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Dynamics and restoration of Australian tropical and subtropical rainforests

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In rain forests with an intact fauna, moderate-scale disturbance of the canopy (large tree-fall gaps to blowdowns of several hectares) typically gives rise to the following successional sequence: pioneers recruit from the seed bank, grow rapidly to form a closed canopy, but do not recruit under their own shade. As the pioneers senesce, they are replaced by slower-growing, shade-tolerant, long-lived trees either present as seedlings at the time of disturbance or subsequently dispersed to the regenerating forest from surrounding areas of rain forest. This successional model has been widely used to describe the ecology of rain forests and often forms the basis of strategies developed for their exploitation Successional models have strongly influenced the development of techniques for restoring rain forests to cleared land. These techniques have been successful insofar as restored sites can develop a rain forest-like structure and provide habitat for some rain forest-dependent biota within one to two decades of establishment. However, few restoration projects have been established long enough to progress to the stage where planted trees have been replaced by species dispersed to the restored site let alone converge on target conditions. As successional models address only some of the factors influencing the dynamics of restored sites in disturbed landscapes, it may be useful to consider alternate models of forest dynamics to inform the long-term management of restored sites. A state-and-transition model of the dynamics of restored sites suggests that these sites will often require long-term intervention, including the control of exotic plants and the addition of native species, to converge on the condition of intact forest. Because the costs of intervention can be high, particularly if a problem such as weed invasion is not controlled at an early stage, the routine monitoring of restoration projects may need to become a necessary part of their management. At present, most Australian rain-forest restoration projects are not monitored in any formal way. Once the long-term outcomes of restoration projects are better known (e.g., through more systematic monitoring), there may need to be a reassessment of what constitutes appropriate “target conditions” for restored sites. It may become apparent, for example, that most restoration projects are so small and isolated and so heavily influenced by the disturbed matrix that the structure, composition, and dynamics of restored sites will always tend to diverge from the condition of intact rain forest. If so, it may be necessary to accept stable assemblages of native and exotic plants (“emerging” or “new forests”) as the target for most restoration projects. Alternatively, investment in projects aimed at reestablishing assemblages typical of intact rain forests may need to be restricted to high-value applications (e.g., habitat for threatened species or plantings strategically positioned to act as future seed sources) rather than being used as the default approach across the landscape, as currently tends to be the case. All models of rain-forest dynamics recognize the importance of landscape context in affecting the trajectory of restored sites. To date, much restoration effort in Australian has been expended on the creation of corridors to facilitate dispersal between remnants. However, corridors do not address other landscape-scale factors affecting the persistence of biota in remnants, such as the influence of the surrounding matrix. A better understanding of the relative importance of the extent and configuration of rain-forest cover at the landscape scale and integration of this knowledge into models of rain-forest dynamics is required to inform future restoration projects aimed at conserving rain-forest biota. Yes Yes
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206
At a stand level and as a generalization, the dynamics of moist tropical and subtropical
rainforests following disturbance can be explained by a few interrelated factors (Swaine and
Whitmore 1988; Hopkins 1990; Finnegan 1996; Richards 1996; Terborgh et al. 2002; Wright
2002). First, mature rainforest has a closed canopy and a shady understory. Second, the long-
lived trees that dominate mature rainforest are relatively shade tolerant and slow growing,
and their seeds typically germinate on dispersal. Third, rainforests also support a suite of
shade-intolerant, fast-growing plants (“pioneers”) whose seeds persist in the soil and whose
germination is stimulated by the increase in heat and light that accompanies disturbance to
the canopy. Fourth, animals are key agents in the pollination, dispersal, and consumption of
rainforest plants.
Consequently, in rainforests with an intact fauna, moderate-scale disturbance of the can-
opy (large tree-fall gaps to blowdowns of several hectares) typically gives rise to the following
successional sequence: Pioneers recruit from the seed bank, grow rapidly to form a closed
canopy, but do not recruit under their own shade. As the pioneers senesce, they are replaced
by slower-growing, shade-tolerant, long-lived trees either present as seedlings at the time of
disturbance or subsequently dispersed to the regenerating forest from surrounding areas of
rainforest (fig. 14.1). We note that this successional model considerably simplifies variation in
plant traits and ecological strategies and does not address issues concerning the maintenance
of diversity in rainforests (Tilman 1994; Westoby et al. 2002; Wright 2002). Nevertheless, this
model has been widely used to describe the ecology of rainforests and often forms the basis
of strategies developed for their exploitation (Hartshorn 1989; Hopkins 1990; Kooyman 1996;
Richards 1996; Lugo 1997; Duncan and Chapman 2003; Ganade 2007; Holl 2007).
In this chapter, we consider the utility of successional and alternate models (e.g., King
and Hobbs 2006; Cramer 2007) for describing forest dynamics and informing the conduct of
restoration projects in rainforest landscapes subject to broadscale anthropogenic disturbance.
Such landscapes are characterized by extensive areas of cleared land dominated by grasses,
crops and other exotic vegetation, patches of remnant forest, areas of secondary forest on mar-
ginal agricultural land, and various types of tree plantations (Richards 1996). Our examples
are drawn largely from Australian rainforests, the focus of our research experience. We begin
with a brief description of intact Australian rainforests before discussing disturbed systems.
Chapter 14
Dynamics and Restoration of Australian Tropical
and Subtropical Rainforests
John Kanowski, Robert M. Kooyman, and Carla P. Catterall
206
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Environmental Determinants and Dynamics of Australian Rainforests
Until the mid-Tertiary, rainforests covered much of Australia. Currently, they cover around
3 million ha, or 0.4% of the continent, mostly in higher-rainfall areas along the east coast
(Cofinas and Creighton 2001). Despite their limited extent, rainforests remain important
repositories of biodiversity, supporting, for example, 60% of Australian vascular plant fami-
lies and 17% of Australian bird species, including many endemic taxa (Lynch and Neldner
2000). From a global perspective, Australian rainforests have some unique characteristics,
including the presence of a number of “primitive” angiosperm families (Adam 1992) and a
distinctive fauna, such as marsupial frugivores and folivores rather than primates (Primack
and Corlett 2005).
