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Conservation status and threats to species associated with oldgrowth forests within the range of the Northern Spotted Owl
(Strix occidentalis caurina) in British Columbia, Canada
S. Yezerinac1 and F. M. Moola2
1
Biology Department
Bishop’s University
100 College Road, P.O.
Box 5000, Lennoxville,
Quebec J1M 2A7
[email protected]
Abstract. Decline of the Northern Spotted Owl (Strix occidentalis caurina) has drawn attention to the management of oldgrowth and late-successional forest habitat in the Canadian Pacific Northwest. Here we expand this focus to include 138
additional species of vertebrates and vascular plants that are found with Spotted Owls in old-growth forests in the
southwestern mainland of British Columbia, Canada. We amassed up-to-date conservation status designations and
enumerated known threats to survival for these species using online databases (NatureServe, COSEWIC, and SARA Public
Registry) in order to assess the taxonomic and trophic breadth of species at risk in this community. Not unexpectedly, we
found species at risk in all major taxonomic groups and that regulatory designations at provincial and federal levels are
inconsistent for many species. 16% of species are classified as endangered, threatened or of special concern by the B.C.
government, yet just 6% are being managed by B.C. Only 50% of species designated at risk by B.C. have been federally
assessed by COSEWIC and only 36% currently have protection under Canada’s Species at Risk Act (SARA), which
mandates protection of critical habitat. The threats to these species are varied, but timber harvesting and its indirect
effects (e.g. road building, forest fragmentation) make up the majority. Although recovery planning for Spotted Owls has
the potential to impact many other species at risk, we argue that the Spotted Owl is not a good Indicator, Keystone or
Umbrella species, and that even as a Flagship species it may be insufficient. So we outline a new flagship fleet concept,
nominate its members and summarize how it relates to conservation reserve planning.
Author for correspondence - The David
Suzuki Foundation,
Suite 219, 2211 West
4th Avenue, Vancouver,
BC V6K 4S2
[email protected]
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INTRODUCTION
Northern Spotted Owls (Strix occidentalis caurina) live
in old-growth and late-successional coniferous and
mixed-coniferous forests from California to British
Columbia (B.C.) (Gutiérrez et al. 1995). Populations
have been declining for over 30 years and the species
is legally listed as endangered in both the USA and
Canada. The primary cause of endangerment is the
loss of old-growth forest habitat necessary for foraging,
nesting and dispersal behaviors (Kirk 1999), though
disease, predation and competition may also be
secondarily operating (Hamer et al. 2001, Peterson and
Robins 2003, Kelly et al. 2003). Since 1993 the United
States has implemented a forest ecosystem management
plan with the goal of conserving suitable habitat,
recovering Spotted Owl populations and protecting cooccurring species (FEMAT 1993, Thomas et al. 1993).
In Canada, the Spotted Owl is found only in the
southwestern mainland of British Columbia (Kirk 1999).
A provincial plan in place since 1997 seeks to manage
“...habitat conditions and to maintain owl populations,
so long as the impact of those prescriptions is no more
than approximately a 10 % reduction in long-term timber
supply over [current]...levels” (SOMIT 1997a,b). Field
surveys indicate the population in B.C. is declining at
about 10% per year and 22 known owls remain
(government sources say that one ‘known’ juvenile died
recently) from an historical population of approximately
500 pairs (Barlee et al. 2005). A recovery strategy for
Northern Spotted Owls is still in preparation as of
December 2005. As one of Canada’s most endangered
species, the Spotted Owl receives much attention.
Here we focus on potential threats to other species that
inhabit late-successional and old-growth forests within
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Northern Spotted Owl (Strix occidentalis caurina). (Copyright Art Wolfe
-www.artwolfe.com)
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(e.g. Crooks & Soulé 1999, Schmidt 2003). Thus, the
pattern and extent of endangerment across a set of cooccurring species indicates the degree of perturbation
of ecological processes in a community. Our second
goal in this study is to compare the pattern of
endangerment across several major taxonomic groups
(vascular plants, amphibians, fish, birds, mammals)
for species that co-occur with Spotted Owls in oldgrowth forests to the pattern of endangerment for these
same taxonomic groups across British Columbia as a
whole. Because this analysis reveals endanger-ment
in this old-growth commu-nity to be taxonomically and
trophically widespread, our final goal is to consider
the utility of the northern Spotted Owl as a focal species
for conservation planning.
