16053
Federal Register
/Vol. 65, No. 58/Friday, March 24, 2000/Rules and Regulations
denning sites with security and thermalcover for kittens (McCord and Cardoza1982; Koehler 1990; Koehler and Brittell1990; Squires and Laurion 1999; J.Organ, U.S. Fish and Wildlife Service,
in litt.
1999). For lynx den sites, the ageof the forest stand does not seem asimportant as the amount of downed,woody debris available (Mowat
et al.
1999). In Washington, lynx used
Pinuscontorta
(lodgepole pine),
Picea
spp.(spruce), and
Abies lasiocarpa
(subalpine fir) forests older than 200years with an abundance of downedwoody debris for denning (Koehler1990). A den site in Wyoming waslocated in a mature subalpine fir/lodgepole pine forest with abundantdowned logs and a high amount of horizontal cover (Squires and Laurion1999). A lynx den site found in Mainein 1999 was located in a forest stand in
Picea rubra
(red spruce) cover type thatwas logged in 1930 and again in the1980s (J. Organ,
in litt.
1999). The siteis regenerating into hardwoods and hasa dense understory (J. Organ,
in litt.
1999). The dominant feature of theMaine site was the abundance of deadand downed wood (J. Organ,
in litt.
1999).The size of lynx home ranges varies by the animal’s gender, abundance of prey, season, and the density of lynxpopulations (Hatler 1988; Koehler 1990;Poole 1994; Slough and Mowat 1996;Aubry
et al.
1999; Mowat
et al.
1999).Documented home ranges vary from 8 to800 square kilometers (3 to 300 squaremiles) (Saunders 1963; Brand
et al.
1976; Mech 1980; Parker
et al.
1983;Koehler and Aubry 1994; Apps 1999;Mowat
et al.
1999; Squires and Laurion1999). Preliminary research supports thehypothesis that lynx home ranges at thesouthern extent of the species’ range aregenerally large compared to those in thenorthern portion of the range in Canada(Koehler and Aubry 1994; Apps 1999;Squires and Laurion 1999).Lynx are highly specialized predatorswhose primary prey is the snowshoehare (
Lepus americanus
), which hasevolved to survive in areas that receivedeep snow (Bittner and Rongstad 1982).Snowshoe hares use forests with denseunderstories that provide forage, coverto escape from predators, and protectionduring extreme weather (Wolfe
et al.
1982; Monthey 1986; Hodges1999a,1999b). Generally, earliersuccessional forest stages have greaterunderstory structure than do matureforests and therefore support higherhare densities (Hodges 1999a,1999b).However, mature forests can alsoprovide snowshoe hare habitat asopenings develop in the canopy of mature forests when trees succumb todisease, fire, wind, ice, or insects, andthe understory grows (Buskirk
et al.
1999b). Lynx concentrate their huntingactivities in areas where hare activity isrelatively high (Koehler
et al.
1979;Parker 1981; Ward and Krebs 1985;Major 1989; Murray
et al.
1994;O’Donoghue
et al.
1997, 1998a).The association between lynx andsnowshoe hare is considered a classicpredator-prey relationship (Saunders1963; van Zyll de Jong 1966; Quinn andParker 1987). In northern Canada andAlaska, lynx populations fluctuate onapproximately 10-year cycles thatfollow the cycles of hare populations(Elton and Nicholson 1942; Hodges1999a, 1999b; McKelvey
et al.
1999b).Generally, researchers believe that whenhare populations are at their cyclic high,depletion of food resources exacerbated by predation cause hare populations todecline drastically (Buehler and Keith1982; Krebs
et al.
1995; O’Donoghue
et al.
1997). Snowshoe hare provide thequality prey necessary to support high-density lynx populations (Brand andKeith 1979). Lynx also preyopportunistically on other smallmammals and birds, particularly whenhare populations decline (Nellis
et al.
1972; Brand
et al.
1976; McCord andCardoza 1982; O’Donoghue 1997,1998a). Red squirrels (
Tamiasciurushudsonicus
) are an important alternateprey (O’Donoghue 1997;1998a; Apps1999; Aubry
et al.
1999). In the Yukon,lynx shifted to red squirrels when harenumbers began to decline (O’Donoghue1998a, 1998b). However, a shift toalternate food sources may notcompensate for the decrease in haresconsumed (Koehler and Aubry 1994). Innorthern regions, when hare densitiesdecline, the lower quality diet causessudden decreases in the productivity of adult female lynx and decreasedsurvival of kittens, which causes thenumbers of breeding lynx to level off ordecrease (Nellis
et al.
1972; Brand
et al.
1976; Brand and Keith 1979; Poole1994; Slough and Mowat 1996;O’Donoghue
et al.
1997).Relative densities of snowshoe haresat southern latitudes are generally lowerthan those in the north, which has ledto differing interpretations of thepopulation dynamics of snowshoe harepopulations. At southern latitudes harepopulations may be—(1) noncyclic, (2)cyclic like northern populations, (3)cyclic with the high and low populationnumbers closer to the averagepopulation numbers, or (4) cyclic witha fluctuating periodicity (length of time between peaks and lows) (Dolbeer andClark 1975; Wolff 1980; Buehler andKeith 1982; Brittell
et al.
1989; Koehler1990; Koehler and Aubry 1994; Hodges1999b). Hodges (1999b) proposes thatnorthern and southern hare populationshave similar cyclic dynamics but that insouthern areas both peak and lowdensities are lower than in the north.Snowshoe hares are generally associatedwith conifer forest cover types (Hodges1999b). Relatively low snowshoe haredensities at southern latitudes are likelya result of the naturally patchy,transitional boreal habitat at southernlatitudes that prevents hare populationsfrom achieving densities similar to thoseof the expansive northern boreal forest(Wolff 1980; Buehler and Keith 1982;Koehler 1990; Koehler and Aubry 1994).Additionally, the presence of morepredators and competitors of hares atsouthern latitudes may inhibit thepotential for high-density harepopulations with extreme cyclicfluctuations (Wolff 1980). If snowshoehare populations in southern borealforests do fluctuate (Hodges 1999b),then southern lynx populations alsomay be expected to fluctuate.Therefore, lynx densities at thesouthern part of the range never achievethe high densities that occur in thenorthern boreal forest (Aubry
et al.
1999). Comparisons between Canadianand contiguous U.S. lynx harvestreturns and snowshoe hare densitiesover time suggest lynx numbers andsnowshoe hare densities for thecontiguous United States aresubstantially lower than those forCanadian provinces (Hodges 1999a,1999b; McKelvey
et al.
1999b). Weconclude that historic and current lynxdensities in the contiguous UnitedStates also are naturally low relative tolynx densities in the northern borealforest.Researchers believe cyclic increasesin historic lynx harvest numbers in thecontiguous United States wereaugmented by dispersal of transientanimals from Canadian populations(Gunderson 1978; Henderson 1978;Mech 1980; McKelvey
et al.
1999b). Theopinion of some individuals andagencies is that presence of lynx insome regions of the contiguous UnitedStates, particularly the Great Lakes, issolely a consequence of dispersal fromCanada (G. Meyer, WisconsinDepartment of Natural Resources,
in litt.
1998; R. Sando, Minnesota Departmentof Natural Resources,
in litt.
1998). Lynxare capable of dispersing extremely longdistances (Mech 1977; Brainerd 1985;Washington Department of Wildlife1993); for example, a male wasdocumented traveling 616 kilometers(370 miles) (Brainerd 1985). Lynxdisperse primarily when snowshoe harepopulations decline (Ward and Krebs1985; Koehler and Aubry 1994;
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