THE IMPORTANCE OF FRAGMENTATION TO THE ECOLOGICAL CONCEPT OF SUCCESSION

Jonathan L. Gelbard
Graduate Group in Ecology
University of California at Davis
Fall, 2000
INTRODUCTION

The ecological concept forest succession has provided ecological theory with an important
framework by which ecologists can study the distribution of plant species and communities
throughout time and space (Picket et al. 1994, Peete and Christensen 1987, Cook 1996,
Waring and Running 1998). However, one area where this concept continues to require
expansion to better predict and explain vegetation dynamics is its ability to account for
anthropogenic changes to the biosphere that may affect which species come to occupy future
forests (Waring and Running 1998). Human impacts that may affect successional dynamics
include land use changes that alter landscape characteristics (Lovejoy et al. 1986, Saunders et
al. 1991, Vitousek 1994, Pearson et al. 1998, Joules et al. 1999), introductions of exotic weeds
and pathogens (Halpern 1989), fire suppression (Baily and Liegel 1998), alteration of nutrient
cycles by atmospheric deposition (Schlesinger 1997), and domestic livestock grazing (Belsky
and Blumenthal 1997). While some of these factors have been incorporated into the
ecological concept of succession (e.g., Mladenoff et al. 1996, Schlesinger 1997), others
remain relatively unaccounted for within its framework. Given the extensive nature of human
induced global environmental changes that may influence vegetation dynamics (Vitousek
1994), it is important that such factors be incorporated into our understanding of succession to
provide ecological theory with a means of accounting for their potential effects.
Habitat fragmentation is one such factor that is often discussed as influencing vegetation
patterns (e.g., Franklin and Foreman 1987, Saunders et al. 1991, Mladenoff et al. 1993, Joules
et al. 1999), but has only recently begun to be incorporated into the framework of the
ecological concept of succession (Mladenoff et al. 1996, Iverson et al. 1999, Laurance et al.
1999, Liu and Ashton 1999 for examples). The importance of fragmentation to forest
successional dynamics is illustrated by the vast extent of forest landscapes that are becoming
increasingly fragmented. For example, Spies et al (1994) found that the size of contiguous
uncut patches in a forested area in Central Oregon decreased from 160 to 62 ha between 1927
and 1988 and that edge density increased from 1.9 to 2.5 km/km-2. Cohen et al (1995)
estimated that 15.3% of the 10.4 million ha of forested land of the Pacific Northwest, USA
was harvested between 1972 and 1991, an average rate double that in the Amazon (Skoale
and Tucker 1993). Clearly, fragmentation has had dramatic impacts on forest landscapes, and
thus may be an important influence on forest succession.
Here, I review research concerning the importance of forest fragmentation to the
ecological concept of succession and show that fragmentation of forest landscapes may
profoundly alter physical, biotic, disturbance, and spatial factors that influence the species
composition of future forests. An extensive review of recent scientific literature revealed four
major types of findings relating fragmentation to successional dynamics. These included
alterations to: (1) patterns and sources of seed dispersal, (2) abiotic habitat conditions that
regulate regeneration and species composition, and (3) disturbance regimes that may
influence succession by (4) altering rates of recruitment, mortality, and extinction of forest
plant species. Below, I report on recent findings concerning these impacts of fragmentation
on forest succession.

