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Abstract

Summary Tropical rain forest fragmentation is one of the most pervasive threats to the conservation of biological diversity, affecting different levels of biological organization including populations, communities and ecosystems. Forest fragmentation involves the creation of "habitat edges" and consequently the so called "edge effects" that generally have a negative impact on the biotic and physical environment. The spatial attributes of fragments in the landscape include fragment size, shape, isolation and the matrix type surrounding the fragments. Although these spatial attributes influence the prevalence and magnitude of the edge effects, they can constitute important threats to biodiversity by themselves. The increment of fragment isolation in highly fragmented landscapes can negatively affect inter-fragment dispersal movements of both plant and animal species, modifying important ecological processes such as pollination and seed dispersal. In this sense, actions to increase the population size and the persistence of several plant and animal species include the establishment of biological corridors. Biological corridors increase landscape connectivity, and may reduce extinction rates by increasing inter-fragment movements and favoring the access to resources available in more than one forest fragment.
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To cite this chapter: Benítez-Malvido J, Arroyo-Rodríguez V. 2008. Habitat fragmentation, edge effects and biological corridors in
tropical ecosystems. In: Encyclopedia of Life Support Systems (EOLSS). Del Claro K, Oliveira PS, Rico-Gray V, Ramirez A,
Almeida AA, Bonet A, Scarano FR, Consoli FL, Morales FJ, Naoki J, Costello JA, Sampaio MV, Quesada M, Morris MR,
Palacios M, Ramirez N, Marcal O, Ferraz RH, Marquis RJ, Parentoni R, Rodriguez SC, Luttge U (editors). International
Commision on Tropical Biology and Natural Resources. UNESCO, Eolss Publishers, Oxford ,UK, [http://www.eolss.net]
[Retrieved August 29, 2008]
HABITAT FRAGMENTATION, EDGE EFFECTS AND BIOLOGICAL
CORRIDORS IN TROPICAL ECOSYSTEMS
Julieta Benítez-Malvido and Victor Arroyo-Rodríguez
Centro de Investigaciones en Ecosistemas, Universidad Nacional Autonoma de Mexico, Antigua Carretera a Patzcuaro
No. 8701, Ex−Hacienda de San Jose de la Huerta, Morelia, Michoacan, Mexico
Summary
Tropical rain forest fragmentation is one of the most pervasive threats to the conservation
of biological diversity, affecting different levels of biological organization including
populations, communities and ecosystems. Forest fragmentation involves the creation of
"habitat edges" and consequently the so called "edge effects" that generally have a negative
impact on the biotic and physical environment. The spatial attributes of fragments in the
landscape include fragment size, shape, isolation and the matrix type surrounding the
fragments. Although these spatial attributes influence the prevalence and magnitude of the
edge effects, they can constitute important threats to biodiversity by themselves. The
increment of fragment isolation in highly fragmented landscapes can negatively affect
inter-fragment dispersal movements of both plant and animal species, modifying important
ecological processes such as pollination and seed dispersal. In this sense, actions to increase
the population size and the persistence of several plant and animal species include the
establishment of biological corridors. Biological corridors increase landscape connectivity,
and may reduce extinction rates by increasing inter-fragment movements and favoring the
access to resources available in more than one forest fragment.
Keywords: Biodiversity loss, connectivity, extinction, fragment size, habitat loss, isolation, tropical
rain forest.
1. Introduction
Deforestation and forest fragmentation have become the most important threats for the
maintenance of biodiversity. Tropical rain forests are one of the most affected ecosystems
with annual rates of deforestation between 100 000 and 150 000 km2. Tropical forests are
also one of the most biodiverse ecosystems of the planet as they contain between 50% and
80% of all the terrestrial species, and they have a critical role on the maintenance of the
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planet homeostasis. Therefore, their destruction may not only threaten the maintenance of
biodiversity, but could also affect climatic and hydrological cycles at local, regional and
global scales. In addition to the loss of forest cover, the process of fragmentation results in
a change on the spatial pattern of the remaining forest (e.g., increase in number of forest
fragments, decrease in fragment size, and increase in fragment isolation), leading to the loss
of ecosystem continuity. These spatial changes produce a wide range of effects across
several levels of biological organization, affecting biological populations and communities,
as well as ecological processes that may modify the overall functioning of the ecosystem.
The magnitude of the effects that tropical rain forest fragmentation has on the biota
and physical environment depend on different elements or aspects that characterize the
fragmented landscapes including: total amount of forest cover, number of forest fragments,
fragment size, shape and isolation, and the characteristics of the matrix (i.e., modified
native vegetation such as deforested areas, cattle pasture, agricultural crops, urban areas,
etc.) that surrounds the fragments. These same elements would also determine the
magnitude of the so called "edge effects", which are an inevitable consequence of forest
fragmentation and imply the influence of processes originated in the matrix that surrounds
the fragments. In this chapter we describe the consequences that tropical rain forest
fragmentation has across different levels of biological organization including populations,
communities and ecosystems, as well as on the physical environment of the remaining
forest. Thereafter, we describe the influence of edge effects and fragment attributes (i.e.,
size, shape, isolation and surrounding matrix) on the biota and physical environment.
Finally, we pointed out the importance and inconveniences of the so called biological
corridors, present in some fragmented landscapes and relevant for the maintenance of the
remaining plant and animal populations within the fragments. For each section, the
examples given on fragmentation effects come from studies conducted in different tropical
forests around the world.
2. Habitat Fragmentation
Habitat may be broadly defined as the range of environments suitable for a given species.
