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Introduction
Tropical rainforests are the most species-rich ecosystems on earth, but these
forests are rapidly disappearing as land is cleared for timber, agriculture and
other uses (Meijaard et al. 2005). The timber trade was a major
incentive for clear cutting in the past. Clear cutting gives a high yield but is not
sustainable. Most logging is now selective, but is still degrading with associated
problems.
Silviculture systems
There are two silviculture systems in the tropics: the polycyclic and monocyclic
systems. Polycyclic systems are based on the repeated removal of selected trees
in a continuing series of felling cycles. The length of the cycle is less than time it
takes the trees to mature (which foresters call the rotation age). It aims to
remove trees before un-harvestable, i.e. before they begin to stagnate and
deteriorate from old age, leaving all appreciating stems to swell the future yield.
Polycyclic systems retain advanced growth of half-grown trees to produce
marketable trees at subsequent intermediate felling cycles within the rotation.
This is done by maintaining young trees at each felling. Under the best
conditions, polycyclic systems can greatly increase the timber yield over a full
rotation. Due to the very species-rich nature of most tropical rainforests, and the
relatively small number of species with timber that is commercial by current
standards, extraction on a polycyclic system tends to result in the formation of
scattered small gaps in the forest canopy. The disadvantage of polycyclic
systems is that the faster growing genotypes are progressively removed,
because at each felling many or all of the stems that have reached a minimum
prescribed girth are removed. The small stems retained may either be young and
potentially fast-growing or old, stunted, and unable to respond to release.
By contrast, monocyclic systems
remove all saleable trees at a single felling. The length of the cycle more or less
equals the rotation age of the trees. Monocyclic systems forego the growth
already accumulated in these adolescents and rely almost entirely on seedlings
to produce the next crop, which will be ready to harvest only at the full rotation
age. It provides maximum yield to satisfy the market. However, damage to the
forest is more drastic than under a polycyclic system, the canopy is more
extensively destroyed, and bigger gaps are formed. Furthermore, new crop trees
all grow up together and this can lead to overcrowding of saplings, competition
between their crowns, and reduced growth rate (Whitmore 1998).
Traffic also has some effect on forest wildlife. The main effect is road-kills.
Some animals appear attracted to the road surface’s warmth, while other
animals learn to use roads for movement. Other species may be attracted to
road kills and learn to feed from them. Increased access also
increases the likelihood of invasion by exotic species associated with humans.
The immigration of exotic species into fragmented forests may threaten some
wildlife populations, and is a cause of concern for the long-term prospects of
tropical biodiversity. Processes such as habitat modification and loss
(i.e. logging) may further promote the establishment of non-native species,
which thrive in disturbed habitats, thus acting synergistically to threaten native
biodiversity. Poorly planned
and implemented logging can result in fragmented and unproductive areas.
Roads and trails also contribute to the total canopy loss during logging (Meijaard
et al. 2005).
Small isolated areas of forest cannot maintain as many species in the long term
as when the same area is part of some larger area.
Some species are un-able to cross open areas and will
even avoid forest margins. This can affect dispersal. Many species require
corridors between forest fragments. Such corridors need to fulfil certain
ecological conditions to make them suitable for dispersal. For instance,
fragmentation effects will be reduced if the distances between intact forest
patches are short (Meijaard et al. 2005).
Bats
Bats are model taxa for logging impact studies as they are a large component of
mammal diversity, have specialist microhabitats, are indicators of forest quality
and disturbance, are easily sampled, are relatively well documented and play key
roles in ecosystem services as seed dispersers and pollinators, maintaining plant
diversity and promoting forest regeneration in degraded land (Clarke et al.
2005).
A study on neotropical bats was carried out by Clarke et al. 2005. The effect of
Trinidad’s periodic block system (PBS) on bat species diversity and community
organization was investigated. PBS is a polycyclic system of selective logging
with a 30-year harvesting rotation. They sampled bats in primary forest and in
PBS-managed forest logged 33, 31, 21, 20 and 10 years previously. They noted
species richness, diversity and community structure.