METHODS
Decades of logging
have taken their toll on
prime Spotted Owl
habitat in the Anderson
Creek Valley in British
Columbia. (Photo by
Jeremy Sean Williams,
2005, courtesy of
Western Canada
A comprehensive list of species associated with oldgrowth and late-successional forests of British
Columbia would ideally derive from extensive field
work, yet limited resources have so far prevented
such a compilation. We used the list of species
generated by the Canadian Spotted Owl Recovery
Team, which used their local expertise to edit the list
of species previously compiled for old-growth and
late-successional forests of the Pacific Northwest
region of the United States (Thomas et al. 1993).
The list for British Columbia includes 71 species of
vertebrates (4 amphibians, 34 birds, 17 mammals, 16
fish) and 67 species of vascular plants that are similarly
associated with late-successional and old growth habitat
as Spotted Owls, as well as 89 species of Arthropods
and 49 species of Molluscs listed as “at risk” (Harper
& Millikin 1994). Because “at risk” was not defined by
Harper and Millikin (1994), and a companion list of
Arthropods and Molluscs not at risk is unavailable, we
do not consider Arthropods and Molluscs further. We
note that 1 of these species is rated as Special Concern
by COSEWIC 2000. It is also important to stress that
while these other species share with Spotted Owls a
common dependence on late-seral forest habitat
conditions, they may not necessarily be ecologically
associated with owls (Harper and Millikin 1994).
the historic range of the Spotted Owl in British Columbia.
Canada has a Species at Risk Act (SARA) in force as
of June 2003. While the SARA prescribes actions for
focal species, it does allow for multi-species and
community level planning and protection if
circumstances warrant (http://www.sararegistry.gc.ca
see SARA Public Registry). A recent national analysis
across major watersheds shows that at risk terrestrial
species are concentrated in three areas of Canada as of
2001: the southwestern mainland of British Columbia,
southern Vancouver Island, and southern Ontario (Kerr
and Cihlar 2004). However, this analysis did not examine
the distribution of at risk terrestrial species among
different ecological communities. Thus, our first goal
in this study is to focus on co-occurring vertebrates and
vascular plants within the old-growth forest community
and assess which species are at risk of extinction and
to enumerate the threats to species survival.
Obtaining up-to-date information on biodiversity and
conservation is difficult. We used the NatureServe portal
(http://www.naturserve.org) to amass current (as of July
2004) conservation status designations. At the Provincial
level these came through the Conservation Data Center
of the BC Ministry of Sustainable Resource Management
(http://srmwww.gov.bc.ca/cdc/). British Columbia uses
the system developed by NatureServe to classify extant
species on a scale extending from 1 (critically imperiled),
through 2 (imperiled), 3 (vulnerable), 4 (apparently
secure), and 5 (secure) determined at a global, national,
or provincial scale. Species are then categorized onto
Red (endangered), Blue (special concern) and Yellow
(apparently secure) lists. Herein we focus on Red-listed
and Blue-listed species at risk. We amassed status
designations at the Federal level through the website of
A group of co-occurring species represents a functioning
ecological community in which interspecific interactions
(e.g. mutualisms, competition, herbivory, predation) affect
the relative abundance of species at different trophic
levels, as well as the flux of nutrients and energy through
food webs. The Northern Spotted Owl, for example, is a
large-bodied predator situated near the top of the food
web (Gutiérrez et al. 1995, Forsman et al. 2001), so its
decline presumably results in increases of its squirrel
prey plus other unspecifiable changes. Although such
trophic cascades are difficult to predict, it is wellsupported that anthropogenically-created endanger-ment
at one level can affect species at other trophic levels
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the Committee on the Status of Endangered Wildlife in
Canada (http://www.cosewic.-gc.ca/). COSEWIC
classifies extant species on a scale extending from
Endangered (facing imminent extirpation or extinction),
through Threatened (likely to become endangered if
limiting factors are not reversed), Special Concern
(particularly sensitive to human activities or natural
events but not endangered or threatened) and Not at Risk.
Herein we focus on those species classified as
Endangered, Threatened or Special Concern by
COSEWIC as being at risk. Finally, we used the
NatureServe Explorer portal to gather citations to the
scientific literature summarizing known threats to species.