That is, it is a species-specific concept. Therefore, to simplify the present synthesis, we
equated "habitat" with "native tropical rain forest", as this vegetation type is very important
for a large number of animal and plant species. Thus, we define habitat fragmentation as a
landscape-scale process in which the continuous habitat is reduced into smaller habitat
remnants. This implies the loss of habitat and its sub-division into a variable number of
remaining fragments scattered within a matrix of modified habitat. In addition to the
changes on habitat pattern described above, the primary effect of fragmentation is the
alteration of the microclimate within and around the fragment. These environmental
changes have major implications on several plant and animal species; modify biotic
interactions and the functioning of the ecosystems. While some changes on the habitat are
visible immediately after fragmentation (e.g., shifts in habitat pattern, changes in
population sizes, forest structure and composition at edges) others may appear in the long
term (e.g., genetic related changes on populations, extinction of species with slow life
cycles).
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The effects of habitat fragmentation on the tropical biota are very variable and
depend on several factors, such as: (1) taxon characteristics (e.g., population size, habitat
and diet requirements, dispersal capacity, etc.); (2) the spatial scale of the analysis; (3) the
ecological processes under study; (4) habitat type; and (5) landscape characteristics (e.g.,
topography, soil type, etc.). Furthermore, there are factors that are not easily detected and
therefore are difficult to evaluate, such as: (1) the response time of populations to
fragmentation (e.g., denominated "extinction debt"); (2) the biogeographical position of the
species under study; and (3) synergisms between different processes (e.g., fragmentation
and hunting; fragmentation and climatic change).
2.1. Impact of Habitat Fragmentation on Populations
Fragmentation involves the decline of many plant and animal populations, altering birth,
mortality and growth parameters, conducting in some cases to the local and/or regional
extinction of particular species. The extinction proneness of a species in a fragmented
habitat is related to factors such as habitat or niche specialization, home range size,
mobility and dispersal capacity, extent of geographic distribution, population density or
rarity, edge sensitivity, body size, trophic level, age-sex class and dietary specialization.
For example, species with restricted geographic distributions and/or reduced population
density are more sensitive to extinction, as these species are the most vulnerable to
stochastic (i.e., unpredictable) threatening processes such as environmental (e.g.,
fluctuations in climate, natural catastrophes), demographic (e.g., year-to-year variability in
reproductive success) and genetic (e.g., genetic drift) stochasticity. In contrast, species with
larger populations are less sensitive to stochastic threatening processes, and therefore, they
have a higher probability to persist in the long term.
In addition to the stochastic threatening processes, populations are confronted with
several deterministic (i.e., predictably lead to population declines) threats; which can be
classified as exogenous (originating independently of the species’ biology) or endogenous
(originating as part of the species’ biology). For example, habitat loss, habitat degradation,
anthropogenic pressures (e.g., logging and poaching), sub-division and habitat isolation are
all exogenous threatening processes that can affect a declining population inhabiting
fragmented landscapes; while disruptions to dispersal, changes in social systems and
physiological stress, and loss of genetic diversity are endogenous threatening processes that
can accelerate population decline in fragmented habitats. Continuing with the factors
described above, species with larger home range size and/or high dietary specialization will
be more vulnerable to exogenous threats such as habitat loss and degradation; while species
with low dispersal capacity through the matrix will be more vulnerable to the sub-division
and isolation of the habitat.
Therefore, fragmentation effects on the population of particular species cannot be
generalized, as each species responds individualistically to a range of processes related to
its requirements for food, shelter, space, suitable climatic conditions and to interspecific
processes (e.g., competition, predation and mutualisms). Although the majority of the
species are directly or indirectly affected by fragmentation, plant and animal species in the
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tropics present three types of responses to the fragmentation of their habitat: (1) a positive
response (e.g., increased population size of pioneer plant species, some invertebrates and
some rodents in fragments compared with continuous forested areas); a negative response
(e.g., decline in population size of many primates species, loss of genetic diversity in
howler monkeys of Belize, death of large trees in Brazil, and local extinction of some dung
and carrion beetles in Brazil); and (3) a neutral response (no effects) (e.g., the population
size of some plant species in 1 000 m2 of continuous forest did not differ from 1 000 m2 of
a fragmented landscape of Los Tuxtlas, Mexico).
2.2. Fragmentation Effects on Communities
The effects of fragmentation on tropical plant and animal communities include changes in
species diversity, composition, abundance, distribution and biotic interactions. Probably the
most important is the loss of biological diversity. However, as forest fragments are prone to
be colonized or invaded by exotic plant and animal species and diseases, in some tropical
forests some taxa (e.g., some frogs, small rodents, and secondary plant species) have shown
an increment in species richness after fragmentation; whereas, in some others, there are not
differences in species richness after fragmentation (e.g., richness of some arthropods was
similar in large and small fragments in a study in south-eastern Australia). Disruptions to
species interactions have particularly severe consequences when keystone species are
involved as these species play an important role in maintaining ecosystem function and
structure. For example, some predators (e.g., jaguar, puma, ocelot) are important regulators
of the density of many herbivore populations, and their disappearance may favor the
overexploitation of the vegetation by herbivores that increase their population sizes due to
the lack of predators. Similarly, some tree species (e.g., the genus Ficus) are important food
sources for many species of birds and mammals and their disappearance or altered
phenology may affect numerous species of animals.
In fragmented tropical communities changes in biotic interactions affect
competition, predation, parasitism and mutualisms (e.g., seed dispersal and pollination).
Nearly 90% of tropical tree species are dispersed by animals. Therefore, plant species
depending on animals for the dispersal of their seeds may be strongly affected in a
fragmented community. The extinction of animal seed dispersers may reduce the
distribution areas and population size of several plant species, or may reduce the possibility
of colonizing new habitats. These effects increase population isolation eventually leading
them to extinction. In Uganda, tree species dispersed by chimpanzees failed to recruit into
forest fragments where this primate species was lacking. In isolated land-bridge islands in
Lago Guri, Venezuela, several islands have populations of howler monkeys at densities up
to more than 30 times greater than those on the mainland, and researchers have
demonstrated that these "hyperabundant" herbivorous have a strong positive influence on
aboveground plant productivity (by increasing the transference of nutrients through their
feces), with a positive, indirect effect on bird species richness.
