A Framework for Monitoring the Effectiveness of
Habitat Management for Mule Deer
DRAFT
Version 1.1
Prepared for:
Ministry of Forest and Range
Forest Practices Branch
272 Fisgard Street
Victoria, BC V8W 1R8
Prepared by:
Steven F. Wilson, R.P. Bio.
EcoLogic Research
406 Hemlock Avenue, Gabriola Island, BC V0R 1X1
&
Christoph Steeger, R.P. Bio.
Pandion Ecological Research Ltd.
532 Park Street, Nelson, BC V1L 2G9
July 2006Executive
Summary
Mule deer (Odocoileus hemionus) are an important feature of British Columbia’s natural heritage,
providing wildlife viewing and hunting opportunities that exceed that of all other ungulate species
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found in the Province. As a result, habitat management for mule deer in forested landscapes is an
important consideration in forest harvesting strategies.
The Province of BC recognizes the importance of forest cover to mule deer populations and, as a
result, allows for the establishment of Ungulate Winter Ranges for mule deer under Section 12 of
the Government Actions Regulation of the Forest and Range Practices Act (FRPA).
We developed a series of indicators to measure the effectiveness of habitat management for mule
deer under the Forest and Ranges Practices Act. Recommended effectiveness indicators address a
number of key monitoring questions and are stratified by the resources required to measure. The
indicators focus on the proportion of suitable habitat captured under management, forest cover
characteristics, forage quality and availability, snow depth and consolidation, and evidence of
sustained winter use.
Although the recommended indicators address all key monitoring questions, not all questions are
addressed comprehensively. Regional and subregional population objectives for mule deer are
rarely explicit, and in general, the required distribution and abundance of Ungulate Winter Ranges
required to meet population objectives is unknown.
Monitoring the effectiveness of habitat management
for mule deer should involve: 1) establishing a
regional or subregional population objective; 2)
implementing procedures to collect, warehouse and
analyze routine indicator data; 3) allocating resources
to collect extensive indicator data in areas of greatest
management concern; and, 4) collaborating with other
agencies to collect intensive indicator data and to
address knowledge gaps. Table of Contents
Excecutive Summary .......................................................................................................................................... 2
Table of Contents ................................................................................................................................................ 3
Acknowledgements ............................................................................................................................................. 3
Introduction ......................................................................................................................................................... 4
Objectives ........................................................................................................................................................... 4
Approach............................................................................................................................................................. 4
Conservation Status & Situation Analysis ........................................................................................................... 4
Conservation Status & Demography ............................................................................................... 4
Population Threats ........................................................................................................................... 5
Wildlife Management ...................................................................................................................... 6
Biology and Life History ...................................................................................................................................... 6
Habitat Management under the Forest & Range Practice Act ............................................................................. 6
Key Effectiveness Monitoring Questions ............................................................................................................. 6
Small Scale (Individual Ungulate Winter Ranges) ................................................................... 6
Medium Scale (Watershed or Management Unit) ..................................................................... 6
Large Scale (Subregional or Regional Populations) ................................................................. 6
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Effectiveness Indicators ...................................................................................................................................... 6
Knowledge Gaps ................................................................................................................................................. 8
Management Recommendations ........................................................................................................................ 8
Literature Cited ................................................................................................................................................. 10
List of Tables
Table 1. Recommended indicators by level of intensity, desired conditions and monitoring
frequency for assessing the effectiveness of Ungulate Winter Ranges (UWR) established for
mule deer in BC. ........................................................................................................................ 10
Table 2. Knowlege gaps related to mule deer ecology that are relevant to the key effectiveness
monitoring questions. ................................................................................................................ 11
List of Figures
Figure 1. Distribution and abundance of the three subspecies of mule deer in British Columbia,
from Shackleton (1999). .............................................................................................................. 6
Acknowledgements
We thank Wayne Erickson (BC Ministry of Forests,
Victoria) for administering this project and Dennis
Hamilton (Nanuq Consulting Ltd., Nelson) for review
and input.Introduction
Mule deer (Odocoileus hemionus) are an important feature of British Columbia’s (BC) natural
heritage, providing wildlife viewing and hunting opportunities that exceed that of all other ungulate
species found in the Province (WLAP 2003). As a result, habitat management for mule deer in
forested landscapes is an important consideration in forest harvesting strategies.