At a landscape scale, the composition and structure of Australian rainforests vary primarily
with temperature, soil nutrient availability, and rainfall (Webb 1968; Floyd 1990; Kooyman
and Rossetto 2006). Distinct forest types, defined in terms of structural attributes or their char-
acteristic floristic composition, are associated with particular combinations of these environ-
mental factors (Webb et al. 1976; Tracey 1982). For example, “complex notophyll vine forests”
occur on relatively fertile soils subject to moderate to high rainfall and a “mesotherm” climate
14. Dynamics and Restoration of Australian Tropical and Subtropical Rainforests 207
FIGURE 14.1. Model of the dynamics of intact rainforests on fertile soils in tropical and subtropical
Australia following disturbance, with sequential stages of recovery dominated by (1) shade-intolerant,
fast-growing, short-lived “pioneer” species; (2) longer-lived pioneer trees; and (3) shade-tolerant, slower-
growing, long-lived “mature-phase” trees. Stages 1 and 2 are transient, and stage 3 may be persistent.
Hobbs-Suding Ch_14-Ch_17.indd 207 11/3/08 8:59:37 AM
208 dynamics and restoration of different ecosystem types
(i.e., lowlands in the subtropics, uplands in the tropics), whereas “simple notophyll vine for-
ests” occur on poorer soils in otherwise similar environments (Webb 1968; Nix 1991).
In the field, transitions between forest types can occur over short distances in conjunction
with abrupt changes in environmental conditions. For example, rainforest types can change
from “complex” to “simple” from one side of a watercourse to the other, with a change in
underlying soil parent materials (Tracey 1982). Similarly, there is often a sharp transition
between rainforest and fire-tolerant eucalypt forest, mediated by topographical features that
act as a barrier to the spread of fire (Ash 1988).
At a stand level, the dynamics of Australian rainforests vary in response to the intensity
and frequency of disturbance events (Hopkins 1990) and also with forest type. In “complex”
rainforests on fertile soils, these dynamics tend to follow the successional model presented in
figure 14.1, at least for disturbances that open the forest canopy sufficiently to stimulate the
regeneration of pioneer species from the soil seed bank (Floyd 1990; Hopkins 1990; Kooy-
man 1991; Kariuki and Kooyman 2005). In contrast, the species that regenerate after distur-
bance in “simple” rainforests on poorer soils include many long-lived trees that also occur in
mature forest (Floyd 1990; Kooyman 1999).
Dynamics of Australian Rainforests in Anthropogenically
Disturbed Landscapes
Australian Aborigines have utilized rainforests for many thousands of years (Horsfall and Hall
1990). Through their use of fire, Aborigines modified the distribution of rainforest, and their
hunting- and-gathering practices presumably influenced the distribution and abundance of
harvested species (Newell 1999; Bowman 2000; Hill et al. 2000). Nevertheless, the tradi-
tional livelihood of the Aborigines relied on the persistence of rainforest and its resources.
European settlers converted extensive areas of rainforest to agricultural land, mainly sugar-
cane plantations in the lowlands and pasture and horticultural crops in the foothills and
uplands (Erskine et al. 2007). Clearing was restricted mostly to forests on fertile soils, and many
cleared areas remain under agricultural production. However, in the subtropics, large areas
of marginal agricultural land have been abandoned in the past few decades and now support
secondary forest (Neilan et al. 2006). There are also around 50,000 ha of monoculture timber
plantations and 5,000 ha of mixed-species plantings established on former rainforest land in
the Australian tropics and subtropics (Catterall and Harrison 2006; Erskine et al. 2007).
In the following paragraphs, we discuss the dynamics of these remnant forests, second-
ary forests, and replanted areas (fig. 14.2). Given the long time scales involved in rainforest
dynamics and restoration and the paucity of relevant longitudinal studies, we are not in a
position to evaluate which type of model “best” describes these systems. Instead, we consider
how successional and alternative models draw attention to different aspects of forest dynam-
ics and how these approaches can inform restoration practices.
Rainforest Remnants
Small patches of rainforest have persisted in extensively cleared landscapes in Australia for 50
to 100 years. For example, there are 30 remnants >1 ha of the former “Big Scrub,” a 75,000-ha
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14. Dynamics and Restoration of Australian Tropical and Subtropical Rainforests 209
lowland subtropical rainforest that was cleared between 1860 and 1900 (Adam 1992). Many
remnant rainforests have been subjected to disturbance from logging, cattle grazing, fire, and
invasion by exotic plants, while all are exposed to an altered physical environment (increased
light, wind and temperature variation), particularly on their edges (Lamb et al. 1997; Laur-
ance 1997). Furthermore, it seems likely that the floristic composition of remnants will be
impacted by changes in the distribution and abundance of fauna involved in plant regenera-
tion, particularly seed dispersers, seed predators, and herbivores (Wright and Duber 2001;
Cordeiro and Howe 2003). For instance, rainforest–dependent frugivorous birds responsible
for dispersing the large, fleshy fruits of mature-phase rainforest trees have declined in some
subtropical remnants (Moran et al. 2004).
Successional models figure prominently in the approaches developed by practitioners to
restore Australian rainforests (Goosem and Tucker 1995; Kooyman 1996; Big Scrub Rainforest
(a) (b)
(c) (d)
FIGURE 14.2. (a) Remnant rainforest, tropical Australia. (b) Mixed-species rainforest plantation,
subtropical Australia. (c) Secondary forest in subtropical Australia dominated by the exotic tree,
camphor laurel (Cinnamomum camphora). (d) Regenerating rainforest, eight years after the camphor
laurel overstory was poisoned.
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210 dynamics and restoration of different ecosystem types
Landcare Group 2005). For example, when applied to remnant rainforests, the model presented
in figure 14.1 predicts that the long-term dynamics of remnants may be adversely impacted by
dispersal limitation. Consequently, the restoration of “corridors” to facilitate dispersal between
remnant forests has underpinned much of the revegetation effort conducted in Australian
rainforest landscapes over the past few decades (Lamb et al. 1997; Tucker 2000).
