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From the Field: Implementing recovery of the red wolf - Integrating research scientists and managers

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The United States Fish and Wildlife Service (USFWS) developed guidelines for the composition and role of endangered species recovery implementation teams, but few teams have been established and their success has not been evaluated. Using the recovery program of the red wolf (Canis rufus) as a model, we describe the genesis, function, and success of the Red Wolf Recovery Implementation Team (RWRIT) in helping guide the establishment of a viable red wolf population in eastern North Carolina. In operation since 1999, the RWRIT meets bi-annually to review USFWS progress and provide recommendations aimed at maximizing success of species recovery. The team is comprised of 8 research scientists from disciplines including population genetics, canid ecology, population ecology, veterinary medicine, and captive management. Representation from each of these disciplines is deemed necessary for proper evaluation of recovery progress and assessment of future needs. Meeting attendance by the USFWS field management team ensures both proper reporting of past progress and future implementation of management recommendations. Over time, RWRIT members have assumed specific assignments for data analyses, further contributing to the recovery effort. Through the combined efforts of the USFWS field team and the RWRIT, the threat of introgression of coyote (Canis latrans) genes into the red wolf population has been substantially curtailed within the recovery area, and red wolf numbers and range have increased. The RWRIT serves as an example of a recovery implementation team that is successfully incorporating the principles of adaptive management and whose template could be adapted to other endangered species.
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From the Field: Implementing recovery
of the red wolf— integrating research
scientists and managers
Michael K. Stoskopf
Karen Beck
Bud B. Fazio
Todd K. Fuller
Eric M. Gese
Brian T. Kelly
Frederick F. Knowlton
Dennis L. Murray
William Waddell
Lisette Waits
This paper is posted at DigitalCommons@University of Nebraska - Lincoln.
http://digitalcommons.unl.edu/icwdm usdanwrc/507
1145
From
the
Field:
Implementing recovery
of
the red
wolf
integrating research
scientists
and
managers
Michael
K.
Stoskopf,
Karen Beck,
Bud
B. Fazio, Todd
K.
Fuller,
Eric M.
Gese,
Brian
T.
Kelly, Frederick F. Knowlton, Dennis
L.
Murray,
William Waddell,
and
Lisette Waits
Abstract
The
United States Fish
and
Wildlife Service (USFWS) developed guidelines
for the com-
position
and
role
of
endangered species recovery implementation teams,
but few
teams
have been established
and
their success
has not
been evaluated. Using
the
recovery
pro-
gram
of the red
wolf (Canis rufus)
as a
model,
we
describe
the
genesis, function,
and suc-
cess
of the
Red Wolf Recovery Implementation Team (RWRIT)
in
helping guide
the
estab-
lishment
of a
viable
red
wolf population
in
eastern North Carolina.
In
operation since
1999,
the
RWRIT meets bi-annually
to
review USFWS progress
and
provide recommen-
dations aimed
at
maximizing success
of
species recovery.
The
team
is
comprised
of 8
research scientists from disciplines including population genetics, canid ecology, popula-
tion ecology, veterinary medicine,
and
captive management. Representation from each
of
these disciplines
is
deemed necessary
for
proper evaluation
of
recovery progress
and
assessment
of
future needs. Meeting attendance
by the
USFWS field management team
ensures both proper reporting
of
past progress
and
future implementation
of
management
recommendations. Over time, RWRIT members have assumed specific assignments
for
data analyses, further contributing
to the
recovery effort. Through
the
combined efforts
of
the USFWS field team
and the
RWRIT,
the
threat
of
introgression
of
coyote (Canis latrans)
genes into
the red
wolf population
has
been substantially curtailed within
the
recovery
area,
and red
wolf numbers
and
range have increased.
The
RWRIT serves
as an
example
of
a
recovery implementation team that
is
successfully incorporating
the
principles
of
adaptive management
and
whose template could
be
adapted
to
other endangered species.
Key words adaptive management, Canis
rufus,
endangered species, implementation, recovery,
red
wolf
Recovery of any endangered species is influ- ceed, it is equally critical that professionals tasked
enced by a range of political, economic, social, as with the responsibility for managing endangered
well as biological issues (Tear et al. 1993, Scott et al. species be able to move forward with timely deci-
1995,
Lundquist et al. 2002). Reconciling disparate sions based on practical management needs and
concerns and perspectives into a cohesive program scientific knowledge (Westrum 1994). The United
requires planning and decision-making processes States Fish and Wildlife Service (USFWS) red wolf
that consider conflicting interests of various stake- (Canis rufus) recovery program is an example of a
holders. However, for a recovery program to sue- program faced with complex issues, where man-
Wildlite Soiietv Bulletin 2005, J3l3i-1145 II32 Peer edited
1146 Wildlife Society Bulletin 2005, 33(3):1145-1152
agement successes have
been strengthened and
accelerated by integrating
active adaptive manage-
ment with careful and
timely scientific inquiry.
This paper describes how
this integration is being
achieved via a designated
"recovery implementa-
tion team."
The red wolf is an
endangered species that
once roamed an extensive
range including the south-
eastern United States, and
possibly the entire wood-
lands of eastern North
America (Wilson et al.
2000,
Nowak 2002,
Grewal et al. 2004).
Although listed as endan-
gered in 1967 (USFWS,
1967),
population decline
and apparent hybridiza-
tion with coyotes (Canis
latrans) were recognized
in the early 1960s
(McCarley 1962,McCarley
and Carley 1979). The remaining red wolves were
removed from the wild in the mid- to late 1970s
with the goals of establishing a captive breeding
program and eventually restoring captive-bred ani-
mals to portions of their historical range (U.S. Fish
and Wildlife Service [USFWS] 1989). In 1987 the
first red wolves were released in easternmost North
Carolina (Figure 1) with the plan to establish a
viable population (Parker 1987). The reintroduc-
tion efforts faced a myriad of social, political, and
biological issues as the Red Wolf Recovery Plan
(USFWS 1989) was implemented (Henry and
Lucash 2000, Phillips et al. 2003).
