Timber Management GuidelinesFor the Provision OfMoose Habitat

February 1988 Wildlife Branch

Ministry ofNaturalResources

© 1988, Queen's Printer for Ontario

Printed in Ontario, Canada

Single copies of this publication are available for$9.00 from the address below.

Ministry of Natural Resources PublicInformation CentreRoom 1640, Whitney Block99 Wellesley Street WestToronto, Ontario M7A lW3

Summary ofTimber Management GuidelinesFor The Provision OfMoose Habitat

Summary Of Timber Management Guidelines ForThe Provision Of Moose Habitat

The purpose of the guidelines is to assist resourcemanagers in maintaining or creating through timbermanagement the diversity of age classes and species ofvegetation that provide habitat for moose.

The attached Background Document indicates morefully the purpose and rationale for the guidelines and themajor components of the Ministry's Moose ManagementProgram. As well, it summarizes moose habitatrequirements and the possible impacts of various timbermanagement operations on moose. Finally, guidelines arerecommended to provide moose habitat in timbermanagement.

This is a summary of those recommendedguidelines.

The guidelines approach the subject in two ways.First, they address the general needs for moose habitat ina manner that acknowledges the fact that moose rangeover the whole forest and require various habitatcomponents throughout that area. Furthermore, theyrecognize that timber management activities can have apositive influence on moose habitat. This aspect of theguidelines provides recommendations for timbermanagers to consider and use when determining whichstands, or portions of stands, should be allocated forharvest during the planning period.

Secondly, the guidelines discuss specific needs formoose habitat that can be identified on discretegeographic areas. These needs allow for the identificationof Areas of Concern to moose habitat management thatare site specific, and the guidelines provide specificdirections for setting timber management prescriptionswithin these areas. Areas of Concern may becomereserves where timber management will not be permittedbecause of the potential for significant adverse effects onmoose habitat. In other instances timber managementmay have to be modified to provide specific habitatrequirements.

Specific Areas of Concern include existing andpotential early and late winter concentration areas,aquatic feeding areas, mineral licks, and calving sites.Within each type of area of concern, prescriptions whichprotect or enhance particular habitat requirements, suchas shelter, or quality and abundance of browse, must bedeveloped. In addition, protection of moose fromdisturbance while in calving areas or using specialfeeding sites may require restrictions on access orscheduling of operations. However, moose habitatrequirements vary with time of day and year, as well astopography and climatic factors, both of which varyacross the province. Timber management practices alsodiffer throughout Ontario.

General Guidelines

To meet the present and future habitat needs of moosethroughout their range the following general guidelinesapply:

1. Boreal Forest Region(a) Potentially large clear cuts, that would not allowmoose utilizing portions of the planned cutover to bewithin approximately 200 m (650 feet) of suitableshelter, should be broken into smaller, irregularly shapedcuts. Clear cut and shelterwood harvesting are best toproduce the desired patterns.

Clear cut in blocks of 80-130 ha (200-320 acres)and leave buffer zones between cuts and scatteredpatches of trees within cutovers. Average cut size isoptimal at about 100 ha (250 acres).(b) Where clearcuts will exceed 100 ha (250 acres) andthe 'edge of cover' to 'edge of cover' distance will bepotentially more than 400 m (1300 feet), provide shelterpatches within the cut area (See General Guideline I (c)).(c) Shelter patches should be of conifer, but could bemixed wood, and be at least 3-5 ha (7-12 acres) in size, atleast 6 m high, have a basal area of about 11 squaremeters/ha (50 square feet/acre) with at least 113 inconifer. The stocking of immature and mature stands withthis basal area will be approximately 70% and 40%respectively. If the objective of the shelter patches is toprovide late winter cover for moose, shelter patchesshould be conifer with stocking of 70% or greater. Wherethese shelter patches are composed of mature conifer,basal areas will be greater than 11 square meters/ha. Theshelter patches should be spaced 300-400 m (1000-1300feet) apart. Leaving shelter patches larger than 3-5 hamay be desirable to encourage future harvest of timberand provide greater protection from predators, includinghunters. Such shelter patches could function as travelcorridors between habitat components.(d) Selection harvesting can occur within mixed woodshelter patches provided that adequate semi-mature ormature conifer cover remains for winter shelter.

2. Great Lakes - St. Lawrence Forest RegionSimilar habitat requirements exist in the southern part ofthe moose range as in the north, but the nature of theforest requires emphasis on protecting and creatingdifferent components. However, few large clear cutsoccur, and hence are seldom if ever a problem, because ofthe nature of the forest and the selection and shelterwoodtechniques usually practised.(a) When cutting in hardwoods stands, cuts should belarge enough to stimulate the growth of early successionalspecies and some conifer regeneration should beencouraged.(b) At least 15% of the total area should have matureconifer cover, preferably in patches or clumps at least 3-5ha in size.

(i)

Specific Area of Concern Guidelines

For the protection of identified site specific habitatcomponents such as early and late winter concentrationareas, mineral licks and calving sites, the followingguidelines apply:1. (a) Access roads should be routed to avoid aquaticfeeding areas, mineral licks, and calving areas.(b) If it is desirable in special circumstances to reducehunting pressure, carefully consider access road locationand measures to limit vehicle use eg. winter extraction,the signing of roads, or the scarification and removal ofroads after timber operations.2. (a) During timber harvest within winter concentrationareas, apply the General Guidelines to maintain thehabitat value of these areas.(b) If required, early winter concentration habitat may becreated or maintained by selection harvesting of somelarge conifers in mature conifer or mixed wood stands.(c) In early winter concentration areas, remaining shelterpatches may be cut when adjacent vegetation has reached2 m in height.(d) In late winter concentration areas cuts should notexceed 400 m (1300 feet) in width. Uncut areas of semi-mature or mature conifer or mixed wood equal in size tocut areas should be left. Remaining shelter patches maybe cut when nearby regeneration has reached 6 m inheight.(e) A reserve is generally recommended around aquaticfeeding areas, mineral licks and calving sites. The shapeand extent of the reserve will be determined by the natureof each site and by the need to maintain its integrity andsafe access to it by moose from surrounding foreststands. In general, a 120 m (400 ft) reserve should be leftaround these areas in certain circumstances somemerchantable trees may be selectively removed.3. The potential impact of site preparation, regenerationand maintenance treatments should be considered in thecontext of the quantity and quality of moose habitat thatsurrounds the harvested site.4. (a) Prescribed burning, if it leaves needed protectiveshelter components, is the preferred site preparationmethod(b) Mechanical site preparation should not destroy shelterpatches of conifer or mixed wood left within the cut area,nor should residual clumps be removed unless theyseriously threaten the success of regeneration.(c) For chemical site preparation or tending, managersshould carefully consider the anticipated effectiveness ofthe herbicide in controlling woody plants (browse), andthe amount and proximity of deciduous growth outsidethe treatment area.5.(a) Natural regeneration of browse species should beallowed where moose browse is, or will be, in shortsupply.

(b) Artificial regeneration to conifer species should beused where shelter is, or will be, in short supply.

Application of Guidelines

It is not feasible to provide too rigid a set of guidelinesspecifying precisely how timber should be harvested tomaintain a good moose population. Local managersmust decide how best to adapt the principles containedwithin the Guidelines to meet the needs of both mooseand the forest industry in their area. As not all wildlifespecies can be managed to maximize populations onthe same land area, neither can all areas be managed ina way that maximizes both moose and timberproduction. Discussion and compromise amonggovernment and industry managers is essential to themanagement process in order to obtain the bestprotection, enhancement and use of both valuableresources.

In general, if individual harvest blocks do noteexceed one hundred hectares, concerns for mooseshould be restricted to known specific areas(concentration areas, mineral lick sites, calving sites,aquatic feeding areas).

If cuts are proposed which exceed the generalguidelines over large areas, the District must receivethe Regional Director's approval prior to agreeing tothe plan. If a Region intends to routinely sanctiondeviation from the guidelines, the Assistant DeputyMinister's approval must be obtained before approvingthe plans.

(ii)

Timber Management GuidelinesFor The Provision OfMoose Habitat

TABLE OF CONTENTS

BACKGROUND DOCUMENT

1.0 Preface ...................................................................................................................2

2.0 Moose Program Development .............................................................................3

3.0 Summary of Moose Habitat Requirements........................................................3

4.0 Impacts of Timber Management on Moose .......................................................44.1 Forest Access.............................................................................................54.2 Harvest Operations ....................................................................................54.3 Site Preparation ........................................................................................54.4 Regeneration..............................................................................................54.5 Maintenance ..............................................................................................5

5.0 Providing Moose Habitat in Timber Management ...........................................65.1 Boreal Forest Regions ...............................................................................65.1.1 Forest Access.............................................................................................65.1.2 Harvest Operations ....................................................................................85.1.3 Site Preparation .........................................................................................85.1.4 Regeneration..............................................................................................85.1.5 Maintenance .............................................................................................85.2 Great Lakes – St. Lawrence Forest Regions .............................................95.2.1 Forest Access.............................................................................................95.2.2 Harvest Operations ....................................................................................95.2.3 Site Preparation .........................................................................................95.2.4 Regeneration..............................................................................................95.2.5 Maintenance ..............................................................................................9

6.0 Application of Guidelines...................................................................................10

7.0 Basis for Guidelines and Sample Plans ............................................................10

APPENDICES

Appendix I Moose Population Targets by WMU..........................................11Appendix II Sample Plans ..............................................................................12Appendix III Moose Habitat in Ontario: A Decade of Change

in Perception...............................................................................18Appendix IV Effects of Forestry on Ungulate Populations in the

Boreal Mixed Wood Forest ........................................................27

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Timber Management Guidelines forthe Provision of Moose Habitat1.0 PrefaceNot all wildlife species can be managed for maximumpopulations on the same land area. Thus managers mustoften make local decisions about which species warrantspecial forest prescriptions. The legitimate concerns forother wildlife will often be accommodated to a largedegree within the prescription for the key species. Theseparticular guidelines deal with moose in the context oftimber management. The timber management terms usedin this document (eg. selection cutting, shelterwoodcutting) are as defined in the "Class EnvironmentalAssessment For Timber Management on Crown Lands inOntario" (MNR, Dec. 1985).

The purpose of this set of guidelines is to assistforest and wildlife managers in planning timbermanagement activities. Virtually all efforts designed tomanage moose habitat will involve working with timbercompanies to manage the forest in order to produce goodmoose habitat with a minimum loss of wood fiber. Inmany circumstances, the practice of good timbermanagement is consistent with good wildlife habitatmanagement. For example, disturbances to the forestcover by timber harvesting will create the kind ofopenings and young growth that are necessary elementsof good moose habitat. Without such disturbance, moosepopulations would be lower. There is not usually aconcern over whether timber is harvested, or how muchtimber is harvested. It is mainly a question of how andwhen harvesting takes place and the relative sizes of cutand uncut blocks that is of concern in moose habitatmanagement. The challenge of integrating timber andmoose management is to retain all of the necessaryhabitat components for moose while extracting theavailable merchantable timber.

These guidelines include general requirements andspecific suggestions for providing habitat, although it isimpossible to foresee every conceivable situation themanager might encounter. The attached papers providemore complete information concerning moose habitat.

Essentially, the guidelines are designed to producegood moose habitat by cooperating with forest managers.It must be emphasized, however, that moosemanagement involves many variables other than theactual habitat work itself. For instance, the best habitatwill not necessarily contain good moose populations ifhunting pressure is excessive or if wolf predation isextensive. The goal should be to achieve a propercombination of moose population management, controlof hunters and careful habitat manipulation.

Moose range in Ontario encompasses a wide varietyof physiographic site conditions. It extends from thewestern boundary of Ontario to the eastern border, andsouth to the edge of the Precambrian Shield. Habitatmanagement for moose in northwestern Ontario may besubstantially different from habitat management ineastern Ontario due to the diversity of site, climaticfactors and other environmental conditions across theProvince. Prescriptions for management thus vary. Theseguidelines provide the principles for moose habitatmanagement which local managers can adapt to meetneeds within their own district or region.

Moose habitat guidelines will also be used inwildlife planning processes. Wildlife plans may notcoincide in time period and area with forest plans, butwildlife objectives and habitat strategies should beconsistent among these plans.

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BACKGROUND DOCUMENT

2.0 Moose Program DevelopmentOn October 22,1980, the Government of Ontario adoptedspecific objectives, targets, guidelines and managementpolicy for moose in Ontario (for a complete listing ofthese items, see Policy WM.3.01.02, 1980 12 15).

