Literature Review - research on MPB and caribou - March …a100.gov.bc.ca/appsdata/acat/documents/r22847/Cicho… · · 2011-04-19Literature Review – Research on Mountain Pine

LITERATURE REVIEW:

RESEARCH ON MOUNTAIN PINE BEETLES AND CARIBOU

MARCH 31, 2011

Submitted to:

Nicola Freeman Ministry of Environment #400 – 640 Borland St.

Williams Lake, B.C. V2G 4T1

Ph: 250-398-4553 email: [email protected]

Submitted by:

Deborah Cichowski Caribou Ecological Consulting

Box 3652 Smithers, B.C.

V0J 2N0 Phone: 250-877-7558

e-mail: [email protected]

Literature Review – Research on Mountain Pine Beetles and Caribou – March 31, 2011

Caribou Ecological Consulting, Box 3652, Smithers, B.C. V0J 2N0 ii

Table of Contents

INTRODUCTION............................................................................................................... 1 POTENTIAL EFFECTS OF MOUNTAIN PINE BEETLES ON CARIBOU ECOLOGY ................... 3 STATUS OF CURRENT RESEARCH..................................................................................... 7 SYNOPSIS OF RESEARCH RESULTS ................................................................................. 19

Effects of Mountain Pine Beetles on Caribou Habitat .........................................................19 Effects of Mountain Pine Beetles on Caribou Habitat Use .................................................23 Effects of Mountain Pine Beetles on Caribou Population Dynamics ...............................25

KNOWLEDGE GAPS ....................................................................................................... 27 REFERENCES.................................................................................................................. 30 APPENDIX 1. SUMMARY OF RESEARCH REPORTS ......................................................... 37 APPENDIX 2. BIOGEOCLIMATIC ZONE UNITS ............................................................... 39

List of Tables Table 1. Timing of potential effects of a mountain pine beetle epidemic on

various aspects of caribou ecology (from Cichowski 2007). ...................................4 Table 2. Summary of research on the effects of mountain pine beetles on

caribou. ...............................................................................................................................8 Table 3. Summary of information collected by ongoing research on the effects

of mountain pine beetles and salvage harvesting on woodland caribou ecology...............................................................................................................28

Acknowledgements This project was initiated by Nicola Freeman, Fish and Wildlife Resource Information Specialist, Ministry of Environment. I would like to thank Nicola for providing advice on the project and for reviewing the draft report. Special thanks to Dale Seip, Ministry of Forests, Lands and Natural Resource Operations (MFLNRO), Elena Jones (ESJ Wildlife Consulting), Harold Armleder (MFLMNRO) and Micheala Waterhouse (MFLMNRO) for contributing unpublished data, providing reports, and reviewing the draft report. I would also like to acknowledge Randy Sulyma, Wildlife Infometrics Inc., for providing reports and clarifying various aspects of his projects. Funding for this project was provided by the Habitat Conservation Trust Foundation.


Caribou Ecological Consulting, Box 3652, Smithers, B.C. V0J 2N0 1

INTRODUCTION

The northern ecotype of woodland caribou (Rangifer tarandus caribou) in British Columbia is found in the west-central and northern portions of the province (Stevenson and Hatler 1985). Northern Caribou typically live in and around mountainous areas during summer and spend the winter either in low elevation forests or on windswept alpine slopes (Stevenson and Hatler 1985, Northern Caribou Technical Advisory Committee 2004). Northern Caribou populations found in west-central and north-central British Columbia in the Southern Mountains National Ecological Area (SMNEA) are designated as Threatened under the federal Species at Risk Act (Northern Caribou Technical Advisory Committee 2004).

At lower elevations, Northern Caribou use mature lodgepole pine (Pinus

contorta) forests, especially during winter when they crater through the snow to find terrestrial lichens. Use of low elevation pine forests varies between Northern Caribou populations and may reflect different ecological conditions, such as snow accumulation or terrestrial lichen availability (Cichowski 2008a). Forest management practices for Northern Caribou also differ between populations.

The current mountain pine beetle (Dendroctonus ponderosae) epidemic in

British Columbia covers over 16 million hectares or 50% of the mature lodgepole pine in the province (Walton 2010), and has affected several Northern Caribou ranges. Prior to the current outbreak, no information was available on the effects of mountain pine beetles on caribou. Although research has been conducted on the effects of forest harvesting and fire on caribou and caribou habitat, impacts of fire and forest harvesting differ from those of mountain pine beetles so results from those studies cannot be applied to the mountain pine beetle epidemic. Fire consumes both canopy and forest floor vegetation, whereas only attacked trees die during mountain pine beetle disturbance. With forest harvesting, canopy trees are removed resulting in an abrupt change in light and forest floor conditions, while mountain pine beetle attack results in a gradual change in light and forest floor conditions over 2-3 years.

Four main questions need to be addressed to assess the effects of mountain

pine beetles on Northern Caribou (Cichowski 2007a, 2010). 1. How will caribou habitat be affected by mountain pine beetle attack? (i.e.

How will terrestrial and arboreal lichens respond? Will snow accumulation increase due to a loss of canopy and/or will eventual treefall lead to impeded movements?)

2. How will caribou habitat use and range use be affected by extensive mountain pine beetle attack? (i.e. Will caribou avoid using mountain pine beetle-attacked areas or stands for traveling or foraging during winter



and migration? Will caribou alter foraging strategies in beetle killed areas or shift their winter ranges to other areas?)

3. How will caribou population dynamics be affected by extensive mountain pine beetle attack? (i.e. Will mortality rates and causes, and/or population growth change following the mountain pine beetle epidemic? Will predator movements be impeded more or less than caribou movements and how will that affect predator efficiency?)

4. How will caribou habitat, habitat use and population dynamics be affected by potential mountain pine beetle management and salvage harvesting?

Since 2001, several research projects have been initiated to investigate the

impacts of mountain pine beetles on caribou and caribou habitat. This report summarizes results from current research on the effects of mountain pine beetles on caribou and their habitat, and identifies knowledge gaps.



POTENTIAL EFFECTS OF MOUNTAIN PINE BEETLES ON CARIBOU ECOLOGY

Although all aspects of caribou ecology are inter-related, potential effects of

mountain pine beetles on caribou can be categorized into: • effects on caribou habitat; • effects on caribou habitat use and range use; and, • effects on caribou population dynamics. The response of Northern Caribou populations and their habitat to the

mountain pine beetle epidemic will depend on the extent of mountain pine beetle attack across their ranges at both the landscape and stand levels. Because ecological conditions vary between caribou ranges, the level and patterns of attack will also likely vary depending on the extent and distribution of susceptible pine on the landscape and climatic conditions affecting beetle mortality. Although responses of caribou to the mountain pine beetle epidemic may differ between populations, information collected on general patterns of caribou behaviour and caribou habitat responses to mountain pine beetles from one caribou range could be used to help predict responses on other caribou ranges.

The effects of tree mortality due to mountain pine beetles on caribou will also

vary depending on the stage of the epidemic. The four stages of the epidemic include green attack, red attack, grey attack, and fall-down. During the green attack stage, attacked trees are reduced in vigour, affecting evapotranspiration and nutrient uptake. At the red attack stage, trees are dead so nutrient uptake and evapotranspiration have ceased, but needles remain on trees so the canopy still functions in rain, snow and light interception. By the grey attack stage, needle loss potentially affects rain, snow and light interception. The fall-down stage results in further changes to light conditions and interception of precipitation, and in accumulations of coarse woody debris.

Table 1 summarizes the characteristics of each mountain pine beetle attack

stage and the potential effects on caribou habitat, caribou habitat and range use, and population dynamics.

Potential effects of extensive mountain pine beetle attack on caribou habitat

include: changes in terrestrial lichen abundance and arboreal lichen abundance; changes in snow conditions following needle loss (potentially affecting winter travel); and accumulation of coarse woody debris (affecting summer and winter travel). Changes in terrestrial and arboreal lichen abundance could be mediated through changes in soil moisture and nutrients, competing vegetation, and light availability. Potential effects on terrestrial and



Table 1. Timing of potential effects of a mountain pine beetle epidemic on various aspects of caribou ecology (from Cichowski 2007).

Mountain Pine Beetle Attack Phase1 Alive Green Red Grey Fall-down

STAND CHARACTERISTICS Nutrient uptake Normal Reduced None None None Evapotranspiration Normal Reduced None None None Needles On On On/Falling Off Off Tree status Standing Standing Standing Standing On ground HABITAT Terrestrial lichen abundance Arboreal lichen abundance Competing vegetation abundance Regeneration Soil moisture Soil nutrients Light availability Snow conditions Coarse woody debris HABITAT AND RANGE USE Forage choice (terrestrial vs. arboreal) Site (stand) selection Landscape (range) selection Elevation selection Winter travel – snow conditions Summer and winter travel – coarse woody debris

POPULATION DYNAMICS Population abundance Population status Calf recruitment Adult mortality Predation risk (other prey abundance/ presence - forage)

Predation risk (other prey abundance/ presence - winter travel)

Predation risk (other prey abundance/ presence - summer travel)

Predation risk (predator winter travel) Predation risk (predator summer travel)

1 Shaded areas indicate phases of attack that could potentially affect various aspects of caribou ecology; dark shading indicates full effects while light shading indicates partial effects



arboreal lichens and competing vegetation will begin during the green attack stage when nutrient uptake and evapotranspiration are compromised and will be fully realized by the red attack stage when trees are dead. These effects will subside when nutrient uptake and evapotranspiration from regenerating forests start affecting availability of nutrients and moisture for understory vegetation. Light availability is expected to increase starting in the red attack stage and to continue increasing into the grey attack stage. Further increases in light availability will occur when trees fall. Arboreal lichen abundance is expected to increase in response to increased light availability resulting from mountain pine beetle attack (T. Goward, pers. comm.). When trees fall, arboreal lichens on the whole tree will become available to caribou; however, arboreal lichens on fallen trees are expected to deteriorate within 1-2 years. Potential changes in snow conditions will start during the grey attack phase and will continue into the fall-down phase. Snow accumulation is expected to change with needle loss, but snow density and crusting conditions could also change with increased insolation. Although fall-down may begin during any phase of the attack, the greatest accumulations will occur after trees have been dead for some time.