FRAGMENTATION EFFECTS ON DISPERSAL AND PROPAGULE AVAILABILITY

Fragmentation may affect the future species composition of forest stands by isolating
patches, thus affecting the natural movements and availability of (1) seeds, spores, and pollen
of plants, and (2) the animals upon which many of them either depend for dispersal or hope to
avoid due to predation (Saunders et al. 1991). Such impacts may influence which species are
able to establish (Bruna 1999, Jules and Foster 1999). For example, Jules et al. (1999) found
decreased recruitment of western trillium (Trillium ovatum) near edges to be highly correlated
with decreased seed production due to changes in pollination and increased seed predation by
rodents. Pollinators, including common beetles and bees appeared to be either less abundant
or less effective near edges (Jules and Rathcke 1999), and increased seed predation probably
resulted from significantly higher densities of deer mice in clearcuts and along edges (Mills
1996). In North Carolina, Pearson et al. (1998) found the coverage and density of 8 of 17
mesic herb species to be significantly greater in large forest patches (> 200 ha) than in small
patches (< 10 ha). Several ant-dispersed species were more likely to be absent from small
patches than from large patches. Furthermore, small patches harbored smaller populations of
herbs that were demographically isolated from other populations, and therefore may have
experienced higher probabilities of local extinction. The authors hypothesized mechanisms
that affected populations to include (a) disruption of population dynamics due to habitat
fragmentation, (b) habitat degradation, and (c) anthropogenic disturbance via land use.
In another example, Liu and Ashton (1989) incorporated effects of fragmentation into a
successional model to predict vegetation composition in fragmented tropical forests. The
authors created a spatially explicit landscape simulation model that considered interactions
between the focal forest and neighboring areas and found that tree species richness of a
tropical forest was related to the interactions between harvest impacts and immigration from
adjacent areas. Adjacent species rich forests increased species richness in a focal forest, but
neighboring single-species plantations had opposite effects. A similar effect was observed in
the field by Matlack (1994). Conducting work in eastern Pennsylvania, the researcher found
that in successional forest stands in that were contiguous with species-rich old-regrowth
forest, understory species richness declined with distance into the successional stands,
implying contagion across the old-regrowth ecotone. In stands isolated from old regrowth,
however, the understory was species poor due to a lack of species dispersed by ants and
spores, and those with no obvious dispersal vector. Understory species richness was greater
in older isolated stands and in stands closer to potential sources of propagules than in young
and isolated stands.
The differences in species composition between isolated and non-isolated stands provides
evidence that fragmentation can affect the trajectory of forest succession where it reduces
accessibility to colonists. Such isolation effects may especially reduce the availability of
propagules, pollen, and spores of plant species that are rare at the landscape scale (Saunders et
al. 1991, Bruna 1999, Jules and Foster 1999). They may also lead to genetic isolation and
inbreeding and drift (Perry 1994), which may be responsible for reductions in seed production
that have been observed for some plant species in habitat fragments (Bruna 1999). Of course,
these effects may depend on the size of a patch, its distance from other large tracks, and the
time since isolation (Saunders et al. 1991). While isolation may in some cases benefit forests
by reducing the spread of contagious diseases, insect infestations, and introduced predators,
and by increasing genetic differentiation among populations (Perry 1994), its overall effect on
successional dynamics has been shown to be detrimental (Saunders et al. 1991, Jules and
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Foster 1999).

FRAGMENTATION EFFECTS ON ABIOTIC HABITAT CONDITIONS

Fragmentation can also affect trajectories of succession because loss of tree canopies and
creation of edge habitat increases daytime air and soil temperatures and radiation, increases
wind and occurrence of frost, and increases soil moisture (Lovejoy et al. 1986, Saunders et al.
1991, Chen et al. 1992). Lovejoy et al. (1986) reported that in the Amazon, differences in
relative humidity between reserve edges and 100 m inside ranged from 5 percent in the early
morning to 20 percent at midday. Differences in air temperature in shaded areas between the
edge and 100 m into a 100 ha reserve were as great as 4.5 °C. The researchers believed that
these changes may have explained elevated tree mortality in small reserves. Janzen (1986)
pointed out that such hotter, drier conditions created by fragmentation may not only increase
mortality, but also prevent regeneration of species whose successful establishment requires
humid conditions or persistent soil moisture. Similar findings have been recorded in
temperate forests. For example, Jules (1998) associated almost complete mortality of western
trillium in clearcuts with radiation-induced moisture stress. Clearly alterations to stand
microclimate by fragmentation may have important direct effects on forest plants, and thus on
succession.
Alteration to abiotic habitat conditions may also indirectly affect the direction of
succession by increasing rates of decomposition, mineralization, and nitrification (Saunders et
al. 1991, Waring and Running 1998). Pulses of nutrients that result from increased rates of
biogeochemical cycling may be lost from the system where plant uptake is reduced, such as
where logging is accompanied by attempts to control the growth of herbaceous and shrub
species through the application of herbicides, grazing, or fire (Waring and Running 1998).
Nitrates and other nutrients may also be lost from the system if the capacity of microbial
organisms and other components of the soil to immobilize nutrients is exceeded (Vitousek
1982). In addition, nutrients may be lost where removal of the tree canopy and/or compaction
results in accelerated erosion (Waring and Running 1998). These changes to nutrient
availability induced by fragmentation’s effects on abiotic habitat conditions may be important
to succession by favoring forest species tolerant of low nutrient conditions (Vitousek 1982,
Tilman 1985, Schlesinger 1997).