Winter is known as a critical period for mule deer, particularly in years when snow cover is deep
and temperatures are well below normal for extended periods. Deep snow affects mobility and
access to forage, which in turn affects energy balance (Ungulate Winter Range Technical Advisory
Team 2005). As a result, malnutrition is a principal mortality factor in winter and can result in
widespread die-offs in severe conditions (e.g., Bishop et al. 2005). To escape deep snow, mule deer
seek out areas where snow is shallow – often under dense forest canopies (e.g., Armleder et al
1994).
The Province of BC recognizes the importance of forest cover to mule deer populations and, as a
result, allows for the establishment of Ungulate Winter Ranges for mule deer under Section 12 of
the Government Actions Regulation of the Forest and Range Practices Act (FRPA).
We developed a series of indicators to measure the effectiveness of habitat management for mule
deer under the Forest and Ranges Practices Act (FRPA), based on a series of key monitoring
questions. These indicators can be used to establish a programme to monitoring the effectiveness of
habitat management in an adaptive management context.
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Objectives
The specific objectives of this report were to:
1. Review existing information related to the conservation status and demography of mule
deer;
2. Assemble information on natural and human-related threats that have the potential to affect
the viability of mule deer populations;
3. Outline current management practices for mule deer;
4. Describe life history requisites of mule deer in relation to management tools available to
protect their habitats;
5. Review methods available to monitor mule deer habitat and populations;
6. Develop indicators to monitor the effectiveness of habitat managed for mule deer under
FRPA; and,
7. Identify knowledge and management gaps in BC’s strategy to sustain mule deer
populations.
Approach
Our approach to developing indicators to monitor the effectiveness of habitat management for mule
deer involved the following steps:
1. Identify limiting habitat and habitat attributes required to fulfil critical life requisites of
mule deer;
2. Outline mechanisms for managing important habitats under FRPA;
3. Identify threats to mountain mule deer habitat and populations;
4. Develop key monitoring questions at small, medium and large spatial scales;
5. Summarize population and habitat monitoring methods;
6. Propose indicators based on a series of desirable criteria that address: a) different levels of
monitoring intensity; b) desired outcomes or results (in relation to a stated management
goal); and, c) required frequency of measurement;
7. Identify of knowledge gaps; and,
8. Develop recommendations for implementation.
Conservation Status & Situation Analysis
Conservation Status & Demography
Three subspecies of mule deer are recognized in BC. Rocky Mountain mule deer (Odocoileus
hemionus hemionus) are the most widespread and are found throughout the southern half of the
Province east of the Coast (and in the south, Cascade) Mountains, and well as the Peace River
district (Figure 1; Shackleton 1999). Two subspecies are associated with coastal BC: Columbian
black-tailed deer (Odocoileus hemionus columbianus), which range throughout Vancouver Island,
smaller coastal islands and the mainland coast north to approximately Rivers Inlet; and Sitka blacktailed deer (Odocoileus hemionus sitkensis), which range farther north, including the Queen
Charlotte Islands, where they were introduced. The black-tailed deer subspecies hybridize along the
respective margins of their range.
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Population Threats
Mule deer are the most common ungulate in BC, numbering in the hundreds of thousands of
animals. Introduced Sitka black-tailed deer on the Queen Charlotte Islands are considered pests. As
with all ungulates, predation is probably the most significant mortality factor affecting mule deer
throughout their range. Predators rarely pose serious threats to prey populations, although some
researchers have speculated that increasing populations of white-tailed deer in areas previously
occupied exclusively by mule deer may be “subsidizing” higher predator numbers (particularly
cougars, Puma concolor) that are in turn affecting the viability of some local mule deer populations
(Robinson et al. 2002).
