fruits: Effects of forest, habitat and species. Australian Journal of Ecology
19:52-64.
Pereira de Souza, R. and I. F. M. Válio. 2001. Seed size, seed germination, and seedling
survival of Brazilian Tropical Tree Species Differing in Successional Status.
Biotropica 33:447-457.
Ranney, J. W., Bruner, M. C. and J. B. Levenson. 1981. The importance of edge in the
structure and dynamics of forest islands. Pages 67-95 in R. L. Burgess and D. M.
Sharpe, editor. Forest Island Dynamics in Man-Dominated Landscapes. Springer
-Verlag, New York.
Renjifo, L. M. 1999. Effect of the landscape matrix on the composition and conservation
of forest bird communities. Ph. D. University of Missouri-St. Louis, St. Louis.
Restrepo, C. and A. Vargas. 1999. Seeds and seedlings of two neotropical montane
understory shrubs respond differently to anthropogenic edges and treefall gaps.
Oecologia 119:419-426.
Restrepo, C., Gomez, N. and S. Heredia. 1999. Anthropogenic edges, treefall gaps, and
fruit-frugivore interactions in a neotropical montane forest. Ecology 80:668-685.
Saunders, S. C., Chen, J. Drummer, T. D. and T. R. Crow. 1999. Modeling temperature
gradients across edges over time in a managed landscape. Forest Ecology and
Management 117:17-31.
Schupp, E. W. 1995. Seed-seedling conflicts, habitat choice and patterns of plant
recruitment. American Journal of Botany 82:399-409.
Sizer, N. C. 1992. The impact of edge formation on regeneration and litterfall in a
tropical rain forest fragment in Amazonia. Ph.D. Cambridge University, London.
230
Sork, V. L. 1983. Distribution of pignut hickory Carya glabra along a forest to edge
transect, and factors affecting seedling recruitment. Bull. Torrey Bot. Club
110:494-506.
Terborgh, J., Losos, E., Riley, M. P. and M. Bolanos Riley. 1993. Predation by
vertebrates and invertebrates on the seeds of five canopy tree species of an
Amazonian forest. Vegetation 107/108:375-386.
Turner, I. M. and R. T. Corlett. 1996. The conservation value of small isolated fragments
of lowland tropical rain forest. Trends in Ecology and Evolution 11:330-333.
Wales, B. A. 1972. Vegetation analysis of northern and southern edges in a mature oakhickory
forest. Ecological Monographs 42:451-471.
Williams-Linera, G. 1990a. Origin and early development of forest edge vegetation in
Panama. Biotropica 22:235-241.
Williams-Linera, G. 1990b. Vegetative structure and environmental conditions of forest
edges in Panama. Journal of Ecology 78:356-373.
Williams-Linera, V., Domínguez-Gastelú, V. and M. E. García-Zurita . 1998.
Microenvironment and floristics of different edges in a fragmented tropical
rainforest. Conservation Biology 12:1091-1102.
Young, A. and N. Mitchell. 1994. Microclimate and vegetation edge effects in a
fragmented podocarp-broadleaf forest in New Zealand. Biological Conservation
67:63-72.
231