However, successional models do not readily predict some aspects of the degradation
of remnant forests. For example, in many subtropical remnants, disturbance of the canopy
has resulted in the vigorous growth of exotic vines that smother trees and further disrupt
the canopy. These conditions also favor the growth of exotic herbaceous ground covers that
can suppress the regeneration of rainforest plants (Floyd 1990; Kooyman 1996; Harden et
al. 2004). In such cases, it may be useful to consider alternative models to describe the
processes involved in degradation and identify restoration strategies. For example, a “biotic
and abiotic threshold” model (Whisenant 2002; King and Hobbs 2006) could be applied to
the degradation of remnants driven by the positive feedback among canopy disturbance, the
growth of exotic vines, and the spread of herbaceous ground covers. In rainforest, the canopy
is considered a key regulator of abiotic conditions, with a closed canopy suppressing light-
demanding vines and ground covers (Floyd 1990; Kooyman 1996). Restoration practices that
have prioritized the reestablishment of a closed canopy and hence the abiotic conditions
characteristic of intact rainforest have proven successful in restoring remnants degraded by
exotic vines (Harden et al. 2004; Big Scrub Rainforest Landcare Group 2005). Earlier resto-
ration practices that focused primarily on the control of the herbaceous ground covers were
unable to halt the decline of these remnants.
Secondary Forests
Intensive agricultural pursuits in rainforest landscapes have frequently been abandoned
because of declining productivity or changes in economic conditions (Richards 1996; Ers-
kine et al. 2007; Ganade 2007). The redevelopment of rainforest following the abandonment
of agriculture may be constrained by the destruction of the rainforest seed and seedling
banks, the limited dispersal of seeds from remnant forests, predation on dispersed seeds, a
reduction in soil fertility, competition from exotic plants, and repeated disturbance from fire
and grazing (Hopkins and Graham 1984; Uhl et al. 1988; Hau 1997; Lamb et al. 1997; Holl
1999; Toh et al. 1999; Holl et al. 2000; Erskine et al. 2007). Under these circumstances,
abandoned agricultural land may remain covered by exotic grasses and shrubs.
Nevertheless, if abandoned agricultural land is protected from fire and other major distur-
bances, secondary forest will often eventually replace the grassland (Richards 1996; Corlett
2002). The transition to secondary forest can occur rapidly if land is abandoned shortly after
clearing and subsequently protected from disturbance, in which case rainforest plants may
regenerate from residual rootstocks and the seed bank, supplemented by dispersal if forests
are nearby (Uhl et al. 1988; Richards 1996; Erskine et al. 2007). In other cases, the transition
may take decades (Toh et al. 1999; Holl 2007). In disturbed landscapes, the initial coloniz-
ers of abandoned land are often exotic plants; consequently, secondary forests dominated by
exotic plants now cover large areas of former rainforest land (Ewel and Putz 2004; Zimmer-
man et al. 2007). In the Australian subtropics, for example, secondary forests dominated by
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14. Dynamics and Restoration of Australian Tropical and Subtropical Rainforests 211
the exotic tree camphor laurel (Cinnamomum camphora) have recolonized around 25% of
cleared land in the former “Big Scrub” (Neilan et al. 2006).
The concepts of facilitation and inhibition, inherent in classical successional models, are
useful in understanding some of the dynamics of secondary forests (Hopkins 1990; Finnegan
1996; Richards 1996; Ganade 2007; Holl 2007). The presence of trees and shrubs in former
agricultural land can facilitate the recruitment of rainforest plants because they attract seed-
dispersing animals and suppress grasses that might compete with recruits (Lamb et al. 1997;
Toh et al. 1999; Aide et al. 2000; Ganade 2007). Conversely, trees and shrubs also tend to
inhibit the growth of recruits through mechanisms such as allelopathy, root competition, and
shading (Bazzaz 1979; Tilman 1994; Ganade 2007; Loh and Daehler 2007).
Exotic tree colonizers in abandoned farmland are a particularly interesting case of the ten-
sion that exists between the contrasting roles of facilitation and inhibition, with implications
for restoration strategies. When exotics are relatively short lived and shade intolerant, as is the
case for Spathodea campanulata, the dominant colonizer of abandoned land in Puerto Rico,
then passive restoration (doing nothing in the expectation that the recruitment of native spe-
cies will be facilitated by the colonizing trees) may be an effective strategy (Zimmerman et
al. 2007). However, when the exotics are long lived, shade tolerant, or capable of altering key
environmental conditions such as soil nutrient status (as is the case for the leguminous tree
Myrica faya in Hawaii; Loh and Daehler 2007), passive restoration is more risky.
In Australia, debate over the management of regrowth dominated by camphor laurel reflects
different perceptions of its role in promoting (or retarding) the development of rainforest on
abandoned land. Some ecologists have argued for the widespread control of camphor laurel
(Floyd 1990), while others have argued for its retention, at least as an interim measure (Neilan
et al. 2006). Camphor laurel clearly facilitates the recruitment of rainforest plants to grass-
land: It establishes among and shades out pasture grasses; when mature, it bears a heavy crop
of fleshy fruits that attracts frugivorous birds and bats that disperse the seeds of rainforest plants
and exotics to camphor stands (Neilan et al. 2006). However, camphor laurel can potentially
live for centuries, during which time it may suppress recruits (Scanlon and the Camphor Lau-
rel Task Force 2000). Given the uncertainty about the long-term trajectory of camphor stands,
restoration practitioners have developed selective or patch-scale culling treatments to acceler-
ate the growth of rainforest plants in camphor stands (Woodford 2000; Lymburner et al. 2006;
fig. 14.2d).
Identifying the factors that drive or inhibit transitions among grassland, secondary forest,
and mature rainforest is an important element in the development of restoration strategies
for disturbed rainforest landscapes (Holl 2002). The model presented in figure 14.1 identi-
fies dispersal limitation as a key factor potentially truncating the development of secondary
forest. The large seeds of mature-phase rainforest trees that are often dispersed by rainfor-
est–dependent fauna are rarely dispersed to secondary forests isolated from remnant forest
(Corlett 2002; Cordeiro and Howe 2003; Moran et al. 2004; Erskine et al. 2007; Holl 2007).