We reviewed each of the 138 species and counted the
number of known threats for each species in the following
categories (Agriculture, Alien Species Introduction,
Disruption of Fire Regimes, Environmental
Contamination, Human Disturbance, Indirect Effects of
Timber Harvesting, Intentional and Accidental Mortality,
Livestock Grazing, Timber Harvesting, Mining + Oil &
Gas, Tourism, Restricted Distribution, Urbanization,
Other, Unknown). We compared the proportion of the
Spotted Owl’s co-occurring taxa at risk with provincewide data for British Columbia available from the
government’s website (http://srmwww.gov.bc.ca/atrisk/
toolintro.html).
concern (Appendix 1), which puts them in the purview of
Canada’s Species at Risk Act. Species that are put forward
by COSEWIC and not denied by Federal Minister(s) pass
onto Schedule 1 of the SARA. Eight of the 138 species we
examined are on Schedule 1, and two species are on either
Schedule 2 or 3 (Appendix 1), which serve as waiting lists
for species assessed by COSEWIC prior to October 1999
and which must be reassessed before consideration for
addition to Schedule 1. Nonetheless, the SARA normally
applies to areas of Federal jurisdiction, which excludes
almost all of terrestrial British Columbia, so over most of
B.C. only those species designated as “Identified Wildlife”
by the B.C. Ministry of Environment receive any special
management attention. Moreover, the Identified Wildlife
strategy contains a limitation on impact to timber supply
(SOMIT 1997a,b). As of July 2004 only 8 of the 22 cooccurring species identified as at risk by B.C. are formally
“Identified Wildlife” and are being managed by B.C.
(Appendix 1). It is important to note that these regulatory
patterns are evident for each of the taxonomic groupings we
examined except for vascular plants, in which 2 species are
Red listed and 3 species are Blue listed in B.C., none have
achieved federal conservation status, and none is “Identified
Wildlife”. Averaging across taxonomic groups, the proportion
of species at risk differs little between the set of species cooccurring with Spotted Owls (26% ± 6.1 S.E.) and all species
in these major taxonomic groupings found throughout British
Columbia (33% ± 6.1); the average percentage of species at
risk is not significantly different between these two groups
(t=0.86, d.f.=8, P=0.42; Figure 1).
RESULTS
Provincial and Federal legislation is such that conservation
listings under the British Columbia Ministry of Environment,
the national Committee on Status of Endangered Wildlife
in Canada (COSEWIC), and the national Species at Risk
Act (SARA) are often inconsistent (Table 1). Typically, a
species would initially be listed provincially; 22 of the 138
species co-occurring species are Red listed (Endangered
or Threatened) or Blue listed (Special Concern) by the
B.C. government (Appendix 1). (available at: http://
www.davidsuzuki.org/WOL/Publications.asp). Provincial
listing does not necessarily lead to any conservation action,
though at the discretion of cabinet the Wildlife Act may be
applied to endangered species. However, at the federal
level COSEWIC has assessed 11 of these 22 species and
also found them to be endangered, threatened or of special
From the scientific literature cited at NatureServe we
were able to identify 119 specific threats for these 138
species. Timber harvesting was the most commonly
stated threat, followed in frequency by a set of threats
that indirectly arise from timber harvesting (road building,
forest fragmentation, and herbicide application to tree
plantations). Timber harvesting plus indirect effects of
timber harvesting comprised 41% and 44% of all identified
threats to species classified as at risk and secure,
respectively. The pattern of threats to species at risk did
not differ noticeably from the threats to species classified
as secure (Figure 2). Moreover, the pattern of threats
Table 1. Percentage of species at risk in different taxonomic groups for species that are found with Spotted Owls (n=138) and that are found
throughout British Columbia (n=2892). British Columbia’s Red and Blue lists contain “critically imperiled”, “imperiled” and “vulnerable”
species. COSEWIC: E = Endangered, T=Threatened, SC=Special Concern. SARA Schedule: 1=protected under law, Schedule 2 or 3 = awaiting assessment.