Although the reintroduction area was initially
considered uninhabited by coyotes, by the mid-
1990s it was apparent coyotes had infiltrated the
area and hybridization with red wolves was recur-
ring (Phillips et al. 2003). Due primarily to the
renewed hybrid threat and termination of the rein-
troduction of red wolves into Great Smoky
Mountains National Park (Henry 1998), the USFWS
decided it needed to re-evaluate its red wolf recov-
ery effort in light of what had been learned over
Figure 1. Changes in management zone boundaries within the Red Wolf Recovery Area of
eastern North Carolina, as made in accordance with Red Wolf Adaptive Management Plans.
The boundaries of the original management Zones 1, 2, and 3 (dashed lines) were first estab-
lished in April 2000. In March 2002, as red wolf recovery proceeded, boundaries in the south-
ern parts of the zones were moved west (solid lines); part of Zone 2 became Zone 1, while part
of Zone 3 became Zone 2 (arrows). In August 2003 some management aspects of canids
(i.e.,
sterilization vs. euthanasia) captured in the eastern half of Zone 3 (thin dotted line) began to
follow guidelines applied to Zone 2.
the previous decade. A key step in this review
process involved a Population and Habitat Viability
Assessment (PHVA) organized by the USFWS in
1999 and facilitated by the Conservation Breeding
Specialists Group of the World Conservation Union,
Species Survival Commission (IUCN SSC) (Kelly et
al.
1999). The diverse assemblage of attendees, rep-
resenting a variety of expertise and interests,
agreed that introgression of coyote genes into the
red wolf population was the principal threat to
recovery success (Kelly et al. 1999). The group also
recognized this issue required urgent attention
before hybridization became so pervasive as to vir-
tually ensure the genetic swamping of the only
extant free-ranging population of red wolves.
However, 2 views of how to address the hybrid
threat emerged from the PHVA; one believed
research was integral to addressing the problem,
and the other expressed concern that research
efforts would distract from the primary goal of
maintaining the only free-ranging population of red
wolves in the world. A consensus agreement was
reached on this debate and resulted in an overarch-
From the Field Stoskopf et al. 1147
ing workshop statement, including:
"...our primary recovery focus must be protect-
ing and promoting the growth of a self-sustain-
ing, non-hybridizing population of red wolves
in the wild and sustaining an active captive
component. Actions to be taken will use an
adaptive management approach that will not
compromise the ability to achieve this goal."
(USFWS 1999:52)
This level of agreement among the diverse par-
ticipants of the PHVA set the stage for designing an
adaptive management plan (cf. Lancia et al. 1996)
that would reduce the threat of wolf-coyote
hybridization. This plan (Kelly 2000) diverged from
conventional endangered species management
because it involved an incremental process tailored
to modify field protocols according to past success
in eliminating the threat of hybridization.
Specifically, it required the release area to be segre-
gated into several defined management zones, each
managed to provide an integrated optimization of
risk reduction •within the resource limitations avail-
able to the project (Figure 1). As nonwolf canids
were removed from given zones and replaced with
red wolves, management options could be adapted
by modifying zone boundaries or adjusting specific
management protocols.
Adoption of this plan, requiring frequent re-eval-
uation of data and attendant management adjust-
ments, spawned close interactions between
USFWS field biologists and scientists with back-
grounds relevant to the work being undertaken. A
Red Wolf Recovery Implementation Team (RWRIT)
was formed to advise USFWS as they implemented
the adaptive management plan; this team was cre-
ated pursuant to Section 4(f)(2) of the amended
Endangered Species Act (ESA), which authorizes
the Secretary of the Interior to procure the servic-
es of appropriate public and private agencies, insti-
tutions, and other qualified persons to help imple-
ment endangered species recovery plans. Other
USFWS-designated species-specific implementa-
tion teams, as opposed to planning teams, have
been formed (e.g., black-footed ferret [Mustela
nigripes], northern right whale [Eubalaena
glacialis], Okaloosa darter [Etheostoma
okaloosae], and southern sea otter [Enhydra
lutris]; USFWS files), but they are rare and no for-
mal description of one's workings or success has
yet been documented.
Recovery implementation team
composition
Selection of the RWRIT scientists and their lead-
ership was important to the success of implement-
ing and evaluating the adaptive management plan.
The PHVA helped the USFWS identify individuals
with the combined expertise and personality con-
sidered important in a functional RWRIT. The PHVA
also provided insight to the breadth of expertise
needed over the long term. This expertise included
such diverse fields as systematics, genetics, popula-
tion modeling, health management, and canid biol-
ogy, behavior, ecology, and management. Social sci-
entists were not required in this case because those
issues were, and continue to be, successfully dealt
with by the USFWS field management team in con-
junction with non-governmental organizations
(Henry and Lucash 2000). Direct experience with
the red wolf was not a requisite criterion for
RWRIT membership. In fact, due to the long and
controversial scientific history of the red
wolf,
some team members were sought for their naivete
of red wolves to minimize preconceived notions
regarding the problems the adaptive management
plan addressed. Thus, a mixture of experienced and
young research scientists •with strong records of sci-
entific productivity and interpersonal skills was
selected. Each member of the RWRIT had to be
willing to use a data-driven approach to decision-
making while remaining open to challenges of
interpretation. Each member also had to be willing
to accept group decisions as well as devote consid-
erable personal time toward solving issues associat-
ed with the red wolf program.
