B I O L O G I CA L C O N S E RVAT I O N 1 3 1 ( 2 0 0 6 ) 4 3 3 –4 4 5

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General management principles and a checklist of

strategies to guide forest biodiversity conservation

D.B. Lindenmayera,*, J.F. Franklinb, J. Fischera

a
Centre for Resource and Environmental Studies, The Australian National University, GPO Box 4, Canberra, ACT 0200, Australia
b
Box 352100, College of Forest Resources, University of Washington, Seattle, WA 98195, USA

A R T I C L E I N F O A B S T R A C T

Article history: Many indicators and criteria have been proposed to assess the sustainable management of
Received 13 September 2005 forests but their scientific validity remains uncertain. Because the effects of forest distur-
Received in revised form bances (such as logging) are often specific to particular species, sites, landscapes, regions
1 February 2006 and forest types, management ‘‘shortcuts’’ such as indicator species, focal species and
Accepted 28 February 2006 threshold levels of vegetation cover may be of limited generic value. We propose an alter-
Available online 18 April 2006 native approach based on a set of five guiding principles for biodiversity conservation that
are broadly applicable to any forested area: (1) the maintenance of connectivity; (2) the
Keywords: maintenance of landscape heterogeneity; (3) the maintenance of stand structural complex-
Forest biodiversity conservation ity; and (4) the maintenance of aquatic ecosystem integrity; (5) the use of natural distur-
General conservation and bance regimes to guide human disturbance regimes.
management principles We present a checklist of measures for forest biodiversity conservation that reflects the
Checklist of strategies multi-scaled nature of conservation approaches on forested land. At the regional scale,
management should ensure the establishment of large ecological reserves. At the land-
scape scale, off-reserve conservation measures should include: (1) protected areas within
production forests; (2) buffers for aquatic ecosystems; (3) appropriately designed and
located road networks; (4) the careful spatial and temporal arrangement of harvest units;
and (5) appropriate fire management practices. At the stand level, off-reserve conservation
measures should include: (1) the retention of key elements of stand structural complexity
(e.g., large living and dead trees with hollows, understorey thickets, and large fallen logs);
(2) long rotation times (coupled with structural retention at harvest); (3) silvicultural sys-
tems alternative to traditional high impact ones (e.g., clearcutting in some forest types);
and (4) appropriate fire management practices and practices for the management of other
kinds of disturbances.
Although the general ecological principles and associated checklist are intuitive, data to
evaluate the effectiveness of many specific on-the-ground management actions are lim-
ited. Considerable effort is needed to adopt adaptive management ‘‘natural experiments’’
and monitoring to: (1) better identify the impacts of logging operations and other kinds
of management activities on biodiversity, and; (2) quantify the effectiveness of impact mit-
igation strategies; and (3) identify ways to improve management practices.
 2006 Elsevier Ltd. All rights reserved.

* Corresponding author: Tel.: +61 02 6241 5358.

E-mail addresses: [email protected], [email protected] (D.B. Lindenmayer).
0006-3207/$ - see front matter  2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2006.02.019
434 B I O L O G I C A L C O N S E RVAT I O N 1 3 1 ( 2 0 0 6 ) 4 3 3 –4 4 5