However, secondary forest may not progress toward the condition of intact rainforest for other
reasons, including dominance of the seed rain by exotic plants (Holl 2007), elevated rates of
seed predation or herbivory (Hau 1997; Holl et al. 2000), changes in underlying soil and/or
hydrological conditions from past land uses (Zimmerman et al. 2007), and repeated distur-
bance (Richards 1996).
Hobbs-Suding Ch_14-Ch_17.indd 211 11/3/08 8:59:40 AM
212 dynamics and restoration of different ecosystem types
Alternative conceptual models of the dynamics of secondary forests, such as state-and-
transition models (King and Hobbs 2006), can help draw attention to the range of factors
driving transitions between different types of vegetation cover in disturbed rainforest land-
scapes (fig. 14.3). For example, the dispersal of plants from the disturbed matrix can strongly
influence the composition of secondary forests, which are usually dominated by small-seeded
pioneers and exotic plants (Duncan and Chapman 2002; Neilan et al. 2006).
Replanted Sites
Extensive areas of rainforest have been converted to plantations of timber trees (Lugo 1997).
In Australia, plantations in rainforest landscapes are predominantly monocultures of the
native conifer Araucaria cunninghamii, with smaller areas of mixed-species timber planta-
tions and restoration plantings (Kanowski et al. 2003; Catterall et al. 2004). Restoration plant-
ings typically use a diverse range of species planted at high densities to rapidly attain canopy
closure and suppress weeds (Kooyman 1991, 1996; Lamb et al. 1997).
Successional concepts have played an important role in the development of replanting
strategies. For example, both timber plantations (when used as part of a restoration strategy)
and restoration plantings are intended to facilitate the recruitment of rainforest plants to
formerly cleared land (Lamb et al. 1997). Restoration plantings are explicitly based on suc-
cessional models in terms of their composition (usually a mix of pioneers and mature-phase
species) and variation in composition with proximity to remnant forests. Planting designs
developed for sites adjacent to remnant forests are comprised mainly of pioneers on the
assumption that mature-phase species will readily disperse and recruit to such sites. In con-
trast, designs developed for more isolated sites include a substantial component of mature-
phase species, particularly large-seeded and wind-dispersed trees with limited potential for
long-distance dispersal (Goosem and Tucker 1995; Kooyman 1996; Freebody 2007).
Nevertheless, it may be useful to consider revegetation strategies within a state-and-transi-
tion framework (fig. 14.3) because successional models do not necessarily address the range of
factors that may influence the long-term dynamics of replanted sites. For example, experience
has shown that pioneer-dominated restoration plantings often do not recruit an understory
of mature-phase rainforest plants as intended, even when located adjacent to remnant for-
ests. Instead, the understory of these plantings often becomes dominated by light-demanding
weeds, either present in the seed bank or dispersed from the surrounding matrix, that then
suppress the recruitment of rainforest trees (Kooyman 1996; Freebody 2007). Similarly, con-
sideration of landscape-scale factors influencing the dynamics of reforested sites may lead to
greater skepticism of claims that plantations can facilitate rainforest regeneration in broadscale
restoration programs (Parrotta et al. 1997; Lamb 1998; Lamb et al. 2005). Many tree species
grown commercially in plantations (e.g., conifers, eucalypts) do not bear fleshy fruits and
hence, when isolated from remnant forests, are unlikely to attract the specialist frugivores that
disperse the seeds of mature-phase rainforest plants (Tucker et al. 2004; Kanowski et al. 2005).
Consequently, although plantations adjacent to remnant forests may recruit a diverse range of
rainforest plants (Chapman and Chapman 1996; Keenan et al. 1997; Lamb et al. 2005), more
isolated plantations are likely to be dominated by small-seeded pioneers and exotic plants dis-
persed from the surrounding matrix (Corlett 2002; Duncan and Chapman 2002).
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14. Dynamics and Restoration of Australian Tropical and Subtropical Rainforests 213
Conclusions
Techniques for restoring rainforests to cleared land have been informed by successional
models developed from observations of intact rainforests (Floyd 1990; Hopkins 1990; Kooy-
man 1996). These techniques have been successful insofar as restored sites can develop a
rainforest–like structure and provide habitat for some rainforest–dependent biota within
one to two decades of establishment (Kanowski et al. 2003, 2006; Grimbacher et al. 2007;
Kanowski and Catterall 2007; Catterall et al. 2008). However, few restoration projects have
been established long enough to progress to the stage where planted trees have been replaced
by species dispersed to the restored site let alone converge on target conditions. As succes-
sional models address only some of the factors influencing the dynamics of restored sites in
disturbed landscapes, it may be useful to consider alternate models of forest dynamics to
inform the long-term management of restored sites (table 14.1).
A state-and-transition model of the dynamics of restored sites (fig. 14.3) suggests that these
sites will often require long-term intervention, including the control of exotic plants and the
addition of native species, to converge on the condition of intact forest. Because the costs of
FIGURE 14.3. Model of the dynamics of secondary forests and replanted rainforests in anthropogenically
disturbed Australian rainforest landscapes. Rectangular boxes = potentially stable states (persisting for
more than one generation of dominant species); parallelograms = transitional states.
Hobbs-Suding Ch_14-Ch_17.indd 213 11/3/08 8:59:42 AM
214 dynamics and restoration of different ecosystem types
Table 14.1.
Models of ecosystem dynamics applicable to forested lands within Australian tropical
and subtropical rainforest landscapes.
Forest Type Successional Models Alternative Models Application to Restoration
Intact rainforest
(extensive areas of rainforest with
intact faunal assemblages)
The recovery of forests on fertile soils
after disturbance is often described by
successional models (fig. 14.1).
None (at the stand scale). Restoration interventions in intact
rainforest are rare, as recovery
through autogenic processes is
usually assumed. Some heavily
logged sites have been treated (e.g.,
by replanting and vine control) to
hasten succession (Kooyman 1999).
Remnant rainforest
(small patches of rainforest in
otherwise disturbed landscapes)
Successional models identify dis-
persal limitation as a factor
influencing the dynamics of
remnant forests.