The RWRIT needed to be large enough to pro-
vide the scientific diversity needed to assess the
broad range of critical issues, but small enough to
support close working relationships among mem-
bers and result in productive meetings (Clark and
Westrum 1989). A basic philosophy was that if the
RWRIT needed expertise from individuals or disci-
plines outside the RWRIT to address specific issues,
guest scientists would be invited to participate in
the appropriate meetings. Initially, a goal of 8 mem-
bers and 4 alternates was considered. Interactions
of the group and reliability of participation in early
meetings were used to identify the core members
of the RWRIT. Since then, the size and composition
of the RWRIT (8 members, no alternates) has
worked well, sustaining effective decision-making
with absences at meetings being rare. The leader of
1148 Wildlife Society Bulletin 2005, 33(3):1145-1152
the RWRIT needed to moderate meetings efficient-
ly while allowing for creative interactions among
RWRIT members. To ensure this, a senior scientist
at a local university was selected due to his demon-
strated scientific and leadership skills.
Experienced and stable field team
The USFWS field team involved in the day-to-day
operation and management of the red wolf recov-
ery program was key to the success of the RWRIT.
The field team attended all RWRIT meetings as non-
voting members and provided the necessary data
and expertise for the meetings to progress effec-
tively. This distinction between the teams initially
caused some anxiety, but this subsided once roles
had been fully elucidated. The field team is remark-
ably stable and has worked cohesively on the red
wolf project for many years (Phillips et al. 2003).
Scientists of the RWRIT recognize the field team as
the most experienced red wolf biologists and
essential for successful functioning of the RWRIT
itself.
In turn, the field team's willingness to listen
to and implement recommendations made by the
RWRIT has been a critical factor in the success of
the program. Open communications between the
2 teams keeps RWRIT scientists aware of the imple-
mentation of recommendations and fosters respect
for the dedication of the field team.
Getting started
The first meeting of the RWRIT was important in
establishing the tenor of group interactions and
future functioning. Subsequent meetings would
focus on examining data related to specific ques-
tions within an established agenda, but the first
meeting focused on developing operating proce-
dures for decisions as well as the types of data and
data formats the team preferred for review and
evaluation. This was a step that helped acquaint
members of the team and recognize proper work-
ing protocols. It also ensured that all members of
the team had a common understanding of the Red
Wolf Adaptive Management Plan (RWAMP).
The charge of the RWRIT was established a pri-
ori by the Team Leader of the Red Wolf Recovery
Program (i.e.,"Red Wolf Program Leader"). This task
was defined specifically as reviewing progress on
the RWAMP and recommending changes to the
plan based on data provided by the USFWS. As the
team gained experience, this charge evolved to
include recommendations for data relevant to
answering specific questions important to the field
team in the day-to-day management of the wild red
wolf population. The 2 charges were closely relat-
ed, frequently blended, seldom distinct, fundamen-
tal to the Adaptive Management Paradigm (Walters
1986),
and are the responsibilities that drive efforts
of RWRIT members. From the beginning, per the
ESA, RWRIT recommendations were strictly adviso-
ry, with decisions for implementation being at the
discretion of the USFWS.
Ground rules established in the first meeting
have rarely been adjusted. Some established the
mechanics of operations. For example, it was
decided a minimum of 6 RWRIT members would
be required as a quorum for a functional meeting.
Failure to achieve quorum would trigger an evalua-
tion by the RWRIT Leader and the Red Wolf
Program Leader to assess whether the RWRIT
remained an appropriate mechanism. To date this
has not been necessary due to continued strong
and enthusiastic attendance.
Other rules provided guidance for RWRIT inter-
actions. To reduce stifling potentially meritorious
but perhaps unconventional ideas, the team adopt-
ed a basic rule indicating that speakers must pres-
ent alternative solutions when challenging or
negating a proposed idea or approach. Ideas would
be withdrawn from consideration only after careful
efforts to refine them failed to produce workable
solutions. To the fullest extent possible, data would
be used to support all positions.
Other procedural mechanisms established in the
first meeting have had a beneficial effect on RWRIT
operations. For example, tentative dates, times, and
location of future meetings are established jointly
early in the agenda of each meeting. In addition,
the agenda of the next meeting is established near
the completion of the current session, which prob-
ably produces a more dynamic agenda than a call
just before the meeting. Opportunities to add agen-
da items at any time remain, but the draft of the
agenda appears in the final minutes; serving as a
reminder for participants as they prepare for the
coming meeting.
An important activity reserved for the end of
each meeting is an exercise in prioritizing "action
items",
which are further classified as either "tasks"
(expected to be accomplished within the time
frame of the meeting or between meetings);
"proj-
ects"
(longer duration activities); or "manuscripts"
(the drafting of information for publication).
From the Field Stoskopf et al. 1149
Individual RWRIT members are recognized as
responsible for addressing each item. With many
issues to consider and an active agenda, many more
action items are identified than can typically be
accomplished with the resources available. The
action items established throughout the meeting
and recognized as "projects" are assembled in a
descriptive list and as a final exercise, each member
of the RWRIT assigns a priority level to each item
and the mean rating is computed. This rating is
offered to the Red Wolf Program Leader as a rec-
ommendation for activities to pursue or fund. At
the first meeting, a pattern was established where
RWRIT members worked to identify key manage-
ment questions and to focus scientific inquiry in
areas of need with constant reference to the adap-
tive management plan. Assets are identified and
resource limitations discussed so recommendations
have a reasonable likelihood of implementation.
Short proposals outlining the objective of projects
and the team member(s) involved in the work are
distributed to the RWRIT via the team's webpage.
The webpage also includes team member contact
information, minutes of meetings (see below), data
sets,
reports, press releases, publications, project
descriptions, manuscripts in progress, and upcom-
ing meeting agendas and related materials such as
reports and summaries.