1. Introduction et al., 1994). The conservation of biodiversity is clearly part

of perpetuating ecosystem integrity (as highlighted in the
Forests support about 65% of the world’s terrestrial taxa above definition). There are numerous definitions of biodiver-
(World Commission on Forests and Sustainable Development, sity – Delong (1996) and Bunnell (1998) reviewed approxi-
1999) and have the highest species diversity for many taxo- mately 90 interpretations of the concept. For this paper,
nomic groups including birds (Gill, 1995), invertebrates (Erwin, biodiversity is considered to encompass:
1982; Majer et al., 1994) and microbes (Torsvik et al., 1990; Cro-
. . .genes, individuals, demes, populations, metapopulations, spe-
zier et al., 1999). Conserving forest biodiversity is therefore a
cies, communities, ecosystems, and the interactions between
critical task (Aanderaa et al., 1996; Hunter, 1999; Putz et al.,
these entities.
2000) and has rightly become a key component of many na-
tional and international forest management agreements Both the numbers of entities (genes, species, etc.), and the
(e.g. Commonwealth of Australia, 1998, 2001; Montréal Pro- differences within and between those entities are empha-
cess Liaison Office, 2000; Food and Agriculture Organisation sized in this definition (see Gaston and Spicer, 2004). This
of the United Nations, 2001). complexity coupled with the inadequate description of biodi-
Most programs to sustain forest biodiversity have focused versity currently available (e.g. Erwin, 1982; Torsvik et al.,
on the creation of protected areas. Reserves are a critical part 1990; Majer et al., 1994) make it difficult to judge whether or
of any credible strategy for conserving forest biodiversity not forests are being managed in an ecologically sustainable
(Norton, 1999), but reserves alone are insufficient to ade- way. There are no cases anywhere in the world where ecolog-
quately conserve forest biodiversity (Sugal, 1997; Daily et al., ically sustainable forestry practices have been demonstrated
2001; Lindenmayer and Franklin, 2002), in part because 92% unequivocally (Bunnell et al., 2003). Moreover, the concept
of the world’s forests are outside formally protected areas of sustainability per se is complex and dynamic since ecolog-
(Commonwealth of Australia, 1999). ical sustainability is an overall direction in conservation and
In this paper, we outline a series of ecological principles to forest management and not all of the movement will neces-
guide how forestry practices might best achieve biodiversity sarily be forward (Lindenmayer and Franklin, 2003).
conservation. A checklist of on-ground management strategies Nevertheless, several international and national initiatives
aimed at operationalising these principles is described. Prob- have sought to develop criteria and indicators of sustainabil-
lems associated with biodiversity conservation ‘‘short-cuts’’ ity in forests (e.g. Arborvitae, 1995) despite the problems of: (1)
are outlined to provide a context for the description of the eco- defining biodiversity; (2) determining what constitutes eco-
logical principles and an associated checklist of strategies. We logically sustainable forestry; and (3) the sheer impossibility
focus on existing areas of native forest rather than exotic plan- of measuring and monitoring the impacts on all species of
tations or farm forests that have been established on former various management practices. Some kinds of indicators of
grazing lands. Biodiversity conservation in plantations and sustainability will be essential, but the scientific validity of
farm forests is considered elsewhere (e.g. Peterken and Ratc- most indicators of biodiversity is poor. This is one reason
liffe, 1995; Moore and Allen, 1999; Lindenmayer and Hobbs, we have proposed the set of guiding principles and the check-
2004; Salt et al., 2004). In addition, much of our focus is on list presented later in this paper.
‘‘well-forested’’ landscapes such as those in North America,
South America and Australia where ‘‘western-style’’ exploita- 3. Problems with ‘‘short-cut’’ methods that
tion has been imposed only relatively recently. However, many attempt to promote ecologically sustainable forest
of the ecological principles and elements in the checklist management practices
should be relevant to forest management in other places such
as Europe (e.g. Peterken, 1996; Angelstam, 1996; Fries et al., ‘‘Short-cuts’’ aimed at promoting biodiversity conservation as
1997). Finally, our discussion is restricted to sustaining popula- part of the sustainable management of natural resources in-
tions of native forest biota, although many other factors (such clude indicator species, focal species, and thresholds in levels
as maintaining soil fertility and productivity) are also funda- of native vegetation cover. All of these (and others; e.g. see
mental aspects of ecologically sustainable forest management. McCarthy et al., 2004) have problems as discussed below.

2. Background – ecologically sustainable 3.1. Indicator species

forest management and biodiversity conservation
Landres et al. (1988, p. 317) defined an indicator species as:
We define ecologically sustainable forestry as:
‘‘an organism whose characteristics (e.g. presence or absence,
‘‘. . .perpetuating ecosystem integrity while continuing to provide population density, dispersion, reproductive success) are used
wood and non-wood values; where ecosystem integrity means as an index of attributes too difficult, inconvenient, or expensive
the maintenance of forest structure, species composition, and to measure for other species or environmental conditions of
the rate of ecological processes and functions within the bounds interest.’’
of normal disturbance regimes.’’
The indicator species approach is widely used as a mea-
Achieving ecological sustainability will often require deter- sure of ecologically sustainable forest management. Many
mining appropriate baselines and ranges of variability for taxonomic groups have been suggested as indicators (re-
natural disturbance regimes against which human distur- viewed by Lindenmayer and Burgman, 2005), while problems
bance regimes can be compared (Hunter, 1993; Rülcker with the indicator species concept have been reviewed
 B I O L O G I C A L C O N S E RVAT I O N 1 3 1 ( 2 0 0 6 ) 4 3 3 –4 4 5 435