Species richness was similar in both forests, but diversity decreased
in logged forest. Assemblage structure differed. They found a decrease in
gleaning animalivores suggesting gleaning animalivores to be adversely affected
by logging. They found an increase in frugivores, indicating that some species
may benefit from changes to the forest brought about by logging. This is due to
an increase in the abundance of a few frugivorous species.
The bat community showed evidence of recovery. In
logged forests the number of years post-logging showed a positive correlation
with abundance and richness of animalivores and a negative correlation with the
proportional abundance of the most common species. Gleaning animalivores
increased in abundance with forest regeneration, and the community became
less dominated by a single generalist frugivore.
Birds
Sometimes avian species richness declines after logging. Marsden (1998) found
73 bird species in unlogged forests in Seram, Indonesia but only 53 in forest
logged 1-5 years previously. Thiollay (1992) found in French Guiana that while
some species increased in abundance, overall species richness was lower in
forest logged one year previously than in unlogged forest.
However, other studies have found an increase in bird species richness following
logging. Johns (1997) recorded more species in heavily logged dipterocarp
forests in Sabah, due principally to the addition of forest edge species.
In a Ugandan forest area, bird
species diversity was similar in selectively logged and unlogged forest, although
the response was species-specific (Owiunji 2000). A study
was carried out in a logged compartment of Budongo Forest, north-western
Uganda to investigate the bird community composition in gaps and adjacent
disturbed forest. Similar sampling efforts showed that forest had more
individuals than gap sites but the species number was similar (Ngabo & Dranzoa
2001). In Belize logging
has been found to have little impact on bird diversity. The 66 most common
species occurred with similar frequency in logged and unlogged forest. The total
numbers of bird species were similar between logging gaps and the logged forest
matrix, and between the logged forest matrix and unlogged forests. A
comparison of numbers of species in 26 guilds based on migration strategy, diet,
foraging substrate, and height strata also showed them to be similar regardless
of logging history (Whitman et al. 1998).
Meijaard et al. 2005 reviewed the impact of logging on 145 Bornean vertebrates
(23 birds, 28 mammals, 54 reptiles and 40 amphibians). Impacts were severe for
some species, but highly variable. The general trend was that logging does not
significantly affect diversity. Logging affects abundances and assemblage
structure. This is broadly
supported by their mammal summary, with 15 out of 41 species declining
significantly following logging, 13 showing litter clear change and 13 increasing.
Their summary of bird species responses appears less consistent with the overall
pattern, but is biased by the available literature focusing more on sensitive
species: more than half (12) of the 23 species they considered declined
significantly following logging. Data on amphibians and reptiles was limited.
Amphibians appear little affected by a light logging regime, at least initially.
Research since the early 1980s by A. D. Johns at three locations in lowland
rainforest, at Sungai Tekam in the Jengka area of Malaya, at the Danum valley,
Sabah, and near Tefe, Brazil, has shown that at those forests bird and mammal
populations were surprisingly robust in the face of a single logging operation.
Similar discoveries are now being made in other rainforests. Species relative
abundances can change markedly but total species numbers and the relative
frequency of species in the different feeding guilds show little change (Whitmore
1998).
Fragmentation studies
Birds
Stouffer & Bierregaard 1995 studied the effects of fragmentation on the diversity and
abundance of understory insectivorous birds at a site in Amazonian Brazil. They sampled
birds before isolation and up to 9 years after isolation. They considered fragments of two
sizes, 1 and 10ha, isolated by 70-650m from the nearest continuous forest, that are
surrounded by two distinct types of vegetation. The matrix was either: Vismia
dominated, cut areas around fragments were burned and used by grazing cattle at
irregular intervals, but too infrequently to prevent regeneration of patches of dense
secondary growth dominated by trees in the genera Vismia;.or Cecropia dominated,
cut areas were not burned so areas regenerated rapidly into a forest with a closed
canopy of Cecropia sciadophylla. Species richness and abundance were noted.