N o. of
Species
Taxon
B.C.1
COSEWIC2
SARA Schedules3
% Red or Blue
% E or T
% SC
%1
% 2 or 3
Amphibians
4
50
25
50
50
0
Fish
16
17
25
25
13
0
Mammals
17
24
0
6
0
12
Birds
35
17
9
14
11
0
Vascular Plants
66
8
0
0
0
0
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TAXON
Figure 1. Conservation status of species associated with late-successional and old-growth forests within the range of the Northern
Spotted Owl in British Columbia, Canada. A=Amphibians, F=Fish, M=Mammals, B=Birds, VP=Vascular Plants.
varied little among taxonomic groups. Timber harvesting
was the most common single threat for amphibians (50%
of 6 threats), birds (38% of 40 threats), vascular plants
(40% of 20 threats), and fish (14% of 35 threats), whereas
human disturbance was the most common threat for
mammals (28% of 18 threats).
data that further timber harvesting of old-growth and latesuccessional forests will negatively affect many species
at risk in this biological community, not only Spotted Owls.
Current conservation planning for old-growth and latesuccessional forests of the southwestern mainland of
British Columbia is focused on Spotted Owls (Chutter et
al. 2003). Seven of the other 21 species at risk are listed
under Schedule 1 of SARA and thus legally required to
have critical habitat defined and protected. The strategy
of habitat conservation being employed for recovery of
Spotted Owls is to use current owl locations to define
particular habitat types and specific areas for protection,
as well as to prescribe particular timber harvesting
techniques to conserve, enhance or create habitat for
owls (Chutter et al. 2003). Notwithstanding considerable
uncertainty about the efficacy of these practices for owls
(Barlee et al. 2005), the results of this study demonstrate
that it is also important to know whether this focal species
approach and associated actions also serve the
conservation needs of other species in this community
and what should be done to improve conservation
planning for all these species at risk.
DISCUSSION
Our analysis reveals that 24% of a sample of vertebrates
(amphibians, birds, mammals, fish) and 8% of vascular
plants in old-growth and late-successional forests of
southwest British Columbia are at risk of extirpation or
extinction. These species at risk are broadly distributed
across the taxonomic groups and trophic levels of this
community. Only 50% of these species have been
assessed by COSEWIC and therefore eligible for some
protection under Canada’s new Species at Rick Act, and
only 36% are being managed by B.C. The causes of
decline and threats to these species are varied, but timber
harvesting, road building, and forest fragmentation are
the most common. Our analysis showed that the
proportion of these co-occurring species at risk was not
significantly different from the province-wide level of
species at risk. In fact, vascular plants in this community
appear to be at relatively little risk compared to the rest
of British Columbia. However, it is possible that formal
assessments of conservation status may be biased
towards particular taxonomic groups, and, as our list of
co-occurring species did not derive from fieldwork in
British Columbia, some species may have been
erroneously omitted or included. It is almost certain that
some ecologically important taxa are missing from our
list. For example, although non-vascular plants were not
considered (Harper and Milliken 1994), data from
Washington and Oregon reveal hundreds of these species
are dependent upon the forests utilized by Spotted Owls
(Thomas et al. 1993). The pervasive pattern of
endangerment in the taxa we have examined suggests
that many of these unexamined species are also at risk,
albeit undocumented. Nonetheless, resolving these
questions could not alter the main conclusion from these
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FOCAL SPECIES CONSERVATION PLANNING
The theory of conservation biology suggests that focal
species that serve as Indicators, Keystones, Umbrellas,
or Flagships in conservation planning can indirectly
benefit other species. An ideal indicator species is easy
to monitor, quickly reflects changes in the physical
environment, and exhibits population changes
corresponding to those of unmonitored species, thereby
revealing the general efficacy of conservation practices
(Simberloff 1998). The northern Spotted Owl fails
as an ideal indicator species for several reasons: its
breeding biology, life history and demography make
it slow to respond (Rosenberg et al. 2003, Forsman
et al. 2003) and thereby indicate effects of environmental changes and management experiments; its recent
population decline has eliminated it from many formerly
occupied sites, so its distribution no longer encompasses
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as an umbrella species, but there are several reasons to
other species at risk; and if, as seems likely, critically
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Y
doubt
the co-occurring species
imperiled populations of Spotted TOwls
arePsubject
in this study have been identified as denizens of old-growth
interventions (e.g. food supplementation, translocations and
forest, their specific habitat requirements are unlikely to
habitat alterations), then natural correlations to other species
exactly coincide with those of owls. For example, several
will be broken. Second, management of keystone species
of the threatened co-occurring species are aquatic.