thoroughly by Landres et al. (1988), Temple and Wiens (1989), sponses for biota is an increasing focus of biodiversity-re-
Niemi et al. (1997), Simberloff (1998), and Lindenmayer et al. lated research (e.g. McAlpine et al., 2002a; Radford and
(2000). The chief difficulties include: Bennett, 2004) including forest management research. They
could be a valuable tool for use in landscape management
• An absence of documented causal relationships between such as in planning the extent and spatial arrangement of
indicator species and the entities they are assumed to use of natural resources, but it seems possible that threshold
indicate. responses for aggregate measures such as species richness
• Major variation between species in their response to envi- may not exist in some ecosystems (Parker and Mac Nally,
ronmental change including members of the same guild 2002; Lindenmayer et al., 2005). Several reasons may account
or closely related species. for this:
• The insensitivity of indicator species to some types of sig-
nificant environmental change, and • Patterns of vegetation loss do not occur in a random fash-
• Insufficient knowledge to guide the selection of indicator ion (Saunders et al., 1987) which results in vegetation
species. For example, although many workers have pro- cover patterns comprised of highly varying levels of habi-
posed particular taxa as indicator species (e.g. Davey, tat quality (e.g. a distinct bias toward low productivity hab-
1989; Johnson, 1994; Hill, 1995), the associated organisms itats of limited value for human uses). Vegetation cover
or other entities for which they are supposed to be surro- levels may therefore equate poorly to levels of habitat suit-
gates often are not identified (Lindenmayer and Cunning- ability for many assemblages and elements of the biota.
ham, 1997). • Each species in an assemblage responds differently to
landscape change. For example, some may be more (rather
Much more work needs to be done to confirm the relation- than less) likely to occur in sub-divided landscapes. Thus,
ships between species chosen as indicators and environmen- many species in an assemblage may not respond in the
tal change (Simberloff, 1999; Fleishman et al., 2002; Kavanagh same way to the same landscape variable (e.g. exhibit a
et al., 2004). This does not mean that particular species sudden change point at 30% of native vegetation cover).
should not continue to be monitored or be the target of man- Notably, some species might be lost at higher levels, and
agement actions (reviewed by Thompson and Angelstam, some at lower levels of native vegetation cover (Radford
1999). However, it should not be assumed that the response et al., 2005).
of a given species (e.g. to logging or targeted habitat manage- • Other factors such as invasive pest and/or weed species
ment) will necessarily be a good surrogate for the responses may have a large impact on the distribution and abun-
of other taxa. dance of many members of an assemblage even at vegeta-
tion levels well above hypothesized vegetation cover
3.2. Focal species thresholds.

Focal species are defined as those most influenced by threat- Thresholds will exist in some landscapes, but it seems un-
ening processes, for example, the taxon or taxa most limited likely there will be generic rules for critical change points or
by dispersal abilities, resources or ecological processes (Lam- threshold levels of vegetation or habitat cover (e.g. 10%, or
beck, 1997). A landscape may then be managed for a suite of 30%, or 70%) that can be applied broadly across different land-
focal species, each of which is thought to be sensitive to a par- scapes and different biotic groups (Parker and Mac Nally,
ticular threatening process. There are serious flaws in the fo- 2002). Major changes may occur across a broad band of points
cal species approach (Lindenmayer et al., 2002a). Like other and the threshold concept might be better re-described as a
taxon-based surrogate schemes, a suite of focal species is pre- regime shift – a phenomenon for which there is good empir-
sumed to act collectively as a surrogate for other elements of ical evidence (Folke et al., 2004).
the biota, but a landscape managed for a given set of focal
species may not meet the requirements of the remaining biota.
The focal species approach also may be difficult to apply 3.4. Overview of problems with ecological ‘‘short-cuts’’
because of the lack of science to guide the selection of a rea-
sonably reliable set of focal species in the majority of land- All of the ‘‘ecological short-cuts’’ exhibit deficiencies which
scapes. Perhaps the real success of the focal species limit their widespread use in gauging the sustainability of
approach will be its ability to act as a ‘‘social hook’’ to moti- forest management. Uncritical use of such short-cuts there-
vate people, communities and governments to tackle the dif- fore may lead forest managers to believe that a forest is
ficult process of landscape management (Lindenmayer and being managed sustainably when it is not. Therefore, we
Fischer, 2003). propose an alternative approach of general ecological princi-
ples for biodiversity conservation and a practical checklist
3.3. Thresholds of vegetation cover of multi-scaled on-ground management practices. Such gen-
eral principles and an associated checklist may, in turn, pro-
With and Crist (1995) defined thresholds as abrupt, non-lin- vide a benchmark against which new codes of practice
ear changes that occur in some measure (such as the rate of might be developed or the efficacy of existing codes as-
loss of species) across a small amount of habitat loss (Rols- sessed and deficiencies in them subsequently addressed.
tad and Wegge, 1987; Andrén, 1994, 1999; Enoksson et al., These principles and a checklist are outlined in the follow-
1995; With and Crist, 1995). The search for threshold re- ing section.
436 B I O L O G I C A L C O N S E RVAT I O N 1 3 1 ( 2 0 0 6 ) 4 3 3 –4 4 5

