BLACK-TAILED DEER
(Odocoi l e u s hemionus c o l umbianus)
General
o Comnon year - round r e s i d e n t i n t h e deciduous and mixed f o r e s t s o f t h e t r a n - s i t i o n zone w i t h i n t h e e c o r e g i o n (Cowan 1956).
o A l s o i n h a b i t s l ow land r i p a r i a n areas o f w i l l o w ( S a l i x spp.) and b r u s h a l o n g streams and wet lands (Cowan 1945, 1956).
o M i g r a t o r y p o p u l a t i o n s use areas above 2,200 f e e t as sumner range (Ruediger and Garc ia 1980).
Food Requirements
o The annual deer d i e t c o n s i s t s o f 65 p e r c e n t browse, 25 p e r c e n t f o r b s , and 10 grasses (Brown 1961).
o Impor tan t browse spec ies a re t r a i l i n g b l a c k b e r r y , v i n e maple, sa lmonberry , and r e d huck leber ry .
o D u r i n g w i n t e r deer feed h e a v i l y on a r b o r e a l l i c h e n s and o t h e r l i t t e r f a l l (Cowan 1945, Jones 1974, R o c h e l l e 1980) and browse (Brown 1961).
o F o r e s t openings and e a r l y success iona l s tages a r e used most commonly f o r f e e d i n g areas i n t h e Cascades o f western Washington and Oregon (Wallmo 1981).
o F o r e s t s tands a r e used as f o r a g e areas i n heavy s n o w f a l l p e r i o d s (Jones 1974).
o Opt imal f o r a g e h a b i t a t s have >50 p e r c e n t herbaceous cover , and 30 t o 60 pe rcen t dec iduous sh rub cover <1.5 meters t a l l .
o H e a v i l y t r a v e l e d roads reduce deer use o f a d j a c e n t f o r a g e and cover areas (Lyon 1983). Closed roads have m inor impac t on deer use, and may a c t u a l l y p r o v i d e fo rage , bedd ing areas, and t r a v e l l a n e s ( W i l l m s 1975, Witmer 1981).
Water Requirements
o Deer r e q u i r e f r e e wa te r i n t h e summer (Lemos and Hines 1974).
o Water i s g e n e r a l l y n o t l i m i t e d w i t h i n t h e home ranges o f deer i n t h e c e n t r a l Cascades o f wes te rn Washington (Taber 1976).
Cover Requirements
Thermal Cover
o F o r e s t s tands a t l e a s t 40 f e e t h igh , w i t h a t r e e canopy o f a t l e a s t 70 p e r - cen t c l o s u r e p r o v i d e s thermal cover (Thomas e t a l . 1979, Witmer e t a l . 1985).
o Deciduous s tands may p r o v i d e sumner b u t n o t w i n t e r the rma l cover (Thomas e t a1 . 1979).
o Old-growth and mature f o r e s t s tands a r e p r e f e r r e d thermal cover s tands (Zahn 1985, Jones 1974).
H id ing-escape Cover
o Shrubs and young c o n i f e r s p r o v i d e v e g e t a t i o n cover f o r s c r e e n i n g o f deer (Wi tmer e t a1 . 1985).
Opt imal o r Winter S t r e s s Cover
o Optimal cover s tands p r o v i d e f o r a g e i n a d d i t i o n t o thermal cover d u r i n g w i n t e r .
o P rov ided by c o n i f e r f o r e s t s tands w i t h f o u r canopy l a y e r s ( o v e r s t o r y , sub- canopy, shrub l a y e r , and herbaceous l a y e r ) .
o Canopy comprised o f t r e e s w i t h OBH o f 21", and crown c l o s u r e o f 70 p e r c e n t o r more, f o r snow i n t e r c e p t i o n (Jones 1974).
Reproduct ion (Fawning)
o Warm exposures, g e n t l e s lopes , w i t h l o w woody v e g e t a t i o n (Lemos and Hines 1974, B l a c k e t a1 . 1976).
o Succu lent f o r a g e and wa te r w i t h i n 600 f e e t (B lack e t a l . 1976).