(that have ecological effects on numerous other species
Protected reserves created to meet the habitat needs of
above what would be expected from their abundance and
Spotted Owls will not provide well for these species or for
biomass) can link focal species and ecosystem
anadromous fish that are central to the ecology of this
management by maintaining a species that has overriding
region (see Franklin 1994). Also, to the extent that the
effects on other species and trophic levels (Davic 2003).
distribution of populations across the landscape is patchy,
Although Spotted Owls are large predators near the top of
remaining locations of other species at risk are unlikely to
their food chains they apparently do not have community
wide impacts indicative of a keystone species (Forsman et
Logging in the Nesikep Creek area near Lillooet, BC. In 2002 this area
al. 2001, Rosenberg et al. 2003). Third, an umbrella species A,
was home to one of BC’s last remaining Spotted Owls. Since logging occurred
is one that has such extensive habitat requirements and the Spotted Owl has vacated this territory and has not been relocated; B,
of the Lillooet logging; C, Spotted Owl biologist Andy Miller, former
large area requirements that protection of its habitat will Detail
member of the BC Spotted Owl Recovery Team, surveys recent logging; D,
incidentally protect sufficient suitable habitat for other This huge red cedar, located in Spotted Owl habitat in the lower mainland of
Columbia (B.C.), is a majestic representative of the massive cedars
threatened species (Lambeck 1997, Roberge & Angelstam British
that once populated the B.C. coast (Photos by Jeremy Sean Williams,
2004). The northern Spotted Owl is on occasion mentioned 2003, courtesy of WCWC).
A
B
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Figure 2. Number of instances of different threats to survival for species that co-occur with northern Spotted Owls in British Columbia, shown
separately for species classified as secure and those at risk by the B.C. Government (see Methods for full names of threats)
B.C. Timber Sales
logging in a Spotted
Owl Special Resource
Management Zone in
the heart of the
Manning/Skagit Park
complex. This recent
logging took place in
the immediate vicinity
of
A an historic Spotted
Owl site, and within the
home range of one of
B.C.’s last remaining
pairs of Spotted Owls
(Photo by Joe Foy,
2004 courtesy of
WCWC)
correspond to remaining populations of Spotted Owls.
Moreover, these species at risk represent several major
taxonomic groups and empirical results from several
other communities in which detailed distributional data
are available for many different taxa demonstrate that
protection of sites for one species or taxonomic group
rarely provides adequate protection for the other taxa
excluded from the site selection process (reviewed by
Roberge & Angelstam 2004). We hypothesize that the
same pattern of weak protection for co-occurring species
will derive from habitat protection designed solely for
Spotted Owls in British Columbia. To do otherwise
requires that the species at risk spatially co-occur on the
landscape at the scale of protected areas. A valuable
research project would be to assess the sites proposed
for protection of Spotted Owl habitat in
B.C. to determine the
occurrence of other
species at risk and
hence the more general value of these
sites for conservation
of this community.
easily understood and which inspire widespread public
sympathy and action (Bowen-Jones & Entwistle 2002).
Our analysis reveals a group of potential flagship
species from which to select a representative species
for old-growth forests in British Columbia, though we
believe that by its size and diversity this group also
shows that expansion of the flagship concept is
appropriate. The nautical roots of flagships trace back
to ancient Athenian mariners who hoisted various flags
as signals to other members of the fleet. Over time, as
maritime tactical complexity increased so did flag
signals. For example, by 1691 British Royal Navy
flagships were using a combination of 11 flags
displayed at 9 different points to convey at least 45
fixed messages. By analogy, in the discipline of
Conservation Biology a single flagship species may
not always be adequate to convey the extent of
conservation needs and the scale of conservation
actions required. Indications are that public
understanding of the scale of conservation needs and
tactical responses is also increasing (e.g. Suzuki &
Dressel 2002) and it is the public at which flagships
are signaling. Thus, we propose a new, larger flagship
fleet for the old-growth forests of British Columbia.
This flagship fleet is not only truer to the maritime
roots of flagship signaling, it better conveys the
community-wide pattern of conservation needs. Our
proposed fleet contains a bird (Northern Spotted Owl),
a frog (Coast Tailed Frog), a mammal (Fisher) and a
plant (Redwood Sorrel) and has 18 other species
available at present (Appendix 1). This new flagship
fleet conveys the message that many species in this
community are at risk and conservation actions must
be directed at the full community.