Since 2000 the RWRIT has met bi-annually, which
is sufficient to respond in a timely manner to ques-
tions from the field and to strengthen collegial
bonds among members. This schedule also allows
sufficient time for the field team to implement rec-
ommendations and to document their progress and
for RWRIT members to work independently on
action items. Other factors affecting meeting
schedules include a need to make recommenda-
tions ahead of budget deadlines and to accommo-
date schedules of the individual RWRIT members
and the field team. The current pattern of meetings
includes 1 meeting in March prior to the denning
season and a second in October prior to intensive
trapping efforts.
cies associated with updating new attendees. The
ability to invite experts in areas not represented on
the RWRIT provides a mechanism to maintain flex-
ibility and adaptability. Periodic review of expertise
needed for specific tasks and projects of the RWRIT
keeps the issue of change before the team. In addi-
tion, there exists ample opportunity to discuss can-
didly both the pros and cons of the teams' efforts,
either formally at the end of each meeting or infor-
mally during meals or after hours. The RWRIT
Leader needs to recognize dissenting views and
address contentious issues promptly and effective-
ly. The fact that for most meetings the entire RWRIT
was communally housed in rented accommoda-
tions further ensured the establishment of favor-
able personal relationships benefiting RWRIT inter-
actions and discussion.
Complete minutes of RWRIT deliberations pro-
vide documentation of the team's discussions and
recommendations. An iterative process of editing
minutes is used by the RWRIT, ensuring important
information developed at each meeting is recorded
accurately and in language deemed appropriate by
the participants. Notes are converted into a draft
each evening and individualized, and printed
copies are distributed to attendees the following
morning for editing. All drafts are synthesized into
the penultimate draft for further comment, which is
followed by a final draft distributed electronically
shortly after completion of the meeting. The
RWRIT members have a week to return any cor-
rections, after which the final minutes are complet-
ed and distributed electronically. The deliberations
of the RWRIT are considered privileged communi-
cation, and all meeting participants are asked to
limit discussions of information received at the
meetings to individuals within their respective
research groups. This policy allows RWRIT mem-
bers access to sensitive and preliminary data and
provides more freedom of discussion without con-
cerns about inappropriate disclosure. Distribution
of the minutes beyond the RWRIT is at the discre-
tion of the Red Wolf Program Leader.
Staying flexible
Any group with dynamic tasks needs a mecha-
nism for adjusting the nature of the group as it
matures and as tasks change (Clark and Reading
1994).
The concept of alternate members soon was
abandoned because of the strong attendance by
RWRIT members and because it reduced inefficien-
How well does it work?
The test of any system is how well it functions to
meet the goals and objectives of the program it
serves. In the 4 years since the first formal meeting
of the RWRIT, key challenges to implementing the
plan developed at the PHVA have been identified
and strategies have been devised to provide practi-
1150 Wildlife Society Bulletin 2005,33(3):1145-1152
cal solutions and evaluate success of recovery
efforts. Perhaps more importantly, all RWRIT mem-
bers and the entire red wolf field team have
become close colleagues who look forward to each
meeting. We enjoy the frank and open exchange of
ideas,
the ability to quickly address both practical
and theoretical problems and make changes in
management practices, and the successes in the
field that result from the collaboration. The details
of these changes and successes are the basis of sev-
eral scientific papers, some already published or in
press and others currently in preparation, but a
brief summary is warranted.
Prior to 1998 all canids captured in the red wolf
recovery area were assumed to be wolves unless
they were so small as to be considered coyotes, if
they were black, or if they looked part dog. If there
was some indication that a single female wolf was
consorting with a coyote or dog, pups she pro-
duced were removed (A. Beyer, USFWS, personal
communication). Thus, the basic challenge of rap-
idly and confidently identifying animals as red
wolves versus hybrids or coyotes, especially young
animals, was identified early as a key concern of the
PHVA and the field team. The RWRIT served as cat-
alyst for developing an enhanced and improved
genomic testing protocol by expanding the ability
to assess alleles at 19 loci (Miller et al. 2003). A pri-
ority placed on obtaining genomic assessments of
the entire group of founders in the captive breed-
ing program, as well for coyotes in the vicinity of
the wolf release zones, greatly improved the confi-
dence in the genomic data now available. Genetic
analyses were integrated with pedigree and mor-
phometric data to develop decision trees for all
captured animals (Table 1). Extension of the DNA
analysis capabilities to fecal samples increased the
potential for assessing presence of red wolves, as
well as undesired non-red wolves, in the field sam-
ples without the need of capturing and handling
animals (Adams et al. 2003). Additional research
efforts were directed at using this technology for
assessing red wolf population size (J. R. Adams and
L.
P.
Waits,
University of Idaho, unpublished data).
To evaluate progress of the adaptive manage-
ment plan, RWRIT scientists wanted detailed and
current descriptions of animal locations, their geno-
types,
and canid inventory efforts in relation to geo-
graphic areas. A coordinated Geographic
Information System (GIS) database system is now
used at all RWRIT meetings to examine recovery
progress. This is steadily approaching the goal of a
real-time data view as data entry and validation
challenges are addressed and data summaries are
refined. These tools help identify areas where data
are insufficient to define the status of canids and
help develop strategies to eliminate so-called "areas
of ignorance" by concentrating efforts in areas
needing more attention. In addition, they have lead
to improved ground telemetry efforts and more
efficient use of resources and personnel.
Modeling effects of coyote genomic intrusion,
using more refined data sets and newer models
than available at the PHVA, provided RWRIT scien-
Table 1. Decision path for genetic results of red wolves (RW) captured in the experimental population area in northeastern North
Carolina, applied in fall of 2003 (explanation of genetic result classifications given in Miller et al. 2003). Decision parameters list-
ed in the following priority: Genetic testing; Pedigree; Morphology; Mate.
Capture location8
Decision parameterZone 1
Release
Consider pedigree (go to
Release
Consider morphology (gc
Euthanize
Consider mate (go to 4)
Release
Euthanize
Consider pedigree (go to
2)
>to 3)
5)
Zone 2
Release
Release
Sterilize
Release
Sterilize 1 mate
1.