The degradation of remnants
following disturbance of the canopy
and invasion by exotic vines could be
described by a “biotic and abiotic”
threshold model, with the canopy
regulating abiotic conditions.
A common landscape-scale target
of restoration is the establishment
of corridors to improve dispersal
between remnants. The primary site-
scale target of remnant restoration
is reestablishment of the canopy, if
disturbed.
Secondary forest
(regrowth forests developed after
clearing, often dominated by
exotic plants)
The concepts of facilitation
and inhibition are useful for
understanding the dynamics of
secondary forests: Established trees
may facilitate plant recruitment by
attracting dispersers and suppressing
grasses but may also inhibit the
growth of recruits.
“State-and-transition” models can
provide a useful framework for
describing the long-term dynamics
of secondary forests, as they focus
on site- and landscape-scale drivers
of the trajectory and stability of
secondary forests and alternate types
of vegetation cover.
Secondary forests can recover
considerable structure and function
by autogenic processes but may
require intervention (e.g., control
of exotics, enrichment planting) to
converge on the condition of intact
rainforest, depending on the longevity
and shade tolerance of colonizing
species and the recruitment of
rainforest plants.
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14. Dynamics and Restoration of Australian Tropical and Subtropical Rainforests 215
Table 14.1.
Models of ecosystem dynamics applicable to forested lands within Australian tropical
and subtropical rainforest landscapes, continued.
Forest Type Successional Models Alternative Models Application to Restoration
Replanted sites
(monoculture and mixed-
species timber plantations, diverse
restoration plantings)
Replanting strategies and techniques
draw heavily on successional
models. “Pioneer” plantings and
timber plantations (when used for
restoration) are explicitly intended to
facilitate the recruitment of mature-
phase rainforest plants.
The long-term dynamics of replanted
sites are likely to be contingent on the
same site- and landscape-scale factors
that influence remnant forests and
secondary forests. These dynamics
may be usefully described by state-
and-transition models.
Techniques for establishing rainforest
trees in Australian tropical and
subtropical landscapes are well
developed. Planting strategies
generally assume that sites will
converge on intact rainforest through
autogenic processes, but long-term
intervention may be necessary to
control weed invasion and to enrich
species composition in cases where
dispersal is limited. To be effective,
intervention needs to be coupled with
a systematic monitoring program.
Hobbs-Suding Ch_14-Ch_17.indd 215 11/3/08 8:59:43 AM
216 dynamics and restoration of different ecosystem types
intervention can be high, particularly if a problem such as weed invasion is not controlled at
an early stage, the routine monitoring of restoration projects may need to become a neces-
sary part of their management (Catterall et al. 2008). At present, most Australian rainforest
restoration projects are not monitored in any formal way (Kanowski et al. 2008; although for
counterexamples, see Kooyman 1996; Tucker and Murphy 1997; Harden et al. 2004).
Once the long-term outcomes of restoration projects are better known (e.g., through more
systematic monitoring), there may need to be a reassessment of what constitutes appropriate
“target conditions” for restored sites. It may become apparent, for example, that most restora-
tion projects are so small and isolated and so heavily influenced by the disturbed matrix that
the structure, composition, and dynamics of restored sites will always tend to diverge from the
condition of intact rainforest. If so, it may be necessary to accept stable assemblages of native
and exotic plants (“emerging” or “new forests”; Ewel and Putz 2004) as the target for most
restoration projects. Alternatively, investment in projects aimed at reestablishing assemblages
typical of intact rainforests may need to be restricted to high-value applications (e.g., habitat
for threatened species or plantings strategically positioned to act as future seed sources; Kooy-
man 1996) rather than being used as the default approach across the landscape, as currently
tends to be the case.
All models of rainforest dynamics recognize the importance of landscape context in
affecting the trajectory of restored sites. To date, much restoration effort in Australian has
been expended on the creation of corridors to facilitate dispersal between remnants. How-
ever, corridors do not address other landscape-scale factors affecting the persistence of biota
in remnants, such as the influence of the surrounding matrix. A better understanding of the
relative importance of the extent and configuration of rainforest cover at the landscape scale
(e.g., Bennett et al. 2006) and integration of this knowledge into models of rainforest dynam-
ics is required to inform future restoration projects aimed at conserving rainforest biota.
Acknowledgments
Thanks to Peter Erskine, Kylie Freebody, Cath Moran, Miriam Paul, Scott Piper, and Tang
Yong for participating in a workshop on “Rainforest Dynamics and Restoration,” where the
ideas presented here were developed. The contributions of JK and CC to this article were
supported in part by funding from the Australian government’s Marine and Tropical Sciences
Research Facility.
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... Rainforest plants with large seeds are particularly prone to dispersal limitation in and around rainforest fragments because the specialised animals they rely on are often restricted to large intact forests (Harrington et al. 1997;Moran et al. 2009). For this reason, the restoration of corridors to facilitate dispersal between remnant forests has been a prominent part of rainforest restoration in Australia over the past few decades (Kanowski et al. 2008b). Similarly, the provision of perches and fast-growing, early successional species to facilitate dispersal into restoration sites can be advantageous (Toh et al. 1999;Shiels and Walker 2003). ...
... Large areas of rainforest have been cleared for agriculture, including the establishment of pasture grasses. When these areas are abandoned and protected from fire and other major disturbances, secondary forest may eventually replace the grassland (Kanowski et al. 2008b). Such secondary forests may contain a mixture of native and exotic tree species, and, in subtropical Australia, areas that once supported wet rainforest are often dominated by the exotic camphor laurel (Cinnamomum camphora) (Big Scrub Rainforest Landcare Group 2005); see section on Weeds below. ...
... Stands of camphor laurel can function as 'nurse' trees for native rainforest trees, and facilitate the conversion of pasture to rainforest (Big Scrub Rainforest Landcare Group 2005), and this can be a relatively economic method of restoring rainforest )(see Encouraging natural regeneration section below). However it is not known whether stands of camphor laurel will eventually revert to native rainforest without intervention (Kanowski et al. 2008a) and this is probably unlikely if most seed rain is composed of exotic species (Kanowski et al. 2008b). Instead, a strategy of selective, progressive culling of camphor laurel trees is recommended, where trees are poisoned but left in situ to provide perches for seed dispersers, some shade for seedlings, and extra carbon for the soil (Big Scrub Rainforest Landcare Group 2005). ...