The plant-pollinator interaction is sensitive to any kind of disturbance. In small and
isolated fragments pollen flux is reduced, affecting fruit and seed set with harmful
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consequences to the quality and quantity of the progeny. Fragmentation affects several
types of pollinators with strong effects in the reproductive success on plant populations and
on the genetic structure of remnant populations. Pollination in fragmented habitats includes
a reduction in the abundance of pollen vectors as consequence of changes in the
environment and in the availability of resources, reduced frequency of visits as the
distribution of floral resources changes, and by competitive exclusion of floral resources by
exotic pollinators. For example, various tropical tree species have shown reduced fruit and
seed set in fragmented habitats (e.g., Samaena saman, Dinizia excelsa, Shorea siamensis,
Clatasetum viridiflavum) due to reduced pollination and increased isolation.
2.3. Fragmentation Effects on Ecosystems
Most research on habitat fragmentation has been focused on plant and animal populations
and communities, and rarely on ecosystem processes. The principal climatic changes
produced by forest fragmentation affecting ecosystem functioning and that are particular
harmful at forest edges and small fragments (< 10 ha) include radiation fluxes, wind
incidence, fire frequency and changes in the hydrological cycle (e.g., evapotranspiration). It
has been suggested that not just species richness, but functional diversity is important to
maintain nutrient and energy fluxes in the ecosystems. Solar radiation, carbon dioxide,
temperature, water and soil nutrients are important for the primary productivity of tropical
rain forests and are strongly modified by habitat fragmentation. The magnitude of such
impacts depends on the size of the fragment as well as on its orientation, slope and matrix
type.
Energy balance within a fragmented forest differs from that of a continuous forest,
especially when the original natural vegetation was denser than that developing after
fragmentation. Vegetation structure is modified in a fragmented habitat altering wind
fluxes, reducing humidity and increasing desiccation. In tropical forests the incidence of
warm and dry winds in fragmented areas increases tree mortality and the incidence of fires.
Strong winds reduce the substrate available for microorganisms and the resources coming
from the soil. The removal of the natural vegetation changes the rates of water interception,
water loss and evapotranspiration altering the hydrological cycle and may also increase soil
erosion.
The kind of matrix (see below) that surrounds the fragment, for example, affects
radiation balance due to increase isolation on the fragment surface. For disturbed forests, in
general, diurnal temperatures are higher and nocturnal temperatures are lower than the ones
present in the unperturbed forest. In Manaus, Brazil, temperatures were around 3 ºC higher
in fragments than in the continuous forest. The alteration on temperature regimes could
affect nutrient cycling as well as the biology of several plants (e.g., seed germination) and
animals (e.g., egg hatching in some insect species) as well as biotic interactions such as
competition, predation and parasitism (e.g., fungal infection on leaves). Furthermore,
fragments may accumulate pollutants and nutrients that are transported by the wind from
urban and agricultural areas, at their edges, affecting nutrient cycling and microbial
activity.
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3. Edge Effects
Forest fragmentation implies the creation of "habitat edges" and consequently the so called
"edge effects". The edge effect could be defined as the interaction of two adjacent
ecosystems separated through an abrupt transition. Forest fragmentation increases the
amount of edges in the landscape producing important physical (e.g., radiation, moisture,
temperature, wind speed, and soil nutrients) and biological (e.g., species composition,
competition, predation, etc.) changes along and close to the edge. Fragment edges are the
most altered area of a fragment and the penetration depth of the edge effects vary widely
from tens of meters (e.g., soil moisture in Manaus, Brazil) to several kilometers (e.g.,
recruitment failure of trees in Borneo). Forest edges may control the flux of organisms
between forest and non-forest habitats. Edges are also the point of entry of external
influences such as fire and the invasion of exotic species including pathogens to the
remaining forest.
The survival and persistence of several animal and plant species, and their
interactions (e.g., herbivory, pathogen infection on plants, nest predation, ant-plant
interactions, etc.) are affected close to the fragment edge. Therefore, ecological processes
close and along forest edges differ from those at forest interior. Naturally, edge effects vary
widely between species, and contrasting edge responses have been found between species
with different life history strategies and habitat requirements. Studies in plant ecology have
found that pioneer species (light demanding) have a speedy growth near forest edges, while
old-growth species (shade tolerant) can establish and growth only under the closed canopy
in the forest interior. Not all edges are necessarily detrimental for all native species,
especially when edges are gradual or of low structural contrast (e.g., developed secondary
forests).
4. Influence of the Spatial Attributes of the Fragments
Here, we grouped fragmentation effects under four categories that together describe the
spatial attributes of individual fragments in fragmented landscapes, and influence the
incidence and magnitude of the edge effects: (1) fragment size; (2) fragment shape; (3)
fragment isolation; and (4) matrix type.
4.1. Fragment Size
Fragment size is considered the most important spatial attribute that affects the maintenance
of biodiversity in a fragmented landscape. Small fragment area imposes a maximum limit
on population size that leaves species vulnerable to local extinction. For many plant and
animal species (e.g., old-growth tree species and large predators or carnivores) habitat
conditions are ideal in large areas of unmodified tropical rain forest vegetation. As
fragmentation increases, the size of the remaining fragments decreases. Smaller fragments
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may have lower number of species than larger ones, for example because resources in
smaller fragments may be more limited. Larger fragments may present larger populations
of different taxonomical groups and larger populations persist for longer time. Furthermore,
larger fragments may have higher colonization to extinction ratio, are more likely to contain
undisturbed areas which are required by some species, are more likely to hold a range of
environmental conditions which constitute adequate habitats for different sets of species,
and are more likely to capture species with patchy distributions. Small fragments may also
be more easily affected by anthropogenic disturbances such as hunting, logging, fires and
cattle grazing. However, in some situations smaller fragments may be more easily
colonized or invaded by exotic species increasing species diversity as small fragments
present greater proportion of edge. Small fragments (< 10 ha in area) may also provide the
last refuge for many native plants and animals for some ecosystems, as in the Atlantic
forest of Brazil.