4. General principles for managing forest (Whelan et al., 2002). In addition, landscapes are character-
biodiversity ized by natural environmental gradients (e.g. in topography,
climate, or soil type and soil depth; see Austin and Smith,
Species loss is predominantly driven by habitat loss (reviewed 1989). Landscape heterogeneity corresponds to the mosaic
by Groombridge and Jenkins, 2002; Primack, 2001; Fahrig, of patches representing different forest composition and
2003). Therefore, the overarching goal of conservation man- age classes within which different structural conditions
agement must be to prevent habitat loss. Forest biodiversity occur (Forman, 1995). Different species inhabit different
conservation will depend on maintaining habitat across the environmental conditions in natural landscapes and the
full range of spatial scales. There are five general principles diversity, size, and spatial arrangement of habitat patches
that can help meet this objective: is important for many taxa (e.g. Hanski, 1994; Saab, 1999;
Debinski et al., 2001).
• The maintenance of connectivity. Connectivity is the linkage of • The use of knowledge of natural disturbance regimes in natural
habitats, communities and ecological processes at multiple forests to guide off-reserve forest management practices. Strate-
spatial and temporal scales (Noss, 1991). Connectivity influ- gies for biodiversity conservation are most likely to be suc-
ences key biodiversity conservation processes such as pop- cessful in cases where human disturbance regimes (such as
ulation persistence and recovery after disturbance (e.g. logging) are similar in their effects to natural disturbance
logging, Lamberson et al., 1994), the exchange of individuals (Hunter, 1993; Korpilahti and Kuuluvainen, 2002); such as,
and genes in a population (Leung et al., 1993; Saccheri et al., for example, the kinds and numbers of biological legacies
1998), and the occupancy of habitat patches (Villard and (sensu Franklin et al., 2000) and the spatial patterns of envi-
Taylor, 1994). ronmental conditions (e.g. patch types) remaining after dis-
• The maintenance of the integrity of aquatic systems by sustaining turbance (Delong and Kessler, 2000). Organisms are likely to
hydrological and geomorphological processes. Aquatic features be best adapted to the disturbance regimes under which
of forest landscapes – streams, rivers, wetlands, lakes and they have evolved (Bergeron et al., 1999; Hobson and Schi-
ponds – are critically important to biodiversity and ecosys- eck, 1999), but are potentially susceptible to novel forms
tem function. A very large proportion of the biodiversity of disturbance (or combinations of disturbances) such as
found in forested landscapes is associated with aquatic eco- those that are more or less frequent and/or more or less
systems – including many terrestrial as well as all aquatic intensive than would normally occur (Lindenmayer and
organisms (Aapala et al., 1996; Calhoun, 1999; Soderquist McCarthy, 2002). Natural disturbance regimes may there-
and Mac Nally, 2000). fore be appropriate baselines and ranges of variability
• The maintenance of stand structural complexity. Structural against which human disturbance regimes can be com-
complexity per se is a common feature of all natural forests pared (Hunter, 1993; Angelstam et al., 1995).
throughout the world (Franklin et al., 1981, 2002; Berg et al.,
1994; Noel et al., 1998). Stand structural complexity embod- 4.1. Relevance to landscapes with a long history of
ies not only particular types of stand attributes, but also the ‘‘western-style’’ human management
way they are spatially arranged within stands (Franklin and
van Pelt, 2004). Attributes contributing to stand structural As set out at the start of this paper, the ecological principles
complexity include: (1) Trees from multiple age cohorts listed in the previous section are directed primarily at land-
within a stand. (2) Large living trees and snags (Linder and scapes with a relatively recent history of ‘‘western-style’’
Östlund, 1998). (3) Large diameter logs on the forest floor exploitation. While some of our principles will be relevant
(Harmon et al., 1986; Berg et al., 1994). (4) Vertical heteroge- to forest landscapes such as those in many parts of Europe
neity created by multiple or continuous canopy layers (Bro- with a prolonged history of management and extensive frag-
kaw and Lent, 1999). (5) Horizontal heterogeneity of which mentation (Fries et al., 1997), others may not. For example,
canopy gaps and anti-gaps are examples (Franklin and the fundamental importance of reserves is recognized in all
van Pelt, 2004). The maintenance of stand structural com- jurisdictions (Margules and Pressey, 2000). Similarly, the crea-
plexity is critical for forest biodiversity conservation tion and/or maintenance of stand structural complexity is
because it may allow organisms to persist in logged areas essential for biodiversity conservation in all forests, including
from which they would otherwise be eliminated. It also those with a long history of management (e.g. Linder and
may facilitate a more rapid return of logged and regener- Östlund, 1998). In addition, landscape heterogeneity is prefer-
ated stands to suitable habitat for species that have been able to intensive forest management resulting in landscape
displaced. The maintenance of stand structural complexity homogeneity (Lindenmayer and Fischer, in press). While the
may enhance dispersal of some animals through a cutover maintenance of landscape heterogeneity is best guided by
area – a ‘connectivity’ function. Finally, structural complex- an understanding of natural heterogeneity in a given land-
ity can provide the within-stand variation in habitat condi- scape, some landscapes have long since lost their natural dis-
tions required by some taxa – a ‘habitat heterogeneity’ turbance regimes (Zackrisson, 1977) and natural patterns of
function. heterogeneity (Gustaffson et al., 1999). Indeed, many Euro-
• The maintenance of landscape heterogeneity. Ecosystems are pean landscapes exemplify this situation (Peterken, 1996).
naturally heterogeneous and landscape heterogeneity is a Notably, landscape heterogeneity tends to benefit native spe-
feature of natural forests worldwide. Disturbance regimes cies richness in such situations, even if it is not based on nat-
may create heterogeneous land cover, such as different suc- ural heterogeneity patterns that once prevailed (e.g. Ferreras,
cessional stages in different locations following a wildfire 2001; Palomares, 2001).
 B I O L O G I C A L C O N S E RVAT I O N 1 3 1 ( 2 0 0 6 ) 4 3 3 –4 4 5 437