Potential effects of mountain pine beetles on caribou habitat use and range

use could include: changes in forage and habitat selection due to potential changes in forage abundance; and, shifts in elevation use or area of use within their range in response to changes in forage abundance or mobility. Changes in habitat selection and forage selection (terrestrial lichen vs. arboreal lichen use) could begin during green attack if lichen abundance is affected at that stage, but will likely become more significant during the red and grey attack stages. If terrestrial lichen abundance decreases, caribou could start relying more on arboreal lichens and consequently could start focusing more on habitats where arboreal lichens are abundant. This could lead to changes in areas used within their winter range, and/or increased use of higher elevations where arboreal lichens are more abundant, as early as during the red attack stage.

During the grey attack stage, canopy loss could lead to changes in snow

conditions (accumulation, hardness), which could further affect caribou winter forage and winter habitat selection, and use of higher elevations and alternate areas within their winter range. Snow depth and snow hardness could affect both cratering and traveling. Increased snow depth and snow hardness could make cratering more difficult in grey-attacked areas, while increased snow depth could make travel more difficult. If snow conditions result in difficult cratering conditions, caribou could focus more on arboreal lichens. If snow conditions result in difficult traveling conditions, caribou could focus more on habitats and areas within the winter range that contain fewer attacked trees, such as higher elevations where pine is less abundant. Changes in cratering and travel conditions could continue into the fall-down phase. While canopy loss could potentially affect conditions for winter travel, fall-down could affect both winter and summer travel, leading to potential shifts in migration patterns, range use, summer habitat use and even further shifts in winter habitat use.



Changes in forage selection, habitat use and seasonal range use will depend

on the extent and distribution of mountain pine beetle attack on the landscape. During winter, caribou could continue to access areas within their winter range if they could move through a matrix of unattacked stands. As long as they could travel through their winter range, they could potentially continue to use grey-attacked or fall-down stands where terrestrial lichens are more abundant. Conversely, caribou could potentially shift their ranges entirely if alternate ranges are available. However, the current mountain pine beetle epidemic is so extensive that alternate ranges will also likely be affected, so switching to alternate ranges would not necessarily result in any benefits to the caribou.

Although changes in the forest landscape due to mountain pine beetles could

lead to changes in forage abundance and travel, the primary limiting factor for caribou is predation, which maintains caribou populations well below food carrying capacity (Seip and Cichowski 1996). Therefore, the effects of the mountain pine beetle epidemic on predation risk will play an important role in the effects of mountain pine beetles on caribou. Population abundance and trend, calf recruitment, and adult mortality of caribou will likely not be directly affected by a mountain pine beetle epidemic; rather, population dynamics will likely be mediated through potential changes in habitat use and abundance of other prey species or potential changes in predator efficiency. The mountain pine beetle epidemic could lead to increased forage production for other prey species such as moose (Alces americanus), which in turn could result in higher moose numbers and higher wolf (Canis lupus) numbers, which could lead to higher predation risk for caribou. The mountain pine beetle epidemic could also affect travel for both other prey and predators. The relative effects on the different species could result in different outcomes. For example, if mountain pine beetles result in treefall conditions that make travel slightly more difficult for caribou but significantly more difficult for wolves, caribou could potentially benefit from reduced predation risk. Conversely, if changes in snow conditions make winter travel more difficult for caribou but less difficult for wolves, predation risk could increase.



STATUS OF CURRENT RESEARCH

The Tweedsmuir-Entiako caribou population in west-central B.C. was the

first caribou population to experience the current mountain pine beetle epidemic. Since then, the Itcha-Ilgachuz, Charlotte Alplands and Rainbow Mountains populations in west-central B.C., and the Kennedy-Siding population in north-central B.C. have also been affected. Mountain pine beetles are also starting to affect other populations in north-central B.C.

Since 2001, four projects have been initiated to investigate the effects of

mountain pine beetles on woodland caribou populations in central British Columbia (Table 2). In addition, three projects have been initiated examining the effects of fire in mountain pine beetle-killed stands, and two other projects are continuing projects that have incorporated the effects of the mountain pine beetle epidemic (Table 2). Of the nine studies currently being conducted or recently completed, seven focus on habitat, one focuses on caribou habitat use, and one includes investigations on both habitat and habitat use.

The following sections summarize the current status of each project listed in

Table 2. Appendix 1 cross-references reports listed in Table 2 with references in the Reference section. Appendix 2 contains a list of full biogeoclimatic unit names for habitat-based studies.



Table 2. Summary of research on the effects of mountain pine beetles on caribou and caribou habitat. Project duration

Project Contact Caribou

Population MPB

Phases Focus Year

Initiated Year

Completed

Pre-MPB data Reports1

PROJECTS INITIATED TO INVESTIGATE THE EFFECTS OF THE MOUNTAIN PINE BEETLE EPIDEMIC ON CARIBOU The response of caribou terrestrial forage lichens to mountain pine beetles and forest harvesting in the East Ootsa and Entiako areas

Cichowski/ Williston

Tweedsmuir-Entiako

Red/ Green to

Grey

• Habitat 2001/02 Ongoing • Annual Reports o 2001/02 – Year 1 o 2002/03 – Year 2 o 2003/04 – Year 3 o 2005/06 – Year 5 o 2007/08 – Year 7 • Final Report 2005/06 (Phase 1)

The effects of a mountain pine beetle epidemic on Northern Caribou habitat use and movements

Cichowski Tweedsmuir-Entiako

Grey • Habitat Use 2005/06 2009/10 1983-2003

• Annual Summaries o 2005/06 – Year 1 o 2006/07 – Year 2 o 2007/08 – Year 3 o 2008/09 – Year 4 • Final Report 2009/10

Effects of a mountain pine beetle epidemic on forest floor vegetation dynamics and regeneration in the Itcha-Ilgachuz caribou winter range in the Quesnel TSA

Youds/ Armleder

Itcha-Ilgachuz (northeastern

portion of winter range)

Red/ Green to

Grey

• Habitat 2005/06 Ongoing • Annual Reports o 2005/06 – Year 1 o 2006/07 – Year 2 o 2008/09 – Year 4

Response of woodland caribou to partial retention logging of woodland caribou ranges attacked by mountain pine beetle

Seip Kennedy-Siding

Alive to Green to Red to Grey

• Habitat • Habitat Use • Harvesting

2006/07 Ongoing 2002-2006

• Annual Reports o 2006/07 – Year 1 o 2007/08 – Year 2 o 2008/09 – Year 3 o 2009/10 – Year 4

The response of caribou terrestrial forage lichens to fire in Entiako Provincial Park

Cichowski/ Batho

Tweedsmuir-Entiako

Fire in MPB Grey

• Habitat 2007/08 Ongoing • Annual Reports o 2007/08 – Year 1 o 2009/10 – Year 3

Identifying factors affecting the succession and availability of terrestrial lichen communities in the Omineca Region of North-central British Columbia: Rehabilitation of caribou winter range following attack by mountain pine beetle: UWR U-7-012

McNay Tweedsmuir-Entiako

Fire in MPB Grey

• Habitat 2008/09 Ongoing • Annual Reports o 2008/09 – Year 1



Table 2. Summary of research on the effects of mountain pine beetles on caribou and caribou habitat. Project duration

Project Contact Caribou

Population MPB

Phases Focus Year

Initiated Year

Completed

Pre-MPB data Reports1

Identifying factors affecting the succession and availability of terrestrial lichen communities in the Omineca Region of North-central British Columbia: Jackfish prescribed burn

McNay Wolverine Fire • Habitat 2009/10 Ongoing • Annual Report o 2009/10

Identifying factors affecting the succession and availability of terrestrial lichen communities in the Omineca Region of North-central British Columbia: Long-term lichen monitoring2

McNay Wolverine Chase Scott

Alive • Habitat 2008/091 Ongoing 2003 2005 2008

• Annual Report o 2008/09

ONGOING PROJECTS ON CARIBOU HABITAT WITH CONTROLS THAT WILL BE MEASURED FOR EFFECTS OF THE MOUNTAIN PINE BEETLE EPIDEMIC3

Itcha-Ilgachuz Alternative Silvicultural Systems: Satah Mountain pilot study

Waterhouse/ Armleder

Itcha/ Ilgachuz (western portion of

winter range)

Alive to Red/ Grey

• Habitat 1995/96 (2008)

Ongoing 1995 1996 1997 2000 2004

• A Pilot Study of Silvicultural Systems for Northern Caribou Winter Range: Lichen Response (Miège et al. 2001a) • Waterhouse et al. in press