Genetic test: 100% RW (pedigree 100% RW)
1.
Genetic test: 100% RW but cannot exclude 75% RW hypothesis
2.
Pedigree is 100% to 87.5% RW
2.
Pedigree is 87% to 75% RW or unknown
3. Morphologically "hybrid-like"
3. Morphologically "RW-like"
4.
Mate is >75% RW
4.
Mate is <75% RW or uncertain
1.
Genetic Test: 75% RW or 75% RW but cannot exclude 50% RW
hypothesis
5. Pedigree is <75% RW
5. Pedigree is >75% RW or unknownEuthanize Sterilize
Consider morphology (go to 3)
a See Figure 1.
From
the
Field
Stoskopf
et al. 1151
tists
new
insights into impacts
of
genomic intrusion
(e.g., Miller
et al.
2003). This allowed
for key
insights
to
establishing acceptable risks defined
in
the decision trees. This also assisted
in the
making
of informed recommendations
for
modifying
approaches
to the
various management zones
for
red wolf recovery. Recently,
the
RWRIT initiated
an
effort
to
conduct detailed analyses
of
home range,
spatial interactions, habitat
use, and
demographic
attributes
of all
radiomonitored
red
wolves since
1986,
with
the
objective
of
developing
a
population
viability model
to
help guide future management
and recovery actions.
Den
management techniques
via implementation
of
early genomic sampling
and
use
of
cross-fostering
of
wild-caught
and
captive
bred pups into wild litters have been developed
(cf. Kitchen
and
Knowlton,
in
press). Methodology
also
has
been enhanced
to
conduct surgical proce-
dures
to
support
the use of
hormonally intact
but
sterile hybrids
and
coyotes
to
serve
as
sterile
buffers (i.e., temporary territory placeholders that
discourage establishment
of
new, intact nonwolves)
in peripheral management zones (Figure
1).
The
net
result
of
such activities
has led to an
increase
in the
area occupied
by red
wolves, total
number
of red
wolves,
and
number
of red
wolf
social units,
as
well
as a
major decrease
in the
total
area where
the
status
of
canids,
in
general,
is
unknown
(B. B.
Fazio, USFWS, unpublished data).
Such changes
in
these metrics were identified
in
the RWAMP
as
key indicators
of
the successful man-
agement
of
wolf-coyote hybridization. Importantly,
coyotes
or
hybrids have essentially been eliminated
from fully half
of the red
wolf recovery area.
To
date,
genetic intrusion into
the red
wolf population
has been largely controlled, albeit through aggres-
sive intervention.
The effective functioning
of the
RWRIT
has
ensured that issues identified
at the
PHVA
as
described
in the
RWAMP have been,
or are
being,
successfully addressed
by
USFWS.
And as
should
be
expected,
the
original
red
wolf adaptive manage-
ment plan
is now
revised
to
include
5
years
of
evolving adaptive management (Fazio
et al.
2004).
The approach taken
by the
RWRIT represents
a
good example
of
successful application
of the
Adaptive Resource Management paradigm
and is
likewise consistent with,
and
respectful
of, con-
cerns raised
by the
participants
at the
PHVA that
the primary goal
of
conserving
the
only free-rang-
ing population
of red
wolves
not be
overshadowed
by
the
desire
to
conduct research. Indeed,
the
USFWS recently highlighted
the
efforts
of the Red
Wolf Recovery Program
in a
videotape
on how the
use
of
sound science
is key to
meeting
its
mission.
We believe
the
recent tangible success
in red
wolf recovery
is a
direct result
of
conducting
the
PHVA, crafting
a
RWAMP, establishing
the
RWRIT,
and
the
cooperation
and
close interaction between
the RWRIT
and the
USFWS field team directly
tasked with
red
wolf recovery. Endangered species
recovery should involve
a
strong linkage between
scientific investigation under
the
rubric
of
adaptive
management
and the
appropriate blend
of
social,
political,
and
economic issues (Clark
et al.
1994).
In
light
of the
mixed past success
in
recovering
endangered species
in the
United States (Crouse
et
al.
2002, Gerber
and
Hatch 2002), we believe, based
on
the
success
of the
RWRIT, that recovery imple-
mentation teams
can
serve
as an
effective vehicle
for helping guide recovery programs
and
actions.
Acknowledgments.
The
authors thank
the
USFWS
and the
Morris Animal Foundation
for
finan-
cial support
of
activities reported
in
this manu-
script
and the red
wolf field team
for its
excellent
efforts
and
cooperation with
the
RWRIT.
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Address
for
Michael
K.
Stoskopf and Karen Beck: Environmen-
tal Medicine Consortium, College
of
Veterinary Medicine,
North Carolina State University, 4700 Hillsborough Street,
Raleigh,
NC
27606,
USA;
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for
Stoskopf:
Michael_Stoskopf@ncsu.edu. Address
for
Bud
B.
Fazio:
Red
Wolf Recovery Program, Alligator River National Wildlife
Refuge, P.O. Box 1969, Manteo, NC 27954, USA. Address for
Todd
K.
Fuller: Department
of
Natural Resources Conservation,
University
of
Massachusetts, Amherst,
MA
01003-9285, USA.
Address
for
Eric
M.
Gese and Frederick
F.
Knowlton: United
States Department
of
Agriculture, Wildlife Services, National
Wildlife Research Center, Utah State University, Logan,
UT
84322-5295, USA. Address
for
Brian T. Kelly: Environmental
Medicine Consortium, North Carolina State University, 4700
Hillsborough Street, Raleigh, NC 27606, USA, and Red Wolf
Recovery Program, Alligator River National Wildlife Refuge,
P.O. Box 1969, Manteo, NC 27954, USA; present address: Eco-
logical Services, United States Fish and Wildlife Service, 4000
Airport Parkway, Cheyenne, WY
82001,
USA. Address for Den-
nis L. Murray: Department
of
Biology, Trent University, 1600 W.