FRAGMENTATION EFFECTS ON DISTURBANCE REGIMES THAT REGULATE SUCCESSION

While disturbances created by landslides, fires, wind, disease, and infestation are
important to maintaining a healthy balance of successional stages in forest landscapes
(Hansen et al. 1991, Cook 1996), fragmented landscapes may experience artificially high
levels of disturbance (Franklin and Forman 1987, Saunders et al. 1991). Such changes to
disturbance regimes may influence trajectories of succession. For example, Franklin and
Foreman (1987) found more fragmented patches to be more susceptible to windthrow and
fire, disturbances that may result in mortality of late successional species, and create habitat
for early successional species. At local scales, increases in susceptibility to disturbance may
affect succession in isolated patches by damaging or even extinguishing populations of fire or
wind-sensitive species, preventing them from becoming important components of future
forests (Saunders et al. 1991, Perry 1994). Disturbances that fragment forests may also
enhance introductions and establishment of exotic weedy species that can inhibit native plant
regeneration by competition or by altering fire regimes (Saunders et al. 1991). Pearson et al.
(1999) speculated that more favorable herb populations in large relative to small plots may
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have been in part related to disturbance regimes in small patches being different than in large
patches. Specifically, they hypothesized that small patches with a greater concentration of
edge habitat had experienced greater levels of both historic and recent disturbance by cattle
grazing and humans. They noted lower coverage of soil litter in small patches to have
indicated that smaller plots were more disturbed. Effects of fragmentation on disturbance
regimes may affect the outcome of succession from local to landscape scales (Franklin and
Foreman 1987, Saunders et al. 1991).

POPULATION LEVEL EFFECTS OF FRAGMENTATION ON FOREST PLANT SPECIES

Possible mechanisms for the above effects of fragmentation on forest vegetation dynamics
have been suggested by studies linking fragmentation with reduced recruitment success,
increased rates of mortality (Jules et al. 1999) and extinction (Tilman et al. 1994, Laurance et
al. 1999), and altered age class distributions (Chen et al. 1992, Bailey and Liegel 1998). For
example, Bruna (1999) found that seeds planted in continuous forest were between three and
seven times more likely to germinate than those in forest fragments. He attributed this finding
to a variety of edge effects on seeds, including hotter, drier conditions and increased light
penetration, all of which can affect the cues required for germination. He also suggested that
plants reproducing in forest fragments may be inbred, and that if inbred seeds are less likely to
germinate, these genetic effects could further reduce the recruitment of seedlings info forest
fragment populations, helping to explain why plant populations in habitat fragments often fail
to persist in the long term.
Similar effects of fragmentation on plant populations have been observed in temperate
forests. Chen et al. (1992) observed that near edges, old growth forests experienced
reductions in stem density and basal area, increases in growth rates of Douglas fir
(Pseudotsuga menziesii) and western hemlock (Tsuga heterophylla), elevated rates of tree
mortality, and increases in seedling and sapling numbers of douglas fir and western hemlock,
but decreases in those of Pacific silver fir (Abies amabalis). The depth of edge influence on
these vegetation dynamics ranged up to 137 m into old growth forest patches, and varied by
species. In the Klamath region of northern California, Jules et al. (1999) found that 7 woody
and 31 herbaceous species sampled in clearcuts, edges, and interiors occurred more frequently
in the interior-most forest plots. Of those 31 species, 12 occurred in forest, but not clearcut or
edge plots. The same effect was observed on both north and south-facing slopes for 69
percent of species that showed a plot location effect. The depth of edge effect was most
dramatic within 30 m of edges, but a number of species, including Taxus brevifolia, Tiarella
trifoliata, and Anemone deltoidea preferred habitats over 60 m from edges. One species,
western trillium was found to experience almost no recruitment within 15 m of edges and 97.6
percent mortality in clear-cuts and plantations. The authors concluded that species with
affinities for forest interior conditions could be experiencing population declines as a result of
fragmentation. They proposed that mechanisms that may explain this response to edge may
include (1) increased water stress for plants adapted to mesic sites, (2) competition with
aggressive disturbance-adapted species that were frequently sampled in clear-cuts, (3)
alterations to interspecific interactions, such as higher herbivory levels, and decreased
effectiveness of pollination, reduced presence of mycorrhizal associates, and increased seed
predation, and (4) plant species with poor tolerance to physical disturbances associated with
logging will decline near edges.
Fragmentation may affect the direction of forest succession by creating habitat for
herbivores that may dramatically reduce recruitment success (Alvorson 1984). The tendency
of ungulates such as deer to favor the edge habitats created by fragmentation may result in
4
increases mortality and reductions in seedling abundance (Perry 1994). Such impacts have
been shown to have altered species compositions in Olympic National Park (Perry 1994) and
in many forested areas of the eastern US (Alvorson 1994). Such effects have been concluded
to contribute to reductions in regeneration of threatened species such as the Pacific yew
(Taxus brevifolia) (Bailey and Liegel 1998). Thus, fragmentation may not only directly affect
forest succession by impacting recruitment and mortality of individual species, but also by
indirect effects on stand demography related to interspecific interactions.