Itcha-Ilgachuz Alternative Silvicultural Systems: Satah Mountain replicated trials



winter range)

Alive to Red/Grey

• Habitat 1995/96 (2008)

Ongoing 1995 1998 2000 2004

• Impact of partial cutting on lichen diversity in lodgepole pine forests on the Chilcotin Plateau of British Columbia (Miège et al 2001b) • Waterhouse et al. in press • Waterhouse 2011

Itcha-Ilgachuz Alternative Silvicultural Systems: Satah Mountain – adaptive management trials



winter range)

Alive to Red/Grey

• Habitat 1996/97 (2007)

Ongoing 1996 2000 2003

• Waterhouse et al. in press

Itcha-Ilgachuz Alternative Silvicultural Systems: Chezacut – adaptive management trials



winter range)

Alive • Habitat 1997/98

Ongoing 1997 2001 2003

•

1 Appendix 1 cross-references reports listed in Table 2 with the references in the Reference section 2 Although this project was initiated in 2008, it includes controls from an adaptive management project of forest harvesting; one control site was measured in 2003 and the other was

measured in 2005; in addition, three new sites were established in 2008 to monitor terrestrial lichen responses to the mountain pine beetle outbreak. 3 This set of projects was installed prior to the current mountain pine beetle outbreak to assess the effects of various silvicultural systems on caribou habitat. Plots in both the no-

harvest controls and partial cuts provide a continuous data set from pre-mountain pine beetle, through the early infestation (2003-2004) to maximum tree mortality (2008). Data collected include microclimate, regeneration growth and treefall rates. The years of pre-beetle data indicate major lichen, and vegetation measurements. For this project, this table lists reports that focus on analyses of lichen trends. Appendix 1 contains a full list of reports associated with the project.



The response of caribou terrestrial forage lichens to mountain pine beetles and forest harvesting in the East Ootsa and Entiako areas.

This project examines the effects of the mountain pine beetle epidemic on

terrestrial lichens and forest floor vegetation on the Tweedsmuir-Entiako caribou winter range. This study is based on 79 permanently marked plots that were established in 2001 in 4 biogeoclimatic subzones within 7 site series (SBPSmc/01a, SBPSmc/01b, SBPSmc/02, SBSdk/03, SBSmc2/01c, SBSmc2/02, ESSFmc/03). Of the 79 permanent plots, 56 were established in red/green mountain pine beetle-attacked stands, and 231 were established in clearcuts that had been recently harvested for mountain pine beetle management. Of the 56 plots established in attacked stands, 17 were earmarked for harvesting so that pre and post harvesting data could be collected; however, harvesting efforts followed the wave of mountain pine beetle attack and moved out of the study area, so those plots were never harvested. In each permanent plot, terrestrial lichens were photographed at 10 permanently marked photoplots. Other measurements at permanent plots included: stand density and composition; status of mountain pine beetle attack; regeneration; canopy openness (based on hemispherical photographs); and, coarse woody debris as an indicator of movement barriers. Plots were established in 2001 in mixed red and green attacked stands and re-measured in 2003, 2005 and 2007. In 2003, attacked trees were mostly in the red or grey attack phase, and by 2005, most trees were in the grey attack phase. In 2004, a wildfire burned 6 plots (4 mountain pine beetle attacked, 2 harvested) providing information on forest floor vegetation dynamics following fire in mountain pine beetle attacked stands.

Summary of Findings:

Overall, terrestrial lichen abundance decreased following mountain pine beetle attack, with a corresponding increase in dwarf shrubs, primarily kinnikinnick (Arctostaphylos uva-ursi). Terrestrial lichen abundance on all plots combined on mountain pine beetle affected forested ecosystems declined from 16% in 2001, to 13% in 2003 to 11% in 2005, to 10% in 2007, while kinnikinnick increased from 19% in 2001, to 30% in 2003, to 37% in 2005, to 39% in 2007 (Cichowski et al. 2008). Other dwarf shrubs that increased in abundance included twin-flower (Linnaea borealis) and crowberry (Empetrum nigrum). The decrease in lichen abundance and increase in kinnikinnick abundance was observed in 2003, but began to level off by 2007. In 2007, kinnikinnick die-back was observed on some plots and twin-flower increased on some of the plots with kinnikinnick die-back, especially in the SBSmc2/01c. In that site series, although kinnikinnick abundance leveled off between 2005 and 2007, total ground cover of vegetation continued to increase, primarily due to an increase in twin-flower. One exception to the overall trends was in the higher elevation ESSFmc/03, where there was no change in terrestrial lichen abundance between 2001 and

1 Initially, 24 permanent plots were established in harvested areas but one plot was disturbed by site preparation and was deleted from the sample.



2005, but a significant decrease between 2005 and 2007. This may have been due to the very low abundance of kinnikinnick in the ESSFmc/03, a greater volume of residual live trees, or a possible delay in the onset of the mountain pine beetle attack. The decline in terrestrial lichens in the ESSFmc/03 corresponded to a sharp increase in mosses and moderate increases in crowberry and twin-flower. Except in the ESSFmc/03, there were no significant changes in moss abundance between 2001 and 2007. Observations from a brief site visit in 2010 suggested that the kinnikinnick die-back had increased and was more widespread. On logged plots, kinnikinnick abundance increased from 2001 to 2007, but lichen abundance was variable. By 2007, tree fall rates in mountain pine beetle attacked stands were still low.


This project was initiated in 2005/06 on the Tweedsmuir-Entiako caribou population and focused on caribou habitat use based on GPS and VHF radio-collared caribou and winter snow tracking. By 2005/06, much of the winter range was in the grey phase of the mountain pine beetle epidemic. Caribou were radio-collared in March 2006, January 2007 and December 2007. Three years of fieldwork were conducted in 2006/07, 2007/08 and 2008/09 and a final report was prepared in 2009/10. Radio-collared caribou were located during monthly radio-telemetry flights in summer months (April to November) and bi-weekly flights in winter months (December to March). During winter months, 3-4 day site investigations sessions were conducted each month to assess foraging in different stand conditions (composition, mountain pine beetle attack status), and snow characteristics. Population status was assessed based on mortality rate of radio-collared caribou, and on calf recruitment rates determined during calf survival surveys. Cause of adult mortalities was investigated when feasible. Caribou locations and winter site investigations data from 1983 to 2003 (pre-MPB) were compiled and compared to data collected during the current study (MPB-Grey).


During the MPB-Grey attack stage, Tweedsmuir-Entiako caribou range use, seasonal movements, winter habitat use and winter feeding site selection patterns were consistent with patterns documented prior to the mountain pine beetle epidemic. In low elevation forests, caribou preferentially cratered for terrestrial lichens and selected craters in open canopy pine stands where terrestrial lichens were most abundant. Percent of MPB-Grey attacked trees in the stand did not influence crater site selection. In 2006/07, caribou use of terrestrial lichens declined by March in response to decreased snow penetrability. Annual variations in caribou winter habitat use and winter foraging were a function of annual variations in winter weather factors (snow accumulation, temperature) rather than a response to the mountain pine beetle epidemic. In 2006/07, an early



heavy snowfall and frequent freeze/thaw conditions resulted in a deep, consolidated snowpack by late winter. In 2007/08 and 2008/09, moderate snowfall throughout the winter and generally cool temperatures for most of the winter resulted in a relatively unconsolidated snowpack both years, allowing caribou to continue foraging for terrestrial lichens until late winter. Continued use of mature pine forests by Tweedsmuir-Entiako caribou on the Entiako winter range suggests that despite needle loss, mature pine forests in the Entiako are continuing to function as winter range and as winter habitat for the Tweedsmuir-Entiako caribou population. From 2006/07 to 2008/09, adult female mortality rates were moderate to high and calf recruitment did not appear to be sufficient to balance adult mortality.


This project was initiated in 2005/06 to examine the effects of mountain pine beetles on caribou habitat in the northeastern portion of the Itcha-Ilgachuz caribou winter range in the Quesnel Timber Supply Area (TSA) in the MSxv and SBPSdc biogeoclimatic subzones. The project includes 10 sample sites (9 MSxv, 1 SBPSdc), each containing 30 permanently marked plots. Seven sites were established in 2005 in mixed green and red attacked stands, and an additional three sites were established in 2006 in mixed green and red attacked stands. Methods for this project follow methods used to assess the effects of forest harvesting on terrestrial and arboreal lichens in other parts of the Itcha-Ilgachuz caribou winter range. Terrestrial lichen and competing vegetation abundance were measured at each permanent plot in a 2 m diameter aluminum hoop with an inlaid triangle. Other data collected included: mountain pine beetle status of canopy trees, regeneration, cones, windthrow, hemispherical canopy photographs, and, arboreal lichen abundance. Plots were established in 2005 and 2006 and re-measured in 2008. In 2005 and 2006, trees were mostly in the green or red phase of the attack. By 2008, most trees were in the grey stage of attack or in the later stages of red attack.


Terrestrial lichen abundance decreased and dwarf shrubs increased during the initial stages of the mountain pine beetle epidemic (2005 to 2008) in the Quesnel TSA (northeastern) portion of the Itcha-Ilgachuz caribou winter range. The primary dwarf shrubs that increased included kinnikinnick, twinflower and Vaccinium sp. The decrease in terrestrial lichen abundance was correlated with changes in overstory (MPB attack) and understory (competing vegetation) conditions. At the start of the study (2005 or 2006), terrestrial caribou forage lichen abundance was significantly positively correlated with the percentage of full sunlight reaching the forest floor. However, the greater the increase in light transmission following the mountain pine beetle epidemic, the greater the



decrease in terrestrial lichen abundance. Data for arboreal lichen abundance was inconclusive.