Abundance and species richness
declined dramatically after isolation. Ant followers were the
first to be lost. Three species of ant followers disappeared or almost disappeared
within the first 2 years after isolation. After their initial decline, ant followers began to
return to fragments surrounded by Cecropia .Thus development of Cecropia second
growth facilitated use of fragments by ant followers. Ant followers were not influenced by
fragment size, but only used fragments surrounded by Cecropia.
Mixed-species flocks disintegrated within 2-3
years after isolation. Flock species decreased after isolation in 1-ha fragments and in the
2nd year in 10ha fragments. This initial decrease continued in 1ha and 10ha fragments
surrounded by Vismia, but capture rate increased beginning 2-6 years after isolation in
10ha fragments surrounded by Cecropia, where they moved in and out of the fragments
through second growth. For mixed-species flocks, fragment size and secondary growth
affected the suitability of fragments for re-colonisation. Although flocks disintegrated
after isolation, three species that dropped out of flocks persisted in fragments.
Solitary terrestrial and aboreal species declined after
isolation. Two edge-specialist species were unaffected by
fragmentation. There were 21 species of insectivores that were not captured before
isolation but were captured after isolation. Most of these were canopy and edge species
that are found in continuous forest, and most were only netted a few times.
Successful species post-fragmentation were edge-specialists, but their
abundance did not compensate for the loss of other species.
There were two factors affecting colonisation.
One was the time since isolation. So fragment size can be a poor predictor of
species persistence. The other was matrix vegetation. Fragments surrounded by
Vismia remained were depauperate. In contrast, many species returned to fragments of
regenerating forest dominated by Cecropia. Both 1- and 10-ha fragments surrounded by
Cecropia were used by ant followers 5 years after isolation. Mixed-species flocks re-
colonised 10-ha fragments surrounded by Cecropia by 7-9 years after isolation.
Michalski & Peres 2005 the effects of forest fragmentation and forest degradation
on the persistence of primate and carnivore assemblages in a highly fragmented
forest landscape of southern Brazilian Amazonia. They studied site occupancy of
5 primates and 14 carnivores. They compared 15 sites in continuous forest with
129 fragments.
1-17 out of the 19 primate and carnivore species were present in
fragments. In contrast, 12-19 species out of the 19 primate and carnivore
species were present in continuous forest sites.
Primates occurred in more fragments than carnivores. Occupied fragments were
significantly larger than unoccupied fragments for 3/5 primates and 8/14
carnivores. Their results indicated that fragment area was
the strongest predictor of species richness, explaining as much as 55% of the
overall variation in primate and carnivore species richness across all forest
fragments. The habitat quality of remaining forest habitat was also an important
predictor of species richness, particularly to those species that rely heavily on
high basal area, closed-canopy forest.
Their results indicated that the recent human
perturbation regime within forest fragments were also significant determinants of
occupancy for some species in fragments. Distance from the town, forest habitat
quality, their measure of cost surface habitat connectivity, presence of water,
and levels of fragment disturbance (fire/logging) were all significant
determinants of occupancy for some species in fragments.
Anthropogenic disturbance, including surface
wildfires, timber extraction and hunting pressure, had negative effects on the
persistence of some species. The severity of surface wildfires was correlated with
species richness and significantly affected the distribution of howler monkeys,
marmosets, and coatis. However, these fires increased the probability of
occurrence of these species, possibly by increasing the amount of high-quality
foliage for arboreal folivores such as howler monkeys, and providing a more
appropriate understorey structure for small insectivores/frugivores such as
marmosets. Unlike many parts of Amazonia, subsistence and recreational
hunters rarely killed primates and carnivores.
Conclusion
References
Meijaard, E. Et al. (2005) Life after logging: Reconciling wildlife conservation and
production foresty in Indonesian Borneo. Centre for International Forestry
Research (CIFOR). Jakarta, Indonesia.
Clarke, F. M., Rostant, L. V. & Racey, P. A. (2005) Life after logging: post-logging
recovery of a neotropical bat community. Journal of Applied Ecology. 42, 409-420.
Whitman, A. A., Hagan, J. M., and Brokaw (1998) Effects of selection logging on
birds in Northern Belize. Biotropica. 30, 449-457.