Diseases do not appear to be a major threat to mule deer populations. Although susceptible to
epizootic haemorrhagic disease, there have been no reports of outbreaks in BC (Shackleton 1999).
Severe winter conditions likely constitute the most significant threat to mule deer populations (e.g.,
Bishop et al. 2005). The effects of severe winters are likely worse when habitats have been
modified to remove important attributes critical for mule deer during severe winter conditions
(Ungulate Winter Range Technical Advisory Team 2005). The winter of 1996-7 was the most
recent winter that was sufficiently severe to significantly affect mule deer populations (Wilson and
Morley 2005), although evidence in some regions suggest that populations were already in decline,
perhaps due to increasing predation (G. Woods, pers. comm.). Mild winters since have allowed
mule deer populations in many areas to recover. Black-tailed deer populations in many parts of
Vancouver Island are still considered to be lower than desirable (K. Brunt, pers. comm.) and mule
deer have not recovered as rapidly as white-tailed deer populations in the Kootenay region (G.
Woods, pers. comm.).
Wildlife Management
Mule deer are an important wildlife species from a socio-economic perspective. In many areas of
the Province (e.g., Vancouver Island) mule deer have readily adapted to human presence and some
of the highest densities now occur near human settlements (Shackleton 1999). As a result they are
an important species for wildlife viewing.
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Figure 1. Distribution and abundance of the three subspecies of mule deer in British
Columbia, from Shackleton (1999).
Mule deer have long been a focus of habitat enhancement activities by government agencies and
public interest groups. For example, the Habitat Conservation Trust Fund alone spent $140 000 on
enhancement projects for mule deer and other ungulates in 2005-6 in the southern interior region.
Demand for mule deer hunting opportunities in BC exceeds that of all other ungulate species. Of
the 85 000 resident hunters in 2002-3, over 63 000 purchased mule deer species tags, resulting in
direct revenue to the Province of $950 000 for the tags alone. Harvest in 2002 was estimated at
more than 16 000 – more than any other ungulate species (WLAP 2003). Most mule deer hunting in
BC is by residents; non-residents harvested only 170 mule deer in 2002-3.
Regionally, the largest harvests of mule deer occur in the Cariboo, Thompson and Okanagan,
respectively (WLAP 2003).
Habitat Use
Mule deer are an adaptable species that inhabit a variety of habitats in BC. In general, Rocky
Mountain mule deer prefer interior Douglas-fir (Pseudotsuga menziesii var. glauca) and Ponderosa
pine (Pinus ponderosa) forest as well as parkland habitats (Shackleton 1999). In contrast, both
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Columbian and Sitka black-tailed deer inhabit dense coastal rainforests (Shackleton 1999). As
generalist browsers, mule deer are generally not food-limited during summer-fall and feed on a
variety of different shrubs and forbs, depending on their availability (Shackleton 1999).
The habitat use of mule deer changes significantly as snow deepens during winter and begins to
restrict mobility. In general, ungulates do not tolerate snow depths greater than chest height and
mobility is affected when snow is knee-deep (Kelsall and Prescott 1971). Serrouya and D'Eon
(2003) found that deer avoided snow depths >50 cm deep near Revelstoke and Woods (1984) found
that white-tailed deer (Odocoileus virginianus) did not appear to select for specific habitats in when
snow was <40 cm deep in the Pend D’Oreille valley of the southern interior.
In addition to snow depth, consolidation and sinking depth are important factors influencing
mobility (Verne 1968). In cold, dry snow conditions, snow depths likely approximate sinking
depths. Conversely, relatively wetter, warmer snow conditions will reduce sinking depths (Bunnell
et al. 1990).
Mule deer winter range characteristics vary throughout the Province, but in general animals move
varying distances to areas where the local snow pack is most shallow – usually warm aspect slopes
at low elevations. The extent of these areas varies broadly with biogeoclimatic characteristics
(Ungulate Winter Range Technical Advisory Team 2005). In shallow snow pack zones, winter
range areas can be large and extensive, while in more rugged areas dominated by deep snow packs,
winter ranges are generally small and spatially restricted.