THE BROAD PROGRAM OBJECTIVE IS: Toprotect and enhance the moose resource and to provideopportunities for recreation from moose for thecontinuous social and economic benefit of the people ofOntario.

The program targets are:1. To increase the moose population from 80,000 to160,000 animals by the year 2000.2. To provide from this herd an annual harvest of 25,000animals by the year 2000.3. To provide a hunter success rate of at least 12 percent.4. To provide 1.2 - 1.4 million viewing opportunities at 24- 30 sites by the year 2000.

The policy for moose habitat management is: "...toensure that the quality of moose habitat is maintained orenhanced by direct involvement in the timbermanagement planning process." and, "to ensure thattimber production will not reduce the quality of moosehabitat, wildlife managers will emphasize upper limits onsizes of clear-cutting operations, within the planningprocess for timber management." Regional targets are:

Moose inventories by wildlife management unit may beobtained by reference to current regional data. Moosepopulation targets by wildlife management unit (WMU)are listed in strategic land use plans for Northwestern andNortheastern Ontario, and are reproduced here with theinclusion of the Algonquin Region (Appendix 1).

Population targets for WMUs may change overtime, but these occurrences are expected to beaccompanied by compensating target revisions in nearbyWMUS.

Much of the target for viewing opportunities may besatisfied from use of wildlife management areas, CrownGame Preserves, Areas of Natural and Scientific Interest(ANSI's) or provincial parks. This will be dictated tosome extent by the presence of visible moose in relationto existing or potential aggregations of people, and ourabilities to provide security for moose and onlookersunder these circumstances.

3.0 Summary of Moose Habitat Requirements

Moose are found predominantly in the Boreal ForestRegion, though they also live in the Great Lakes-St.Lawrence Region. Moose are generally absent insouthern Ontario because of the clearing of land andperhaps overlap with white-tailed deer that carry abrainworm that is fatal to moose. Moose are generallyanimals of the forest edge, living in proximity to youngdeciduous stands which provide food, and semi-matureand mature conifer which provide shelter from weatherand predators, including hunters. They are well adaptedto extreme cold and snow when food and shelterconditions are adequate. Their large bodies are wellinsulated and their long legs make movement throughsnow relatively easy. Unless they are hampered by verydeep snow (greater than 80 cm), their size and strength issufficient to cope with most factors in their environment.

In spring and early summer, moose feed extensivelyon selected species of aquatic plants whenever they areavailable. These plants contain important dietary itemsand may supply certain nutrients (e.g., sodium), notfound in other items of their diet. Travel to these aquaticfeeding areas is often along well-defined routes orcorridors. Mineral licks are also used at this time of year.In summer, fall and early winter, most feeding occurs inearly successional, terrestrial plant communities.Cutovers and burns are especially important. In winter,moose seek out areas of conifer for shelter, and may useportions of nearby cutovers or burns for feedingprovided that snow is less than about 80 cm (35 inches)and is not heavily crusted. During the winter months,moose feed almost exclusively on twigs and branches ofwoody plants, such as willow, birch, aspen, hazel andmountain ash. In addition, their metabolic rate is lowerthan in summer and they use their own stored body fat tosupplement food sources. Moose conserve energy bydecreasing their movements to a minimum. Theseadaptations to cold and snow help them to survive innorthern forest areas.

Moose and other wildlife are active throughout theentire forest. The ability of the forest to support moosechanges through time. These changes can occur slowly asa forest develops and matures, or they may occur quicklyas a result of such events as fire, insect damage orlogging. Such factors acting throughout moose rangeaffect the type, quantity and quality of vegetation, andthus affect the numbers, location and physical conditionof moose that the forest will support.

Prime moose areas are those that produce or attract,or have the potential to produce or attract, a significantlyhigher number of moose than surrounding areas atcertain times of the year. These areas can be identified asbeing key components of moose habitat on a local basis.

Boreal Forest RegionThere are two main types of prime moose areas in theBoreal Forest Region. The first type is seasonal high-use

NorthwesternNorth CentralNorthernNortheasternAlgonquin

Population47,50044,50037,50030,500

5,000

Harvest7,3007,0005,6004,600

500

165,000 25,000

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or winter concentration areas which are known to beimportant to moose for a wide variety of reasons (see theattached papers for details). The second type includesspecial sites such as mineral licks, calving areas, andaquatic feeding areas that may require reserves of timberto protect the special nature of the site. The second typewill remove a small percentage of the land base fromtimber production. The first type will not remove anyland from timber production, but will require modifiedharvesting techniques and may require removing theallowable cut for the 5 year operating period from alarger planning area.

Early winter concentration areas may be typified bymature or over mature, open canopy, mixed-wood standsof relatively low stocking (less than 60 percent). Stockingis an expression of the relationship between actual basalarea as measured in the field and normal basal areaobtained from normal yield table. The need is to leaveportions of these stands uncut for a period of time toallow the animals to continue to use them. As well, bumsand cutovers, usually from 5 to 20 years of age, are alsooften used. Because of the open canopy, early winterconcentration areas usually have considerable browse.These sites are also important to moose as they providesome lateral protection from winds as well as predators.The shape, abundance and nature of these areas is sovariable that each must be treated on an individual basis.

Late winter concentration areas, usually fairly largein size, are those where the average moose density ishigher than the surrounding area. Generally, they are wellstocked stands of mature conifer (greater than 70 percentstocking) with complete crown closure which providesoverhead protection from snow accumulation and severecold. These areas are most functional when near earlywinter or other feeding habitat so that travel distancebetween food and shelter is minimal.

Aquatic feeding areas, mineral licks and calvingsites are important to moose because they attract mooseand contain critical components of their diet or importantlife history features. Identification of these areas, theirshape and importance must be determined by districtstaff. It is important to maintain both the integrity of thesites and sheltered access by moose to them.

Great Lakes - St. Lawrence Forest RegionMoose populations in the Great Lakes - St. LawrenceForest Region have the same basic habitat requirements asthose in the Boreal Forest Region, a diverse series of plantcommunities in early to late successional stages. Toleranthardwood forests may have relatively little browseavailable to either moose or deer, and often insufficient,semi-mature and mature conifer shelter. In the mixedwood (eg. poplar-pine) areas browse and shelter are moreabundant. Group selection cutting and shelterwood cuttinghave contributed to browse production in both toleranthardwood and mixed wood forests.

SummaryAlthough much remains to be learned about the ecologyof moose, here is a brief summary of their known needs:1. Moose populations need the early successional plantcommunities which follow a major disturbance such as aforest fire, insect damage or a logging operation.Populations may expand after disturbance provided thatexcessive hunting or predation does not occur andadequate shelter remains.2. Moose also require semi-mature or mature stands ofconifer in winter. These stands provide protection fromsevere weather and predation, and minimize snow depthand crusting thereby allowing easier access to food.3. Aquatic plant communities in certain waterbodies areused extensively during spring and summer. Bothpreferred emergent and submergent vegetation is utilized.4. Mineral licks are important in certain areas.5. Calving sites such as islands and peninsulas areimportant in certain areas.6. The best moose habitat contains food (earlysuccessional plant communities) and cover (semi-matureand mature conifer) in close proximity such that theanimals need not travel far between these important items.

4.0 Impacts of Timber Management on Moose

In many situations the practise of good timbermanagement is consistent with good wildlife habitatmanagement. For example, disturbances of forest coverby timber harvesting will generally create young growththat is a necessary element of moose habitat. If anadequate amount of shelter eg. unallocated areas,protection forest, remains nearby, then good moosehabitat can be provided. The challenge of integratingtimber and moose management is to retain all of thenecessary vegetation components for moose whileextracting the available merchantable timber. There isnot usually a conflict over whether timber is harvested.It is a question of how and when harvesting occurs andthe relative sizes of cut and uncut blocks that is ofconcern in moose habitat management.

The timber management undertaking involves fivebasic processes. These are: (i) forest access, (ii) harvestoperations, (iii) site preparation, (iv) regeneration, and(v) maintenance operations. The last four steps (ii-v) arereferred to as the silviculture system. Each of theseprocedures may directly impact the quality of moosehabitat and indirectly affect the size of the moosepopulation. The impacts cited below are normally relatedto the immediate areas of the treatment. They areconcerns of a general nature and in practice the impactmay be substantially mitigated by vegetation surroundingthe area of timber operations. The real potentialsignificance of these impacts must be assessed within theentire context of each operating plan. Local managersmust try to balance poten-

4

tial negative impacts by positive ones.As well, the implications to moose of nearby, past

and future silvicultural operations may also influencedecisions when developing an operating plan.

4.1 Forest AccessForest access, principally by roads, may have bothpositive and negative impacts on moose management.While roads may subject newly accessed moosepopulations to local over exploitation, they also allow fordistribution of hunting pressure over a wider geographicarea. As permanent access stabilizes, within a WildlifeManagement Unit, the moose harvest will stabilize andthe benefits of road access to moose management willoutweigh the adverse effects. Within the concept ofOntario's Selective Moose Harvest Program generaloverharvest of moose within a Wildlife ManagementUnit (WMU) should not occur. If local overharvestoccurs in one part of the WMU it should be offset byunderharvest in another part. If an overharvest of moosefrom the entire WMU occurs or is anticipated the harvestquotas for the Unit can be established to correct or avoidthe problem.

Road construction and use within or near aquaticfeeding sites, mineral licks, calving site and winterconcentration areas could destroy habitat or disruptnormal moose activities, and possibly result in anincrease in vehicle accidents.

In the Great Lakes-St. Lawrence Forest Regionconcerns relating to access and silviculture are similar tothose in the boreal forest, although the impacts areprobably less significant because of the shelterwood andselection harvesting systems more commonly practicedin the southern portion of the moose range.4.2 Harvest OperationsThe effects of harvest operations may also be eitherbeneficial or detrimental to moose populations,depending on the manner in which food (youngdeciduous vegetation) and shelter (semi-mature tomature conifer) are left, or Produced, by timberharvesting. When cutting operations produce irregularlyshaped cuts, scattered shelter patches, and a highdiversity of age-class and species composition, moosepopulations will benefit. In the Boreal Forest whereclear-cutting is a common timber harvesting technique,some standing timber with its associated subordinatevegetation should be retained to provide a variety ofplant communities close to each other.

Generally, the greater the amount of edge producedbetween food and shelter habitat components, the betterwill be the quality of habitat.43 Site PreparationPreparing the site to accept seeds or seedlings may havean impact on moose, particularly in the Boreal Forestwhere clear cutting and site preparation are commonlypracticed. The objective of site preparation is to baresome mineral soil and if possible reduce potentialcompetition from

broadleaf species. These are often preferred browsespecies and an important source of nutrition to moose.Except on more infertile soils, site preparation oftenencourages the establishment of herbaceous ordeciduous plants.

Mechanical preparation may remove residualclumps of vegetation within a cut, which could contributeto good wildlife habitat by providing diversity and visualbarriers as protection from hunters and predators. Thevalue of residual vegetation to wildlife increases with thesize of clear cut. Some types of mechanical preparationencourage coppicing or root suckering which increasebrowse.

The effect of chemical site preparation dependslargely on the chemical being used. Chemicals, such as 2,4-D used at approved rates, suppress growth butgenerally do not kill most deciduous woody plants andthey can encourage root suckering. Recently approvedchemicals, such as "Roundup/Vision" (glyphosate),appear to be very effective at killing herbaceous andwoody plants and may substantially reduce browsespecies for an extended period.

Prescribed bunting, where it leaves needed shelterand does not damage the soil, benefits moose by quicklyreturning nutrients to the soil thereby increasing thenutritional quality of the browse growing on the site.Mechanical site preparation may have advantages wherethe retention of needed shelter components cannot beassured by prescribed burning.4.4 RegenerationThe objective of forest regeneration is to return the cutover area to desirable commercial species in a mannerthat minimizes competition and maximizes growth of thedesired tree species. Regeneration in the boreal foreststrives for even-aged stands of coniferous species.Artificial regeneration, along with tending, attempts toincrease the growth of the crop species by reducingcompeting vegetation. Where regeneration is veryeffective, there could be a negative impact on moose inthe initial stages. This could be partially compensated forby leaving residual or nearby stands of young deciduousvegetation.4.5 MaintenanceMaintenance of the forest includes tending, protectionand improvement activities.

The objective of tending is to further reducecompetition and in most respects the impacts of tendingare the same as for site preparation. Manual, mechanicaland some chemical treatments do not kill most deciduousgrowth, but will set it back. Later, coppicing and rootsuckering may occur. Forest improvement by convertingmixed wood stands to more pure conifer may createwinter shelter but remove a significant source of browsefor moose.