This project was initiated in 2006/07 on the Kennedy-Siding caribou population to assess the effects of mountain pine beetles and partial retention logging for mountain pine beetles on both caribou habitat and caribou habitat use. Fieldwork was conducted in 2006/07, 2007/08, 2008/09, 2009/10 and has continued into 2010/11 (Seip and Jones 2007, 2008, 2009, 2010). Most trees were attacked by mountain pine beetles in 2006 (green attack phase) and progressed to red attack in 2007. In 2008, most trees were in the grey phase of the attack and by 2009, over 90% of the canopy had been attacked. The caribou habitat use portion of the project was based on GPS radio-collared caribou, bi-weekly radio-telemetry flights and winter tracking investigations. Mortality rates of radio-collared caribou and calf recruitment rates based on calf survival surveys were also assessed. Caribou were captured and collared in March 2006, 2007, 2008, 2009 and 2010. The caribou habitat portion of the study was based on snow conditions, and terrestrial and arboreal lichen abundance on the low elevation winter range in the SBSwk2. Snow conditions were measured at permanent sample points in mountain pine beetle killed stands, a salvage harvesting block (clearcut with leave patches cut in 2005) and in a pre-existing clearcut (cut in the 1990s). Terrestrial and arboreal lichen abundance were measured at permanently marked plots along transects in mountain pine beetle killed stands, salvage harvested areas and in the pre-existing clearcut. Terrestrial lichen and other vegetation abundance were measured at each 10- x 10-cm intercept within a square-meter plot.


Initially, terrestrial lichen (Cladina sp.) abundance was greatest in the old cutblock (47%), followed by the pine forest (24%); terrestrial lichen abundance was lowest in the salvage block (11%). From 2006 to 2009, Cladina sp. abundance had decreased in the pine forest and increased in the salvage block so that by 2009, Cladina sp. abundance was similar on the two sites (12% in forest; 14% in salvage block). Abundance of dwarf shrubs (Arctostaphylos uva-ursi, Vaccinium myrtilloides) increased in the pine forest from 19% in 2006 to 28% in 2009 and in the salvage block from 26% in 2006 to 33% in 2009, and decreased in the clearcut from 23% in 2007 to 17% in 2009.

As observed prior to the mountain pine beetle epidemic, all of the collared

caribou used the Kennedy Siding low elevation winter range during early winter each year, but left the low elevation winter range earlier in 2006/07 and 2007/08 compared to other years, presumably due to deep snow conditions developing earlier in the winter than usual. Caribou were located primarily in the pine



forest, and to a lesser extent in the clearcut and salvage block in all years. In 2009/10, use of the clearcut was higher and use of the pine forest was lower than in previous years.

In 2006/07, 2007/08 and 2008/09, caribou foraged in the pine forest, salvage

block and clearcut when snow was shallow, then foraged only in the pine forest when snow depths exceeded 40-50 cm. In 2009/10, caribou foraged in the salvage block, clearcut and pine forest throughout the entire duration of use of the low elevation winter range. In all 4 years, snow depths were typically similar in the pine forest and the clearcut while the caribou were using their low elevation winter range. Snow hardness was lower in the pine forest than in the clearcut, except in 2009/10 when it was similar in both habitats. In the pine forest, caribou generally fed on both terrestrial lichen and arboreal lichens when snow was shallow, but switched exclusively to arboreal lichens when snow depth and hardness increased.

Caribou continued to use grey attacked forests, which continued to provide

terrestrial and arboreal lichens for forage, and usually had softer snow conditions that facilitated cratering for lichens. Although feeding on terrestrial lichens in the forest may have declined since mountain pine beetle attack, over half the caribou locations were in the dead pine forest and caribou continued to feed on arboreal lichens in the forest. Recent salvage-logged portions of the low elevation winter range supported some terrestrial lichens, while the older clearcut supported abundant terrestrial lichens, suggesting that terrestrial lichens will recover quickly in salvage-logged blocks.

From 2006/07 to 2009/10, adult female mortality rates were low to moderate

and calf recruitment appeared sufficient to balance adult mortality (D. Seip, unpubl. data). In 2010/11, adult female mortality was high and calf recruitment was low.

The response of caribou terrestrial forage lichens to fire in Entiako Provincial Park

This project examines the effects of fire in mountain pine beetle-killed stands

on terrestrial lichens and other forest floor vegetation on the Tweedsmuir-Entiako caribou winter range. Of particular interest is how quickly terrestrial lichens will re-establish following fire, and how well kinnikinnick, which had increased substantially following mountain pine beetle attack, survived fire. This study is based on 15 permanently marked plots that were established in 2007 in 3 biogeoclimatic subzone/site series (SBPSmc/01a, SBPSmc/01b, SBPSmc/02), 1 year following a wildfire. The wildfire burned 700 ha of mountain pine beetle-killed forest in the grey phase of the epidemic in 2006. Methods for this project followed methods used in the study on the effects of mountain pine beetles and forest harvesting on terrestrial lichens in the East



Ootsa and Entiako areas. In each permanent plot, terrestrial lichens were photographed at 10 permanently marked photoplots. Other measurements at permanent plots included: stand density and composition; burn severity; regeneration; canopy openness (based on hemispherical photographs); and coarse woody debris as an indicator of movement barriers. Plots were established in 2007 and re-measured in 2009.


Terrestrial lichens and kinnikinnick were nearly absent on plots in 2007 and 2009. In 2007, Cladina rangiferina and kinnikinnick were both recorded in the same 2 plots, and were survivors in unburned patches. By 2009, kinnikinnick had resprouted on four more plots and had increased on one of the two plots where it was present in 2007. Overall, kinnikinnick abundance was still low in 2009 (3 years following fire). Lodgepole pine regeneration was present on 14 of 15 plots, spruce regeneration was found on 7 of 15 plots and aspen regeneration was present on 6 plots. Post-disturbance regeneration densities were higher in mountain pine beetle killed forests following fire than in mountain pine beetle killed forests without fire. The abundance of forbs, gramminoids (grasses and sedges), dwarf shrubs and shrubs increased from 2007 to 2009. Forbs were the most abundant vascular plant group on SBPSmc/01a (mesic) plots while shrubs and dwarf shrubs were the most abundant vascular plant group on SBPSmc/02 (xeric) plots. The most common vascular plants on all plots were bunchberry (Cornus canadensis), fireweed (Epilobium angustifolium), bluejoint reedgrass (Calamagrostis canadensis), bronze sedge (Carex aenea), twin-flower, dwarf blueberry (Vaccinium caespitosum), prickly rose (Rosa acicularis), soopolallie (Shepherdia canadensis) and birch-leaved spirea (Spirea betulifolia). Coverage of all those plants increased between 2007 and 2009 except for birch-leaved spirea, which remained the same or decreased. Bryophytes also increased on all plots between 2007 and 2009.


This project was initiated in 2008/09 to investigate the effects of prescribed

burning in mountain pine beetle-killed forests on terrestrial lichens and competing vegetation on the Tweedsmuir-Entiako caribou winter range in the low elevation Ungulate Winter Range (UWR) U-7-012. In 2008/09, pre-burn monitoring plots were established in a red/grey mountain pine beetle attacked stand in the SBPSmc. Each plot contained 4-6 quadrats (0.71m x 0.71m square), which were used to measure % cover of herbs, mosses and lichens. Other measurements included: stand density, composition and mountain pine beetle status; regeneration; canopy closure using a concave densiometer; and coarse woody debris. Prior to the burn, trees were knocked down with a cable strung



between two bulldozers in March 2009 to allow the fire to carry in the prescribed burn area when fire hazard in the adjacent area was low. The prescribed burn was conducted on September 4, 2009 and post-burn measurements were collected in summer 2010 but have not yet been analyzed.


This project was initiated in 2009/10 to provide a replicate for the UWR U-7-

012 prescribed burn, which is investigating the effects of prescribed burning in mountain pine beetle-killed forests on terrestrial lichens and competing vegetation. The Jackfish prescribed burn site is located in the BWBSdk1 biogeoclimatic subzone/variant on the Wolverine caribou range. In 2009/10, a burn plan was developed and pre-burn monitoring plots were established in a mostly live pine stand with <10% green and red mountain pine beetle attack. Each plot contained quadrats (0.71m x 0.71m square), which were used to measure % cover of herbs, mosses and lichens. Other measurements included: stand density, composition and mountain pine beetle status; regeneration; canopy closure using a concave densiometer; and coarse woody debris. Knockdown of trees was proposed for 2010 and the prescribed burn was proposed for fall 2010. Neither the knockdown of trees nor the prescribed burn were conducted in 2010.

Identifying factors affecting the succession and availability of terrestrial lichen communities in the Omineca Region of North-central British Columbia: Long-term lichen monitoring

This project was initiated in 2008/09 to assess the impacts of the mountain

pine beetle epidemic on terrestrial lichens in a range of biogeoclimatic zones. Two sites used in this study were controls that were established for an adaptive management trial for forest harvesting in the SBSmk2 biogeoclimatic subzone/variant (Sulyma and Sulyma 2006). One site was established in 2003 and the other site was established in 2005. Three additional sites were established in the BWBSdk1 biogeoclimatic subzone/variant in 2008 to broaden the range of ecological conditions being assessed. At the time of establishment, pine stands at all sites were still alive. Permanently marked quadrats (0.71m x 0.71m square) were established in random locations along transects. Quadrats were used to measure % cover of herbs, mosses and lichens. Other measurements included: stand density, composition and mountain pine status; and regeneration. No post-mountain pine beetle sampling has taken place.