Energy balance is the key to mule deer survival in winter; therefore, the highest quality winter
ranges are those that not only provide the lowest mobility costs but also provide access to abundant,
preferred forage. The rooted forage accessible to mule deer is largely a function of snow depth:
where snow is shallow, forage is generally more accessible. The forest canopy can intercept snow
and reduce snow depths on the ground, which in turns increases access to forage. In areas
dominated by very shallow snow packs (e.g., the bunchgrass biogeoclimatic zone), little forest
cover is required to moderate snow, while in areas dominated by deep snow (e.g., coastal western
hemlock and interior cedar hemlock biogeoclimatic zones), extensive forest cover is required to
access to rooted forage (Ungulate Winter Range Technical Advisory Team 2005).
Forest overstorey is critically important during severe winters (e.g., Pauley et al. 1993, Doerr et al.
2005). As snow depths increase, the proportion of rooted forage in mule deer diets decreases and is
replaced by more accessible forage, such as cedar buds and litterfall in coastal areas (Hanley and
McKendrick 1985) and Douglas-fir needles and litterfall in the southern interior (Waterhouse et al.
1993).
The ability of the forest overstorey to intercept snow increases with canopy closure, although the
relationship is influenced by a number of variables (Bunnell et al. 1985, McNay 1985, Kirchoff and
Schoen 1987, D’Eon 2004). Forest age influences snow interception and also influences the
potential of the understorey to produce forage. Older-aged stands with multi-storied canopies are
associated with higher forage values than younger, even aged stands (Ungulate Winter Range
Technical Advisory Team 2005). Older forests are also associated with more litterfall (Waterhouse
et al. 1991).
Columbian and Sitka black-tailed deer are smaller than Rocky Mountain mule deer (Shackelton
1999) and inhabit areas where deep and unconsolidated conditions are common. As a result, snow
interception on winter ranges is even more critical for these subspecies than for Rocky Mountain
mule deer (Nyberg and Janz 1990).
Based on these and other results, The Ungulate Winter Range Technical Advisory Team (2005)
recommended a range of forest cover objectives or winter ranges of the southern interior, from very
limited snow interpretation cover in NDT 4 ecosystems to dense canopies with mature and old
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characteristics in deep snow zones of the southern interior. Similarly, Nyberg and Janz (1990)
recommended objectives for coastal winter ranges that depended on snow zones.
Spring is also a critical time for mule deer as they attempt to recover from the nutritional stress of
winter. Early green-up areas that provide abundant, high quality forage near winter ranges are
critically important (Ungulate Winter Range Technical Advisory Team 2005).
Overview of Inventory and Monitoring Methods
Monitoring habitat conditions on ungulate winter ranges is typically restricted to office procedures
involving geographic information systems. Leading species characteristics and age class
information are readily available in forest cover databases and provide sufficient information for
coarse-level interpretation of winter range conditions.
Ground-based assessments of habitat capability, suitability and use by mule deer can be
accomplished by developing Wildlife-Habitat Ratings Models (RISC 1999). Resulting models can
be mapped where Terrestrial Ecosystem Mapping or Predictive Ecosystem Mapping is available
and structural stage can be estimated.
Habitat use by mule deer has been investigated most commonly using winter track surveys
(Armleder et al. 1998, D’Eon 2001, Hebert and Halko 2001, D’Eon et al. 2006). Pellet counts have
been used successfully to quantify winter-use by white-tailed deer (Boulanger et al. 2000). Habitat
use has also been determined via radio telemetry studies (e.g., Kremsater and Bunnell 1992).
BC has developed standards for both aerial (RISC 2002) and ground-based inventory methods
(RISC 1998). Aerial surveys for mule deer are not recommended except in the Boreal Plains
ecoprovince (BWBS biogeoclimatic zone), where mule deer winter in areas of sparse coniferous
and deciduous cover and sightability has been demonstrated to be close to 100%, based on markresight methods (RISC 2002). Elsewhere in the Province, sightability it too low to use aerial survey
methods reliably.