In the Great Lakes-St. Lawrence Region there areoften too few stands of semi-mature and mature coniferto provide shelter. Silvicultural treatments which producethese will often benefit both moose and deer.

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5.0 Providing Moose Habitat in TimberManagement

The objective of habitat management is to provide all ofthe necessary habitat components within the area ofactivity normally inhabited by moose. The size of thisarea will be dictated to a large extent by topography, thenature of the forest, and the size of the moose population.The purpose of these Guidelines is to demonstrate how toproduce good vegetation patterns necessary to meetmoose requirements.

Moose are animals of the forest edge requiringyoung deciduous growth for food, and semi-mature andmature coniferous forest as shelter from weather andpredators. To benefit moose, timber management shouldproduce irregularly shaped cuts with scattered shelterpatches and a high diversity of age classes and species ofvegetation.

Following are a set of general principles which willlead to the maintenance or improvement of moose habitatin Ontario, recognizing prevalent timber harvestingpractices. Moose habitat needs vary during the day,different times of the year, and across their range. Also,the topography and climatic conditions in Ontario are notuniform and timber management practices vary widelyacross the Province. Because of this variation, theGuidelines for use in planning timber management are setdown in a general way to ensure that average habitatconditions are provided. Local managers and plannerswill decide how to best apply the principles to meet localsituations. In addition, not all areas can be managed insuch a way that maximum timber production willcoincide with maximum wildlife production.Compromise and discussion among managers is essentialto the management process.

Figure 1 illustrates a possible scenario resultingfrom the implementation of these principles.

5.1 Boreal Forest Region

5.1.1 Forest Access. Where new access is created toharvest the forest, the potential for local overharvestof moose exists. Although legislation (eg. PublicLands Act, Game and Fish Act), may be used toinhibit or prevent hunting within these areas foreither short or long periods of time, it tends topostpone problems of overharvest rather thansolving them. In special cases where it is desirableto minimize hunting by controlling access, roadsmay be closed by signing or they may be kept awayfrom the area of concern, or wood may be extractedusing winter roads. As well, in some circumstancesit may be appropriate to scarify and remove accessroads after extraction is complete.

Access roads should avoid mineral licks,aquatic feeding areas, and calving sites to protectthese important habitat features and minimizedisturbance and accidents to moose using theseareas.Road use and location must be addressed as early aspossible in the planning process so that fieldexaminations can identify possible alternatives.

6

5.1.2 Harvest Operations. The moose managementobjective of maintaining or enhancing the quality ofmoose habitat includes the concept of protecting keyfeatures (eg. aquatic filling areas) and providingfood (early successional plant communities) close toshelter (semi-mature or mature conifer stands).

This objective may be met by no or modifiedcutting in the vicinity of key features, by reducingthe size of planned clear cuts or by providingshelter patches within cutovers. Additionally, adiverse vegetative pattern may be obtained ifcutting is dispersed among all eligible stands ratherthan cutting them in a contiguous manner duringthe planning period.

In some areas, clear-cutting in blocks of 80-130 ha (200-320 acres) with buffer zones betweencuts, and scattered clumps of trees within thecutovers, will provide the desired conditions. Clear-cuts greater than 100 ha (250 acres) should havescattered shelter patches within the cut area. Thiswould keep the overall vegetative diversity of thearea high and still provide a reasonable timberharvest.

The best habitat should provide conditionsenabling a moose to be within 200 m (650 feet) ofshelter patches or other cover. These shelterpatches should preferably be of conifer but couldbe of mixed-wood, with at least 1/3 in conifer.They should be at least 3-5 ha (7-12 acres) in size,be spaced 300-400 m (1000-1300 feet) apart, be atleast 6 in (20 feet) high, and have about 11 squaremetres/ha basal area (50 square feet/acre). Thestocking densities of immature and mature standswith this basal area will be approximately 70% and40% respectively. If the objective of the shelterpatches is to provide late winter over for moose,shelter patches should be conifer with stocking of70% or greater. Where these shelter patches arecomposed of mature conifer, basal areas will begreater than 11 square meters/ha. It may bebeneficial to moose and advantageous to the timberindustry to leave shelter patches large enough toinhibit blowdown problems and to warrant futureharvest (eg. > 8 ha).

If late winter habitat will be adequate in thearea, a return cut of shelter patches can occur whennearby regeneration has reached 2 metres in height.Regeneration of Us size will provide lateral shelter,and function as early winter habitat if theregenerated site contains sufficient browse.

If late winter habitat will be inadequate in thearea after an early return cut, the cutting of shelterpatches should not occur until nearby regenerationhas reached 6 metres in height, thereby providingoverhead cover for moose.

Clear cut and shelterwood harvestingtechniques can produce these patterns, but selectioncutting, seldom practiced in the Boreal Forest, maynot disturb the forest canopy enough to create sig-

nificant successional growth. Some selectionharvesting of conifer could be practiced withinmixed wood shelter patches provided adequateprotection remains. There may be a need oropportunity to provide early winter habitat where itdoes not currently exist. This can be achieved byselection cutting within mature conifer andmixedwood stands to remove some of the largerconifers.

In late winter concentration areas, width ofindividual cuts should not exceed 400 m (1300 feet).Uncut areas equal in size to cut areas should be left.

To protect aquatic feeding areas, mineral licksand calving sites, generally reserves are requiredwith the shape and extent dictated by surroundinghabitat conditions. Usually a 120 m reserve shouldbe left around these areas. Some merchantableconifer may be removed by selection cuttingprovided the general nature of the reserve remainsintact.

Figure 2 is a stylized illustration of some ofthe principles of timber harvesting impacts onwildlife and timber production.

5.1.3 Site Preparation To benefit moose, mechanicalpreparation should not destroy shelter patches.Residual clumps of conifer or mixed wood withinthe cut should not be destroyed unless theseseriously threaten the success of regeneration.Chemical site preparation is acceptable providedthere is adequate browse in nearby stands.

5.1.4 Regeneration Natural regeneration, on suitablesites that produce deciduous woody growth, is ofbenefit to moose where food supplies areinadequate. Harvest methods which facilitate thisshould be encouraged in these areas. Artificialregeneration to conifer may be best where mooseshelter is in short supply.

5.1.5 Maintenance This aspect of the silviculturesystem, as those above, should be considered inrelation to the vegetation surrounding the treatmentarea.

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Figure 2.Stylized Illustration of Principles of TimberHarvesting Impacts on Wildlife andTimber Production1.

2.

3.

4.

5.

6.

Large clear cut. Total timberutilization, no shelter of any kind forwildlife.

Narrow travel corridor or uncut block(eg. <120m) Some blowdown possible.Possible return for remaining wood.Shelter good for visual barrier, windand possibly snow depth. Some edgeprovided.

Wide corridor or uncut block (eg.400m). Timber volume warrants futureharvest. Shelterwood for visual barrier,wind, snow and radiation. Same edge as#2.

Large cut block pattern. Timber volumewarrants future harvest. Shelter goodfor visual barrier, wind, snow andradiation. Much more edge than #2 and#3. Browse and shelter nearly equal andavailable.

Small block pattern. Same timbervolume as #4, but small blocksparticularly if scattered may makefuture harvest impractical. Shelter goodfor visual barrier wind, snow andradiation. Strip and shelterwood cuts arevariations of this. Much more edgethan#4.

Large uncut, irregular patches (eg. >5ha.) Some loss of Umber unless patchesare connected. Shelter good for visualbarrier, wind, snow and radiation. Largeamounts of edge.

Note: A rigid checkerboard harvest pattern is not desirable to producethe best wildlife habitat nor is it practical on most sites. This is astylized representation.

Where browse is abundant, most tending isacceptable. Where shelter is or will be in short supply,and browse is adequate, tending is encouraged.

Tending efforts should not destroy deciduous growthwithin shelter patches. Treatments which increase browse

in these areas are beneficial. The use of herbicides thatsuppress deciduous growth for long periods of timeshould be carefully considered for their potentialnegative impacts on the quantity of moose browse.

Some chemicals, notably 2, 4-D, can have the effectof encouraging browse production (coppice growth).

5.2 1 Great Lakes - St. Lawrence Forest Region5.2.1 Forest Access. There are similar concerns about

access in both the Boreal and Great Lakes - St.Lawrence Forest Regions. See section 5.1.1 forappropriate recommendations.

5.2.2 Harvest Operations. In the Great Lakes - St.Lawrence Forest Region the emphasis in forestoperations is primarily on natural regenerationthrough the selection or uniform shelterwoodsystems. Since these systems obtain regenerationunder the shelter of a residual stand, they normallyprovide optimum moose habitat.Cutting in tolerant hardwoods can create goodhabitat if it produces sufficient disturbance tostimulate early succession species growth. Cuttingshould not remove, and it may help regenerate,conifer that is necessary for shelter from extremeweather.At least 15 per cent of the total area should havemature conifer cover at all times, preferably inpatches or clumps at least 3-5 ha in size. Thisobjective can be met using the shelterwoodharvest system and, if feasible, by undertakingsome artificial regeneration to conifer.The principles for protecting winter concentrationareas, aquatic feeding areas, mineral licks, andcalving sites are the same as in northern forestregions. Each area should be identified, itsimportance determined and the site treated on anindividual basis by district staff. See Section 5.1.2for appropriate harvesting recommendations.

5.2.3 Site Preparation. Recommendations for sitepreparation in the Boreal Forest Region (Section5.1.3) are applicable in a few isolated cases to theGreat Lakes - St. Lawrence Forest Region. Asharvest blocks are generally small, site preparationoperations are seldom implemented over largeenough areas to have a significant impact onmoose habitat.

5.2.4 Regeneration. Because conifer shelter isfrequently lacking in the Great Lakes - St.Lawrence Forest Region, regeneration to coniferwill generally benefit both moose and deer. Alsosee Section 5.1.4.

5.2.5 Maintenance. The recommendations for tending,protection and improvement operations in theBoreal Forest Region apply here as well. SeeSection 5.1.5. Most maintenance in the GreatLakes - St. Lawrence Region is in the form ofthinning and improvement of established standswhich has little impact on moose habitat.9

6.0 Application of Guidelines

Moose habitat needs vary during the day, different timesof the year, and across their range. Also, the topographyand climatic conditions in Ontario are not uniform andtimber management practices vary widely across theProvince. Because of this variation, the Guidelines foruse in planning timber management are set down in ageneral way to ensure that average habitat conditions arepro- vided. Local managers and planners will decide howto best apply the principles to meet local situations. Inaddition, not all areas can be managed in such a way thatmaximum timber production will coincide withmaximum wildlife production. Compromise anddiscussion among managers is essential to themanagement process.

In general, if the individual harvest blocks in theproposed five year allocation do not exceedapproximately one hundred hectares, there should be noor few moose concerns. In such cases concerns should berestricted to known specific areas (concentration areas,mineral lick sites, calving sites, aquatic feeding areas).

If cuts are proposed which exceed general guidelinesover large areas, the district must consider existing andpotential moose habitat requirements prior to approvingthe plan. When a district proposes a cut that greatlyexceeds the general guidelines they must, in advance,receive the Regional Director's approval. In addition, if aregion intends to routinely sanction deviation from thegeneral guidelines, the Assistant Deputy Minister'sapproval must be obtained in advance of approving theplans.

7.0 Basis for Guidelines and Sample Plans

These guidelines are based upon a body of scientificliterature which is summarized in two appended papers,Thompson and Euler (Swedish Wildlife Research:Viltrevy. in press), and McNicol and Timmermann (1981.in Boreal Mixedwood Symposium Proc. p. 141 - 154).Examples of plans are provided in Appendix 11.

10

Moose Population Targets for the year 2000.By Wildlife Management Unit*

APPENDIX I

WMU No.1234567891011121314151617181920

Population Target21,092

4,5754,5363,8253,3401,3932,6621,7482,655

1502,9933,5924,386

4659,6107,6773,0364,2074,015

152

WMU No.2122232425262728293031323334353637383940

Population Target10,3372,7223,6272,8331,1113,9773,1333,7692,6985,1693,9484,5002,565

7203,4952,0151,8954,0102,1703,475

WMU No.41424647484950515354555657585960

Population Target3,5602,210

50110800

60300

3,000 30260210

4030303050

*From: Northwestern Ontario Strategic Land Use Plan, May 1982; Northeastern Ontario Strategic Land Use Plan, April1982; Algonquin Region, 1984

11

APPENDIX 11

The maps that follow illustrate some actual examples offish and wildlife concerns that have been addressedthrough alteration of timber management plans. For eachexample a description of the area, the concerns involvedand their solutions are given. These examples are of aspecific area and may not be duplicated exactlyelsewhere, but they am typical of solutions that areencountered in timber management planning.