Itcha-Ilgachuz Alternative Silvicultural Systems This project was initiated in 1994 to investigate the impacts of various

silvicultural systems: irregular group shelterwood (50% retention), group selection (70% retention), clearcutting with high retention (30% dispersed and islands) and clearcuts, on terrestrial and arboreal lichens in the western (Williams Lake TSA) portion of the Itcha-Ilgachuz caribou range in the MSxv and SBPSxc biogeoclimatic subzones. It consists of four studies:

• Satah Mountain pilot study; • Satah Mountain replicated trial; • Satah Mountain adaptive management trial; and, • Chezacut adaptive management trial. Although the focus of these studies was on forest harvesting, they are now

also being used to assess the impacts of the mountain pine beetle epidemic on terrestrial and arboreal lichens. The studies are based on permanently marked plots. Terrestrial lichen and competing vegetation abundance (by layer and species) were measured at each permanent plot in a 2 m diameter aluminum hoop with an inlaid triangle. Within the plots, data was also collected for natural regeneration, cones, canopy closure and light transmittance (hemispherical photos), and arboreal lichen abundance.

The Satah Mountain pilot study was initiated in 1994 with forest harvesting

conducted in April 1995 and measurements conducted post-harvesting in 1995, 1996, 1997, 2000, 2004 and 2008. The Satah Mountain replicated trial was initiated in 1995 with pre-harvesting measurements conducted in 1995, harvesting conducted in April 1996, and post-harvesting measurements conducted in 1998, 2000, 2004 and 2008. This trial includes five blocks with four treatments where data is collected on treefall, microclimate (air and soil temperature, rainfall and soil moisture), planted stock, long-term site productivity (woody debris), growth and yield, and breeding birds. The Satah Mountain adaptive management trial was initiated in 1996 with pre-harvest measurements conducted in 1996, harvesting conducted in 1997-1998, and post-harvesting measurements conducted in 2000, 2003 and 2007. The Chezacut adaptive management trial was initiated in 1997 with pre-harvest measurements conducted in 1997, harvesting conducted in March 1999, and post-harvesting measurements conducted in 2001 and 2003. All measurements conducted in 2008 were in the red/grey phase of the mountain pine beetle epidemic. Mountain pine beetle mortality was 5% in 2003, 21% in 2004, 49% in 2006 and 68% in 2008 on the no-harvest control plots (Waterhouse 2011).


On no-harvest control plots at the Satah Mountain replicated trial, terrestrial lichen abundance declined from pre-mountain pine beetle cover of 11.3% in 2004 to 9.7% in 2008, representing a 14% relative drop in cover, but the decrease was not significant (Waterhouse 2011). Changes were variable between blocks with



smaller declines in the two MSxv blocks than in two of the SBPSxc blocks, and an increase in lichen cover in one SBPSxc block with a very open canopy.

For partial cuts, in the group selection treatments (30% of the overstory cut in 15 m diameter openings) lichen cover decreased from 10.3% in 2004 to 7.4% in 2008 (Waterhouse 2011). In the irregular group shelterwood treatments (where about 50% of the overstory was cut in 1996) lichen cover was 7.3% in 2004 and 7.8% in 2008 (Waterhouse 2011). Within the irregular group shelterwood treatment, lichen cover in openings increased from 7.5% in 2004 to 9.5% in 2008 while lichen cover in the residual forest remained relatively constant (7.6% in 2004 and 7.7% in 2008).

Moss cover remained unchanged pre- and post- beetle in all treatments. Dwarf shrubs and herbs increased in both partial cuts and no-harvest controls, with the greatest relative increase in the no-harvest controls (Waterhouse 2011).

The density of natural regeneration stems increased in both the no-harvest control and partial cut treatments from 2004 to 2008 (Waterhouse 2011). Regeneration densities were lower on the higher elevation MSxv sites (average densities of 2284 to 3333 stems/ha in 2008) than on the lower elevation drier SBPSxc sites (average densities of 6073 to 11308 stems/ha in 2008). Height growth rates of regeneration in the small openings in partial cuts from 2006 to 2008 was greater than that of regeneration growing in no harvest control plots and in the forested portions of the partial cuts (Waterhouse 2011).

Average annual treefall rates were similar prior to and following mountain pine beetle attack (Waterhouse 2009). The difference between soil water content in clearcuts versus partial cuts decreased from 1999 to 2010, but it is not known if this decrease was due to reduced evapotranspiration in the forested portions of the partial cuts because of mortality due to the mountain pine beetle epidemic or to increased water use in clearcuts by regenerating pine stands and other regeneration (Sagar 2011).



SYNOPSIS OF RESEARCH RESULTS

EFFECTS OF MOUNTAIN PINE BEETLES ON CARIBOU HABITAT Currently, information is available from four projects assessing the effects of

mountain pine beetles on caribou habitat (Cichowski et al. 2008, Cichowski et al. 2009, Seip and Jones 2010, Waterhouse 2011). Habitat characteristics addressed by these projects include: intensity of attack, terrestrial lichen abundance, competing vegetation abundance, light availability, regeneration, coarse woody debris, snow conditions, and arboreal lichen abundance.

Mountain pine beetle attack intensity was moderate to high in all four areas.

On the Tweedsmuir-Entiako winter range, mortality due to mountain pine beetles averaged 78-96% of pine trees >7.5 cm dbh. Lodgepole pine was the dominant tree on all sites and by 2005, 80-90% of pine trees were killed by mountain pine beetles on most sites except for smaller diameter stands that experienced 74% mortality (Cichowski et al. 2008). On the Kennedy-Siding winter range, all trees on plots were lodgepole pine and 89% of pine canopy trees were killed (Seip and Jones 2010). Mountain pine beetle mortality was slightly less on the Itcha-Ilgachuz winter range with 46-75% of pine trees killed in the northeastern (Quesnel TSA) portion (Cichowski et al. 2009), and 68% of pine trees killed in the western (Williams Lake TSA) portion (Waterhouse 2011).

The most dramatic change in caribou habitat following mountain pine beetles

was a decrease in terrestrial lichen abundance, and an increase in competing vegetation, primarily dwarf shrubs. The species of dwarf shrubs that increased varied by study area. On the Tweedsmuir-Entiako caribou winter range, kinnikinnick was the primary dwarf shrub that increased in abundance; however, twin-flower and crowberry had also increased on some sites (Cichowski et al. 2008). In the northeastern (Quesnel TSA) portion of the Itcha-Ilgachuz caribou winter range, twin-flower, kinnikinnick and Vaccinium sp. (V. scoparium, V. caespitosum) all contributed to the increase in dwarf shrubs (Cichowski et al. 2009). In the western (Williams Lake TSA) portion, the primary dwarf shrubs that increased were kinnikinnick in the SPPSxc, and twinflower, crowberry and V. scoparium in the MSxv (Waterhouse and Armleder, unpubl. data). On the Kennedy-Siding caribou winter range, Vaccinium myrtilloides and to a lesser extent kinnikinnick, were the primary species of dwarf shrubs that increased (Seip and Jones 2010).

On the Tweedsmuir-Entiako winter range, although the decline in terrestrial

lichen abundance was negatively correlated to mountain pine beetle mortality, it was better correlated to abundance of competing species (kinnikinnick, feathermoss, twin-flower, crowberry). Abundance of competing species was the single best predictor of terrestrial lichen abundance and was strongly associated



with terrestrial lichen declines (Cichowski et al. 2008). The degree of mountain pine beetle attack also contributed to the decline in terrestrial lichens (over and above the increase in competing species); the more mountain pine beetle mortality, the greater the decrease in terrestrial lichen cover.

The response by dwarf shrubs and terrestrial lichens was rapid, occurring

within a few years of attack and continuing into the grey attack stage. On the Tweedsmuir-Entiako winter range, changes began levelling off by 2007, about 7-8 years after attack (Cichowski et al. 2008). Some of the levelling off may have been due to kinnikinnick die-back observed on some of the plots. Although the kinnikinnick increase levelled off by 2007, total vegetation cover continued to increase in 2007, partly due to an increase in twin-flower. Casual observations on some of the plots in 2010 suggested that kinnikinnick die-back continued and was more widespread (pers. observ.). Cichowski et al. (2008) suggested that the increase in dwarf shrubs was likely due to increased availability of resources (soil nutrients, soil moisture, light) that were freed up when canopy trees died.