4.2. Fragment Shape
The negative effects of fragment shape are decreased in large fragments as larger fragments
present a greater proportion of forest interior relatively free of edge effects. The proportion
of a forest fragment affected by edge effects depends on the relation between fragment size
and shape; being the smallest and most irregularly shaped fragments the ones with a greater
area affected by edge effects. The survival and persistence of several animal (e.g., beetles,
birds) and plant species (e.g., all life stages of tree species) and their interactions (e.g.,
herbivory, pathogen infection on plants, nest predation, ant-plant interactions, etc.) are
affected close to the fragment edge. The magnitude of such effects depends on the species
attributes, species habitat requirements and matrix type. However, in some cases large
irregular fragments may have a positive effect on the populations (e.g., howler monkeys),
for example, some large irregular fragment along rivers or living fences (fencerows) may
function as dispersal corridors or "stepping stones". In these large fragments, shape
complexity may have a positive effect on population persistence, as they can be colonized
more frequently than are compact fragments. Increased colonization of complex fragments
occurs because fragments with high shape complexity (e.g., ameba shape) have a
proportionally greater amount of edge, increasing the likelihood that a fragment will be
encountered by a moving individual. In Los Tuxtlas, Mexico, the local people maintain
native vegetation along streams and delimit their properties with living fences (i.e., trees).
These management practices produce more irregularly shaped fragments but reduce
considerably their isolation facilitating the movement of animals (i.e., primates) between
them. Nevertheless, the opposite trend has been observed for some animal species, as shape
complexity can also facilitate emigration. Thus, the combination of increased emigration
and colonization may lead to greater variability in the population size of the most complex
shaped fragments.
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4.3. Fragment Isolation
Fragmentation increases the isolation of the remaining habitat fragments. Isolation can
negatively affect day-to-day movements (e.g., between foraging resources) of a given
species, and dispersal of plants (e.g., seeds) and animals (e.g., juveniles and sub-adults).
The distance among the remaining forest fragments and the continuous forested area is also
important for the maintenance of biodiversity because the dispersal capacity of several
plant and animal species may be affected by increasing isolation. Some species could be
negatively affected by distances of less than 100 m between forest fragments, or between
forest fragments and continuous forest, as they are incapable of crossing cleared areas (e.g.,
some birds, euglossine bees, beetles, frogs, etc.). However, the negative effects of distance
may vary depending on the trait considered (e.g., population size, species richness, genetic
diversity), the taxa (e.g., births and bats may be less affected than arboreal mammals) and
habitat configuration. The negative effects of isolation may be reduced when there are
several small fragments between large fragments, as small fragments may act as "stepping
stones" for several species; or when two forested areas or fragments are connected by the so
called biological corridors that are formed by the remaining vegetation. Furthermore, some
matrix types disrupt dispersal more than others, as particular species can use some matrix
types for their mobility than others. For example, some plant species dispersed by gravity
(e.g., the palm Astrocaryum mexicanum in Los Tuxtlas, Mexico), could be drastically
affected by fragment isolation; whereas some others like those dispersed by animals and
wind are less affected.
In some cases, isolated populations may utilize the resources of more than one patch
for foraging, shelter, reproduction, etc. (a process named "landscape supplementation").
The ability or capacity of a species to utilize more than one patch may favor its persistence
in fragmented landscapes where resources may be scarce and of low quality. Other effect
related to fragment isolation is the so called "rescue effect" that occurs when a given
population is declining within a fragment and receives immigrant individuals from a
neighboring fragment "rescuing" the population that is in the verge of extinction.
4.4. Matrix Type
Matrix type (e.g., cattle pastures, secondary forest, urban and agricultural areas, mining,
etc.) and the related physical factors determine the magnitude of fragmentation effects on
the biota and the physical environment. The type of matrix affects the dispersal of plants
and animals; may provide food and space for the species capable to survive in it, and may
determine the magnitude of edge effects within the fragment. For example, in the Central
Amazon if a forest fragment is surrounded by a secondary vegetation dominated by large
and dense Cecropia trees, edge effects on the vegetation are less detrimental than when
surrounded by a secondary vegetation dominated by the more slender and sparse Vismia
trees. Furthermore, a number of studies have demonstrated that generalist species - which
can use also resources from the matrix - maintained higher populations in fragmented
landscapes than specialist species that depended on resources available only in fragments.
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The structure and composition of the matrix ("effective isolation") can resist, hinder
or enhance movement behavior, affecting colonization/extinction dynamics within the
fragments, and population size. Therefore, fragment isolation does not solely depend on the
Euclidian distance between fragments but on the matrix type surrounding them. The matrix
may also provide suitable habitat for some native species, especially if it is structurally
similar to the native vegetation within the fragments, enhancing food availability and
habitat connectivity for these species. For example, in Mexican coffee plantations ants were
actively foraging in the surrounding matrix and some species were even able to survive in
the matrix habitat in perpetuity. Similarly, in some fragmented landscapes of Africa, some
mammals species that inhabit forest fragments, can take some resources from the matrix.
5. Biological Corridors
Biological corridors refer to linear elements or strips of vegetation (but not necessarily of
native vegetation) in the landscape, connecting two or more habitat patches that have been
linked in historic time and that function as passageway for the biota. Thus, biological
corridors increase connectivity between different elements of a landscape. Connectivity
determines the connectedness between fragments of suitable habitat (commonly equated to
native vegetation), as well as the connectedness of ecological processes (e.g., biotic and
abiotic interactions, hydrological flows) across multiple scales. Therefore, as previously
discussed for the opposite process (i.e., fragment isolation), lack of connectivity may be a
major threat for wildlife, and the establishment and maintenance of landscape
linkages/connectivity of native vegetation networks or wildlife corridors, is a common
conservation strategy.