I n t e r s p e r s i o n
o Opt imal sumner range h a b i t a t c o n t a i n s 60 p e r c e n t i d e a l f o o d p r o d u c i n g areas, and 40 p e r c e n t i d e a l the rma l o r h id ing -escape cover (Thomas e t a l . 1979).
o Opt imal w i n t e r range h a b i t a t c o n t a i n s 50 p e r c e n t i d e a l f o o d p r o d u c i n g areas, 30 p e r c e n t i d e a l the rma l cover, and 20 p e r c e n t o p t i m a l the rma l cover ( w i n t e r s t r e s s c o v e r ) .
o Grea tes t use o f cover i s f rom 200 t o 800 f e e t f rom f o r e s t - c o v e r edge (wisdom e t a l . 1985).
o Use o f fo rage dec l i nes a f t e r 600 f e e t f rom t h e forage-cover edge, w i t h l i t t l e use o f forage more than 1,200 f e e t f rom cover (Witmer e t a l . 1985, Hanl ey 1983).
o Optimal forage openings a re 1,200 t o 1,500 f e e t wide. Use i s g rea tes t w i t h i n 600 f e e t o f cover edge (Hanley 1983).
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LITERATURE CITED
Black, H., R. Scherz inger , and J. Thomas. 1976. R e i a t i o n s h i p s o f Rocky Mounta in e l k and Rocky Mounta in mule deer h a b i t a t t o t i m b e r management i n t h e B lue Mountains o f Oregon and Washington. pp. 11-13 In. Heib, S. (ed). Proc. E l k - l o g g i n g roads Symp. Univ. o f Idaho, Moscow. 142 pp.
Brown, E.R. 1961. The b l a c k - t a i l e d deer of wes te rn Washington. B i o l . B u l l . 13. Wash. Dept. Game, Olympia. 124 pp.
Cowan, I. McT. 1956. L i f e and t i m e s o f t h e c o a s t b l a c k - t a i l e d deer. pp. 623- 617 In . T a y l o r . W.P. (ed) . The deer of N o r t h America. Stackpo le , H a r r i s b u r g , Pa. 668 pp.
- . 1945. The e c o l o g i c a l r e l a t i o n s h i p s o f t h e f o o d o f t h e Columbian b l a c k - t a i l e d deer (Odoco i leus hemionus columbianus R ichardson) i n t h e c o a s t f o r e s t r e g i o n o f sou the rn Vancouver I s l a n d . B r i t i s h Columbia. Eco l . - Monogr. 15:109-139.
Hanley, T. 1983. B l a c k - t a i l e d deer , e l k , and f o r e s t edge i n a wes te rn Cascades watershed. J. W i l d l . Manage. 47(1):237-242.
Harshman, E., and R. Jubber. 1977. Rooseve l t e l k and b l a c k - t a i l e d deer g u i d e l i n e s f o r t i m b e r , r e c r e a t i o n , and road management. Unpubl. W i l l i a m e t t e N a t i o n a l Fo res t .
Jones, G. 1974. I n f l u e n c e o f f o r e s t development on b l a c k - t a i l e d deer w i n t e r range on Vancouver I s l a n d . pp 139-148 In. B lack , H. (ed) . W i l d l i f e and f o r e s t management i n t h e P a c i f i c Nor thwest . Symp. proceed. Oregon S t a t e Univ., C o r v a l l i s . 236 pp.
Lemos, J., and W. Hines. 1974. E c o l o g i c a l s tudy of Rooseve l t e l k . P r o j . No. W-70-R-4. Progress Repor t . Oregon W i l d l . Corm.
Lyon, L. 1983. Road d e n s i t y models d e s c r i b i n g h a b i t a t e f f e c t i v e n e s s f o r e l k . J. For. 8 1 ( 9 ) :592-595.
Roche l le , J. 1980. Mature f o r e s t s , l i t t e r f a l l and p a t t e r n s o f f o r a g e q u a l i t y as f a c t o r s i n t h e n u t r i t i o n o f b l a c k - t a i l e d deer on n o r t h e r n Vancouver I s l a n d . D o c t o r a l d i s s e r t a t i o n , Univ. o f B r i t i s h Columbia, Vancouver. 295 pp.
Ruediger, W., and E. Garc ia . 1980. C o o r d i n a t i n g d e e r and e l k w i n t e r range and t i m b e r h a r v e s t i n g . USDA F o r e s t Serv i ce , G i f f o r d P i n c h o t N a t i o n a l F o r e s t , Vancouver, MA., 14 pp.
Taber, R. 1976. Seasonal landscape use by e l k i n t h e managed f o r e s t s o f t h e Cedar R i v e r d ra inage , wes te rn Washington. Rept. t o USDA F o r e s t S e r v i c e , C o n t r a c t No. PNW-14. 145 pp.
Thomas, J., H. B lack , R. Scherz inger , and R. Pedersen. 1979. Deer and e l k . pp 104-172 In . Thomas, J. (ed) . W i l d l i f e h a b i t a t s i n managed f o r e s t s - t h e B lue Mountains o f Oregon and Washington. USDA A g r i c . Handbook No. 553. 553 pp.