Bank Drive, Peterborough, Ontario K9J 7B8, Canada. Address
for William Waddell: Point Defiance Zoo and Aquarium, 5400
North Pearl Street, Tacoma,
WA
98407, USA. Address
for
Lisette Waits: Department
of
Fish and Wildlife Resources, Uni-
versity
of
Idaho, Moscow,
ID
83844-1136, USA.
The collaborators
on
this paper have
a
variety
of
academic
degrees, work experiences, and publications
in
a wide selection
of fields and disciplines and have studied species from
A to Z.
They include (seated, from left
to
right) Karen
Beck
(ecological
epidemiologist), Buddy Fazio (fiscal conservationist), Todd
Fuller (quasi-experimental theoriologist), Eric Gese (investiga-
tive carnivologist),
and
Brian Kelly (politico-ecologist),
and
(standing,
left
to
right) Fred
Knowlton
(historical
cani-
dilist),
Dennis Murray (taxon-free numero-ecologist),
Michael Stoskopf (conservation metabonomist), Will
Waddell (ex-situ zoologist),
and
Lisette Waits (pan-
molecular faecologist).
... To thwart a presumed imminent extinction of red wolves caused by hybridization with coyotes (C. latrans) and small population size, the U.S. Fish and Wildlife Service (FWS) conducted intensive capture and removal efforts in the area during the 1970s, which led to the creation of a red wolf captive breeding (FWS, 2018c;Hinton et al., 2013;Hinton, White, Rabon, & Chamberlain, 2017;Stoskopf et al., 2005). Although application of an intensive placeholder strategy successfully mitigated coyote genetic introgression in the NEP to <5% (Gese & Terletzky, 2015), recent reviews questioned the longterm sustainability and necessity of this conservation action (FWS, 2018b;Wildlife Management Institute, 2014). ...
... Red wolf ancestry values for three and 13 individuals based on STRUCTURE and BAPS analyses, respectively, were within the range of that expected for 50% red wolf or F1 hybrid (i.e., q = 0.24-0.43; Bohling et al., 2013;Stoskopf et al., 2005). One individual had red wolf ancestry values of 0.78 from STRUCTURE and 1.00 from BAPS, the former of which was within the range observed for 75% red wolves or red wolf backcrosses, and the latter of which was representative of a pure red wolf (Bohling et al., 2013;Stoskopf et al., 2005). ...
... Bohling et al., 2013;Stoskopf et al., 2005). One individual had red wolf ancestry values of 0.78 from STRUCTURE and 1.00 from BAPS, the former of which was within the range observed for 75% red wolves or red wolf backcrosses, and the latter of which was representative of a pure red wolf (Bohling et al., 2013;Stoskopf et al., 2005). We note that STRUCTURE is more likely to detect admixture and correctly assign true ancestry, whereas BAPS is less likely to misclassify pure individuals as admixed hybrids (Bohling et al., 2013). ...
Article
Full-text available
Concerns over red wolf (Canis rufus) extinction caused by hybridization with coyotes (C. latrans) led to the capture and removal of remnant wild wolves from southwestern Louisiana and southeastern Texas, United States, during the 1970s. Here we show that despite decades of unmitigated hybridization, and declaration of endangered red wolves as functionally extinct in the wild, red wolf mitochondrial or nuclear DNA ancestry persists in ∼55% of contemporary wild canids sampled in southwestern Louisiana. Surprisingly, one individual had 78–100% red wolf ancestry, which is within the range for 75% red wolf, red wolf backcross, or putative red wolf, depending on estimation method. Our findings bolster support for designation of red wolves as a distinct species, demonstrate a critical need for the United States Government to consider adopting an existing but unimplemented hybrid policy, and suggest that immediate reassessment of canid management and taxonomic designation in southwestern Louisiana may be warranted.
... Low genetic diversity, increased genetic drift, and inbreeding depression can be significant issues whenever low numbers of breeding animals are involved, such as captive breeding programs (Rabon and Waddell 2010) or some wild populations (Brzeski et al. 2014), which can result in reduced litter sizes (Lockyear et al. 2009;Rabon and Waddell 2010). In addition, the expansion of coyotes (Canis latrans) into the northeastern North Carolina experimental population area and the risk of hybridization represented another threat to red wolf restoration (Kelly et al. 1999;Stoskopf et al. 2005;Fredrickson and Hedrick 2006;Bohling and Waits 2015). To mitigate introgression of coyote genes into the red wolf population, an adaptive management plan was implemented that 1) sterilized and released coyotes to serve as nonbreeding placeholders (Gese and Terletzky 2015), 2) removed coyotes from the area, 3) translocated red wolves (McLellan and Rabon 2006), and 4) cross-fostered captive-born pups into wild litters (Stoskopf et al. 2005). ...
... In addition, the expansion of coyotes (Canis latrans) into the northeastern North Carolina experimental population area and the risk of hybridization represented another threat to red wolf restoration (Kelly et al. 1999;Stoskopf et al. 2005;Fredrickson and Hedrick 2006;Bohling and Waits 2015). To mitigate introgression of coyote genes into the red wolf population, an adaptive management plan was implemented that 1) sterilized and released coyotes to serve as nonbreeding placeholders (Gese and Terletzky 2015), 2) removed coyotes from the area, 3) translocated red wolves (McLellan and Rabon 2006), and 4) cross-fostered captive-born pups into wild litters (Stoskopf et al. 2005). Combined with intensive field management and genomic testing protocols (Miller et al. 2003), wild-born pups could be identified as red wolf, red wolf-coyote hybrid, or coyote, leading to the option of replacing hybrid individuals with red wolf pups from the captive breeding population. ...