Biological corridors are aimed to increase the exchange/movement of individuals
between forest fragments, promote new foraging areas for many animals, act as refuges for
several plant and animal species, act as wind barriers diminishing edge effects, and can stop
the desiccation of rivers and streams (e.g., riparian corridors). However, biological
corridors may not be used by all native species, not all ecological processes are effectively
facilitated though biological corridors (e.g., disturbance processes), and depending on their
shape they are prone to edge effects. Furthermore, they facilitate the spread of introduced
species and diseases, and increase the vulnerability to catastrophic events. Nevertheless,
biological corridors are usually more likely to have beneficial effects on native species and
ecological processes than detrimental effects favoring, on the whole, the persistence of the
species at local and regional scales.
The inclusion of biological corridors in reserve design has become an important
conservation tactic for protecting biodiversity. The Mesoamerican Biological Corridor
(MBC), for example, is a region-wide initiative covering 768 990 km2 and comprises
southern Mexico, Guatemala, Belize, El Salvador, Honduras, Nicaragua, Costa Rica and
Panama. Although the region contains only 0.5 % of the world’s land surface, it contains
around 7 % of the planet’s biological diversity. Mesoamerica does not only include about
22 eco-regions (e.g., lowland rainforest, grasslands, semi-arid woodlands, pine savannas,
etc.) but is also considered to be one of the world’s most important centers of the origin of
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agriculture. The MBC is intended to conserve biological and ecosystem diversity in a
manner that promotes sustainable social and economic development in the region.
6. Conclusions
Forest edges and fragments are becoming dominant features in many tropical landscapes
threatening native populations and communities and the functioning of tropical rain forest
ecosystems. The magnitude of fragmentation effects on the different levels of biological
organization depends on a complex interplay of factors related to fragment spatial
attributes. On the whole, despite of the fact that each species responds individualistically to
fragmentation, forest fragmentation is detrimental for the maintenance of biodiversity
because tropical forest species need functional ecosystems to survive. Fortunately,
conservation actions, such as the implementation of biological corridors at a landscape and
regional scales, are being undertaken to lessen fragmentation effects on tropical
ecosystems.
Acknowledgments: The authors thank the Biological Dynamics of Forest Fragments Project, in Manaus,
Brazil; the Los Tuxtlas Biosphere Reserve, Veracuz, Mexico; the Montes Azules Biosphere Reserve, Chiapas,
Mexico; and to their people, for providing the opportunity to gain knowledge and expertise on the fascinating
and challenging topic of tropical rain forest fragmentation.
Glossary
Habitat: The range of environments suitable for a particular species.
Habitat loss: Loss of habitat for a particular species.
Habitat fragmentation: The breaking apart of sub-division of continuous habitat for a particular species.
Landscape: Following Fischer and Lindenmayer (2007), a human-defined area ranging in size from ca. 3 km2
to ca. 300 km2.
Matrix: In modified landscapes usually not native vegetation (e.g., deforested areas, cattle pasture,
agricultural crops, urban areas, etc.) that surrounds the forest fragments of native vegetation.
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Biographical Sketches
- Julieta Benitez-Malvido received the Bachelors degree in Biology (cum laude) from Universidad Autonoma Metropolitana-
Iztapalapa, in Mexico City in 1988; the Master in Science degree in Ecology from Durham University, at the United Kingdom; and
the PhD. from Cambridge University, United Kingdom in 1994. She is a principal researcher in the Center for Ecosystem Research,
National Autonomous University of Mexico (UNAM, 1996-present); assistant professor in Population Ecology and Conservation;
and Research Associate, to the Biological Dynamics of Forest Fragments Project, National Institute for Research in the Amazon
(INPA)-Smithsonian Institution (1991-present) and at the Long Term Ecosystem Research (LTER), international program at the
Chamela site (Mexico). Current research sites include the Brazilian Amazon and several locations at tropical Mexico (Los Tuxtlas,
Chajul, Cozumel and Chamela). Research interests: tropical ecology; tropical forest recovery after human disturbances (e.g.,
fragmentation, deforestation and impact of roads); biotic interactions of plants with herbivores and pathogens in disturbed tropical
habitats, and tropical forest restoration.
- Victor Arroyo-Rodriguez received the B.Sc. (biology) degree from Universidad Autonoma de Madrid, Spain, in 2002, and both the
M.Sc. and Ph.D. degrees from Instituto de Ecologia A.C., Mexico, in 2005 and 2007, respectively. Nowadays he is a postdoctoral
fellow at the Centro de Investigaciones en Ecosistemas (Universidad Nacional Autonoma de Mexico). His main research interest is
conservation biology in human-modified tropical rain forests. He has published ca. 20 papers in wildlife ecology, forest ecology and
biodiversity conservation in fragmented landscapes, and has scientific presentations in numerous national and international
congresses and symposia.
... Additionally, it is vital to understand that the disruption of plantinsect relationships due to fragmentation can have cascading effects on ecosystem processes and services (Ewers and Didham, 2005;Rossetti et al., 2017). Changes in pollination, herbivory, and seed dispersal can alter plant community composition, leading to shifts in species dominance and diversity (Collinge and Palmer, 2002;Fahrig, 2003;Benítez-Malvido and Arroyo-Rodríguez, 2008). These shifts can affect higher trophic levels, including predators and parasitoids, further influencing ecosystem dynamics (Tscharntke et al., 2012). ...
... Insects like beetles and ants play critical roles in breaking down organic matter (Gormley et al., 2007), and their decline in fragmented habitats can slow nutrient cycling processes, affecting soil fertility and plant growth (Didham et al., 1996;Flores-Rentería et al., 2018). As such, habitat fragmentation poses significant challenges to plant-insect relationships, affecting pollination, herbivory, and seed dispersal (Benítez-Malvido and Arroyo-Rodríguez, 2008;De Carvalho Guimarães et al., 2014). This means that understanding the impacts of fragmentation on insect biodiversity and plant-insect interactions is essential for developing effective conservation and management strategies for habitats (Gormley et al., 2007). ...