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 In Queensland, rainforest includes wet rainforest and dry rainforest (some of which are also known as vine-thicket or softwood scrub).  Temperature, moisture, soil fertility and drainage exert a strong influence on the carbon accumulation rates and standing carbon stocks in rainforest, but management can also have a large effect.  Based on sites in Queensland, wet rainforest has the potential to store from 300 to more than 1700 tonnes of carbon dioxide equivalent (CO2-e) per hectare, and dry rainforest has the potential to store from 170 to more than 800 tonnes CO2-e per hectare.  The maximum accumulation rate of wet rainforest can exceed fifteen tonnes/ha of CO2-e per year, while the rate is likely to be slower for dry rainforest.  Weeds, fire, vines, grazing pressure and seed limitation can all limit the establishment and growth of rainforest plants. Careful management of these threats is required in the early stages of restoration.  Restoring rainforest will be of great benefit to biodiversity, as rainforests are diverse ecosystems and support a high number of plant and animal species.
... It also promotes botanical supplementation (Li et al., 2012). As other research on reforestation in Australia and elsewhere has shown, this system appears to have the lowest economic recovery costs (Kanowski et al., 2008). The relative effectiveness of mollisols quality restoration on cultivated lands through ecological corridors depends on their ability to restore ecosystem properties and their relative costs (Paul et al., 2010). ...
... Studies in other regions also support such observations (Cheng et al., 2021). Conversely, a smaller corridor width is not conducive to supporting large birds and mammals (Kanowski et al., 2008). We found that, based on the same restoration period, within the width range of 125-150 m, the growth rate of the TN indices was the fastest in the entire observation period, and TN could recover to the level of cultivated land outside the corridor where fertilizers were applied. ...
Article
Full-text available
Artisficial ecological corridors (AECs) are internationally recognized as a standard method for restoring the regional ecological environment. However, the coupling relationship between AECs and soil quality has rarely been studied. Harbin, a typical mollisols region in the cold area of China, has severe soil problems and remediation is urgently needed, yet AEC research in this region is lacking. Based on the perspective of soil restoration, the construction factors of ecological corridors are quantitatively evaluated. It can predict the long-term impact of AECs already built along Harbin’s Ashi River on soil chemical indices. This research studied the ecological restoration of secondary woodland, cultivated land within the ecological corridor, and cultivated land outside the influence range of the corridor under the influence of continuous recovery time and different locations in the corridor (distance from the Ashe River). Soil samples were taken from 5 plots, with a total of 161 samples, and 12 indices of soil ecological characteristics were monitored. The result are as follows: It is believed that the quality restoration of mollisols through ecological corridors has great application potential. Based on the low-cost natural restoration of ecological corridors, the highest values of total phosphorus (TP) and soil organic matter (SOM) in soil indices were detected in corridors (restored for more than 10 years). In addition, after ten years of recovery, pH and electrical conductivity (EC) in the ecological corridor returned to normal from high levels in cultivated land that far exceeded the reference values. The recovery process of mollisols mass begins to decrease, then increases, and finally reaches and exceeds the reference value of standard mollisols. The redundancy analysis of soil samples found the distance to be a key factor affecting soil total nitrogen (TN), SOM, and cation exchange capacity (CEC). Recovery time is a crucial factor affecting soil total organic carbon (SOC), pH and EC. According to the TN, SOM, and CEC mollisols indices, the ecological corridor’s unilateral width is 125-150m. According to the SOC, pH, and EC indices of mollisols, the AECs should complete a natural recovery cycle of a minimum of 13 years. This study reveals the change mechanism of soil quality in mollisols area corridors based on recovery time and location. This research offer ideas and a scientific basis for worldwide governments in mollisols to formulate mollisols restoration policies.
... These factors include distance to the nearest vegetation and the quality of that vegetation Zimmerman et al. 2000), the suitability of microclimate on site for seedling establishment and growth (Uhl 1987), competition between grasses and seedlings (Uhl 1987), and previous land use history (Rasiah et al. 2004). Some abandoned sites may recover vegetation structure within a few decades, but the resultant species composition may be completely different from that of the original rainforest (Holl et al. 2000;Kanowski et al. 2009). Previous studies have also shown that active restorative intervention is essential if the system's natural recovery process is slow or the damaged ecosystem is in a state of arrested development (Zahawi & Augspurger 1999;Zimmerman et al. 2000;Florentine & Westbrooke 2004b;Sampaio 2007). ...
... A positive effect of species richness in restoration plantings for restoration success has been observed in the Atlantic rainforest of Brazil (Massad et al. 2011), in the Brazilian Amazon (Parrotta & Knowles 1999), in northeastern and southeastern Queensland (Kanowski et al. 2009), in tallgrass prairie in the Indiana, U.S.A. (Rowe & Holland 2013), and in dry forests in Hawaii (Ammondt & Litton 2012). This study, the two framework treatments provided very similar results to the maximum diversity method, and these plots need to be monitored over a longer period of time to determine which method or methods actually provide the best restoration outcomes. ...
Article
A long-term rainforest restoration experiment was established on abandoned pasture in northeastern Queensland in 1993 to examine the effectiveness of five different restoration planting methods: (T1) control (no plantings); (T2) pioneer monoculture (planting seedlings of one pioneer species, Homalanthus novoguineensis, Euphorbiaceae); (T3) Homalanthus group framework method (H. novoguineensis and eight other pioneer species); (T4) Alphitonia group framework method (Alphitonia petriei, Rhamnaceae, with eight other pioneer species); and (T5) maximum diversity method (planting pioneers, middle-phase species, and mature-phase species). We investigated temporal patterns in the (1) fate of seedlings originally planted in 1993; (2) natural recruitment of native plant species; and (3) current habitat structure (canopy cover and ground cover of grasses and invasive plants) within each restoration treatment. A total of 97% of seedlings planted in T2 died within the first 13 years and all had died by 2014. A total of 72% of seedlings planted in T3, 55.5% of seedlings planted in T4, and 55% of seedlings planted in T5 also died by 2014. By 2014, 42 species from 21 families had recruited across the experimental site, and the abundance of recruits was almost twice that recorded in 2001 and 2006. Overall, T3, T4, and T5 had the greatest diversity and abundance of recruits. By 2014, canopy cover was greatest in T3, T4, and T5 but grass cover was least in T5. It is concluded that some restoration success measures increase with planting diversity, but overall the rate of recovery is similar in framework species and maximum diversity method.