On the Tweedsmuir-Entiako winter range, lichen abundance on all plots

combined on mountain pine beetle affected forested ecosystems declined from 16% in 2001, to 13% in 2003 to 11% in 2005, to 10% in 2007 (Cichowski et al. 2008). This represents a 38% relative decline in lichen abundance in a 6 year period from the mixed green/red attack stage to the grey attack stage. On the Itcha-Ilgachuz winter range (Quesnel TSA portion), lichen abundance for all plots decreased from 20.5% from 2005/06 to 16.4% in 2008, for a relative decrease of 20% in 2-3 years from the mixed green/red attack stage to the mixed red/grey attack stage (Cichowski et al. 2009). In the western portion of the Itcha-Ilgachuz winter range, terrestrial lichen abundance declined from an average of 11.3% in 2004 to 9.7% in 2008 in attacked stands, representing a relative decrease of 14% over about 4 years (Waterhouse 2011). For the Tweedsmuir-Entiako and Itcha-Ilgachuz (Quesnel TSA) study areas, the % decrease in terrestrial lichen abundance was correlated to the % of the canopy killed by mountain pine beetles; the more mountain pine beetle mortality, the greater the decrease in lichen abundance. The Kennedy-Siding caribou winter range experienced the greatest decline in terrestrial lichen abundance. On that winter range, lichen abundance decreased from 24% in 2006, to 17% in 2007, to 12% in 2008 and 2009 (Seip and Jones 2007, 2008, 2009, 2010) for a relative decrease of 50% over a 4 year period from green attack to grey attack. Similar to the Tweedsmuir-Entiako caribou winter range, the greatest decline on the Kennedy-Siding winter range occurred soon after mountain pine beetle attack and began leveling off in the grey phase of the attack.

The greater decrease in terrestrial lichen abundance and the faster

progression to the leveling off of the decrease in terrestrial lichen abundance on the Kennedy-Siding caribou winter range relative to the Tweedsmuir-Entiako winter range may have been due to several factors, including: a more synchronous mountain pine beetle attack, a greater intensity of attack with fewer



remaining live trees of all species, and, a narrower range of ecological conditions. For the Tweedsmuir-Entiako study, sites were selected across a spectrum of ecological conditions, including higher elevation sites in the ESSFmc, and spread out over an area of almost 100,000 ha. Sites in the Kennedy-Siding study were relatively uniform in ecological condition and were all located within a 1500 ha area. On the Tweedsmuir-Entiako winter range, up to 25% of the canopy on plots consisted of other species, primarily hybrid white spruce (Pinus glauca x engelmanii) and subalpine fir (Abies lasiocarpa), whereas the tree canopy in plots on the Kennedy-Siding winter range consisted almost entirely of lodgepole pine. Because the Tweedsmuir-Entiako study covered a wider variety of ecosystems and canopy species over a larger area, greater variability in timing and intensity of attack, and residual canopy, may have extended the duration and dampened the degree of the terrestrial lichen decline in comparison to the Kennedy-Siding study.

In the northeastern (Quesnel TSA) portion of the Itcha-Ilgachuz winter range,

terrestrial lichen abundance at the beginning of the study was correlated with % light transmission to the forest floor; the greater the amount of light, the greater the abundance of lichens (Cichowski et al. 2008). Similarly, in the western (Williams Lake TSA) portion of the winter range, terrestrial lichen abundance prior to mountain pine beetle attack was greater in areas with less overhead canopy (Waterhouse et al. in press). In the northeastern (Quesnel TSA) portion of the winter range, plots with high initial mountain pine beetle attack underwent greater decreases in terrestrial lichen abundance. Plots with a greater increase in light transmission experienced a greater decrease in terrestrial lichen abundance; and the greater the increase in dwarf shrubs, the greater the decrease in terrestrial lichens. Waterhouse (2011) suggests that the initial decline of lichen in the western (Williams Lake TSA) portion of the Itcha-Ilgachuz winter range may be due to increased light directly affecting lichens adapted to low light conditions under the tree canopy.

Although overall lichen abundance declined from 2001 to 2007 on the

Tweedsmuir-Entiako winter range and the rate of decline slowed in the last two years of the study, decline trends for individual ecosystems varied (Cichowski et al. 2008). The SBSmc2/01c did not appear to be leveling off by 2007. In the ESSFmc/03, the trend differed in that there was no change in abundance from 2001 to 2005, then a steep decline from 2005 to 2007. This may have been due to less kinnikinnick on the site, greater residual live trees continuing to use resources, and a greater proportion of green attacked trees at the beginning of the study resulting in a delay in decrease in terrestrial lichen abundance. Also, from 2005 to 2007, mosses, crowberry and twin-flower increased. On the SBSmc2/01c, although the increase in kinnikinnick abundance slowed between 2005 and 2007, total vegetation continued to increase, mainly due to twin-flower. This increase corresponded with continued decrease of terrestrial lichens.



There was no detectable change in moss cover from 2001 to 2007 on the Tweedsmuir-Entiako caribou winter range, except on ESSFmc/03 sites where moss cover increased (Cichowski et al. 2008). On the Kennedy-Siding caribou winter range, moss cover decreased in mountain pine beetle killed stands from 2006 to 2009 (Seip and Jones 2010). Moss cover also decreased in mountain pine beetle killed stands in the northeastern (Quesnel TSA) portion of the Itcha-Ilgachuz winter range during the first few years following attack (Cichowski et al. 2009), whereas it remained unchanged in the western (Williams Lake TSA) portion (Waterhouse 2011).

In clearcuts on the Tweedsmuir-Entiako winter range, there was a general

increase in terrestrial lichen abundance on harvested plots following mountain pine beetle attack, but because of significant variability from year to year, there were no significant differences between ecosystems and no significant time trends between ecosystems (Cichowski et al. 2008). Kinnikinnick and other ground vegetation generally increased and moss decreased in abundance. On the Kennedy-Siding caribou winter range, terrestrial lichen abundance increased in the salvage block (cut in 2005) between 2006 and 2009 and decreased in the old cutblock (clearcut in 1990s) between 2007 and 2009 (Seip and Jones 2010). Kinnikinnick and Vaccinium myrtilloides both increased on the salvage block from 2006 to 2009, while kinnikinnick decreased on the old cutblock, and Vaccinnium myrtilloides and moss increased slightly on the old cutblock from 2007 to 2009. In clearcuts in the western (Williams Lake TSA) portion of the Itcha-Ilgachuz caribou winter range, terrestrial lichen abundance remained relatively unchanged while abundance of dwarf shrubs and herbaceous vegetation increased slightly (Waterhouse 2011). All three studies established permanent plots in clearcuts following forest harvesting so no pre-harvesting data was available to assess initial changes in lichen or other vegetation abundance following salvage harvesting.

In partial cuts in the western (Williams Lake TSA) portion of the Itcha-

Ilgachuz winter range, both terrestrial lichens and dwarf shrubs increased after most mountain pine beetle mortality occurred (Waterhouse 2011). While terrestrial lichen abundance increased in openings in the partial cuts, abundance in the residual forest remained the same.

On burned sites on the Tweedsmuir-Entiako winter range, the dominant

vegetation species were twin-flower, Vaccinim caespitosum, Vaccinium membranaceum, soopolalie, sedges (Carex sp.), fireweed (Epilobium angustifolium) and bunchberry (Cornus canadensis). Although kinnikinnick was largely absent from fully burned plots in 2005, some re-sprouting was observed on those plots in 2007, and kinnikinnick cover on the partially burned plot was 30% (Cichowski et al. 2008). Resprouting of kinnikinnick was also observed in some plots in the Entiako Fire (Cichowski and de Groot 2010). Originally, it was expected that kinnikinnick would be consumed during fire disturbance and that it would have



to re-establish through seed dispersal, rather than re-sprouting. Twin-flower had the most aggressive growth form following fire.

No change in arboreal lichen abundance has yet been detected on either the

Itcha-Ilgachuz winter range (Quesnel TSA portion) or the Kennedy-Siding winter range (Cichowski et al. 2009, Seip and Jones 2010). If terrestrial lichen abundance continues to decline, caribou could potentially increase their use of arboreal lichens. Eventually, when trees fall, arboreal lichens on the whole tree, rather than just the lower portion, will become available to caribou. However, arboreal lichens become very sparse on fallen trees within a year of tree-fall, especially if they become covered by snow during winter (D. Seip, pers. comm.).

Pine seedlings are the primary regeneration species in all four study areas but

no substantial increase in regeneration has occurred in mountain pine beetle-killed stands in any of the study areas, other that the western (Williams Lake TSA) portion of the Itcha-Ilgachuz winter range (Waterhouse 2011). Unlike fire, mountain pine beetle disturbance may not result in dense second growth pine stands that could potentially affect caribou movements, but will likely progress to multi-aged stands with varying densities of pine, spruce and fir. However, there is sufficient regeneration at most sites to develop into a forest.

Coarse woody debris levels were similar throughout the study periods in all

four study areas. On the Tweedsmuir-Entiako caribou winter range, although coarse woody debris levels were higher on harvested plots than on mountain pine beetle-killed plots, they were generally low on all plots and did not affect caribou movements (Cichowski et al. 2008). In the western (Williams Lake TSA) portion of the Itcha-Ilgachuz winter range, average annual treefall rates were similar prior to and following mountain pine beetle attack (Waterhouse 2009).

On the Kennedy-Siding caribou winter range, snow depth was typically

similar in the forest and the clearcut in early winter when caribou were on their low elevation winter range (Seip and Jones 2007, 2008, 2009, 2010). Snow hardness was lower in the pine forest than in the clearcut, except in 2009/10, when snow hardness was variable, but mostly similar in both habitats. In the pine forest, caribou generally fed on both terrestrial lichen and arboreal lichens when snow was shallow, but switched exclusively to arboreal lichens when snow depth and hardness increased.