Table 1 – Management strategies to achieve general biodiversity conservation principles (based on Lindenmayer and
Franklin, 2002)

Principle Strategy

Principle 1 – maintenance of connectivity Riparian and other corridors

Protection of sensitive habitats within the matrix
Vegetation retention on logged areas throughout the landscape
Careful planning of roading infrastructure
Landscape reconstruction

Principle 2 – maintenance of landscape heterogeneity Riparian and other corridors

Protection of sensitive habitats within the matrix
Mid-spatial-scale protected areas
Spatial planning of cutover sites
Increased rotation lengths
Landscape reconstruction
Careful planning of roading infrastructure
Use of natural disturbance regimes as templates

Principle 3 – maintenance of stand complexity Retention of structures and organisms during regeneration harvest
Habitat creation (e.g. promotion of cavity-tree formation)
Stand management practices
Increased rotation lengths
Use of natural disturbance regimes as templates

Principle 4 – maintenance of intact aquatic ecosystems Riparian corridors

Protection of sensitive aquatic habitats with off reserve areas
Careful planning and maintenance of roading infrastructure

Principle 5 – adoption of natural disturbance regimes as Ensuring that strategies are varied between different between
templates to guide human disturbance regimes stands and landscapes (‘do not do the same thing everywhere’)

4.2. Spatial and temporal variation in conditions as a (e.g. protected areas within production forests) and stand-
risk-spread strategy level silvicultural approaches (e.g. stand structural
retention).
The maintenance of habitat is the overarching goal of forest
biodiversity conservation, but what constitutes suitable habi-
tat is different for each species (Morrison et al., 1992; Block 5.1. Large ecological reserves
and Brennan, 1993; Guisan and Zimmerman, 2000). Similarly,
what constitutes suitable connectivity, stand complexity, land- Large ecological reserves are an essential part of all compre-
scape heterogeneity and aquatic ecosystem integrity will also hensive biodiversity conservation plans and are critically
be defined on a species-specific basis and can vary markedly important for at least five key reasons (after Lindenmayer
between species. Enabling or creating spatial and temporal var- and Franklin, 2002):
iation in a range of conditions at multiple spatial scales is a
practical response to the problem of defining these variables • They support some of the best examples of ecosystems,
for a large set of species. Conditions needed by different species landscapes, stands, habitat, and biota and their inter-rela-
should then be provided in at least some parts of a forest land- tionships as well as opportunities for natural evolutionary
scape (Table 1). Management for diversity calls for diversity of processes.
management (Evans and Hibberd, 1990) and this a risk-spread- • Many species find optimum conditions only within large
ing approach to forest management to ensure not doing the ecological reserves which become strongholds for these
‘‘same thing everywhere’’ (Lindenmayer and Franklin, 2002). species.
• Some species are intolerant of human intrusions, making it
5. A checklist for forest biodiversity imperative to retain some areas which are largely exempt
conservation from human activity.
• Large ecological reserves provide ‘‘control areas’’ against
The conservation of forest biodiversity embodies a contin- which the impacts of human activities in managed forests
uum of conservation approaches from the establishment can be compared.
of large ecological reserves through to an array of off-re- • The effects of human disturbance on biodiversity are poorly
serve conservation measures including the maintenance of known and some impacts may be irreversible. Others such
individual forest structures at the smallest spatial scale as synergistic and cumulative effects can be extremely dif-
(Fig. 1). In Table 2 the elements are arranged in approximate ficult to quantify or predict. These factors make large eco-
hierarchical order progressing from regional-scale strategies logical reserves a valuable ‘safety net’ relatively free from
(large ecological reserves) through landscape-level strategies human disturbance.
438 B I O L O G I C A L C O N S E RVAT I O N 1 3 1 ( 2 0 0 6 ) 4 3 3 –4 4 5

THE FOREST ESTATE

DESIGNATED PROTECTED AREAS OFF-RESERVE MANAGEMENT

• National parks
• Nature reserves
Special Protection Areas Other Areas (incl. Potential Harvest Areas)

• Special habitats (e.g.

lakes, caves, Landscape level Stand level
wetlands,
biological
hotspots)
• Corridors (e.g. • Arrangement of timber management units in • Structural retention e.g.
riparian, wildlife) time and space potential den trees,
- Dispersion versus aggregation coarse woody debris
- Rotation length • Habitat creation
- Size of harvest units • Habitat retention e.g.
• Transport systems (roads) understorey islands
• Landscape goals for specific structural
features e.g. retention of large snags

Fig. 1 – A framework for biodiversity conservation across protected areas (typically in public ownership) and off-reserve areas
(including public and private native forests) (redrawn from Lindenmayer and Franklin, 2002).