Map 1- Big Ghee Lake and Mowe Lake

SiteThis area is predominantly mixed jack pine, blackspruce, poplar, and white birch. There are many largeand small lakes and other wetlands. A large number ofmoose make use of this area.

ConcernsThe main concerns for this site were: that after cuttingthe area would be susceptible to overhunting because ofthe large waterbodies to the east and west, that acomplete clearout would greatly reduce the moosepopulation inhabiting the area, and that early and latewinter habitat would be eliminated.

SolutionsReserves of 30.5m (33 yards) were left along the waterto tie into residual stands and to provide a partial east-west corridor between Mowe Lake and North Channel.The combination of reserves and residuals createsanother partial corridor in the north and northwest thatbreaks up the cut, "buffers" an older cut in the northwest,and protects winter habitat. The area was closed tohunting for 3 years to provide protection fromoverhunting.

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Map 2 - Watershed Lake and Squeers Lake

SiteThis large tract of predominantly poplar was to beclearcut for kraft pulp. The areas to the east provide agood mix of species and age classes, there is a naturaltravel corridor between Watershed and Squeer lakes, andthere are 2 small ponds that are suspected lick sites. BothWatershed and Squeers lakes provide habitat for laketrout.

ConcernsThe objectives for this tract were to: break up the large cutto maintain a diversity of age classes, maintain the travelcorridor between the takes, protect the suspected lick sites,and protect the lake trout habitat.

SolutionsA 120m (130 yard) area of concern is required aroundlake trout lakes. Three corridors, one of which includesthe lick sites, were added to break up the cut, provide ageclass diversity, and serve as travel lanes. After 10 years,when the cuts have regenerated, the corridors can be cut.

14

Map 3 - Three Island Lake and Jack Lake

SiteEssentially a mature to oven-nature conifer monoculturewith little mixedwood, Us area is quite different from theprevious two and generally less productive for moose.Clearcutting in large blocks was the proposed treatment.

ConcernsAge class diversity in this area is lacking both beforecutting and with a plan for large clearcuts. The objective,therefore, is to provide some diversity, paying particularattention to areas with some mixedwood because anymoose that are in the area will tend to centre theiractivity at these sites.

SolutionsA combination of residual stands and conifer reservescontaining some mixedwood were used to address the ageclass diversity problem. The area was also closed tohunting. Subsequently, numbers of moose and their use ofthe area improved.

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APPENDIX III

Moose Habitat in Ontario: A Decade ofChange in Perception

Ian D. Thompson, Canadian Forestry Service,Box 6028 St. John's NFLD. A1C SX8David L. Euler, Ontario Ministry of Natural Resources,Wildlife Branch, Queen's Park, Toronto, M7A 1W3

IntroductionMoose (Alces alces) management has changeddramatically in Ontario both in philosophy and substancein the past decade. An integral part of this change hasbeen an attempt to understand what constitutes goodhabitat for moose and to apply this knowledge within theframework of a comprehensive moose managementprogram. Timber and land use management planninghave also changed during the period and have oftenplaced conflicting interests on the land base. The task ofintegrating these activities is difficult and moosemanagers still have much to learn before these programssuccessfully complement each other.

In the early 1950s the basic question in moosemanagement was ... can the moose herd support anannual hunt? Based on the best data available, unlimitedhunting was permitted and habitat or predatormanagement was not a major activity. By 1974, itseemed clear that the possibility of over-hunting wasremote and, if it did occur, the herd could quicklyrecover from any excessive exploitation (Cummings174a, b). However, several local managers expressedconcern over the welfare of the population with regard toboth hunting pressure and the impact of timber-harveston moose habitat. From 1975 to 1980 an examination ofavailable data revealed a decline in the Provincialpopulation. Over the past twenty-five years, a 35 percentreduction in herd size occurred (Euler 1984). Thereasons for the decline have been controversial, but thereis general agreement that hunting was a major factor. Anew management program was developed to increase theherd based on manipulation of hunting pressure,improvement of habitat and selective control ofpredators (Euler 1983).

In this paper we will summarize knowledge gainedabout moose habitat in Ontario during the past ten yearsand try to portray moose habitat attributes as we now viewthem.

Description of Ontario Moose RangeMoose inhabit two forest regions in Ontario: thenorthern part of the Great Lakes-St. Lawrence Forestregion and the Boreal Forest Region (Rowe 1972, Fig.1). The first region forms the southern area of mooserange from the Manitoba border to Thunder Bay andcontinues east of Lake Superior through Sudbury to theQuebec border. Moose are generally absent in southernOntario due to the clearing of land in these areas andprobable overlap with white-tailed deer (Odocoileusvirginianus) which carry brainworm(Paralaphostrongylus tenuis) (Anderson 1963, 1972).Most moose range in this Province is the Boreal Forest.

The Great Lakes-St. Lawrence forest region is atransitional type from southern deciduous areas into theboreal region and has components of both. Importantconiferous species include: white (Pinus strobus) and redpine (P. mesa), hemlock (Tsuga canadensis), white(Picea glauca) and black spruce (P. Mariana), balsam fir(Abies balsamea) and jack pine (P. banksiana). Abundantdeciduous species are yellow birch (Betulaalleghaniensis), sugar maple (Acer saccharum), redmaple (A. rubrum), red oak (Quercus rubra), beech(Fagus grandifolia), trembling aspen (Populustremuloides), balsam poplar (P. balsamifera) and whitebirch (B. papyrifera). The entire Great Lakes-St.Lawrence forest type is rolling to rugged with areas ofexposed bedrock. Logging in this region has beenextensive since the late 1800s, although it has beenlargely selective for white pine saw logs and veneerquality hardwoods. Large scale clear-cutting has notgenerally been employed, except in the western areasbetween Thunder Bay and Manitoba and to some extenteast of Lake Superior. Regeneration of stands ispredominantly to boreal hardwood types (aspen andwhite birch) in northern areas and to sugar maple andyellow birch on deep, dry soils in southern areas.

Lowland areas are characterized by black spruce,white cedar (Thuja occidentalis) and tamarack (Larixlaricina). The common regeneration pattern in lowlandsis to speckled alder (Alnus rugosa) with black spruce andbalsam fir on better drained sites.

The boreal region can be subdivided into three mainsections based on soils:1) the clay belt in the east;2) the southern and central area of glacial tills andsandy soils; and3) a large northern section of more or less poorlydrained marine clays. The latter area is poor mooserange and is generally not managed for habitat; it willnot be discussed in this paper.

The clay belt is an area of flat terrain lying in thebasin of post glacial Lake Objibway. Tree cover isdominated by black spruce, both on uplands and on theextensive poorly drained lowlands. Less abundant treesspecies include balsam fir, balsam poplar, white birch,trembling aspen, jack pine and white spruce (Carletonand Maycock18

1978). Regeneration after logging in upland areas is toaspen, balsam fir, mountain maple (Acer pensylvanicum),beaked hazel (Corylus cornuta) and to a lesser extentwhite birch. There are large areas of organic soils overlygleysols, and in upland sites both humo-ferric podzols,and grey soils have developed. Bedrock outcrops aregenerally absent.

The southern and central area of the Boreal zonevaries considerably, ranging from an area of somesouthern influence in the east, over a central plateau areanorth of Lake Superior, through a large sandy areanorthwest of Thunder Bay to an intensely glaciated areaof thin glacial soils east of the Manitoba border. Bedrockexposure is common throughout. The central and easternsection is well-drained by large rivers and the westernarea contains many lakes. Soils are predominately humo-ferric podzols, overlying Precambrian granites andgneisses. Tree species are the same as in the clay belt butsince there is more relief, upland species are dominant.Mixed woods are common in the east and central area,jack pine and spruce covers much of the area west ofThunder Bay, and black spruce is again dominant to thenorthwest. Cutover areas regenerate to a mixture of shrubsincluding mountain maple, beaked hazel, white birch,trembling aspen, mountain ash (Sorbus americana), greenalder (Alnus crispa), some balsam poplar and balsam fir.Pin cherry (Prunus pensylvanica) and willows (Salix spp.)are common in drier areas.

In both the claybelt and the south-central area of theboreal forest logging tends to convert stands from spruceand mixed wood to aspen, balsam poplar and balsam firon uplands or mesic sites, and to speckled alder inlowlands. In the past, scarification and planting operationscovered less than 30 percent of the cut area, although thishas improved over the past five years to about 60 percentof the annual cut.

Management ProblemsAs late as 1960, much of Ontario was still being loggedwith horses. These operations produced excellent moosehabitat much like that described by Telfer (1978).However the change to mechanical logging whichoccurred in the early 1960s probably resulted in habitatof poorer quality. By the 1970s, when the mechanicallylogged habitat was regenerating, it was probably not ascapable of supporting as many moose as habitat createdprior to 1960. From approximately 6,000,000 hectares ofmoose range in Ontario in the general area of forestexploitation, about 200,000 ha each year are cut usingthe clear-cut technique. No statistics exist to ascertainhow many of these are large cuts and how many aresmall. Cutting thus affects 2 to 3 percent of the mooserange each year.

The most commonly assumed habitat managementproblem during this period concerned large clear-cuts bythe forest industry. Many hunters and wildlife managersobserved extensive large clear-cuts and wondered how

they affected the moose population. Moose require thesecondary succession produced by disturbance, yet someevidence suggests that large disturbed areas may not beas useful as smaller ones (Telfer 1978; Hamilton et al.1980, Peek et al. 1976). If large areas of moose range arechanged into poor habitat by logging operations, adecline in the moose herd would be the result.

Evidence to support the concern that largeclearcuts, by themselves, have reduced Ontario's moosepopulation is not convincing. The most instructiveexample is in the Chapleau Crown Game Preserve whereintense logging and large clearcuts both occur. Aerialinventories show this area supports one of the highestmoose densities in Ontario (NINR unpublished aerialsurveys).

Bergerud et al. (1983) suggested that wolves wereresponsible for a moose decline in Pukaskwa NationalPark. A reassessment of the aerial survey data reportedby these authors does not indicate a decline moosenumbers or support the idea that wolves wereresponsible for keeping the herd from increasing (ParksCanada 1980).

The only other evidence to give instruction in thiscase is the Management Unit 23 and Unit 31 example. Ifwolves were the prime factor in keeping those moosepopulations from expanding, as was found in. Alaska(Gasaway et al. 1983), then the moose herd should nothave increased when hunting pressure was reduced byapproximately half (Table 2, Euler 1983). While theincrease in moose in these units was not statisticallysignificant, the general trend over a four year periodseems to be increasing. In the light of evidenceavailable, most of which is not as definitive as would bedesirable, it is reasonable to base a management decisionon the assumptions that predators played a relativelyminor role in the decline of moose in Ontario during thistime period.

Thus, while moose habitat in general may not havebeen the very best, the major causes of the moose declineundoubtedly involved hunting (Euler 1983), climate(Thompson 1980) and perhaps predation (Euler 1983)rather than habitat loss. Even though managers did notbelieve habitat loss was the most important cause of theherd decline in Ontario, habitat management is still animportant technique for future moose management.

Despite the foregoing conclusion, it is important toremember that logging activities affect moosepopulations in two ways. They change vegetation butthey also provide access to animals which would nototherwise be available to hunters. Three separate studiesin Ontario have documented the problem of the impact oftimber harvest on moose because of access roads, and allconcluded that the access provided by logging roads wasimportant in the herd decline (Flemming 1976, Eason etal. 1982, Timmerman and Gollat 1983).

An important component of the effort to changemoose management was to establish how many moosecould be carried on the available land area. The ChapleauCrown Game Preserve has not had hunting for some 50years, but has an active forest operation, involving some

19

clear cuts over 4,000 hectares in size. Further, the wolfdensity is about normal for that part of Ontario(Kolenosky 1981).