EFFECTS OF MOUNTAIN PINE BEETLES ON CARIBOU HABITAT USE Changes in habitat resulting from mountain pine beetle attack could

potentially affect caribou use of attacked pine stands (see Potential Effects of Mountain Pine Beetle Use on Caribou Ecology). Terrestrial lichens declined following mountain pine beetle attack on three caribou winter ranges in central British Columbia (Cichowski et al. 2008, Cichowski et al. 2009, Seip and Jones



2010, Waterhouse 2011). Eventual needle loss by the grey phase of the attack could potentially lead to changes in snow condition (snow depth, snow hardness). Consequently, changes in snow conditions and reduced terrestrial lichen abundance could lead to changes in winter foraging strategy.

Despite habitat changes resulting from mountain pine beetle attack,

Tweedsmuir-Entiako caribou range use, seasonal movements, winter habitat use and winter feeding site selection patterns during the grey attack phase were consistent with patterns documented prior to the mountain pine beetle epidemic (Cichowski 2010, Cichowski and MacLean 2005). Caribou continued to use the same summer and winter ranges, and continued to use low elevation pine forests (now in the grey attack phase) during winter, where they preferred to crater for terrestrial lichens (Cichowski 2010). Caribou also used arboreal lichens throughout the winter, especially when snow conditions precluded use of terrestrial lichens. Annual variations in caribou winter habitat use and foraging were a function of annual variations in winter weather factors (snow accumulation, temperature) rather than a response to the mountain pine beetle epidemic (Cichowski 2010).

On the Kennedy-Siding caribou winter range, caribou continued to use their

low elevation winter range during early winter throughout the green, red and grey attack stages of the mountain pine beetle epidemic (Seip and Jones 2007, 2008, 2009, 2010). Prior to mountain pine beetle attack, in early winter caribou used low elevation winter range where they preferred to forage on terrestrial lichens in both a recent clearcut (cut in the 1990s) and in pine forests when snow was shallow and light (Jones 2007). Caribou stopped using the clearcut when snow depth and hardness increased and focussed their foraging on terrestrial and arboreal lichens in the pine forest. When snow hardness further increased, caribou left the low elevation winter range to use alpine and subalpine habitat for the duration of the winter. Following mountain pine beetle attack, this pattern of use continued (Seip and Jones 2007, 2008, 2009, 2010). In 2009/10, caribou left the low elevation winter range before snow depth and hardness reached levels that triggered migration to higher elevations in previous years (Seip and Jones 2010). However, in 2010/11, caribou exhibited typical low elevation winter habitat use and migration behaviour (D. Seip, pers. comm.). After a windstorm in December 2010, which resulted in scattered blowdown on the low elevation winter range, caribou focussed their foraging on arboreal lichens on fallen trees (D. Seip, pers. comm.).

Because winter habitat use and foraging patterns differ between populations,

some differences in how caribou respond to the mountain pine beetle epidemic may also occur. On the Tweedsmuir-Entiako caribou winter range, caribou forage in low elevation habitat throughout the winter (Cichowski 2010, Cichowski and MacLean 2005). During some years, a portion of the population moves into subalpine and alpine habitat in response to snow conditions at lower elevations (Cichowski 2010). Although some caribou move to higher elevations,



the majority of caribou usually stay at lower elevations and forage for arboreal lichens during those years. In contrast, on the Kennedy-Siding caribou winter range, caribou typically forage for terrestrial lichens at lower elevations in early winter, and by December or January each year, move to higher elevation subalpine and alpine habitat (Jones 2007, Seip and Jones 2007, 2008, 2009, 2010). The common feature of both winter range use patterns is that caribou prefer to forage on terrestrial lichens until snow conditions make it difficult for them to do so. Initially, both caribou populations showed the same response to the mountain pine beetle epidemic – they did not significantly alter their winter habitat use and foraging strategies.

On the Itcha-Ilgachuz caribou winter range, some caribou winter in lower

elevation forests, while other caribou winter in subalpine and alpine habitat (Cichowski 1993, Young and Roorda 1999). On the low elevation winter range, caribou forage for terrestrial lichens in large fescue-lichen meadows and in mature pine forests during early winter. In mid and late winter, caribou use primarily mature pine stands. Although no radio-collared caribou studies have yet been conducted on the effects of the mountain pine beetle epidemic on Itcha-Ilgachuz caribou habitat use, a late winter survey was attempted in March 2010. No caribou were seen on the winter range during that survey, despite over 5 hours of search effort (N. Freeman, pers. comm.). It is unknown whether the lack of caribou seen during the survey was due to poor sighting conditions, a shift in range use due to snow conditions, a shift in range due to the effects of the mountain pine beetle epidemic, or some other factor.

EFFECTS OF MOUNTAIN PINE BEETLES ON CARIBOU POPULATION DYNAMICS

Both the Tweedsmuir-Entiako and Kennedy-Siding caribou studies included

calf recruitment surveys and adult female mortality rate; otherwise, no other information is available on the effects of mountain pine beetles on population dynamics. For the Tweedsmuir-Entiako caribou population, from 2006/07 to 2008/09, adult female mortality rates were moderate to high and calf recruitment did not appear to be sufficient to balance adult mortality (Cichowski 2010). The general declining population trend during the 3-year study was consistent with the population trend prior to the mountain pine beetle epidemic. For the Kennedy-Siding caribou population, from 2006/07 to 2010/11, adult female mortality rates were low to moderate and calf recruitment appeared sufficient to balance adult mortality, except in 2010/11 when the adult female mortality rate was high and calf recruitment was low (D. Seip, unpubl. data). In 2010/11 the adult female mortalities occurred in high elevation habitat during summer, so the increase in mortalities that year was not related to the mountain pine beetle epidemic (D. Seip, unpubl. data).



Although terrestrial lichen abundance declined on both the Tweedsmuir-Entiako and Kennedy-Siding caribou winter ranges following mountain pine beetle attack, caribou numbers are not expected to respond to the reduction in food since caribou populations are limited by predation and persist at densities far below food carry capacity (Seip and Cichowski 1996). However, any changes to habitat that could potentially affect predation risk could affect caribou population dynamics. Changes to predation risk could result from changes in: other prey abundance/presence; predator abundance/presence; and/or, predator efficiency on caribou ranges (see Potential Effects of Mountain Pine Beetles on Caribou Ecology).

No information is available on changes to predator or other prey abundance

or predator efficiency in response to a mountain pine beetle epidemic. Observations during winter fieldwork suggest that moose and wolves continue to use the Tweedsmuir-Entiako caribou winter range but changes in abundance or ease of travel have not been assessed. In Utah, Stone (1995) documented a positive relationship between moose use of mountain pine beetle attacked stands and degree of attack, based on fecal pellet counts in 1 ha plots. Moose use was higher in heavily attacked stands where grasses were abundant. In British Columbia, abundance of grasses was low and did not increase substantially following mountain pine beetle attack on caribou ranges (Cichowski et al. 2008, Cichowski et al. 2009, Seip and Jones 2010). Moose browse species were also low or non-existent in permanent plots on all three ranges. Because dwarf shrubs present prior to mountain pine beetle attack increased in abundance in response to increased site resources (i.e. light, nutrients, moisture), and because the sites did not support moose browse prior to the mountain pine beetle epidemic, it is unlikely that moose browse species will colonize those sites and capitalize on the increase in resources.

Although we have been learning more about how caribou are responding to

the mountain pine beetle epidemic, it will also be important to collect information on responses of other prey and predators to understand the full implications of the mountain pine beetle epidemic on woodland caribou populations.



KNOWLEDGE GAPS

Table 3 summarizes information collected through the existing studies on the

effects of mountain pine beetles on caribou, and information that will be collected through ongoing studies where control plots will be used to monitor the effects of the mountain pine beetle epidemic on lichens.

In all four studies assessing the effects of the mountain pine beetle epidemic

on terrestrial lichens, terrestrial lichens declined following mountain pine beetle attack while competing forest floor vegetation increased (Cichowski et al. 2008, Cichowski et al. 2009, Seip and Jones 2010, Waterhouse 2011). The response of forest floor vegetation varies with ecological conditions (Cichowski et al. 2008); therefore, effects of mountain pine beetles should be assessed across a wide range of ecological conditions, including on ranges in different biogeoclimatic zones/subzones, and on sites that do not support terrestrial lichens. The research that is currently underway focuses on high value terrestrial lichen sites. It will be important to also assess vegetation responses on sites that do not support terrestrial lichens, especially where forage for other prey species could increase, which in turn could lead to increased predation risk for caribou.

Although the decrease in terrestrial lichen abundance and increase in

competing vegetation abundance has been well documented (Cichowski et al. 2008, Cichowski et al. 2009, Seip and Jones 2010, Waterhouse 2011), no information has yet been collected on the mechanism for the increase in competing vegetation. Information on changes in soil moisture and soil nutrients starting at the green attack phase are needed to determine what resources are limited prior to mountain pine beetle attack, and how long competing vegetation will be able to capitalize on those resources following mountain pine beetle attack. Changes in soil moisture could vary depending on site drainage properties and should be evaluated at the site level as well as the landscape level. Rex and Dubé (2006) found that poorly drained or receiving areas may experience increased water tables. However, well drained or shedding areas (e.g. where terrestrial lichens are more abundant) could potentially be less affected by water table increases. Data from the western (Williams Lake TSA) portion of the Itcha-Ilgachuz caribou winter range suggests that soil moisture is increasing on lichen sites (Sagar 2011).

Other important information gaps include: changes in habitat and range use

as a function of the degree of mountain pine beetle attack; the response of caribou and caribou habitat to the fall-down phase of the mountain pine beetle epidemic; and, changes in caribou population dynamics.