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Fragmentation is threatening insect biodiversity and intricate interactions in various ecosystems. Ecological interactions – especially those involving plants and insects – are significantly impacted by fragmented habitats. Because of fragmentation, edge effects and reduced habitat connectivity and quality affect insect species diversity, abundance, behavior, movement, life cycles, and interactions with plants, e.g., pollination, herbivory, and seed distribution. To a large degree, ecosystem services or processes are mediated by these interactions. While fragmented habitats create suitable conditions for invasive alien plants (IAPs), such invasions modify native plant composition and herbivorous insect communities because they cause a decline or loss in insect biodiversity. A systematic review was conducted by reviewing eighty-eight (88) articles to gather evidence for fragmentation effects on insect biodiversity, insects’ behavior and adaptations, plant-insect interactions (i.e., pollination, herbivory, and seed dispersal), and its influence on IAP invasions. This review deduced that any change in insect community composition and diversity due to fragmentation can have cascading effects on ecosystem processes within habitats. It further contends that successful conservation and management of fragmented habitats requires an understanding of the intricate dynamics of plant-insect interactions. However, the long-term resilience and health of ecosystems can be guaranteed by supporting sustainable land use, improving connectivity, and restoring habitats. These actions may help stop and/or reduce the effects of fragmentation on insect biodiversity and support the livelihoods and well-being of millions of people.
... The mobility of the species from one region to another depends on various environmental factors, including climate conditions, habitat fragmentation, species competition, etc. [10,11]. It has a massive impact on the ecosystem. ...
... One of the major concerns is the fragmentation of the ecological habitats, which was considered an invasive threat to biodiversity. Habitat fragmentation can define as a landscape-scale process in which the continuous habitat is reduced into more minor habitat remnants [10]. The size, shape, edge of the habitat fragments, and habitat isolation are some significant factors having huge implications on the species interaction and species survival [7,11]. ...
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Spatio-temporal pattern formation over the square and rectangular domain has received significant attention from researchers. A wide range of stationary and non-stationary patterns produced by two interacting populations is abundant in the literature. Fragmented habitats are widespread in reality due to the irregularity of the landscape. This work considers a prey-predator model capable of producing a wide range of stationary and time-varying patterns over a complex habitat. The complex habitat is assumed to have consisted of two rectangular patches connected through a corridor. Our main aim is to explain how the shape and size of the fragmented habitat regulate the spatio-temporal pattern formation at the initial time. The analytical conditions are derived to ensure the existence of a stationary pattern and illustrate the role of the most unstable eigenmodes in determining the number of patches for the stationary pattern. Exhaustive numerical simulations help to explain the effect of the spatial domain size and shape on the transient patterns and the duration of the transients.
... Habitat loss and fragmentation decrease wildlife habitat availability and quality, leading to increased edge effects, human-wildlife interactions, and movement barriers (Padmakumar and Shanthakumar, 2023;Benitez-Malvido and Arroyo-Rodríguez, 2008). Edge effects significantly alter temperature, humidity, light, and wind conditions in inland forests (Vermont Department of Forests Council, 2015), impacting species composition and community structure (Andriatsitohaina et al., 2020). ...
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Indonesia, a global biodiversity hotspot, confronts escalating threats from land-use change, triggering extensive forest fragmentation and threatening biodiversity. This review synthesizes existing literature on Indonesian forest fragmentation, highlighting key findings, methodologies, and knowledge gaps. It assesses spatial fragmentation patterns, evaluates current conservation efforts, and identifies areas for enhancement. Utilizing the Driving Forces-Pressures-State-Impacts-Responses (DPSIR) framework, we comprehensively examine the intricate dynamics influencing forest fragmentation, emphasizing the role of driving forces, pressures, conditions, impacts, and responses in biodiversity conservation. Pressures such as agricultural expansion and infrastructure development induce changes in forest conditions and biodiversity, resulting in diverse impacts such as habitat destruction, altered animal behaviors, and human-wildlife conflict. These findings accentuate the pressing need for adaptive conservation strategies addressing the root causes of fragmentation. We propose a comprehensive biodiversity conservation strategy for fragmented landscapes, encompassing integrated land use planning, habitat connectivity, restoration, wildlife-friendly infrastructure, agroecology, community-based conservation, buffer zones, invasive species management, education, outreach, transboundary cooperation, translocation, monitoring, research, and innovation.
... The impact of habitat transformation (Ritchie and Roser, 2013;Winkler et al., 2021) on ecological characteristics of species has been well-studied from diverse perspectives, such as life histories characteristics (Kolb and Diekmann, 2005;Bruna et al., 2009), extinction probabilities (Fréville et al., 2007), plant animal interactions (Benítez-Malvido and Arroyo-Rodríguez, 2008;Benitez-Malvido et al., 2016) and reproductive success (Brudvig et al., 2015;Vellend et al., 2017). ...
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The tropical Andean landscape has been dramatically transformed over the last century with remaining native forest limited to small fragments within a heterogeneous matrix of crops, cattle pastures, and urban environments. We aimed to explore the impact of habitat transformation on the population dynamics in an endemic twig epiphytic orchid located within the undisturbed forest and within modified matrix habitat in two regions with contrasting landscape structures: with a dominant shade coffee matrix and a dominant grassland matrix. Over 2 years, we surveyed 4,650 individuals of the Colombian endemic orchid, Rodriguezia granadensis. We undertook four post-breeding censuses in three sites in each region in both native forest and pasture sub-sites (12 sub-sites; 48 censuses in total), and constructed demographic transition matrices (n = 36). The transition probabilities were calculated using a Bayesian approach and population grow rates were evaluated using asymptotic models and elasticities using transient dynamics. Between regions, higher population growth rate and inertia (defined as the largest or smallest long-term population density with the same initial density distribution) was seen in the shade coffee-dominated landscape. Additionally, population growth rate and damping ratio was higher in forest compared with pasture, with lower convergence time for the forest subsites. These demographic patterns reveal the contrasting levels of population resilience of this orchid in different landscape structures with the more connected shade-coffee dominated landscape permitting some healthier populations with greater population growth and survival in forest than pasture. This study highlights that twig epiphyte colonization of isolated phorophytes in pastures should not be interpreted as a sign of a healthy population but as a temporal transitory period.