It has been observed for some time from OntarioMinistry of Natural Resources aerial surveys that moosedensity on the Preserve is about 0.30 - 0.35 moose persquare kilometer, about two times the density in mosthunted areas outside the Preserve. Thus, managersconcluded that although this might not be the maximumcarrying capacity, it did represent a reasonablejudgement as to the ability of the land to support mooseover the long term in the presence of predators.Similarly, Quetico Provincial Park in NorthwesternOntario showed moose densities in good habitat, withnaturally regulated wolf populations but withouthunting, approximately equal to Chapleau Preserve.From the 600,000 square kilometers of moose rangeavailable, using a figure on the low side of knowncapacity to carry moose, we expect a moose potential ofat least (600,000 x 0.3) = 180,000 animals. In an effortto be realistic, conservative and allow for unexpectedevents, the goal for the moose population was set at160,000 animals (approximately double the 1982 herdsize), but somewhat less than the present habitat canprobably support.Moose Habitat UseMoose have a range of habitat requirements whichchange with the season and life history events. Habitatshould be viewed as a series of components, each withintrinsic variables. Winter habitat is probably the mostcritical component, and the winter severity varies withsnow accumulation and crust conditions (Des Meules1964, Poliquin et al. 1977, Welsh et al. 1980, Thompsonand Vukelich 1981).

Winter habitat can be divided into early and latecomponents, and use may differ for separate age and sexclasses of moose. A strong desire for sodium in springand early summer indicates that aquatic feeding areas andmineral licks constitute extremely important aspects ofmoose habitat (Jordon et al. 1973, Fraser and Reardon1980). Less known but possibly significant features ofgood range also include calving habitat, areas to escapefrom wolves, and rutting and post-rutting habitat. Thesecomponents of moose habitat are best examined at 3scales:1) a broad scale approach which considers the needs ofan entire local population,2) a smaller scale which includes basic needs of theanimals, and3) an individual scale which refers to foodrequirements.Broad Scale Habitat RequirementsRather than view moose habitat on a local basis whichconsiders few animals as the working group (forexample, an annual cut plan of 700 ha), the larger scaleneeds of a population must be considered. That is, at ascale of

several hundred square kilometers.Seasonal movements by moose appear universal in

North America (LeResche 1974) and are common inOntario (Goddard 1970, Saunders and Williamson 1972,Addison et al. 1980). Although movements, betweensummer and winter ranges are known, the extent of thesemovements and broad scale characteristics of winterrange has only been examined for Ontario in theunhunted Chapleau Crown Game Preserve (Welsh et al.1980). They clearly demonstrated a movement by themajority of the local population (about 260 animals)from cutover areas to a 200 square kilometre tract ofuncut mixed and coniferous forest adjacent to thecutovers. Residual stands of timber, even those occurringin 30 year old cutovers, were not heavily used in the latewinter. Movement appears associated with time of yearbut is enhanced by heavy short-term snow falls (Peek1971, Welsh et al. 1980, Addison et al. 1980). However,not all moose move long distances to late winter habitat;for example, some cows with calves were found to over-winter in cutover habitat in the claybelt (Thompson andVukelich 1981).

Although not specified by their authors, otherstudies in Ontario suggest movement to large blocks ofuncut forest. For example, McNicol and Gilbert (1980)found fewer moose in the cutovers during a year of deepsnow and crust compared to February of the previousyear when the snow was not so deep. Residual stands ofconifer within cuts were equally used in both years.Adjacent uncut tracts of coniferous forest were probablyused by the missing moose. Also aerial surveys formoose flown after late February generally result in fewermoose seen than earlier because of the shift to heaviercover. This is not new, but the size of the areas used inlate winter and the extent of movement as demonstratedby Welsh et al. (1980) was not previously documented.

Since natural regeneration of cutover moose rangein Ontario is to deciduous species or a deciduous-dominated mixed association, and because moose uselarge uncut areas in late winter, it is possible that localpopulations are limited by an insufficient amount of latewinter habitat. Further research on movements andwinter requirements on a macro-scale need to be carriedout to resolve this question.

Basic Habitat RequirementsOver the past decade, investigations into use of. minerallicks and/or aquatic feeding areas for sodium, summerfeeding areas, early winter and late winter habitat andearly and late winter concentration areas have occurred.Since not all animals change areas in different seasons,the late winter component can be subdivided into uncutforest and preferred habitat in cutover areas. Similarly,early winter areas may or may not be areas ofconcentrated and traditional use.

20

Aquatic Feeding Areas and Mineral Licks - Moose inOntario are believed to have a strong annual requirementfor sodium (Fraser and Reardon 1980). To fulfill theirneeds, mineral licks and/or aquatic macrophytes are usedas sources of this element (Fraser et al 1980).Apparently, few mineral licks exist in Ontario,undoubtedly due to the igneous and metamorphic rockwhich underlies most moose range. Where licks dooccur, they are used most heavily in late May to earlyJune when aquatic plants are not yet available. Certainspecies of aquatic macrophytic plants provide a bettersource of sodium than mineral springs, and moose switchto aquatic feeding in June once these plants are largeenough. These areas are frequented until late July whenuse declines markedly. Males apparently predominate atboth licks and in aquatic feeding areas early in theseason, with females arriving slightly later (Cobus 1972,Fraser et al. 1980, 1982).

As would be expected, not all waterbodies arepreferred equally and, even within preferred lakes orstreams, certain bays are used more gm others as a resultof differing concentrations of minerals (Fraser et al.1980, 1982). Preferred lakes generally appear to have amineral soil substrate and are traditionally used. Whilemoose may deplete preferred plants in these areas, theycontinue to use them probably because of the superiorquality of plants which remain (Fraser and Hristienko,1983).

Aquatic feeding areas and mineral licks draw moosefrom considerable distances. Fraser et al. (1980)estimated 35 different moose used a single lake whichthey studied in the Chapleau Crown Game Preserve.While it is impossible to directly assess distances movedby these animals, an estimate of up to 30 kilometres isreasonable based on distribution and densities of moosein this area.

In Ontario, many waterbodies are protected by theuse of shoreline reserves (strips of uncut timber 30 to 150metres wide) left along edges of lakes or streams duringlogging of surrounding areas. A study designed, in part,to assess the value of these reserves to moose using lakesfor aquatic feeding failed to detect any differencebetween use of lakes with or withoutstanding timber atthe shores (Brusnyk and Gilbert 1983). However, thesample sizes were very small, it was unknown if all lakeswere used equally before logging of the area, no sex andage information was presented, and there wasconsiderable variability in the amount and species ofplants available in the lakes studied. More work needs tobe done on this subject. At the population level, no dataexist as to the potentially limiting nature of sodium orpreferred aquatic macrophytes as has been suggested forIsle Royale by Jordan et al. (1973).

Summer Habitat - This is perhaps the least knowncomponent of moose habitat due in part to the difficultyin studying moose at that time of the year. It appears thatuntil mid-July habitat use is strongly influenced by areasused for aquatic feeding (Keamey and Gilbert 1976,loyal and Scherrer 1978, Brusnyk and Gilbert 1983).Kearney and Gilbert (1976) showed increased use ofopen, upland

Deciduous habitat types in late summer, as did Addisonet al. (1980_. In the latter case, 3 radio collared animalsmade, use of a ten-year old bum with deciduous and jackpine regeneration.

Although several studies have indicated onlylocalized movement in summer, large summer ranges of90 square kilometres for an adult male and 32 squarekilometres for a yearling in northwestern Ontario havebeen shown (Addision et al. 1980).

Early Winter Habitat - Early winter habitat is definedhere as habitat used until snow and/or crust forces mooseinto heavy coniferous areas. It is not necessarily a simplefunction of time of year.

Generally early winter habitat differs from earlywinter concentration areas. These latter sites are smallareas (2-10 km2) used traditionally by large numbers ofmoose during the November to early January period (eg.densities of up to 10moose/km2) (Thompson et al.1981). Concentration areas are topographically discrete,upland open sites with abundant browse. An apparentearly winter concentration area in the Quetico Parkshowed heavy traditional use over at least eight years.Them, moose were depleting the available browse(McNicol et al. 1980) indicating such areas wouldbenefit from management for shrub species.

In general, early winter moose habitat may not differfrom areas used in late summer and fall, although few dataexist for the latter period. The majority of data on winterhabitat comes from logged areas and indicate open,upland areas are used most. Welsh et al. (1980) showedlittle selection for age of cut in November but heaviest useof younger cuts in early winter with a progressivetendency to use older cuts and in cut areas as winterprogressed. Areas used most heavily were those with mosttopographic relief and with less than 40 percent of theoriginal time cover removed. McNicol and Gilbert (1980)found greatest use of stands characterized by a relativelyopen canopy with scattered conifers and deciduous treeswith a basal area of 2.5 metres square per hectare,averaged for the cutover.

The presence of coniferous trees as uncut residual(McNicol and Gilbert 1980, Thompson and Vukelich1981), in shoreline reserves (Brusnyk and Gilbert 1983) oras uncut border (McNicol and Gilbert 1980, Hamilton etal. 1980) appears to influence use of logged areas in mostyears. The influence may be more important in huntedthan unhunted populations. While this interspersion ofedge and open area appears important, Hamilton et al.(1980) found browsing by moose was unaffected by coverin the second year of their study. They attributed this tomore difficult snow conditions in the first year. SimilarlyMcNicol and Gilbert (1980) found little pattern in use ofcutovers when snow did not impede movements. Cowswith calves are more restricted in movement from covereven in easy winters, generally wandering less than 60metres from edge either as uncut or residual stand.

21

(Thompson and Vukelich 1981).Novak and Gardner (1975) noted fewer cows with

calves in moose concentration areas than were known tobe in their population near Kirkland Lake. This indicationthat different social classes of moose may have differinghabitat requirements was suggested also by Peek (1971)and Peterson (1977). Thompson and Vukelich (1981)working in the claybelt region of the boreal forest zonewere able to discriminate between areas used by cowswith calves in early winter, late winter, and from sitesnever used by this social group. Cutovers used by cowswith calves in early winter averaged 64 ha and were mostoften in lowlands, differing somewhat from large cuts onupland sites generally reported used by other moose inOntario. Best early winter habitat was in areas cut morethan 18 years ago containing several small residual standsof mixed or coniferous trees with an average coniferousbasal area of 9.5 metres square per hectare (mostly in treesunder 10 metres) and a maximum distance between uncutedges of half a kilometre.

Late Winter Habitat -Late winter habitat is definedhere as those areas used by moose once movements arerestricted by snow conditions. Animals may still usecutover areas during Ns period, but most abandon thementirely in favour of uncut forest, especially in March.Some areas used in late winter contain high moosedensity and have variously been called yards, winterconcentration areas or high density areas. As with earlywinter concentration areas, late winter yards aregenerally found in upland habitat but are more densityforested (Telfer 1967, Poliquin et al. 1977). These areasmay be more or less heavily used in successive years,although the reason for this different rate of occupancy isnot understood.

There is in general, reduced use of open areas inlate winter caused apparently either by a rapidaccumulation of snow and/or crusted deep snowconditions (Des Meules 1964, Hamilton et al. 1980,McNicol and Gilbert 1980, Welsh et al. IP80). Even useof residual stands within cutovers decreases during thisperiod and Hamilton et al. (1980) reported browsingonly up to 80 metres from over. Cows and calves aregreatly affected, moving only about 12 metres fromcover (Thompson and Vukelich 1981).

Welsh et al. (1980) showed use of older cuts onmore rugged terrain in later winter, and finally amovement out of cutovers altogether into an extensivearea of uncut forest. Late winter ranges in an uncut areareported by Addision et al. (1980) were in black spruceswamp for 2 yearlings and an adult male, and in a rollingupland area of tall aspen over younger, dense balsam firfor an adult female.

Cows with calves in the claybelt area may notmigrate out of cutovers during later winter, althoughthere is no empirical evidence to support this assumption.Habitat used in cutover areas by cows with calves duringlate winter had several small cut areas averaging about 16ha, with a maximum distance between cover edges of 370

metres. They were generally in lowland areas with mesicor upland components on sites cut at least 25 years ago.Residual stands of uncut or partially cut timber werelarger than those used in early winter and had moreconifer, mostly large and small balsam fir (Thompson andVukelich 1981).

Late winter is an extremely sedentary period formoose with a substantial reduction in the size of homerange (Goddard 1970, Phillips et al. 1973). Addision et al.(1980) using radio telemetry reported winter ranges of 2,3and 12 square kilometres for an adult female, a yearlingmale and an adult male respectively. Cows with calvesaveraged only 29 ha activity areas in late winter(Thompson and Vukelich 198 1).

No effort has been made on a broad scale to assesswhether adequate late winter habitat is available to moosegenerally in Ontario or for cows with calves in particular.Also there remains a need to characterize uncut standsused by moose in March and April.