The Tweedsmuir-Entiako and Kennedy-Siding caribou habitat use studies

suggest that caribou did not significantly alter their winter habitat use and foraging strategies following the mountain pine beetle epidemic. However,



Table 3. Summary of information collected by ongoing research on the effects of mountain pine beetles and salvage harvesting on woodland caribou ecology.

Mountain Pine Beetle Attack Phase1

Alive Green Red Grey Fall-down

MPB/ Salvage

harvesting HABITAT Terrestrial lichen abundance KS2, IIW,

W TEL, KS, IIQ, IIW, W

TEL, KS, IIQ, IIW, W

TEL, KS, IIQ TEL, KS

Arboreal lichen abundance KS, IIW KS, IIQ, IIW KS, IIQ, IIW, W KS, IIQ, IIW KS Vegetation abundance IIW, W TEL, IIQ, IIW, W TEL, IIQ, IIW, W TEL, KS, IIQ TEL, KS Regeneration IIW, W TEL IIQ, IIW, W TEL, IIQ, IIW, W TEL, KS, IIQ TEL, KS Soil moisture IIW Soil nutrients IIW Air and soil temperatures IIW IIW IIW IIW Light availability TEL, IIQ, IIW TEL, IIQ, IIW TEL, IIQ TEL Snow conditions KS KS KS KS KS Treefall KS, IIW KS, IIW TEL, KS, IIQ, IIW TEL, KS, IIQ, IIW Coarse woody debris W, IIW TEL, IIQ, IIW, W TEL, IIQ, IIW, W TEL, II TEL HABITAT USE AND RANGE USE Forage choice (terrestrial vs. arboreal) KS KS TEC, KS KS Site (stand) selection KS KS TEC, KS KS Landscape (range) selection KS KS TEC, KS KS Elevation selection KS KS TEC, KS Winter travel – snow conditions KS KS TEC, KS KS Summer and winter travel – coarse woody debris POPULATION DYNAMICS Population abundance Population status Calf recruitment TEC Adult mortality KS KS KS TEC Predation risk (other prey abundance/presence - forage) Predation risk (other prey abundance/presence - winter travel) Predation risk (other prey abundance/presence - summer travel)

Predation risk (predator winter travel) Predation risk (predator summer travel) 1 Shaded areas indicate phases of attack that could potentially affect various aspects of caribou ecology 2 IIQ= Itcha-Ilgachuz lichen project (Quesnel TSA area); IIW= Itcha-Ilgachuz lichen project (Williams Lake TSA); KS=Kennedy-Siding study; TEC=Tweedsmuir-Entiako collared caribou project;

TEL=Tweedsmuir-Entiako lichen project; W=Wolverine continuing lichen study



neither study was able to document how caribou responded to different intensities of attack. For the Tweedsmuir-Entiako caribou habitat use study, a reliable map showing the intensity of attack was not available, while for the Kennedy-Siding caribou habitat use study, intensity of attack was relatively uniform across the winter range. Consequently, the relative importance of varying degrees of partially killed stands has not yet been assessed. On the Itcha-Ilgachuz caribou winter range, the lack of caribou observed during the March 2010 survey could imply a shift in habitat or range use. Further investigations are needed to determine whether there has been a shift in use, and if so, whether it is a response to the mountain pine beetle epidemic.

The response of caribou and caribou habitat to the fall-down phase of the

mountain pine beetle epidemic cannot be assessed until trees fall-down. Measurements for the current lichen projects should continue at scheduled intervals into the fall-down phase to assess: when fall down occurs, whether fall-down is gradual or rapid, how quickly fall-down will decay, and how fall-down affects forest floor vegetation dynamics. Monitoring of radio-collared caribou should resume when the fall-down phase begins to assess whether fall-down affects caribou movements and habitat use.

One of the most important knowledge gaps is how mountain pine beetles

affect caribou population dynamics. Information on how other prey and predators respond to the mountain pine beetle epidemic will be important for understanding whether predation risk to caribou will increase, decrease or stay the same following mountain pine beetle attack.

Information on the effects of salvage harvesting on caribou is also important

to address. The Kennedy-Siding project addresses vegetation responses in and caribou use of salvage harvested areas (Seip and Jones 2010). Additional studies on other caribou ranges would help to assess whether the trends observed in the Kennedy-Siding area are consistent across other caribou ranges. Although some information is available on the effects of harvesting in unattacked stands on terrestrial lichens, landscape level changes in hydrology due to widescale mountain pine beetle attack may result in different ecological responses in salvage harvested areas in mountain pine beetle killed stands than in harvested areas in unattacked stands.



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APPENDIX 1. SUMMARY OF RESEARCH REPORTS

Project Reports Reference PROJECTS INITIATED TO INVESTIGATE THE EFFECTS OF THE MOUNTAIN PINE BEETLE EPIDEMIC ON CARIBOU

2001/02 – Year 1 Williston and Cichowski 2002 2002/03 – Year 2 Cichowski and Williston 2003 2003/04 – Year 3 Williston and Cichowski 2004 2005/06 – Year 5 Williston and Cichowski 2006

Annual Reports

2007/08 – Year 7 Cichowski et al 2008

The response of caribou terrestrial forage lichens to mountain pine beetles and forest harvesting in the East Ootsa and Entiako areas

Final Report (Phase 1 – 2001-2005) Williston et al 2006 2005/06 – Year 1 Cichowski 2006a 2006/07 – Year 2 Cichowski 2007b 2007/08 – Year 3 Cichowski 2008b

Annual Summaries

2008/09 – Year 4 Cichowski 2009


Final Report Cichowski 2010 2005/06 – Year 1 Cichowski 2006b 2006/07 – Year 2 Cichowski 2007c


Annual Reports

2008/09 – Year 4 Cichowski et al 2009

2006/07 – Year 1 Seip and Jones 2007 2007/08 – Year 2 Seip and Jones 2008 2008/09 – Year 3 Seip and Jones 2009


Annual Reports

2009/10 – Year 4 Seip and Jones 2010

2007/08 – Year 1 Cichowski and Williston 2008 The response of caribou terrestrial forage lichens to fire in Entiako Provincial Park

Annual Reports 2009/10 – Year 3 Cichowski and de Groot 2010


Annual Reports 2008/09 Sulyma 2009


Annual Reports 2009/10 Sulyma et al. 2010

Identifying factors affecting the succession and availability of terrestrial lichen communities in the Omineca Region of North-central British Columbia: Long-term lichen monitoring

Annual Reports 2008/09 Sulyma and McNay 2009 Sulyma and Sulyma 2006



Project Reports Reference ONGOING PROJECTS ON CARIBOU HABITAT WITH CONTROLS THAT WILL BE MEASURED FOR EFFECTS OF THE MOUNTAIN PINE BEETLE EPIDEMIC Itcha-Ilgachuz Alternative Silvicultural Systems: Satah Mountain pilot study

Itcha-Ilgachuz Alternative Silvicultural Systems: Satah Mountain replicated trials

Itcha-Ilgachuz Alternative Silvicultural Systems: Satah Mountain – adaptive management trials

Itcha-Ilgachuz Alternative Silvicultural Systems: Chezacut – adaptive management trials

Armleder and Waterhouse 1994 Armleder and Waterhouse 2008 Armleder et al. 1996 Daintith et al 2005 Miège et al. 2001a Miège et al. 2001b Sagar 2011 Sagar et al. 2005 Steen et al. 2007 Waterhouse 2001 Waterhouse 2009 Waterhouse 2011 Waterhouse and Armleder 2004 Waterhouse and Armleder 2005 Waterhouse and Armleder 2007 Waterhouse et al. 2010 Waterhouse et al. in press Wei, et al. 2000 Wiensczyk and Waterhouse 2005 Youds et al. 2002



APPENDIX 2. BIOGEOCLIMATIC ZONE UNITS

Project Biogeoclimatic Subzone/Variants HABITAT-BASED PROJECTS

SBPSmc Sub-Boreal Pine Spruce – moist, cold SBSdk Sub-Boreal Spruce – dry, cool SBSmc2 Sub-Boreal Spruce – moist, cold, Nechako variant

The response of caribou terrestrial forage lichens to mountain pine beetles and forest harvesting in the East Ootsa and Entiako areas

ESSFmc Engelmann Spruce-Subalpine Fir – moist cold SBPSdc Sub-Boreal Pine Spruce – dry, cold Effects of a mountain pine beetle

epidemic on forest floor vegetation dynamics and regeneration in the Itcha-Ilgachuz caribou winter range in the Quesnel TSA

MSxv Montane Spruce – very dry, very cold


SBSwk2 Sub-Boreal Spruce – wet, cool Finlay-Peace variant

The response of caribou terrestrial forage lichens to fire in Entiako Provincial Park SBPSmc Sub-Boreal Pine Spruce – moist, cool


SBPSmc Sub-Boreal Pine Spruce – moist, cool


BWBSdk1 Boreal White and Black Spruce – warm, cool Stikine variant

SBSmk2 Sub-Boreal Spruce – moist, cool Williston variant Identifying factors affecting the succession and availability of terrestrial lichen communities in the Omineca Region of North-central British Columbia: Long-term lichen monitoring

BWBSdk1 Boreal White and Black Spruce – wet, cool, Stikine Variant

SBPSxc Sub-boreal Pine Spruce – very dry, cold Itcha-Ilgachuz Alternative Silvicultural Systems MSxv Montane Spruce – very dry, very cold

Documents

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