... Cross-fostering, the raising of young by nonbiologically related individuals, has the potential to increase population sizes of threatened or endangered species (Kitchen and Knowlton 2006;Stoskopf 2012). Fostering of offspring from natal to surrogate parents has been conducted with Columbian ground squirrels (Spermophilus columbianus- Murie et al. 1998), captive coyotes (Kitchen and Knowlton 2006), many bird species (Powell and Cuthbert 1993;Drewien et al. 1997;Oswald et al. 2013), and has been observed without human facilitation in black bears (Ursus americanus- Benson and Chamberlain 2006). ...
Article
Full-text available
Cross-fostering offspring with nonbiological parents could prove useful to augment populations of endangered carnivores. We used cross-fostering to augment captive-born and wild-born litters for the endangered red wolf (Canis rufus). Between 1987 and 2016, 23 cross-fostering events occurred involving captive-born pups fostered into captive litters (n = 8 events) and captive-born pups fostered into wild recipient litters (n = 15 events). Percentage of pups surviving 3 and 12 months was 91.7% for captive-born pups fostered into captive recipient litters. For pups fostered into wild litters, percentage of pups surviving 5 months was > 94% among fostered pups (pups fostered into a wild red wolf litter or replaced a hybrid litter), pups in recipient litters (wild-born litters receiving fostered pups), and pups in control litters (wild-born litters not in a fostering event) when using pups with known fates. Including pups with unknown fates as deaths, percentage of pups surviving 5 months was > 54% among fostered pups, pups in recipient litters, and pups in control litters. Among wild litters, percentage of pups surviving 12 months was > 82% among fostered pups, pups in recipient litters, and pups in control litters when using pups with known fates. Including pups with unknown fates as deaths, percentage of pups surviving 12 months was > 48% among fostered pups, pups in recipient litters, and pups in control litters. Although survival to 12 months was similar among the groups, average life span was different with pups in control litters living 3.3 years, pups in recipient litters living 4.6 years, and fostered pups living 5.6 years. Of fostered pups surviving > 12 months in the wild, 9 animals whelped or sired 26 litters. Cross-fostering was successful at augmenting litter size for red wolves without any deleterious effects on recipient litters, illustrating fostering as a tool for increasing populations of endangered carnivores.
... Annual tallies of red wolves fitted with radio-collars and pups counted at dens conducted by the USFWS Red Wolf Recovery Program (Recovery Program) peaked at 131 known individuals in 2001 and then fluctuated between 90 and 125 until 2014 (USFWS 2007(USFWS , 2014. Because the PHVA reported red wolf hybridization with coyotes to be the primary threat to recovery, the Red Wolf Adaptive Management Plan (RWAMP) was initiated in 2000 to prevent coyote introgression into the wild red wolf population (Stoskopf et al. 2005, Gese and Terletsky 2015. However, since 2004, the number of red wolves killed by gunshot increased approximately 2.75 times when compared to years prior to 2004 . ...
... Anthropogenic mortality and hybridization were identified as the 2 primary threats to red wolves (Kelly et al. 1999, USFWS 2007. Despite measures taken to address hybridization via the RWAMP (Stoskopf et al. 2005, Gese and Terletzky 2015, the USFWS has not addressed threats of anthropogenic mortality for red wolves. Ultimately, understanding how causes of red wolf mortalities change over time will allow the USFWS to respond with effective management to reduce excessive mortality and achieve population sizes essential to recovery. ...
... Standard errors represent unknown red wolves in the wild population. introgression via sterilization of coyotes paired with red wolves (Stoskopf et al. 2005, Hinton et al. 2013, Gese and Terletzky 2015. Regardless of whether coyote mates are fertile or sterile, congeneric pairings with coyotes represents lost reproductive effort by the red wolf population (Brzeski et al. 2014, Hinton et al. 2015a). ...
Article
Evaluating anthropogenic mortality is important to develop conservation strategies for red wolf (Canis lupus) recovery. We used 26 years of population data in a generalized linear mixed model to examine trends in cause-specific mortality and a known-fate model in Program MARK to estimate survival rates for the reintroduced red wolf population in North Carolina, USA. We found the proportion of mortality attributable to anthropogenic causes, specifically mortality caused by gunshot during fall and winter hunting seasons (Oct–Dec), increased significantly since 2000 and became the leading cause of red wolf death. Mortality rates were greatest for red wolves <4 years of age, and we suspect inexperience with human activities (e.g., hunting) likely caused younger wolves to be more susceptible to opportunistic killing by hunters. Since 1987, the red wolf population steadily grew and peaked at an estimated 151 individuals during 2005 but declined to 45–60 by 2016. To reduce the negative effects of anthropogenic mortality and ensure long-term persistence of red wolves, the United States Fish and Wildlife Service (USFWS) will need to re-implement previous long-standing and proven management practices (e.g., Red Wolf Adaptive Management Plan) on public and private lands and cease issuing take permits. The USFWS will also need to establish an effective management response to mitigate gunshot mortality through stronger regulation of coyote (Canis latrans) hunting and provide adequate ecologically and biologically supported regulatory mechanisms to protect red wolves. Finally, the USFWS should enhance recovery by providing information and education about red wolves to hunters and the general public.
... The red wolf "once roamed an extensive range including the southeastern United States, and possibly the entire woodlands of eastern North America" (Stoskopf et al., 2005(Stoskopf et al., , p. 1146. Due to anthropogenic factors (hunting, habitat destruction, economic development, etc.) and consequent red wolf population decline, the species was listed as endangered in 1967 and extinct in the wild in 1980. ...
... This version of the red wolf story is based on others told elsewhere (e.g., Adams et al., 2003;Bohling & Waits, 2011;Manganiello, 2009;Roth et al., 2008;Stoskopf et al., 2005) and I intentionally include specific terms repeatedly occurring in those accounts, such as threaten, fear, genetically swamped, hybrid swarm, pure. Let us focus on the words "pure" and "threat" as an example. ...