CONCLUSIONS AND IMPLICATIONS

Current ecological knowledge concerning effects of fragmentation on successional

dynamics clearly demonstrated that in order for the concept of succession to continue to
provide ecological theory with a framework by which to explain changes in vegetation
patterns through space and time, it must embrace human environmental impacts such as
habitat fragmentation. Fragmentation has been shown to affect succession by (1) isolation
effects that influence dispersal and propagule availability, (2) direct and indirect impacts to
plants associated with altered abiotic habitat conditions, and (3) alteration of disturbance
regimes. These impacts have been associated with both direct and indirect impacts to
recruitment, mortality, and age class distributions of forest plant species. Clearly, many of
these effects appear to be intercorrelated (e.g., effects of altered microclimate on increased
susceptibility of fragmented areas to fire; Saunders et al. 1991), a state of knowledge that is
not surprising given that many of the ideas presented above have been quantified only in
models and descriptive studies (and should therefore be taken in that light).
Many of the effects of fragmentation on successional changes in forest species
composition are likely scale and system specific. For example, while fragmentation may
constitute a widespread influence on succession at the landscape scale by affecting abiotic and
disturbance conditions that in turn effect plant survival and reproduction, at local scales the
severity of these effects may vary with the size of a fragment. Furthermore, moist tropical
and temperate deciduous forests are characterized by rather mild, gap-type disturbance
regimes that seldom (if ever) disrupt the essential interior forest habitat, and species of these
systems may therefore be more vulnerable to fragmentation (Perry 1994). On the other hand,
the mix of species within forest types such as boreal and temperate coniferous forests, which
are adapted to fragmentation caused by wildfire, may be better adapted to fragmentation and
thus more resistant to its effects. Nonetheless, it has become so widespread that it can be an
important influence in all types of forests (Perry 1994).

FUTURE RESEARCH DIRECTIONS

Important areas for future studies exploring effects of fragmentation on succession include
experimental verification the patterns described above elucidation of mechanisms responsible
for these patterns. An important first step is sorting through confounding correlations that are
readily apparent among the different effects of fragmentation on succession reported above.
This may be accomplished by first using experiments to ask how the recruitment, growth, and
mortality of different forest plant species varies across different combinations of propagule
availability/isolation, microclimate, and disturbance type, frequency, and severity. Once
effects and patterns are elucidated, multivariate analyses accompanied by path analyses may
provide useful means of illustrating both the relative importance of different fragmentation
effects and the potential pathways by which these effects may influence the future species
composition of forests, from stands to landscapes.
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