Wallmo, O.C. 1981. Mule and b l a c k - t a i l e d deer o f N o r t h America. Univ. Nebr. Press, L i n c o l n . 605 pp.
Wi l lms, W.O. 1975. The i n f l u e n c e o f f o r e s t edge, e l e v a t i o n , aspect , s i t e index, and roads on deer use o f logged and matu re f o r e s t , n o r t h e r n Vancouver I s l a n d . MSc. t h e s i s . Univ. o f B r i t i s h Columbia. Vancouver, B.C. 184 pp.
Witmer, G. 1981. Rooseve l t e l k h a b i t a t use on t h e Oregon coas t range. D o c t o r a l d i s s e r t a t i o n , Oregon S t a t e U n i v e r s i t y , C o r v a l l i s . 104 pp.
Wisdom, J., L. B r i g h t , C. Carey, W. Hines, R. Pedersen, 5. Smithey, J. Thomas, and G. Witmer, 1985. D r a f t : A model t o e v a l u a t e e l k h a b i t a t i n western Oregon. US Dept. I n t e r i o r , P o r t l a n d , OR. 45 pp.
Witmer, G., M. Wisdom, E. Harshman, R. Anderson, C. Carey, M. K u t t e l , I. Lurnan, J. Roche l le , R. Scharpf , and 0. Smithey. 1985. Deer and e l k . pp. 131-158 In . Brown, E. (ed ) . Management o f w i l d l i f e and f i s h h a b i t a t s i n f o r e s t s o f western Oregon and Washington. USDA F o r e s t S e r v i c e Pub. No. R6-F and WL-192-1985.
Zahn, H. 1985. Use o f thermal cover by e l k (Cervus e laphus ) on a wes te rn Washington sumner range. D o c t o r a l d i s s e r t a t i o n , Univ. o f Wash., S e a t t l e . 176 pp.
Appendix A
Word model used i n o r i g i n a l HEP s t u d y ( W D G 1982)
BLkCK-TgILED D E E R Coni fercus Forests
The Coi umbian bi ack- t a i led deer ( ~ d o c o i l eus hemicnus colunbianus) i s a comon herbivore wnich occurs in most n a b i t a t types i n t h i s Ecoregion.
Food 2eouirements Ear ly successiofial s t aaes of f o r e s t comnunities genera l ly provide
both increased d i v e r s i t y a n d hiorr2ss of forage spec ie s f o r b l ack - t a i l ed deor (Cowan 1956). ~ o u t h e a s t exposures wi th-adequa t e shrub cover a r e p re fe r r ed feeding areas cozpared t o no r theas t exposures, s i n c e more d e s i r a b l e fzod spec ies z re produczd i n the f o r m r s i t u a t i o n (Cowan 1556; Meehan 1973). T h ? b?ack- t a i l climbs well 2nd nay u t i l . i ze p l a n t s on p rec ip i tous rock ledges (Cowan 1 9 5 6 ) . Deer feed on vegeta t ion up t o 50 inches (127 cm) above :he ground.
Scuthern exposures a t l obe r e i evz t ions a r e p re i e r r ed foraging areas f o r deor i n e a r l y spr ing because o f :he new emergent vegeta t ion (Mil ler 1966). Dee; forage on the leaves of a l d e r (Alnus %), red huckleberry - (Vrccinium parv i f iorum) , vine maple (Acer c i r c i n a t u m ) , t r a i l ing blackberry ( E U C U S u r s i n u s ) , and the new growth o m r b s a s they become ava i l ab le .
Thiz3;ecerry (4ubus i x r v i f l o r u s ) i s the p r i n c i p l e food item during the s u m e r and e a r l y fall (Mi l l e r 1066). I t i s most va luzble t o black- t a i l s when fsgnd i n a vine rxp le c x z u n i t y . Use of b i a l ea f maple (Ac.r macroohyllum) comuni:ies incre25es t t~roagh the s u m e r and f a l l with p e z k use occurr ing i n October.
P r i o r t o l e a f - f a l l deer f i e d sx:ensively on t h e f o l i a g e of broadleaf t r e e s and shrubs. T r a i l i n g b l a c k k r r y , red huckleber ry , s a l a l (GBultheria s n a l l o n ) and crasses m k e U D the bulk of : h ~ deer d i e t during f a i l . w e 2nd e a r l y spring (v ine maple and huckleber ry-sz la l c o m u n i t i e s ) ( M i 11 e r 1965).