... Approximately half the pine-oak forest and tropical dry forest (TDF) fragments measure around 21 ha and < 3 ha, respectively . The condition of the TDF is alarming, since fragments below 10 ha have limited ecological functionality, mainly leading to biodiversity loss (Benitez-Malvido & Arroyo, 2008). However, data are limited and there are no studies on the faunal diversity within these remnants. ...
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La Montaña, one of seven regions making up the state of Guerrero, Mexico, has high social-ecological vulnerability. Its extreme poverty and marginalization levels are among the country’s highest, superimposed in a context of severe ecosystem degradation. We evaluated the social-ecological resilience of two indigenous communities of this region engaged in agroecological projects by integrating the Resilience Assessment and MESMIS methodological proposals, using qualitative and quantitative indicators. The most critical aspects were the low quality of forest fragments, crop losses through hurricanes, lack of access to information, low socioeconomic infrastructure and lack of gender equity. On the other hand, formal and informal social organization is a major strength. Crop losses due to drought or pests are not significant, but a more critical situation is foreseen in the face of climate change. Local efforts drive the main strategies to maintain a minimum level of resilience but will not be sufficient if unaccompanied by structural changes.
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Urbanization has profound effects on wildlife. Although some species benefit or even thrive in urban environments, most species respond differently to the varying degrees of disturbance that can be found across an urbanized landscape. Quantifying the effects of urbanization in wildlife distributions, however, is complicated: species vary in their patterns of presence/absence, abundance, and detectability across spatial and temporal scales, e.g., daily, seasonal, or throughout the annual cycle. Here, we use occupancy models to offer a realistic approach to the study of populations of urban owls. Most studies on owl population dynamics have not considered temporal variation in occupancy between seasons or assess the uneven effects of urbanization along a habitat gradient. We investigated the seasonal habitat associations of mottled owls along an urban gradient in the Neotropical city of Xalapa, Veracruz, Mexico. Using high-resolution satellite images and object-based image classification techniques, were analyzed the relationship between different vegetation and environmental characteristics with the occupancy of mottled owls. We employed different sampling techniques, including playback surveys and silent listening periods, to detect the presence or absence of owls along a gradient from highest to lowest urbanization. Environmental data and different vegetation types were used to analyze the habitat associations of mottled owls during January (late non-breeding season) and May (late breeding season) of 2023. In total, we detected 68 mottled owls during the non-breeding season and 102 during the breeding season, with higher detection rates in areas with >28 % forest Global Ecology and Conservation 55 (2024) e03243 2351-9894/© 2024 Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/). surface. Our results revealed that the best occupancy model included forest and forest division (occupancy), ambient noise and moonlight (detection) for the non-breeding season, as well as urban, forest, grass, and forest division (occupancy), and noise (detection) for the breeding season. The percentage of forested and grass areas positively influenced mottled owl occupancy while the percentage of urbanization and forest division influenced it negatively. Moonlight was positively related to mottled owl detection, while ambient noise had a negative effect on detection probabilities of mottled owls during both seasons. Forested areas emerged as pivotal for owl occupancy, indicating their sensitivity to forest changes along the urban gradient. With urban areas increasing, the interplay between forest division, ambient noise, and moonlight unveils critical insights into mottled owl behavior and habitat dynamics, underscoring the necessity for informed conservation strategies amidst urban expansion. Future research should survey different years to provide a more robust assessment of the dynamic occupancy of mottled owls in Neotropical urban gradients.
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This study describes the characteristics of forest fragments occupied by a Critically Endangered endemic Peruvian primate, the San Martín titi monkey, Plecturocebus oenanthe (Pitheciidae; Platyrrhini). We selected 45 fragments; 20 had already been surveyed in 2015 by the Proyecto Mono Tocón (six of these had been further split, resulting in 27 fragments); an additional 18 fragments were randomly selected from satellite images. We surveyed these fragments for the presence of P. oenanthe and determined characteristics of the fragments (size, shape, tree density, canopy height) and of the landscape (distance to nearest fragment and road). We also examined changes in the number of fragments and in forest cover between 2015 and 2019. We encountered P. oenanthe in all surveyed fragments except for the smallest one (0.2 ha). Our findings suggest that P. oenanthe can persist in fragments with a wide range of characteristics, particularly with regard to size and tree density. Unless fragmentation continues and overall forest cover in the area diminishes further, the species may be able to persist even in a fragmented landscape, provided that the matrix allows for movements between fragments. However, persistence might not be long-term if groups are not reproductive, populations become too small, and reduced gene flow results in inbreeding.
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A close phylogenetic relationship between nonhuman primates (NHPs) and humans is recognized as a causal subjacent factor to the sharing and coevolution of zoonotic pathogens. A relevant barrier to limit the transmission of infectious agents between humans and NHPs is the reduction of exposure and interspecific encounters. Neotropical NHPs occur from southern Mexico through northern Argentina and southern Brazil, presenting high species diversity. All Neotropical NHPs live in forests and are eminently arboreal, although they can descend to the forest floor to cross canopy gaps, to move or disperse between forest fragments, to access food patches located in the anthropogenic matrix, or to drink water from ground sources. A significant proportion of NHP populations have been encroached by human activities, a scenario that has favored contact with humans. The spillover of infectious agents is also possible whenever humans come into close contact with NHPs. This bidirectional pathogen exchange is only successful when the infectious agent adapts to the new host and reaches a high propagation density. In this review, we assess the risk of bidirectional transmission between humans and Neotropical NHPs facilitated by anthropogenic, ecological, and biological variables that may favor their spread in a fragmented Neotropical landscape.