Ground-based inventory methods used to estimate relative abundance of mule deer include winter
track count surveys (D’Eon et al. 2006), pellet counts and spotlight surveys (RISC 1998). Annual
spotlight surveys have been used to index black-tailed deer populations on Vancouver Island for
>15 years (K. Brunt, pers. comm.). Absolute abundance can be estimated using mark-resight
methods where marked (e.g., radio-collared) animals are available (RISC 1998). Total counts via
ground transects have also been suggested for estimating absolute abundance of black-tailed deer
on small coastal islands (RISC 1998).
Habitat Management under the Forest & Range
Practice Act
Habitat management for mule deer under the FRPA is restricted to the establishment of Ungulate
Winter Ranges under Section 12 of the Government Actions Regulation. Ungulate Winter Ranges
are areas established to ensure the over-winter survival of mule deer, recognizing that this is a
critical season for ungulates due to nutritional deprivation and high energy expenditure related to
thermoregulation and mobility in snow.
As of 31 March 2006 there were 35 approved winter ranges for all ungulate species covering >2.6
million ha in BC. Associated with each ungulate winter range plan are approved objectives or
General Wildlife Measures that outline forestry management intended to sustain critical habitat
characteristics. These objectives and General Wildlife Measured differ among plans due to different
management approaches and biological requirements in different parts of the Province.
Monitoring compliance with the legal objectives in ungulate winter range plans is an important
aspect of ungulate winter range management; however, beyond compliance, there is a need to
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investigate whether ungulate winter ranges are meeting the intent of the FRPA. Effectiveness
monitoring assesses whether the outcomes of management are consistent with the future desired
conditions that the legal objectives were designed to achieve.
Key Effectiveness Monitoring Questions
The following are proposed key monitoring questions related to assessing the effectiveness of
ungulate winter ranges established for mule deer under the FRPA.
Small Scale (Individual Ungulate Winter Ranges)
1. Does the winter range provide the habitat elements required to fulfil the life requisites of
wintering mule deer?
2. Is the winter range receiving sustained use by mule deer?
3. Is human-related disturbance affecting use of the winter range?
Medium Scale (Watershed or Management Unit)
1. Are there barriers outside of the Ungulate Winter Range that prevent or reduce the potential
use of the range?
2. Is the distribution and abundance of Ungulate Winter Ranges established for mule deer
sufficient to ensure over-winter survival in typical and severe winters?
Large Scale (Subregional or Regional Populations)
1. Is the distribution and abundance of Ungulate Winter Ranges established for mule deer
sufficient to sustain the regional population?
Effectiveness Indicators
Indicators can measure the function (e.g., snow depth, forage accessibility), structure (e.g., forest
canopy characteristics) or use (e.g., direct inventory) of Ungulate Winter Ranges. Indicators should
be:

Measurable (accurately and precisely);

Cost-effective to monitor; and,

Sensitive to management or stressor change.
Different indicators require different levels of resources to measure and monitor. Indicators can be
classified as:

Routine: Generally measured through office procedures from data collected for other
purposes (e.g., map interpretation);

Extensive: Requires low-intensity qualitative or quantitative field assessments (e.g., blowdown assessments); and,

Intensive: Requires detailed, quantitative fieldwork (e.g., population inventory).
Assessing the effectiveness of habitat management at medium and large spatial scales also requires
interpretation in relation to broad management objectives:

Population maintenance: Maintain the current approximate pattern of habitat occupancy
(medium scale) and abundance and distribution (large scale) of mule deer. Use currently
occupied habitat and indices of abundance as the basis for defining desired conditions.
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Population recovery: Increase rates of occupancy of suitable habitat and increase the
abundance and possibly the regional distribution of mule deer. Use habitat capability and
recovery objectives related to abundance and distribution as the basis for defining desired
conditions.
Broader management objectives will differ in space and time throughout BC, depending on a
variety of socio-economic and biological factors.
Recommended indicators are presented in Table 1 and general recommended methods for collecting
indicator data are outlined in Table 2.