Large ecological reserve systems are rarely comprehen- • The selection of the spatial and temporal pattern for har-
sive, representative and adequate for all elements of biodi- vest units or other management units (Franklin and For-
versity (Margules and Pressey, 2000; Scott et al., 2001). In man, 1987).
other cases, past land management means there are few • The application and/or management of appropriate distur-
or no opportunities to set aside large ecological reserves bance regimes such as those involving fire (Rülcker et al.,
(e.g. in parts of southern Sweden; Gustaffson et al., 1999). 1994; Keith et al., 2002) or grazing (Vera, 2000).
Hence, credible plans for forest biodiversity conservation • The protection of aquatic ecosystems and networks (such
must incorporate off-reserve approaches that complement as rivers, streams, lakes and ponds), specialized habitats
reserve-based approaches – i.e. conservation strategies at (e.g. cliffs and caves), wildlife corridors, biological hotspots
the landscape and stand levels (Lindenmayer and Franklin, (e.g. spawning habitats, roosting areas for birds or camps
2002). for flying foxes), and remnants of late-successional or old-
Mapping of forest types across all tenures and assessing growth forest and disturbance refugia found within off-
representativeness of forest types in the formal (public) reserve forests (McCarthy and Lindenmayer, 1999).
protected area system should reveal the extent to which
off-reserve conservation strategies are needed and the It is important to distinguish between large ecological re-
kinds of conservation management activities that will serves (Noss and Cooperrider, 1994) and the protection of
be required. For example, forest types that are poorly smaller areas within landscapes broadly designated for wood
protected in a reserve system will need to be managed dif- production (Gustaffson et al., 1999). Such systems of scattered
ferently than forest types already well represented in small reserves provide: (1) increased protection of habitats,
reserves. vegetation types, and organisms poorly represented or absent
in large ecological reserves; (2) protection for aquatic and
5.2. Off-reserve conservation measures at the semi-aquatic ecosystems; (3) refugia for forest organisms that
landscape-level subsequently provide propagules and offspring for recolonis-
ing surrounding forest areas as they recover from timber har-
The five broad categories of approaches to landscape-level vesting; and, (4) ‘stepping stones’ to facilitate the movement
off-reserve forest management are: of biota across managed landscapes.
The management of disturbances such as fire is an addi-
• Establishment of landscape-level goals for retention, main- tional key aspect of landscape-level sustainable forest man-
tenance, or restoration of particular habitats or structures agement and biodiversity conservation. This may involve
as well as limits to specific problematic conditions (e.g. both the suppression of unwanted (wild)fires and ignition of
the amount of a forest landscape subject to prescribed prescribed fires (Gill, 1999). Issues associated with the impacts
burning (Gill, 1999)). of disturbances by fire and its effects on biota are complex.
• The design and subsequent management of transportation This is because in some landscapes, such as those in Sweden,
systems (generally a road network) to take account of problems like a lack of regeneration of particular plant spe-
impacts on species, critical habitats, and ecological pro- cies have been created by the absence of fire (Zackrisson,
cesses (Forman et al., 2002). 1977) whereas in others such as Ponderosa Pine (Pinus ponder-
 B I O L O G I C A L C O N S E RVAT I O N 1 3 1 ( 2 0 0 6 ) 4 3 3 –4 4 5 439

Table 2 – Checklist of factors for off-reserve conservation management (based on Lindenmayer and Franklin, 2002)
Large ecological reserves
CARR principles (comprehensive, adequate, representative, replicated) for large ecological reserves and implications for adjacent Private Native
Forest lands

Maps of vegetation types for cross-tenure assessment of land uses

Landscape-level conservation strategies within off-reserve forest

Protected habitat within the landscape – protected areas at intermediate-spatial scales
Special habitats
Cliffs, caves, rockslides etc
Remnant patches of late-successional forest
Biological hotspots
Source areas for coarse woody debris, populations of rare species
Fire, wind and other disturbance refugia
Aquatic ecosystems and riparian buffers
Springs, seeps, lakes, ponds, wetlands, streams and rivers and associated buffers
Wildlife corridors

Other landscape-level considerations

Transportation systems (e.g. roading networks)
Landscape-level goals for specific structural features (e.g. large trees with hollows)
Spatial and temporal patterns of timber harvesting
Dispersed versus aggregated
Size of harvest units
Rotation lengths
Restoration and re-creation of late-successional (old growth) forests or other habitat features
Appropriate fire management regimes (e.g. maintenance of a range of post-fire age classes), and varied prescriptions between stands
Management strategies for particular species (e.g. Swift Parrot)
Control strategies for unwanted species (e.g. weed management, feral animal control)
Consideration of natural disturbance regimes as template for logging regimes (e.g. identification of natural disturbance refugia as places for
logging exemption)