... For eastern wolves (Canis lycaon) and red wolves (Canis rufus), increased rates of human-caused mortality resulting in collapsed territories facilitates coyote (Canis latrans) encroachment and increased rates of hybridization between coyotes and wolves [3,14,[16][17][18]. Consequently, recovery of the critically endangered red wolf has been challenging because of its vulnerability to extinction via anthropogenic mortality and hybridization with coyotes [3,16,[19][20][21][22][23]. ...
... The goal of recovery efforts was to ensure that all Canis breeding pairs within the Recovery Area were red wolves [22]. To accomplish this, the Red Wolf Adaptive Management Plan was implemented to minimize hybridization by monitoring red wolf and coyote breeding pairs throughout the Recovery Area, and replacing coyotes and hybrids with red wolves until the Albemarle Peninsula was saturated with red wolf packs [20][21][22]. In this context, the presence and space use of transients has a profound effect on recovery of red wolves via the ability of transients to replace lost residents and deter coyote encroachment in the Recovery Area. ...
Article
Full-text available
Recovery of large carnivores remains a challenge because complex spatial dynamics that facilitate population persistence are poorly understood. In particular, recovery of the critically endangered red wolf (Canis rufus) has been challenging because of its vulnerability to extinction via human-caused mortality and hybridization with coyotes (Canis latrans). Therefore, understanding red wolf space use and habitat selection is important to assist recovery because key aspects of wolf ecology such as interspecific competition, foraging, and habitat selection are well-known to influence population dynamics and persistence. During 2009–2011, we used global positioning system (GPS) radio-telemetry to quantify space use and 3rd-order habitat selection for resident and transient red wolves on the Albemarle Peninsula of eastern North Carolina. The Albemarle Peninsula was a predominantly agricultural landscape in which red wolves maintained spatially stable home ranges that varied between 25 km2 and 190 km2. Conversely, transient red wolves did not maintain home ranges and traversed areas between 122 km2 and 681 km2. Space use by transient red wolves was not spatially stable and exhibited shifting patterns until residency was achieved by individual wolves. Habitat selection was similar between resident and transient red wolves in which agricultural habitats were selected over forested habitats. However, transients showed stronger selection for edges and roads than resident red wolves. Behaviors of transient wolves are rarely reported in studies of space use and habitat selection because of technological limitations to observed extensive space use and because they do not contribute reproductively to populations. Transients in our study comprised displaced red wolves and younger dispersers that competed for limited space and mating opportunities. Therefore, our results suggest that transiency is likely an important life-history strategy for red wolves that facilitates metapopulation dynamics through short- and long-distance movements and eventual replacement of breeding residents lost to mortality.
... The red wolf "once roamed an extensive range including the southeastern United States, and possibly the entire woodlands of eastern North America" (Stoskopf et al., 2005(Stoskopf et al., , p. 1146. Due to anthropogenic factors (hunting, habitat destruction, economic development, etc.) and consequent red wolf population decline, the species was listed as endangered in 1967 and extinct in the wild in 1980. ...
... This version of the red wolf story is based on others told elsewhere (e.g., Adams et al., 2003;Bohling & Waits, 2011;Manganiello, 2009;Roth et al., 2008;Stoskopf et al., 2005) and I intentionally include specific terms repeatedly occurring in those accounts, such as threaten, fear, genetically swamped, hybrid swarm, pure. Let us focus on the words "pure" and "threat" as an example. ...
Article
This essay is about the interdependence of story and action with respect to cultural and natural heritage. It is also about the inexorability of change and its relationship to heritage conservation. In the following paragraphs I share several stories and excerpts, some heroic, others less so (I leave it to the reader to decide which is which), to make the case that the traditional, Western perspective on heritage does not hold up well under scrutiny—there is now an emerging paradigm for heritage conservation, one that both realizes its “empty” nature and guides us in developing a conservation approach that aligns with this recognition.
... Coyotes and hybrids (coyote  red wolf) were captured in conjunction with red wolves (Canis rufus Audubon and Bachman, 1851) to determine the spatial extent and amount of coyote genetic introgression in red wolves (Stoskopf et al. 2005;. Coyotes and hybrids were sterilized to determine the effectiveness of the placeholder concept on reducing hybridization between coyotes and red wolves (Gese and Terletzky 2015). ...
Article
Fertility control among carnivores has been used to reduce depredations on livestock and wild neonates, population control, modify behavior, inhibit genetic introgression, and reduce human–wildlife conflicts. Although there is considerable knowledge on techniques to sterilize carnivores, there is little information concerning how the absence of gonadal hormones influences behavior, space use, and survival of wild canids. We examined territorial fidelity, home-range size and overlap, and survival of 179 surgically sterilized free-ranging canids (124 coyotes (Canis latrans Say, 1823), 55 coyote – red wolf (Canis rufus Audubon and Bachman, 1851) hybrids) with gonadal hormones present (tubal-ligated females (n = 70), vasectomized males (n = 73)) versus absent (spayed females (n = 22), neutered males (n = 14)). The absence of gonadal hormones did not influence annual home-range size and home-range overlap, territory fidelity, and annual survival rates. Additionally, no differences were detected across sexes and hormonal treatments in annual home-range size, percent home-range overlap, annual home-range residency rates, and annual survival rates. Methods of fertility control that do not keep gonadal organs intact may prove useful for management without concern for changes in behavior, mainly territoriality, space use, and survival.
... To maintain genetic viability, socially disagreeable control measures may be taken as well. Such is the case with red wolves and northern spotted owls (Strix occidentalis caurina), whereby hybrid wolf-coyote (Canis latrans) pups and spotted-barred (Strix varia) owlets are euthanized or sterilized, respectively (Livezey, 2010;Stoskopf et al., 2005). Additionally, not all species will be wild once again and some will remain in captivity (Scott et al., 2010), perhaps designated as zoo curiosities or breeding stock. ...
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