Food RequirementsWithin the broader context of major habitat units andcomponents of these, food requirements must also beconsidered. This is the most refined, or third scale, ofunderstanding moose habitat with which managers mustwork.Aquatic Plants - There is strong evidence that moosefeed on aquatic plants due to a seasonal sodium hunger(Jordan et al. 1973, Fraser et al. 1982). Species which arehighest in sodium content or most abundant, althoughcontaining slightly less sodium, are most important. Insome traditionally well-used lakes certain preferredspecies, such as Potamogeton filiformis and Nupharvariegatum, have largely been eliminated. Sodium richspecies from Sibley Provincial Park include Utriculariavulgaris, Sparganium angustifolium, Myriophyllumexalbescens, Potamogeton epihydrus, P. gramineus, P.filiforimis and Nuphar variegatum. One other species,Eleocharis acicularus, was reported as high in sodium inthe Chapleau Game Preserve but low at Sibley, (Fraser etal. 1980, 1982) indicating some regional differences inspecies ability to concentrate this element.

Aquatic plants are richer in protein early in theseason and, although this declines later, they maintainvalues equivalent to woody species. On Isle Royale itappears that sodium may be a critical and limiting factorto moose (Belovsky and Jordan 198 1) but Fraser et al.(1981) suggested this is not likely for Ontario. However,the claybelt region is relatively poor in aquatic feedingareas and further studies are needed to determine ifperhaps sodium is limiting in that area.

Browse Species - Preferences, Importance andAbundance - Recent studies where use of availablebrowse by moose was assessed are summarized in Table1. While the methods of collection differed among allstudies, prohibiting direct comparison, certaingeneralizations can be22

made. In early winter, when snow does not impedemovement and does not cover many stems, Salix spp. arethe most important (% of total eaten) browse species. Inlate winter shade tolerant species become important,including Corylus cornuta and Amelanchier spp. Thewide range of other abundantly eaten species reflects thevariety of micro-site types and cosmopolitan nature of thediet of the moose. Among the preferred browse species(% removed greater than % available), Sorbus americanaand Salix spp. are commonly eaten over Ontario's borealrange.

Generally, percentage of stems removed of what wasavailable is low indicating no shortage of food (Table 1).In two cases percentage use was high: an apparent earlywinter yard in Quetico Park (McNicol et al. 1980) andlate winter sites used by cows with calves in the clay belt.In the latter study, data represent areas where feedingoccurred and did not randomly assess availability. Thework in Quetico Park suggests extensive use and the needto manage traditional early winter areas if managers wishto maintain them. However, in general it does not appearthat the amount of browse is limiting to moose in Ontario.This is the same conclusion reached by Crête and Jordan(1983) in Quebec.

Research is needed to assess browse availability intraditionally used late and early winter areas, and tocompare this to areas where animals are not found. Whilebrowse over the entire range is not limiting animals, theymay need specific nutrients; no data exist for this difficultproblem. Also little information exists on use of leaves insummer or on the observed importance of dogwood tomoose after the rut period.

Summary of Moose HabitatMoose in Ontario apparently can be grouped into thosewhich undergo seasonal movements and those which donot; those which concentrate and those which do not;and, within the latter, a subgroup of cows with calveswhich require isolation

Movements by moose are influenced by severalfactors during the year. In the spring their requirementfor sodium may cause the animals to move to atraditionally used mineral lick and/or a particularwaterbody with preferred aquatic plants. Use of habitat atthis time is strongly influenced by location of these licksand lakes. A general spring movement from winter tosummer range probably also occurs. After early summer,a more gradual movement likely occurs to upland ormesic sites when feeding is largely on leaves of preferredspecies. Feeding patterns and preferred plant types duringthis period are not well known. If specific habitatrequirements exist during the rutting period, they are alsounknown as are habitat use and browse preferencesimmediately after the rut. However, if animals are activeand eat less during this period, post-rut feeding may beimportant in building nutrient reserves prior to winter.Early winter concentration areas and use of dogwood(Cornus stolonifera),

especially along rivers, may actually be post-rut feedingareas. Movement of animals which concentrate in theselatter sites probably begins in late October with amovement out by mid-January. Use of this type of yarddiffers from the use of late winter yards, as habitatoccupancy in late winter concentration areas is notconsistent among years. Further, it is unclear whether latewinter areas in cutover sites are well used until April, or ifthese animals also move into the uncut forest. Use andcharacteristics of uncut forest in late winter have not beenstudied.

Not all moose concentrate or move and some animalswill always be found in apparently less than optimalhabitat, such as cutovers in late March. Cows with calvesmay be one exception to generalized movements. Theyappear to select suitable habitat in isolated areas andcontinue to use cutovers in late winter. In both early andlate winter habitat in logged areas, the amount of coniferpresent influences the amount of use by moose. Fewestimates are available as to how much conifer left in acutover after logging is optimal, but Welsh et al. (1980)found higher use in areas where less than 40% of alltimber had been removed.

Habitat in general in Ontario is probably not limitingto moose populations. However, in local situations latewinter habitat and sodium availability may be critical.Other micro-aspects of moose range, such as nutritionalquantity of browse, are important locally in influencinghabitat use and perhaps population levels and deservefurther attention.

Managers in Ontario should view moose habitat atthree scales:1) the population level which encompasses large areas ofmanagement units to ensure late winter habitat,particularly maintaining large blocks of uncutpredominantly coniferous forest in rolling upland areas inassociation with early seral areas;2) a general level which considers individual cut plans toameliorate these into producing good moose habitatthrough both general knowledge of where upland mixedridges occur and specific knowledge of early and latewinter concentration areas, mineral licks and primeaquatic feeding areas, and,3) an individual level which reflects knowledge ofpreferred foods and where these am likely to be found in amanagement area before or after cutting.

It is our view that while habitat is not presentlyknown to limit the moose population, when new huntingregulations permit a population increase then habitat andhabitat-predator interactions will become key elementsinfluencing moose density.

23

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Addison, R.B., J.C. Williamson, B.P. Saunders and D.Fraser. 1980. Radio-tracking of moose in the borealforest of northwestern Ontario. Can. Field-Natur.94:269-276.

Anderson, R.C. 1963. The incidence, development andexperimental transmission of Pneumostrongylustenuis (Dougherty) of the meninges of the white-tailed deer (Odocoileus virginanus borealis) inOntario. Can. J. Zool. 41:775-792.

Anderson, R.C. 1972. The ecological relationships ofmeningeal worm and native cervids in NorthAmerica. J. Wildl. Dis. 8:304-310.

Anon. 1977. Manual of Forest Management PlanRequirements for the Province of Ontario. SecondEd. Ont. Min. Nat. Res. 56pp.

Armson, K.A., F.C. Robinson, and J.E. Osborn. 1980.Forest management manual for the Province ofOntario. Ont. Min. Nat. Res. 132pp.

Belovsky, G.E. and P.A. Jordan. 1981. Sodium dynamicsand adaptations of a moose population. J. Mammal.62:613-621.

Bergerud, A.T., W. Wyett, and B. Snider. 1983. The roleof wolf predation in limiting a moose population. J.Wildl. Manage. 47(4):977-988.

Brusnyk, L.M. and F.F. Gilbert. 1983. Use of shorelinetimber reserves by moose. J. Wildl. Manag. 47:673-685.

Carleton, T.J. and D.F. Maycock. 1978. Dynamics of theboreal forest south of James Bay. Can. J. Bot. 56:1157-1172.

Chamberlin, L.C. 1981. Managing for fish and wildlifevalues within the forest management planningprocess. Alces 17:193-228.

Cobus, M. 1972. Moose as an aesthetic resource and theirsummer feeding behaviour. Proc. N. Amer. MooseConf. and Workshop. 8:224-275.

Crête, M. and P.A. Jordan. 1982. Population consequencesof winter forage resources for moose. Alces alces, insouthwestern Quebec. Can. Field-Natur. 96(4):467-475.

Cummings, H.G. 1974a. Annual yield, sex and age ofmoose in Ontario as indices to the effects of hunting.Naturaliste can., 101:539-558.

Cummings, H.G. 1974b. Moose management in Ontariofrom 1948 to 1973. Naturaliste Can. 101:673-687.

Des Meules, P. 1964. The influence of snow on thebehaviour of moose. Trans. Northeast Wildl. Conf.21:138-160.

Eason, G., E. Thomas, R. Jerrard, and K. Oswald. 1982.Moose hunting closure in a recently logged area.Alces 17:111-125.

Euler, D.L. 1981. A moose habitat strategy for Ontario.Alces 17:180-192.

Euler, D.L. 1983. Selective harvest, compensatorymorality and moose in Ontario. Alces. 19: (in press).

Euler, D.L. 1984. Moose and man in northern Ontario.Forestry Chronicle (in press).

Flemming, S.T., and K. Koski. 1976. Moose habitatstudies of moose management unit 40 with particularreference to the effects of roads and cutovers.Unpubl. - Ontario Ministry of Natural Resources 20pp. (Available from Wildlife Branch. ON4NR.Toronto M7A lW3).

Fraser, D. and E. Reardon. 1980. Attraction of wildungulates to mineral-rich springs in central Canada.Holartic Ecol. 3:36-AO.

Fraser, D., D. Arthur, J.K. Morton and B.K. Thompson.1980. Aquatic feeding by moose in a CanadianLake. Holartic Ecol. 3:218-223.

Fraser, D., B.K. Thompson and D. Arthur. 1982. Aquaticfeeding by moose: seasonal variation in relation toplant chemical composition and use of mineral licks.Can. J. Zoology. 60:3121-3126.

Fraser, D. and H. Hristienko. 1983. Effects of moose,Alces alces, on aquatic vegetation in SibleyProvincial Park, Ontario. Can. Field - Natur. 97:57-61.

Gasaway, W., R.O. Stephenson, J.L. David, P.K.K.Shepperd and R.O. Burris. 1983. Interrelationshipsof wolves, prey and man in Alaska. Wild. Monog.84.51 pp.

Goddard, J. 1970. Movements of moose in a heavilyhunted area of Ontario. J. Wildl. Manage. 34:439-445.

Hamilton, G.D., P.D. Drysdale and D.L. Euler. 1980.Moose winter browsing patterns on clear-cuts innorthern Ontario. Can. J. Zool. 58:1412-1416.

Houser, A.M. 1972. Basis for assessing land capabilityand degree of effort for wildlife in Ontario. Proc. N.Amer. Moose Conf. and Workshop. 8:209-221.

Jones, R.K., G. Pierpoint, G.M. Wickware, J.K. Jeglum,R.W. Arnup, and J.M. Bowles. 1983. Field guide toforest ecosystem classification for the clay belt, siteregion 3e. Ontario Min. Nat. Res. Toronto. 123pp.

Jordan, P.A., D.B. Botkin, A.S. Dominski, H.S. Lower-forf and G.E. Belovsky. 1973. Sodium as a criticalnutrient for the moose of Isle Royale. Proc. N.Amer. Moose Conf. and Workshop. 9:13-42.

Joyal, R. and B. Scherrer. 1978. Summer movements andfeeding by moose in western Quebec. Can. Field-Natur. 92:252-258.

Kearney, S.R. and F.F. Gilbert. 1976. Habitat use bywhite-tailed deer and moose on sympatic range. J.Wild. Manage. 40:645-657.

Kolenosky, G.B. 198 1. Status and management of wolvesin Ontario. pp. 35-40, in Wolves In Canada AndAlaska (Ed. L. Carbyn). C.W.S. Report Series No. 5.

LeResche, R.E. 1974. Moose migration in North America.Natur. Can. 101:393-415.

McNicol, J.G., and F.F. Gilbert. 1980. Late winter use ofupland cutovers by moose. 1. Wildl. Manage.44:363- 371.

24

McNicol, I.G., H.R. Timmerman and R. Gollat. 1980. Theeffects of heavy browsing pressure over eight yearson a cutover in Quetico Park. Proc. N. Amer. MooseConf. and Workshop. 16:360-373.

Novak, M., and ].F. Gardner. 1975. Accuracy of aerialmoose surveys. Trans. North Am. Moose Conf.11:154-180.

Parks Canada. 1980. Moose management plan forPukaskwa National Park.

Peek, J.M. 1971. Moose-snow relationships innortheastern Minnesota. In Proceedings of theSymposium on snow and ice in relation to wildlifeand recreation. Edited A.0. Haugen. Iowa StateUniv. Press, Ames. pp. 3(,-49.