Stand-level conservation strategies within off-reserve forest

Habitat within management units or stands
Retention of structures and organisms at time of regeneration harvest
Trees with hollows (and recruits), large decaying logs, understorey thickets, gaps and anti-gaps
Creation of structural complexity through stand management activities
Lengthened rotation times
Application of novel silvicultural systems to meet stand-level goals
Variable retention harvest system (VRHS), novel thinning systems
Appropriate fire management regimes and varied prescriptions between stands
Consideration of adjacency to other vegetation/stands (= landscape context)
Management of additional kinds of disturbances (e.g. grazing)
Targeted management strategies for particular species
Control strategies for unwanted species (e.g. weed management, feral animal control)
Consideration of natural disturbance regimes as template for logging regimes
Stand level patterns and quantities of biological legacies that remain after natural disturbance events

osa) stands in south-western USA, fire suppression together ‘‘aim at achieving suitable proportions of landscape with a vari-
with livestock grazing mean that wildfires are too intense ety of times-since-fire stages within appropriate intensity levels
(Covington et al., 1997; Moore et al., 2004). The objectives of at appropriate times of the year and within appropriate fre-
fire management will vary depending on the proximity of quency range’’.
people and property and the relative importance of values like
timber resources, conservation and water production (Keith 5.3. Conservation measures at the stand-level
et al., 2002). The most appropriate fire regime also will depend
on the characteristics of the system (Rülcker et al., 1994). Dif- The objective of off-reserve management at the stand level is
ferences between vegetation communities and individual ele- to increase the contribution of harvest units to the conserva-
ments of the biota in their response to fire mean that there tion of biodiversity. Harvest units can be managed to: (1) sus-
are no simple management recipes (Whelan, 1995). Fire re- tain species; (2) increase habitat diversity; (3) improve
gimes may be varied between and within landscapes, to cre- connectivity; (4) buffer sensitive areas, and, (5) sustain eco-
ate a range of conditions. Therefore, if unsuitable habitats system processes including site productivity.
are created in one area for a given species, there will be other The internal structure and composition of harvested units
places it can survive. Gill (1999, p. 47) argued that the manage- can have a significant influence on the degree to which a man-
ment of fire for biodiversity conservation should: aged forest can sustain biodiversity and maintain ecosystem
440 B I O L O G I C A L C O N S E RVAT I O N 1 3 1 ( 2 0 0 6 ) 4 3 3 –4 4 5

processes. Several broad types of strategies can contribute to It is notable that many current codes and standards of for-
the maintenance of structural complexity: estry practice usually do not take into account some of the is-
sues and management needs/considerations identified in the
• Structural retention at the time of regeneration harvest (e.g. checklist such as:
large hollow trees and associated recruit trees (Fries et al.,
1997); understorey thickets (Ough and Murphy, 1998), and • The importance of using natural disturbance regimes as
large fallen logs (Harmon et al., 1986)). In other cases, spe- templates to guide human (logging) disturbance regimes
cifically targeted strategies may be required to add or create to better ensure that forest ecosystems are managed within
particular structures such as girdling trees to increase the natural bounds of disturbance intensity and variability
quantities of dead wood (Bull and Partridge, 1986) or install- (Rülcker et al., 1994; Lindenmayer and McCarthy, 2002).
ing nestboxes (Petty et al., 1994). • The importance of maintaining natural disturbance refugia
• Management of regenerated and existing stands to create for biodiversity conservation (e.g. multi-aged stands in Vic-
specific structural conditions (e.g. through novel kinds of torian ash forests; see Mackey et al., 2002; patches of rem-
thinning activities (Carey et al., 1999a)). This may include nant rainforest in south-east Asia (Johns, 1996)), and the
the maintenance of open areas as well as heath and grass- careful management of these areas to ensure their integrity
land habitats within forests that can be critical for some key is not impaired by additional human disturbances (Van Nie-
elements of biota. For example, in the forests of the Swiss uwstadt et al., 2001).
Jura, a reduction in the cover of trees and shrubs is consid- • The need to limit multiple cumulative impacts on biodiver-
ered critical for the survival of populations of the Asp Viper sity and stand structural complexity in areas subject to for-
(Vipera aspis) (Jäggi and Baur, 1999). est management (e.g. the combination of logging, fire and
• Long rotations or cutting cycles (Seymour and Hunter, grazing) (see McAlpine et al., 2002b).
1999). • The need to ensure that post-disturbance (salvage) logging
• Application of appropriate disturbance management does not cause negative impacts on biodiversity and forest
regimes such as prescribed burning to reduce fuel loads structure, potentially magnifying effects of wildfires (Sha-
and reduce the risk of a high-intensity fire. kesby et al., 1996; Lindenmayer et al., 2004; Donato et al.,
2006).
The various stand-level strategies can often be effectively
combined to address a broader range of objectives as part of
innovative silvicultural systems that address the twin objec- 7. Other key issues
tives of commodity production and biodiversity conservation
(Taulman et al., 1998; Carey et al., 1999b; Hickey et al., 1999; 7.1. The need for multiple, multi-scaled conservation
Beese et al., 2003). For example, the advantages of long rota- measures
tions are multiplied when accompanied by structural reten-
tion at the time of harvest. Conversely, rotation times may Biodiversity is multi-scaled concept. Therefore, attempts to
be shortened if greater levels of retention characterise logged conserve forest biodiversity must also be multi-scaled – with
stands at the time of harvest. appropriate conservation strategies at the level of individual
trees through to landscape and regional levels. Multi-scaled
6. Making the checklist operational management is needed because:

Each landscape is unique in terms of physical and biological • There are multiple ecological scales for different ecological
conditions, human developments (such as roads), the objec- processes (Urban et al., 1987; Poff, 1997; Elkie and Rempel,
tives of the landowner(s), regulatory and social directives, 2001). For example, the ecological process of habitat loss
and the taxa targeted for conservation. Thus, it is impossible can occur at regional and landscape levels by activities such
to provide generic solutions to landscape- and stand-level as forest clearing (Angelstam, 1996). Particular age classes
prescriptions for the on-ground implementation of the check- (e.g. old growth) can be subject to habitat loss and fragmen-
list. As an example in an Australian context, the kinds of sil- tation within landscapes of formerly continuous forest
vicultural options and strategies relevant to the wet eucalypt cover. Finally, structural and floristic attributes can be lost
forests of Tasmania and its associated biota will be quite dif- from individual stands (Angelstam, 1996).
ferent, even for broadly similar forest types on the Australian • There are multiple ecological scales for different species
mainland (e.g. in the Central Highlands of Victoria). This is, in (Allen and Hoekstra, 1992). For example, the spatial require-
part, because of the significant differences in biodiversity (e.g. ments of invertebrates requiring decayed logs with particu-
hollow-dependent vertebrates) between the two regions (Gib- lar sorts of attributes (e.g. Velvet Worms [Phylum
bons and Lindenmayer, 2002). What constitutes suitable hab- Onycophora] Barclay et al., 1999) are markedly different
itat or connectivity for a given species in a particular from the spatial requirements of wide-ranging predators
landscape dominated by a particular forest type or set of for- such as large forest owls (Lamberson et al., 1994).
est types can be markedly different in another landscape • There are multiple ecological and management scales for
even for the same species (e.g. greater glider [Petauroides vo- the same species (Forman, 1964; Hokit et al., 1999). This
lans] Lindenmayer, 2002). Forest managers must therefore be was demonstrated for Leadbeater’s possum (Gymnobelideus
quite clear about the objectives they have for a forest land- leadbeateri) in the Central Highlands of Victoria – key man-
scape and for the stands which comprise that landscape. agement actions were required at the individual tree level,
 B I O L O G I C A L C O N S E RVAT I O N 1 3 1 ( 2 0 0 6 ) 4 3 3 –4 4 5 441

the stand level, the patch level, the landscape level and the tions are limited. Considerable effort is needed to implement
regional level (Lindenmayer, 2000). This outcome is paral- trule adaptive management ‘‘natural experiments’’, and mon-
leled by many other examples for forest landscapes around itoring to: (1) better identify the impacts of logging operations
the world ranging from tropical forests in New Guinea (Dia- and other kinds of management of biodiversity, and, (2) quan-
mond, 1973) to the temperate forests in the Bavarian Alps of tify the efficacy of impact mitigation strategies and ways to
Germany (Storch, 1997). improve practices where necessary. True adaptive manage-
ment involves rigorous monitoring and a commitment to
change when negative impacts of current practices are identi-
7.2. The need for multiple, multi-scaled conservation fied. Unfortunately, the record on forest monitoring (and par-
measures – risk-spreading ticularly for biodiversity management) is generally poor in
forests around the world (Lindenmayer, 1999) and this needs
Implementing an array of strategies at different scales is a to be rectified as part of attempts to make transitions to eco-
risk-spreading approach. That is, if one strategy subsequently logically sustainable forest management.
proves to be ineffective, others will be in place that might bet-
ter conserve the entities targeted for management. Risk- Acknowledgements
spreading reduces the over-reliance on any one particular con-
servation strategy and attempts to deal with the considerable This project was supported through the Joint Venture Agrofor-
uncertainty regarding the effectiveness of current conserva- estry Program and guidance from Dr. Rosemary Lott, David
tion management strategies. Risk-spreading is particularly Thompson, Phil Norman and Warwick Ragg was greatly
appropriate for biodiversity conservation because it is often appreciated. Insightful comments on the manuscript by Alex
extremely difficult to accurately forecast the response of Jay, David Thompson, Warwick Ragg and several anonymous
species to processes such as landscape modification (see referees were greatly appreciated.
Mac Nally et al., 2000), stand simplification (Lindenmayer
et al., 2002b), prescribed fire (Moore et al., 2004) and climate
change (McCarty, 2001).
R E F E R E N C E S

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