A FLAGSHIP FLEET
A fourth role that a
single focal species
has served in conservation planning is
as a flagship species, a compelling
species whose lifestyle, and plight are
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Owl management plan. Strategic component. B.C. Ministry of Forests
and Ministry of Environment Victoria, B.C.
SOMIT (Spotted Owl Management Inter-Agency Team) 1997b. Managing
Spotted Owl habitat: Operational Guidelines Component of the Spotted
Owl Management Plan. B.C. Ministry of Forests and Ministry of
Environment Victoria, B.C.
Suzuki, D., and H. Dressel 2002. Good news for a change: how everyday
people are helping the planet. Greystone Books, Vancouver.
Thomas J., M.G. Raphael, R. Anthony, E.D. Forsman, A. Gunderson, R.
Holthausen, B. Marcot, G. Reeves, J. Sedell, & D. Solis. 1993. “http:/
/pnwin.nbii.gov/nwfp/viability/”. Viability Assessments and
Management Considerations for Species Associated with LateSuccessional and Old-Growth Forests of the Pacific Northwest. U.S.
Department of Agriculture.
Finally, a flagship fleet also suits the first of the three
central tasks of conservation reserve planning (Noss
1996, Noss et al. 1999) mapping the locations of special
elements for conservation. As outlined by Noss et al.
(1999), Special Element Mapping involves identifying
the locations of threatened species and geographical
formations that require protection. For old-growth
forests in the southwestern mainland of British
Columbia we can point to 22 threatened species that
could be mapped. The second step is to mesh these
special element locations with the output from a
separate mapping of habitat types that serves to
proportionately protect the full variety of habitat types.
Finally, the third track of conservation reserve planning
focuses on the spatial arrangement of conservation
areas and explicitly considers species whose
movement and life history make them most dependent
on large areas. The northern Spotted Owl and fisher
are appropriate for this analysis. Our ultimate
conclusion is that such a design process for
conservation areas of old-growth forest in the
southwestern mainland of B.C. can prevent the flagship
fleet and its associated vessels from sinking, that is
this process can prevent well-known species and their
little-known and unknown co-occurring species from
going extinct.
ACKNOWLEDGMENTS
We thank our dedicated colleagues Andy Miller and
Michelle Connolly for assistance with data collection,
Derek Martin, Ann Rowan, Cheri Burda, Panos Grames
and Gwen Barlee for discussions, and Devon Page for
helping us understand legislation. Photographs appear
courtesy of the Western Canada Wilderness Committee
and were previously published in the report “In Defense
of Canada’s Spotted Owl” (Barlee et al. 2005).
NOTES
Appendix 1 is available at: http://www.davidsuzuki.org/
WOL/Publications.asp Spotted Owls in Washington. Journal
of Raptor Research 35:141-150.
REFERENCES
Barlee, G., D. Page, & A. Miller. 2005. In Defense of Canada’s Spotted
Owl. 2005. Western Canada Wilderness Committee and Sierra Legal
Defense Fund. Vancouver, B.C. “http://www.wildernesscommittee.org”.
Blackburn, I.R. & S. Goodwin. 2003. Status of the northern Spotted Owl
in British Columbia. Ministry of Water, Land and Air Protection. Surrey, B.C.
Bowen-Jones, E., & A. Entwistle. 2002. Identifying appropriate flagship
species: the importance of culture and local contexts. Oryx 36:189195.
Chutter, M., I. Blackburn, D. Bonin, J. Buchanan, B. Costanza, D.
Cunnington, A. Harestad, T. Hayes, D. Heppner, L. Kiss, J. Surgenor,
W. Wall, L. Waterhouse, & L. Williams. 2003. National recovery strategy for the Northern Spotted Owl. Report. Recovery of Nationally Endangered Wildlife Committee, Ottawa, ON.
COSEWIC 2000. COSEWIC assessment and update status report on the
Northern Spotted Owl Strix occidentalis caurina in Canada. Committee
on the Status of Endangered Wildlife in Canada. Ottawa. vii + 14 pp
Crooks, K. R., & M. E. Soulé. 1999. Mesopredator release and avifaunal
extinctions in a fragmented system. Nature 400:563-566.
Davic R. D. 2003. Linking keystone species and functional groups: A new
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Received: 20 September 2005; Accepted: 5 January 2006
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