Peek, J.M., D.L. Urich, and R.J. Mackie. 1976. Moosehabitat selection and relationships to forestmanagement in northeastern Minnesota. Wildl.Monogr. 48. 65pp.

Petersen, K.L. 1955. North American Moose. Univ. ofToronto Press. 280 p.

Peterson, R.O. 1977. Wolf ecology and prey relationshipson Isle Royale. U.S. Natl. Park. Serv. Sci. Monogr.Ser. No. 11.

Peterson, R.O. and D.L. Allen. Snow conditions as aparameter in moose-wolf relationships. NaturalisteCan. 101:481-492.

Phillips, R.L., W.E. Berg, and D.B. Siniff. 1973. Moosemovement patterns and range use in northwesternMinnesota. J. Wildl. Manage. 37:266-278.

Poliquin, A., B. Scherrer, and R. Joyal. 1977.Characteristics of winter browsing areas of moose inwestern Quebec as determined by multivariateanalysis. Proc. N. Amer. Moose Conf. andWorkshop. 13:128-143.

Rowe, J.S. 1972. Forest Regions of Canada. Can. For.Serv. Publ. No. 1300. 172pp.

Saunders, B.P. and J.C. Williamson. 1972. Moosemovements from ear-tag returns. Proc. N. Amer.Moose Conf. and Workshop. 8:177-184.

Telfer, E.S. 1978. Habitat requirements of moose - Theprincipal taiga rouge animal. Proc. First InternationalRangeland Congress: 462-465.

Telfer, E.S. 1967. Comparison of a deer yard and a mooseyard in Nova Scotia. Can. J. Zool. 45:485-490.

Thompson, I.D. 1980. Effects of weather on productivityand survival of moose in northeastern Ontario.Proceedings North. Am. Moose Conf. and Workshop.16:463-481.

Thompson, I.D., and M.F. Vukelich. 1981. Use of loggedhabitats in winter by moose cows with calves innortheastern Ontario. Can. 1. Zool. 59:2103-2114.

Thompson, I.D., D.A. Welsh and M.F. Vukelich. 1981.Traditional use of early-winter concentration areasby moose in northeastern Ontario. Alces. 17:1-14.

Timmerman, H.R. and R. Gollat. 1983. Age and sexstructure of harvested moose related to seasonmanipulation and access. Alces 18:-

Welsh, D.A., K.P. Morrison, K. Oswald and E.R.Thomas. 1980. Winter habitat utilization by moosein relation to forest harvesting. Proc. N. Amer.Moose Conf. and Workshop. 16:398-428.

25

TABLE 1:

Summary of browse species preference and most important (abundant) species in diets of moose inOntario.Percentage of stems available that were browsed is in brackets. From published studies, 1974-1983. Author/Area (methods)

Author/Area (Methods)

Hamilton and Drysdale(1975) - south andcentral (2m2 plots every25 m along transects)

McNicol et al. (1980)- south and central(.6m x 20m plots 80mintervals on transects

McNicol and Gilbert (1980)- south and central(12.2 x 20m plots on lines)

Thompson and Vukelich(1981)-Claybelt(100 x 1 m plots placedin feeding areas)

Brusnyk and Gilbert (1983)-south central(2m radius circuits plots)

Time of Winter Important Species (% use) Preferred Species (% use)

early

1) early

2) earlyand late

1) early

2) late

unknown

Corylus cornuta (62.3)Acer pensylvanicum (49.7)Salix spp. (84. 1)Betula papyrifera (60.7)

Betula papyrifera ()Salix spp-. ()Sorbus americana

Betula papyriferaSorbus americanaCorylus corrutaSalix spp.

Salix spp. (37.1)Cornus stolonifera (56.6)

Amelanchier spp. (65. 1)Salix spp. (48.3)Corylus cornuta (22.5)

Corylus cornuta (24.3)Acer pensylvanicum (12.7)Salix spp. (43.7)

26

Sorbus americana (73.3)Salix spp. (84. 1)Prunus pensylvanica (50.0)Betula papyrifera (60.7)

Salix spp.Sorbus americanaPrunus pensylvanica

Corylus corrutaSorbus americanaBetula papyriferaSalix spp.

Cornus stolonifera (56.6)Acer pensylvanicum (5 1.7)Sorbus americana (40.7)Salix spp. (37. 1)

Sorbus americana (78. 1)Amelanchier spp. (65. 1)Salix spp. (48.3)Prunus spp. (37.5)

Sorbus americana (24.8)Salix spp. (43.7)Corylus cornuta (24.3

Salix spp. (41.4)Prunus spp. (32.4)Populus tremuloides (14.8)Amelancier spp. (13.4)

n/a

Effects of Forestry Practices onUngulate Populationsin the Boreal Mixed Wood Forests*

J.G. McNicol, Regional Habitat Biologist and H.R.Timmermann, Regional Moose Biologist North CentralRegion, Ontario Ministry of Natural Resources ThunderBay, Ontario

Moose, white-tailed deer and woodland caribou allinhabit the boreal forest but, for a variety of reasons, themoose is the most likely ungulate to be found commonlyutilizing boreal mixedwoods. The paper examines howvarious forestry practices affect the suitability of borealmixedwoods as moose habitat.

Si la forêt boréale héberge aussi bien l'Original et leCerf de Virginie que le Caribou des bois, l'Originalest l'ongulé que l'on rencontre le plus communémentdans les peuplements mélangés de cette forêt. Dansle présent document, les auteurs étudient lesrépercussions des différentes mé'thodes d'exploitationsur ces peuplements mélangés en tant qu'habitat del'Original.

Three distinct species of ungulates-white-tailed deer(Odocoileus virginianus), woodland caribou (Rangifertarandus caribou) and moose (Alces alces), all membersof the deer family, utilize portions of the boreal forestecotone. The utilization of the mixedwood cover type inOntario by these species is governed by their habitatrequirements and by ecological and environmentalfactors.

Deer expanded their range northward into the borealforest in the early 1900s following logging and landclearing (Cumming and Walden 1970). During extendedperiods of mild winters (i.e., winters with below-averagesnowfall), and where logging provided abundant food,deer moved as far north in Ontario as Kirkland Lake,Sioux Lookout and Red Lake (Euler 1979) (Fig. 1).

Leaves, stems and buds of woody plants areconsidered by many to be the mainstay of deer diet,mainly because they are available through the year (Halls1973). The greater the variety of plants, the greater thechances for the whitetail to achieve its productivepotential (Halls 1978). Deer feed on the leaves and tipsof shrubs and trees including aspen (Populus spp.),cherry (Prunus spp.), mountain maple (Acer spicatumLain.), willow (Salix spp.), bearberry (Arctostaphylosspp.), hazel (Corylus spp.), honeysuckle (Lonicera spp.),juneberry (Amelanchier spp.) and a great variety ofherbaceous plants.

Figure 1. Changes in the white-tailed deerdistribution from 1620 to 1975 (Smith andBorczon 1977) and ft present woodland cariboudistribution (Bergerud 1978) in Ontario.

* Boreal Mixedwood Symposium Proc. p. 141-154)27

APPENDIX IV

In winter (the most critical season), they browse on thewoody vegetation of trees and shrubs. During thisperiod, the average deer requires approximately 2.3 kg ofgood quality browse per day (Smith and Borczon 1977).

Logging opens the forest canopy, and as a resultthere is more food available for deer. Adjacent maturestands of eastern white cedar (Thuja occidentalis L.),spruce (Picea spp.), balsam fir (Abies balsamea [L.]Mill.) and pine (Pinus spp.), if left uncut, provide theshelter needed in winter. However, the populationdensities of deer in the mixedwood component of theboreal forest depend primarily on the depth to whichsnow accumulates during winter, the duration of the snowcover and the suitability of range conditions followinglogging. Winter weather and the increasing size ofclearcuts have been generally unfavorable for deerinhabiting the boreal forest areas since 1955, and as aresult both the range (Fig. 1) and local populationdensities have decreased.

Most woodland caribou in Ontario, of which anestimated 13,000 remain (Simkin 1965), are found in thenorthern, unlogged boreal forest and the Hudson Baylowlands. A few scattered local herds survive in thecommercial forest zone along the north shore of LakeSuperior, north of Lake Nipigon and in the Irregular Lakearea northwest of Kenora (Bergerud 1978) (Fig. 1). Formany years, the disappearance of woodland caribou inthis area has been attributed to environmental changessuch as forest fires, settlement and logging. More recentevidence suggests that hunting mortality, disease andpredation also play an important role.

Woodland caribou are gregarious and prefer to usethe more mature boreal forest habitats (Cringan 1957).Tree and ground lichens are important winter foods.Caribou eat a wider variety of plants than do other deerspecies and are highly adapted, as well as adaptable, totheir environment (Bergerud 1978). A partial list of plantseaten includes 282 kinds of seed plants and 62 lichens aswell as green vascular plants and mushrooms.

During summer, caribou concentrate on the leaves ofdeciduous trees and shrubs, especially those of willow,birch (Betula spp.) and blueberry (Vaccinium spp.).During the fall, after abscission, the animals switch toterrestrial lichens and leaves of coniferous species(Bergerud 1978). In winter, caribou frequently "crater"(dig down through the snow) to reach lichens and vascularplants. They also seek the fine twigs of browse speciessuch as birch, willow, cherry and blueberry, and mayutilize closed canopy habitats, when these are available, insearch of arboreal lichens growing on coniferous andsome deciduous trees.

The moose is well adapted to the boreal forest (Fig.2), where important habitats are produced in the earlyseral stages of plant succession (Krefting 1974). Habitat,according to Franzmann (1978), is the primary limitingfactor of moose populations throughout their NorthAmerican range. The reproductive potential of moose isadapted to maintenance of the species at low densities(115 km2), rapid expansion into larger areas of goodhabitat following fire and logging (1/km2) and slowcontraction following forest maturation (Geist 1974).

Figure 2. Moose distributionin Ontario (Cumming 1972).

Large quantities of forage (18-25 kg/day) arenecessary to maintain an adult moose (Gasaway andCoady, 1974). Mixed stands of conifers and hardwoods incontinuous succession are favored (Pimlott 1953, Peterson1955, Dodds 1974). In winter, moose tend to select siteswhere snow depths allow for bedding in comfort and donot restrict movement to feeding areas (Dodds 1974).

Key browse species in eastern North American mooserange are balsam fir, white birch (Betula papyriferaMarsh.) and trembling aspen (Populus tremuloidesMichx.). Hazel, mountain maple, mountain ash (Sorbusamericana Marsh.), willow, Juneberry, dogwood(Cornus spp.) and cherry are other major browse foods.In summer and early fall, habitat use is centred on uplanddeciduous tree stands (Krefting 1974).

In early winter, young successional mixedwoodstands created by disturbances such as fire, budworm orlogging are preferred. As the winter advances and snowdepths increase, moose gradually shift their habitat fromopen mixedwood stands to denser coniferous cover (DesMeules 1965, Telfer 1970, Peek 1971). The latter,consisting primarily of balsam fir and spruce, providesthe moose with shelter during this critical period of theyear.

Wildfire and logging have generally benefitedmoose by providing large areas of early successionalgrowth containing a much greater variety and quantity offood than mature forests (Cumming 1972). Moosepopulations generally increase in such disturbed areas(Aldous and Krefting 1946, Krefting 1951, Spencer andChatelain 1953, Peterson 1955, Peek 1971). Over the past15 years, wildfire has generally become a less importantfactor in the creation of early successional stages becauseof man's increasing efforts to prevent and suppress forestfires.

Timber harvesting, on the other hand, has increasedin importance as a major disturbance factor in the borealforest over the same period. Timber harvesting iscurrently the most predictable major disturbance in theboreal forest. Meaningful habitat management of borealungulates, primarily moose, is dependent on therecognition by forest managers of the need in some areasto maintain a variety of habitats through the modificationor restriction of some forestry practices. An examinationof how such practices in boreal mixedwoods affectmoose will illustrate how forest management can eitherbenefit or hinder habitat management goals.Timber HarvestingThe preceding discussion on the habitat requirements ofmoose indicated the importance of a diversity of woodyvegetation types and age classes within the animal'snormal range. Timber harvesting in boreal mixedwoodtypes can enhance the inherent species and age classdiversity of these stands. The removal of the matureconiferous component from conifer-dominated stands,while leaving the mature deciduous component and ad-

vance coniferous regeneration, results in good moosehabitat for several reasons:1. Opening the canopy allows the establishment ofimportant shade-intolerant browse species such as whitebirch and trembling aspen, which augment shade-tolerantbrowse species already in the understory.2. The residual deciduous component providesimmediate age class diversity on the disturbed site inaddition to escape cover and increased "edge".3. Advance coniferous regeneration in the cutover pro-vides immediate early winter cover, escape cover andvegetational species diversity.