Table 1. Recommended indicators by level of intensity, desired conditions and monitoring frequency for
assessing the effectiveness of Ungulate Winter Ranges (UWR) established for mule deer in BC.
Minimum
Monitoring
Frequency
Once when
winter ranges
are being
legally
established
Level of
Intensity
Routine
Desired
Condition/Result
All suitable winter
ranges under
management
Forest cover
characteristics
Extensive
Every 5 years
and following
any
harvesting
Forage availability
Extensive
Characteristics
consistent with
objectives when
winter ranges were
established (will
vary depending on
subspecies and
geographic area.
Abundant and
available preferred
rooted forage and
litterfall
Snow depth and
consolidation
Intensive
Snow depths <25
cm in typical
winters, <50 cm in
severe winters
Sufficiently
frequent to
capture
variation in
winter
conditions
Evidence of
sustained winter
use by mule deer
Extensive
Evidence of browse,
presence of pellet
groups, tracks,
direct observations
Every 3 years
Indicator
Proportion of area
in established
Ungulate Winter
Range relative to
available suitable
winter range
habitat
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Every 5 years
Rationale and Comments
Additional habitat will be required
if population recovery rather than
maintenance is an objective. In
some areas significant winter
range is also found outside the
Crown Forest Land Base on
private land. Its contribution and
security should be considered in
the examination of this indicator.
Aerial or ground-based
assessments are required to
assess results of any allowable
harvest treatments, blowdown
and forest health issues.
Requires qualitative ground
assessments. Forage availability
is influenced by a variety of site
characteristics (e.g., soil and
moisture regime, forest canopy
characteristics, slope and aspect,
snowfall regime. Quantitative
forage assessments could also
be considered.
The importance of winter ranges
increases with winter severity;
therefore, snow conditions
should be assessed during
average and severe winter
conditions. General qualitative
assessments can be made from
the air or on the ground, but
quantitative assessments,
requiring ground reconnaissance
(e.g., actual snow depth and
sinking depth measures), are
recommended.
Requires aerial surveys (where
forest cover is sparse) or ground
reconnaissance.
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Table 2. Methods available to collect data related to recommended indicators for assessing the effectiveness
of Ungulate Winter Ranges (UWR) established for mule deer in BC.
Indicator
Proportion of area in
established Ungulate Winter
Range relative to available
suitable winter range habitat
Forest cover characteristics
Forage availability
Snow depth and consolidation
Evidence of sustained winter
use by mule deer
General Methods
Maps of suitable winter range based on empirical (e.g., Mowat et al.
2002) or expert-based (e.g., Wilson and Hamilton 2002) habitat models
are available for many areas of the Province. Areas of suitable habitat
captured in UWR can be compared to the distribution of suitable habitat
on a landscape unit basis using GIS office procedures.
Forest cover characteristics can be assessed using approved standard
methods (MELP and MOF 1998).
Forage availability can be assessed using approved standard methods
(MELP and MOF 1998).
No standards currently exist for measuring snow depth and consolidation;
however, methods have been proposed by Wilson (2006) based on
MSRM (2002).
Stand level assessments of use by mule deer can be measured using
approved standard methods (RISC 1998, 2002, D’Eon et al. 2006; see
Inventory and Monitoring Methods above).
Knowledge Gaps
Mule deer are one the most-studied wildlife species in BC and, in fact, North America. However,
there are still knowledge gaps related to habitat management that are best addressed through
adaptive management trials (Table 3).
Table 3. Knowledge gaps related to mule deer ecology that are relevant to the key effectiveness monitoring
questions.
Monitoring Question
Does the winter range provide the
habitat elements required to fulfil the life
requisites of wintering mule deer?
Is the winter range receiving sustained
use by mule deer?
Is human-related disturbance affecting
use of the winter range?
Knowledge Gap

None

None

Tolerance of mule deer to human-related activities is
variable; however, evidence of sustained use
addresses the monitoring question adequately
Are there barriers outside of the
Ungulate Winter Range that prevent or
reduce the potential use of the range?