... These factors include distance to the nearest vegetation and the quality of that vegetation Zimmerman et al. 2000), the suitability of microclimate on site for seedling establishment and growth (Uhl 1987), competition between grasses and seedlings (Uhl 1987), and previous land use history (Rasiah et al. 2004). Some abandoned sites may recover vegetation structure within a few decades, but the resultant species composition may be completely different from that of the original rainforest (Holl et al. 2000;Kanowski et al. 2009). Previous studies have also shown that active restorative intervention is essential if the system's natural recovery process is slow or the damaged ecosystem is in a state of arrested development (Zahawi & Augspurger 1999;Zimmerman et al. 2000;Florentine & Westbrooke 2004b;Sampaio 2007). ...
... A positive effect of species richness in restoration plantings for restoration success has been observed in the Atlantic rainforest of Brazil (Massad et al. 2011), in the Brazilian Amazon (Parrotta & Knowles 1999), in northeastern and southeastern Queensland (Kanowski et al. 2009), in tallgrass prairie in the Indiana, U.S.A. (Rowe & Holland 2013), and in dry forests in Hawaii (Ammondt & Litton 2012). This study, the two framework treatments provided very similar results to the maximum diversity method, and these plots need to be monitored over a longer period of time to determine which method or methods actually provide the best restoration outcomes. ...
... For example, selection of species for initial planting or later enrichment could more explicitly incorporate slow-recruiting taxa, in both plantings and regrowth. Such concepts have been previously discussed (Elliott et al. 2013;Goosem & Tucker 2013;Kanowski et al. 2008), but firm evidence about which taxa actually fail to recruit into biodiversity plantings has largely been lacking (although see (2003)). Based on our findings, species with large diaspores and those from nearbasal evolutionary lineages are prime candidates. ...
... Low diversity and monoculture timber plantations are considerably less costly to establish (e.g. AUD$4-8,000 in Erskine et al. (2007)), but with inferior biodiversity outcomes (Catterall et al. 2008). ...
Article
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There is current debate about the potential for secondary regrowth to 'rescue' tropical forests from an otherwise inevitable cascade of biodiversity loss due to land clearing, and scant evidence to test how well active restoration may accelerate recovery. We use site chronosequences to compare developmental trajectories of vegetation between self-organised forest regrowth and biodiversity plantings (established for ecological restoration, with many locally native tree species at high density), in the Australian wet tropics uplands. Across 28 regrowth sites aged 1-59 years, some structural attributes reached reference rainforest levels within four decades, whereas wood volume and most tested components of native plant species richness (classified by species' origins, family and ecological functions) reached less than half reference rainforest values. Development of native tree and shrub richness was particularly slow among species that were wind-dispersed or animal-dispersed with large (>10 mm) seeds. The latter group includes many species from near-basal evolutionary lineages that contribute to recognised World Heritage values of the study region. Faster recovery was recorded in 25 biodiversity plantings aged 1-25 years, in which: wood volume developed more rapidly; native woody plant species richness reached values similar to reference rainforest and was better represented across all dispersal modes; and species from near-basal plant families were better (although incompletely) represented. Both plantings and regrowth showed slow recovery in species richness of vines and epiphytes, and in overall resemblance to forest in species composition. These findings contribute to informed decision-making about when and where to invest in active restoration, and provide strong evidence that protecting old growth forest is crucially important for sustaining tropical biodiversity. This article is protected by copyright. All rights reserved.
... Kondisi sebuah bentang alam dengan berbagai tipe ekosistem didalamnya tidaklah stabil karena di dalam ekosistem hutan, contohnya, akan terjadi proses suksesi. Bahkan hutan dalam kondisi klimaks masih berproses secara dinamis dan mengalami perubahan dari waktu ke waktu (Kanowski et al., 2009) dan kejadian kebakaran seringkali menjadi pemicu proses suksesi (Hartman dan McCarthy, 2008). Proses suksesi akibat gangguan kebakaran maupun gangguan lainya disebut dengan suksesi sekunder. ...
Technical Report
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Kajian riset yang dilakukan atas kerja sama antara Badan Restorasi Gambut, Kedeputian Penelitian dan Pengembangan Kelompok Kerja Penelitian dengan konsorsium yang terdiri dari Pusat Penelitian dan Pengembangan Hutan, Badan Litbang dan Inovasi, Universitas Sriwidjaya, Universitas Riau dan Universitas Palangka Raya dilakukan selama kurang lebih lima bulan sejak bulan Juli hingga November 2020. Topik riset Penentuan dan Pengembangan Kriteria Kerusakan dan Indikator Pemulihan Ekosistem Rawa Gambut merupakan topik riset nomor 5 dari 9 topik yang ditawarkan oleh BRG. Tujuan riset adalah mengembangkan kriteria kerusakan dan indikator pemulihan ekosistem gambut di tiga KHG di tiga Provinsi: KHG Pulau Tebing Tinggi, Provinsi Riau; KHG Sungai Mendahara-Batang Hari, Provinsi Jambi; dan KHG Sungai Kahayan-Sebangau, Provinsi Kalimantan Tengah. Ada lima (5) luaran yang akan dihasilkan yakni: 1) Peta klasifikasi tipologi dan sebaran ekosistem gambut atas dasar kondisi biofisik kawasan di tiga KHG, 2) Kriteria kerusakan berdasarkan data dan informasi mengenai tipe dan sebaran KHG (area gambut) yang sudah terdegradasi, 3) Paket ilmu pengetahuan dan teknologi (iptek) untuk restorasi ekosistem gambut dan pencegahan kebakaran lahan dan hutan, 4) Indikator pemulihan kerusakan di ekosistem gambut dan 5) Indikator kelembagaan restorasi dan perbaikan tata air secara permanen pada kesatuan hidrologis gambut dengan pola partisipatif.