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Aim of study: This study focuses on creating a secondary forest succession (SFS) map between 1972 and 2014 according to the Clementsian theory based on land cover, assessing the spatio-temporal pattern of forest succession change, and determining the factors affecting the forest ecosystem. Area of study: This study was conducted at the Çermik Forest Enterprise (FE) in Diyarbakır city, located in the Southeastern Anatolia Region of Türkiye. Material and methods: Clementsian theory, Remote Sensing (RS), and Geographical Information System (GIS) were used to generate the SFS map. Patch Analyst 4.0 was used to determine changes in spatiotemporal patterns with landscape indices. Main results: The total forested area increased from 32405.1 ha (13% of the study area) in 1972 to 45054.7 ha (18% of the study area) in 2014, with a net increase of 12649.6 ha. It was determined that the progressive succession area was 87736.7 ha, the regressive succession area was 39216.5 ha, and the unchanged succession area was approximately 129989.6 ha. The number of patches increased over a 42-year period. Research highlights: The forest ecosystem was more fragmented, with patches becoming more irregular, complex, and edgy
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Research on fragmented ecosystems has focused mostly on the biogeograpbic consequences of the creation of habitat “islands” of different sizes and has provided little of practical value to managers. However, ecosystem fragmentation causes large changes in the physical environment as well as biogeograpbic changes. Fragmentation generally results in a landscape that consists of remnant areas of native vegetation surrounded by a matrix of agricultural or other developed land. As a result fluxes of radiation, momentum (La, wind), water, and nutrients across the landscape are altered significantly. These in turn can have important influences on biota within remnant areas, especially at or near the edge between the remnant and the surrounding matrix. The isolation of remnant areas by clearing also has important consequences for the biota. These consequences vary with the time since isolation distance from other remnants, and degree of connectivity with other remnants. The influences of physical and biogeographic changes are modified by the size, shape, and position in the landscape of individual remnant, with larger remnants being less adversely affected by the fragmentation process. The Dynamics of remnant areas are predominantly driven by factors arising in the surrounding landscape. Management of, and research on, fragmented ecosystems should be directed at understanding and controlling these external influences as much as at the biota of the remnants themselves. There is a strong need to develop an integrated approach to landscape management that places conservation reserves in the context of the overall landscape
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We reviewed empirical data and hypotheses derived from demographic, optimal foraging, life-history, community, and biogeographic theory for predicting the sensitivity of species to habitat fragmentation. We found 12 traits or trait groups that have been suggested as predictors of species sensitivity: population size; population fluctuation and storage effect; dispersal power; reproductive potential; annual survival; sociality; body size; trophic position; ecological specialisation, microhabitat and matrix use; disturbance and competition sensitive traits; rarity; and biogeographic position. For each trait we discuss the theoretical justification for its sensitivity to fragmentation and empirical evidence for and against the suitability of the trait as a predictor of fragmentation sensitivity. Where relevant, we also discuss experimental design problems for testing the underlying hypotheses. There is good empirical support for 6 of the 12 traits as sensitivity predictors: population size; population fluctuation and storage effects; traits associated with competitive ability and disturbance sensitivity in plants; microhabitat specialisation and matrix use; rarity in the form of low abundance within a habitat; and relative biogeographic position. Few clear patterns emerge for the remaining traits from empirical studies if examined in isolation. Consequently, interactions of species traits and environmental conditions must be considered if we want to be able to predict species sensitivity to fragmentation. We develop a classification of fragmentation sensitivity based on specific trait combinations and discuss the implications of the results for ecological theory.
His main research interest is conservation biology in human-modified tropical rain forests. He has published ca
  • Victor Arroyo-Rodriguez Received The
  • B Sc Ecologia
  • A C Mexico
Victor Arroyo-Rodriguez received the B.Sc. (biology) degree from Universidad Autonoma de Madrid, Spain, in 2002, and both the M.Sc. and Ph.D. degrees from Instituto de Ecologia A.C., Mexico, in 2005 and 2007, respectively. Nowadays he is a postdoctoral fellow at the Centro de Investigaciones en Ecosistemas (Universidad Nacional Autonoma de Mexico). His main research interest is conservation biology in human-modified tropical rain forests. He has published ca. 20 papers in wildlife ecology, forest ecology and biodiversity conservation in fragmented landscapes, and has scientific presentations in numerous national and international congresses and symposia.
This paper introduces a conceptual framework for understanding the effects of landscape modification on species and communities, discusses how species and communities are affected by landscape modification, and provides management recommendations for biodiversity conservation
  • J Fischer
  • D B Lindenmayer
Fischer J., Lindenmayer D.B. (2007). Landscape modification and habitat fragmentation: a synthesis. Global Ecology and Biogeography 16, 265-280. [This paper introduces a conceptual framework for understanding the effects of landscape modification on species and communities, discusses how species and communities are affected by landscape modification, and provides management recommendations for biodiversity conservation].
finding from the Biological Dynamics of Forest Fragments Project, the world's largest and longest-running experimental study of habitat fragmentation
  • W F Laurance
  • T E Lovejoy
  • H L Vasconcelos
  • E M Bruna
  • R K Didham
  • P C Stouffer
  • C Gascon
  • R O Bierregaard
  • S G Laurance
  • E Sampaio
Laurance W.F., Lovejoy T.E., Vasconcelos H.L., Bruna E.M., Didham R.K., Stouffer P.C., Gascon C., Bierregaard R.O., Laurance S.G., Sampaio E. (2002). Ecosystem decay of Amazonian forest fragments: a 22-year investigation. Conservation Biology 16, 605-618. [This paper synthesizes key finding from the Biological Dynamics of Forest Fragments Project, the world's largest and longest-running experimental study of habitat fragmentation, which has been carried out in Brazil].