The boreal mixedwood forest type alone has thepotential to offer moose this variety of habitatadvantages, provided that only the mature coniferouscomponent is harvested. Welsh et al. (1980) found onstudy areas in the Chapleau Crown Game Preserve that,in midwinter, with a choice of a variety of cut and uncuthabitats, moose preferred cutover mixedwood standsfrom which up to 40% of the original composition hadbeen removed.

McNicol and Gilbert (1980) reported that within 16boreal mixedwood cutovers they studied in the ThunderBay area, moose habitat utilization during January andFebruary was concentrated in areas supporting ascattered residual coniferous and deciduous component(basal areas equal, at approximately 2.5 m2/ha). Resultsfrom this study showed that the scattered residual covertype produced a much greater abundance and diversity ofbrowse species (10) than did the three other availablecover types delineated within the cutovers. Except forthe dense conifer cover type (residual blocks ofunmerchantable conifer), the other less productive covertypes were characterized by the absence of a mature orsemimature residual component.

An abundant and diverse browse source is animportant element of good moose range, particularly inwinter. Browse consumption by moose at this time isgenerally lower than in summer (Le Resche 1970, Verme1970), probably because food is less digestible and istherefore retained longer in the rumen, as had been notedfor other ruminants (Corbett 1969). Under winterconditions, about one third of the energy metabolized bymoose may have to be provided through thecatabolization of body fat and protein reserves (Gasawayand Coady 1974). The movements necessary to collectenough browse to maintain rumen-fill may requiresignificant energy expenditures, especially if snowconditions are limiting to moose movement. Anabundant browse source reduces the energy expenditurenecessary to collect browse.

A selection of different browse species is importantsince changes in snow conditions can cause majorchanges in the species which moose prefer to browse(Peek et al. 1976, McNicol and Gilbert 1980). A varietyof browse species consumed in the winter diet probablyenhances the digestibility of this fibrous food source aswell (Mellen-berger et al. 197 1).

The coniferous component which often develops in29

the understory of boreal mixedwood stands seems to beimportant as both escape and survival cover for moosefollowing the harvest of the mature conifer component inthese stands. Moose bed an average of five times a day(Franzmann et al. 1976), probably to ruminate, as dodomestic ruminants. Studies in Quebec (Des Meules1965) and Ontario (McNicol and Gilbert 1978) show thatmoose prefer to bed within 2mof coniferous cover whenutilizing open habitats during the winter. Indirectevidence indicates that moose select bedding sites inclose proximity to immature coniferous cover on openhabitats to reduce wind chill (McNicol and Gilbert1978). Accumulated snow in the lee of these windbreaksoften tends to be softer and more compressible since itlacks a wind crust (Des Meules 1965).

Advance coniferous regeneration, primarily balsamfir, is often found in close association with goodbrowsing opportunities in mixedwood cutovers (McNicoland Gilbert 1980). Since browsing, bedding andruminating are related activities, close proximity of foodand vertical protective cover is an attractive feature ofmixedwood cutovers.

The preceding discussion on the value ofmixedwood stands after timber harvesting assumes apartial cut where only merchantable conifers are removed.This type of timber harvest has been common in borealmixedwood stands in northern Ontario and hasaccidentally resulted in excellent moose habitat overmany areas. Quite simply, the lack of deciduous fibremarkets in many areas of Ontario has improved mooserange. However, the same global demand for wood fibrewhich caused the liquidation of Ontario white pine (Pinusstrobus L.) and red pine (P. resinosa Ait.) and thecontinued depletion of this province's spruce and jackpine (P. banksiana Lamb.) stands, has now focused onboreal hardwoods.

The results of the inevitable growth in the deciduouswood fibre market will be a substantial loss of age classand species diversity on boreal mixedwood sites. The lossof the mature hardwood component and destruction ofbrowse and cover species in the understory duringharvesting, will prove as negative for moose as previouscutting practices proved positive.

Clearcutting of boreal mixedwood stands will provedetrimental not only on a small scale (individual cutovers)but also on a large scale (license areas). Wildlifemanagers will no longer be able to rely on mixedwoodstands to buffer large coniferous clearcuts. The result wingive new meaning to the word clearcut, provided thattimber companies do not adopt modified cutting practiceson a large scale. The adverse temporal and spatialdevelopment of moose habitat on contiguously clearcutareas would limit moose populations.

Silvicultural TreatmentsBoreal mixedwood forest types are often found onproductive sites. A variety of silvicultural treatmentsmay be applied by forest managers to encourage theregrowth of merchantable coniferous stands on thesesites. The following comments relate to the beneficial ordetrimental effects various treatments may have onmoose.ScarificationThe practice of scarification after logging is common innorthern Ontario. On mixedwood cutovers, scarification isgenerally carried out in preparation for the planting ofjack pine, or more commonly black spruce (Picea marian[Mill.] B.S.P.) and/or white spruce (P. glauca [Moench]Voss) seedlings.

Scarification followed by planting of coniferousseedling stock is an attempt by forest managers to truncatethe normal successional sequence following disturbanceon mixedwood sites. It is the early stage of successionfollowing fire (3-20 years) which historically hassupported growth in moose populations (Geist 1974, Irwin1975). Logged mixedwood stands have the potential forsupporting increased populations of moose for a similarperiod. However, scarification and planting tend toshorten this early successional stage and in turn mayreduce the period during which these areas can supporthigh moose densities (Telfer 1976).

Boreal mixedwood cutovers which have beencompletely scarified and artificially regenerated differfrom those which follow a natural course of successionand retain unmerchantable trees (Telfer 1976). Resultsfrom studies in Alberta (Stelfox et al. 1976) and Ontario(McNicol and Gilbert 1980) indicate that in scarified areasthe production of deciduous browse species is reducedand much delayed. Not only is the browse component onscarified mixedwood sites altered but conifer cover isaffected as well. The coniferous species which are plantedwill eventually produce both early and late winter coverfor moose. However, artificial regeneration does notduplicate the immediate habitat advantages of interspersedadvance coniferous regeneration which would havesurvived if scarification had not taken place.

In the past, many mixedwood cutover sites were notextensively scarified and planted because of heavyresidual concentrations left after the harvest of theconiferous component. If we assume an increased demandfor deciduous fibre, there will no longer be suchconcentrations on many mixedwood sites after harvest.More extensive scarification on mixedwood sites willlikely follow, com- pounding the degradation to moosehabitat initiated with the removal of the deciduouscomponent. Forest management agreements (FMAs)between private pulp and paper companies and thegovernment provide incentives for the companies toincrease conifer stocking on cutover sites. Sincemixedwood sites are often the most productive, it wouldseem logical for the companies to concentrate

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silvicultural efforts, including extensive scarification andplanting, on sites that could have been good mooserange.

'Tunneling", a form of scarification that is used lesscommonly, normally improves moose range. Thistechnique is used by forest managers to introduce amerchantable coniferous component into hardwoodstands. Heavy scarification equipment is initially driventhrough the stand, usually in a strip pattern, and conifersare planted in the scarified strips. Such plantingintroduces diversity and will eventually provide earlyand some late winter cover, improving the stand asmoose habitat (Telfer 1976).HerbicidesThese have proved useful in forestry for controllingstand composition, particularly in the early stagesfollowing logging or fire when forest managers areattempting to establish commercial species (Telfer1976). Boreal mixedwood cutovers, because of theinherent productivity of the sites and usually abundantdeciduous regeneration after disturbance, are oftenprime candidates for treatment. Chemical defoliants arealso used on mature or semimature mixedwood stands inan effort to release advance coniferous regeneration inthe understory.

The impact of herbicide applications and theutilization of treated sites by ungulates, primarily deer,have been examined by a number of researchers in theUnited States (Krefting and Hansen 1969, Beasom andScifres 1977, Tanner et al. 1978). Apart from atemporary reduction in use (usually just the year oftreatment), utilization of sprayed areas was either thesame as, or better than, utilization of similar untreatedsites.

Krefting and Hansen (1969) believed that aerialapplications of 2,4-D could be effective in increasingbrowse production in areas where access or labour costsfor cutting browse were problems. In Ontario, assessmentof a bum in the White River area which was subsequentlytreated with a combination of 2,4-D and 2,4,5-T showedno selection by moose for sprayed or unsprayed areas 3to 4 years after treatment (Barker and Malone 1972).

The effects of recommended application rates ofherbicides such as 2,4-D and 2,4,5-T differ according tothe browse species treated. White birch and pin cherry(Prunus pensylvanica L.f.) are often killed immediately,and there is little or no suckering to compensate for theloss. Mountain maple and mountain ash remain relativelyunchanged, since suckering of these species replacesstems which are killed within three or four years (Barkerand Malone 1972). The most common effect on themajority of browse species is a temporary setback,followed by subsequent regrowth.

The loss of pin cherry and especially white birchfrom treated mixedwood cutovers could significantlyaffect winter browsing activities by moose. McNicol andGilbert (1980) found that white birch and pin cherrywere two of five species most often browsed by mooseutilizing the 16 mixedwood cutovers they studied. Whitebirch twigs alone accounted for approximately 31% ofall

browse consumed by moose over two winter studyperiods.

Discussion thus far has centered on the use ofherbicides to set back deciduous regeneration temporarilyand "release" conifers. A less common use of herbicidesis for stand conversion, where all deciduous competitionin a mixedwood stand is killed through repeatedapplications of herbicides or where mixedwoodcutovesites are completely "sterilized" with herbicidesbefore conifers are planted. The ramifications of thisdrastic intervention into natural succession for moose anda variety of other wildlife species dependent on mid-successional forests are obvious. Mixedwoodmanipulation of this type must be considered carefullywith regard to wildlife requirements within the affectedarea.Prescribed burningAccumulated slash and logging debris can be removedeconomically and effectively from cutover sites throughthe use of fire. Bunnell and Eastman (1976) suggest that,after prescribed burning, site damage is generally lesssevere than it is after wildfire and that slash burningshould extend the early successional stages. In Maine,prescribed burning of logging debris in clearcuts resultedin greater production of hardwood stems, shrubs andherbaceous plants than in clearcuts where slash was leftundisturbed (Rinaldi 1970). Stoeckeler (1948) reportedthat root suckering of trembling aspen, an importantbrowse species for moose, was stimulated by light burns.

The benefits of a more nutritious browse source, asa result of the rapid return of nutrients to the soil, areprobably short term (Einarsen 1946, Taber 1973).However, the rapid invasion of fire-disturbed sites byungulates (Stenlund 197 1) and heavy initial utilizationof post-fire vegetation (Randall 1966) may be a responseto an increase in quality as well as quantity of browse(Cumming 1972).

The beneficial effects for ungulates of prescribedburning in the Lake States may last 10-15 years beforethe canopy closes over (Krefting 1962). Ahlgren (1973)suggested infrequent (once per tree generation) use offire in the Lake States forests because of thinner soils,slower accumulation of organic material, shorter growingsea- sons for competing vegetation and increasedvulnerability to fire damage.

The use of prescribed burning as a silvicultural toolon boreal mixedwood cutovers could be beneficial tomoose provided that most residual elements were notdestroyed by the fire. The residual deciduous componentwith attendant understory is an important factorinfluencing the early use of boreal mixedwood cutoversby moose. Prescribed burning should take place whenresidual patches are relatively incombustible. In this way,the residuals could also help to improve fire control byensuring discontinuity of combustible slash.

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ConclusionsAll forest harvesting and management activities in theboreal mixedwood forest type have an effect on borealungulates. The extent to which these activities arebeneficial or detrimental to species such as moose orwhite-tailed deer is directly related to how radically andextensively the natural age class and species diversity ofmixedwood stands are altered. Efforts by forest managersto short circuit natural successional sequences over largeareas in boreal mixedwood stands to create monotypicconiferous stands will prove detrimental to ungulatespecies dependent on successional forest types. On theother hand, a forest management approach whichrecognizes the value to the boreal ecology of a vigorous,prominent mixedwood component in the boreal forestwill prove beneficial to a wide variety of boreal wildlife.

As the demand for wood fibre and recreationopportunities on the same land base increases,competition between resource managers because ofdifferent objectives will increase. It is extremelyimportant that wildlife and timber managers be aware ofeach other's objectives for the boreal mixedwood timbertype so that conflicts can be avoided.

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