... Kooyman 1996;Tucker and Murphy 1997;Freebody 2007; and see Acknowledgments) and from relevant research (e.g. Grundon et al. 2002;Catterall et al. 2004Catterall et al. , 2008Kanowski and Catterall 2007;Kanowski et al. 2008aKanowski et al. , 2008b2008c;2008d). ...
... Alternative frameworks such as the State and Transition model have been developed for systems such as the rangelands (Whalley 1994, Bestelmeyer et al. 2011, Schooler et al. 2011. Kanowski et al. (2009), in reviewing a number of vegetation models applied to primary and secondary tropical rainforest, note the widespread use of state and transition models to describe rainforest restoration. There is the potential that this could be a useful model to apply to mature rainforests and associated vegetation. ...
Technical Report
Full-text available
This technical report provides an overview of the evolution and ecology of cool temperate rainforests in Tasmania. Associated ecosystems such as tall eucalypt forests are also included. The report was a valuable input to the assessment of World Heritage status of these forests.
Article
Fragmentation presents a threat to tropical forest biodiversity and restoration can be expensive. Secondary forests regenerating on abandoned pasture are widespread, represent an opportunity to restore rainforest at minimal management cost, but can become arrested in a state dominated by a single tree species. Species richness and diversity was assessed from 26 sites in Acacia secondary forests in the Australian Wet Tropics of varying age since abandonment and the influence of rainfall and soils, and the context of remnant mature forest on succession were assessed. Stand structure indicated a lack of Acacia recruitment. Late successional species richness and diversity increased with age indicating recruitment under the Acacia canopy. The species richness of late successional tree species with fruit size 10 mm or larger also displayed an increasing trend with age, although it was statistically not significant. Forest succession progresses in Acacia secondary forest and large seeded tree species are able to recruit. The enhancement of rainforest succession with fertile geology, increased rainfall or with more remnant forest in the vicinity was not evident in this study. Secondary forest even when dominated by a single species, particularly a nitrogen fixing legume, represents a viable means of tropical forest restoration provided there is sufficient mature forest in the region to act as a seed source.
Article
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An important aim of plant ecology is to identify leading dimensions of ecological variation among species and to understand the basis for them. Dimensions that can readily be measured would be especially useful, because they might offer a path towards improved worldwide synthesis across the thousands of field experiments and ecophysiological studies that use just a few species each. Four dimensions are reviewed here. The leaf mass per area-leaf lifespan (LMA-LL) dimension expresses slow turnover of plant parts (at high LMA and long LL), long nutrient residence times, and slow response to favorable growth conditions. The seed mass-seed output (SM-SO) dimension is an important predictor of dispersal to establishment opportunities (seed output) and of establishment success in the face of hazards (seed mass). The LMA-LL and SM-SO dimensions are each underpinned by a single, comprehensible tradeoff, and their consequences are fairly well understood. The leaf size-twig size (LS-TS) spectrum has obvious consequences for the texture of canopies, but the costs and benefits of large versus small leaf and twig size are poorly understood. The height dimension has universally been seen as ecologically important and included in ecological strategy schemes. Nevertheless, height includes several tradeoffs and adaptive elements, which ideally should be treated separately. Each of these four dimensions varies at the scales of climate zones and of site types within landscapes. This variation can be interpreted as adaptation to the physical environment. Each dimension also varies widely among coexisting species. Most likely this within-site variation arises because the ecological opportunities for each species depend strongly on which other species are present, in other words, because the set of species at a site is a stable mixture of strategies.
Book
Why do Australian rainforests occur as islands within the vast tracts of Eucalyptus? Why is fire a critical ecological factor in every Australian landscape? What were the consequences of the ice-age colonists use of fire? In this original and challenging book, David Bowman critically examines hypotheses that have been advanced to answer these questions. He demonstrates that fire is the most critical factor in controlling the distribution of rainforest throughout Australia. Furthermore, while Aboriginal people used fire to skilfully manage and preserve habitats, he concludes that they did not significantly influence the evolution of Australia's unique flora and fauna. This book is a comprehensive overview of the diverse literature that attempts to solve the puzzle of the archipelago of rainforest habitats in Australia. It is essential reading for all ecologists, foresters, conservation biologists, and others interested in the biogeography and ecology of Australian rainforests.
Article
There are many peoples living in the tropical forested part of our planet today. A few of these subsist entirely on the products of the rainforest without significant detrimental impact while many others live where the rainforest was after first destroying it. Between these extremes are others who gain their livelihood from this ecosystem. It is also possible that people have lived in rainforests or off their resources for a very long time and it is the task of archaeology to understand and explain the development of human-rainforest adaptation. Since archaeologists require information from the present to build explanatory models of past human behaviour it follows that they require current data from not only ecological studies of modern rainforests but also data from anthropological studies of human adaptations to this ecosystem. The scientific value of this ecosystem cannot be overestimated; without tropical rainforests there are no tropical rainforest peoples and without both, our inferences concerning the human past must be severely handicapped. We underscore this point through a review of the antiquity of rainforest adaptation and then examine in more detail the archaeological quest for understanding about the prehistoric use of the rainforest by the peoples of northeast Queensland. -from Authors
Article
Outlines some ideas on the nature of species succession, focusing on contemporary views on the relationships between time, space and direction. The components of disturbance which relate to intensity, size and, for rain forests especially, the extent are of fundamental importance. Secondary successions which follow moderately intensive disturbances are reconstructive or progressive. Regenerating secondary forest is completely dependent on external seed sources for all recruits of the mature phase rain forest trees; the role of pioneer species is not properly understood. Most natural disturbances are of local significance, and do not affect the rain forest as a whole, except integratively over time. Man has severly affected rain forest biomass. Management of rain forest biological systems needs to be designed to accommodate biological constraints, and economic considerations should take second place. -S.J.Yates