The extent to which clearcuts and associated slash
(and deep snow in winter), roads, immature forests and
other landscape modifications create barriers to mule
deer movements is not clearly understood
Is the distribution and abundance of
Ungulate Winter Ranges established for
mule deer sufficient to ensure overwinter survival in typical and severe
winters?

The distribution and abundance of winter ranges
required to sustain local or regional populations through
moderate and severe winters is generally unknown. In
addition, regional population objectives are rarely
articulated.
Is the distribution and abundance of
Ungulate Winter Ranges established for
mule deer sufficient to sustain the
regional population?
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Management Recommendations
We recommend five indicators to monitor the effectiveness of habitat management within Ungulate
Winter Ranges legally established under the FRPA. These indicators address small-, medium- and
large-scale monitor questions, and standard and/or well-developed methods for data collection are
available.
Considerable differences exist in the amount of time and resources required to monitor the proposed
indicators and in the strength and conclusiveness of evidence they may provide related to the
effectiveness of habitat management. As a result, adaptive management based on monitoring
outcomes will require an assessment of general trends among the different indicators.
Monitoring the effectiveness of Ungulate Winter Range management for mule deer should involve
the following components:
1. Establishing a regional or subregional population objective;
2. Implementing procedures to collect, warehouse and analyze routine indicator data;
3. Allocating resources to collect extensive indicator data in areas of greatest management
concern; and,
4. Collaborating with other agencies and researchers to collect intensive indicator data and to
address knowledge gaps.
Literature Cited
Armleder, H. M., M. J. Waterhouse, D. G. Keisher, and R. J. Dawson. 1994. Winter habitat use by
mule deer in the central interior of British Columbia. Canadian Journal of Zoology 72:17211725.
Armleder, H. M., M. J. Waterhouse, R. J. Dawson, and K. E. Iverson. 1998. Mule deer response to
low-volume partial-cutting on winter ranges in Central Interior British Columbia. BC Ministry
of Forests Research Report No. 16.
Bishop, C. J., J. W. Unsworth, and E. O. Garton. 2005. Mule deer survival among adjacent
populations in southwest Idaho. Journal of Wildlife Management 69:311-321.
Boulanger, J. G., K. G. Poole, J. Gwilliam, G. P. Woods, J. Krebs, and I. Parfitt. 2000. Winter
habitat selection by white-tailed deer in the Pend D’Oreille Valley, southeastern British
Columbia. Columbia Basin Fish and Wildlife Compensation Program, BC Hydro and BC
Ministry of Environment, Lands and Parks, Nelson, BC.
Bunnell, F. L., R. S. McNay, and C. C. Shank. 1985. Trees and snow: the deposition of snow on the
ground. BC Ministry of Forests Integrated Wildlife Intensive Forestry Research Report No. 17.
Bunnell, F. L., K. L. Parker, R. S. McNay, and F. W. Hovey. 1990. Sinking depths of black-tailed
deer in snow, and their indices. Canadian Journal of Zoology 68:917-922.
D’Eon, R. G. 2001. Using snow-track surveys to determine deer winter distribution and habitat.
Wildlife Society Bulletin 29:879-887.
D’Eon, R. G. 2004. Snow depth as a function of canopy cover and other site attributes in a forested
ungulate winter range in southeast British Columbia. BC Journal of Ecosystems and
Management 3:2.
D’Eon, R. G., S. F. Wilson, and D. Hamilton. 2006. Winter track count survey standards. Prepared
for: BC Ministry of Environment, Resources Information Standards Committee, Victoria.
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Doerr, J. G, E. J. Degayner, and G. Ith. 2005. Winter range habitat selection by Sitka black-tailed
deer. Journal of Wildlife Management 69:322-331.
Hanley, T. A., and J. D. McKendrick. 1985. Potential nutritional limitations for black-tailed deer in
a spruce-hemlock forest, southeastern Alaska. Journal of Wildlife Management 49:103-114.
Hebert, D., and R. Halko. 2001. 2000-2001 Boundary Forest District mule deer winter range
project. Prepared for: Pope & Talbot Ltd., Midway, BC.
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