Crouch (1966) reported t h a t red huckieberry i s the most valuable winter food of t h e b l ack - t a i l . In the Tillamook burn. Mi l l e r (1966) considered h u g l a s - f i r (Pseudotsuaa nenzies i i), vSne mapie, a l d e r , sword f e rn (Polvstichum s ~ p . ) , a n d Cregon grape ( E e r b e r i s aouifol ium) t o be winter su rv iva l foods. The buds and twigs of hazel m r y l u s c w n u t a ) , vine maple, and a l d e r a r e used when l ea fy forage i s u n a v a i l a b ~ c r o u c h 1968). As highly prefer red f o l i a g e becomes unavai lzb ie 'dur in deer inc rease t h e i r use of Osuglas-f i r s eed l ings (Crouch 1966
Rater Reoui rements Daily and even more f requent t r i p s t o a s t ream o r pond become
important during the heat of summ?r (Cowan 1956). Free water i s l e s s important t o t h e b l ack - t a i l ed deer i n the win te r 2 n d sp r ing .
Cover Reauirements BlacK-railed deer p re fe r dens? arezs of t a l l shrubs and herbaceous
vegeta t ion (Dealy 1959) . Open areas without s u f f i c i e n t cover a r e not used by deer even when food i s a v z i l & l e t h e r e (Hines and Lands 1974). Deer i n Arizona did not u t i l i z e are?$ t n a t were more than 600 f e e t (i83 n ) from t h e f o r e s t ~ d g e ( R e y n ~ l d s 19662,h).
D e ~ r spend much of t h e i r time in shaded drzas, t h i c k e t s , ins ide :he f o r e s t edge during the hot s t imer months (Anderson 2ro tec t ion from s u m e r heat i s provided hen crown c losure i s
and i u s t
60&+ g rea te r (Oregon Dept. Fish and wild1 i f e unpubl. ins.). ~ i l l e r (1966) reported t h a t deer o f t e n make bedding grounds in the b ig l ea f maple cornuni t ies during t h e s i imer.
P ro tec t ion from winter weather i s provided when t h e r e a r e 250 t o 303 t r e e s / a c r e (617 t o 7 4 1 t rees /ha . ) with a mean dbh of 7. t o 8 inches (17.8 to 20.3 cn) in Douglas-fir c o n u n i t i e s (Oreson Dept. F ish and N i l d l i f e unpubl. ins:)- These s i t e s d e t e r deep snow accumulation. 3 l a c k - t a i l s show a s t rong preference f o r a r e z s . she1 t e red from p reva i l ing winds (Wi l l e r 1970). Black-tai led deer may d i e of ma lnu t r i t i on because ibjey s e n e r a l l y do n o t fo rage - fa r from wint2r bedding and f e e d i n g areas even w i x n a v a i l a b l e food beccmes Scarce ( Y i l l e r 1?66). Oeer winter i n zreas of b i g l e a f n a p l e , vine qaple , and red huckleberry a s s o c i a t i o n s .
3 ~ g r o d u c t i v e Fecuirerrients S ? e c i f i c in ionnat ion on the reproduct ive requirements o f black-
t a i l 4 deer was not found i n the l i t ? r a t u r e ; hoirever, reproduct ive cover requir.=e.its a r e 1 i k e l y t o be s i m i l a r t o c t h e r cover requirerrients.
Soecial % n i t a t ? e a ~ < r s n e ? t s No spec3,al h z b i ~ a t r e q u i r e w n t s dere found in the l i t e r a t u r e .
, , ??tllrs 'cersion ? e ~ u i ' r ~ ~ ? n t S Glack- ta i led deer p re fe r d i v e r s i f i e d h ? b i t ? t t h a t con ta ins f o r e s t
edges, dense cover , and openings (Lindz2y 1944; Dealy 1959). Black- t l i i s p r e f e r a reas where these h a b i t a t elements ? r e in c l o s e jux tapos i t ion (Taber 1960). Adequate food supply, in q u a l i t y 2nd q u z n t i t y , i,s of c r i t i c a l importance, e spec ia l ly during periods of lowest a v a i l z b i l i t y and n u t r i t i v e worth (Cowan 1956). The ideal deer range should have f o r e s t s t ands of var ious age c l a s s e s .
Annual h o ~ e ranges of the S lack - t a i l ed deer vary from 0 .1 t o 0.5 mi 2
( 0 . 3 to 1.3 k m ) ( M i l l e r l97O), depending on the a v a i l a b i l i t y o f fooh, water, cave r , and d i s c o n t i n u i t y o f . t e r r a i n in c lose a s s o c i a t i o n (Miller J
1965). Does sha rp ly reduce the s i z e of t h e i r home range during fawning (Mi 1 l e r 19iO).
SoecizT C o m i d e r i t i o n s T ~ P b lack-ca l led deer i s d e ~ e n d e n t ucon e a r l y success ional s tzqes
(~cwan i956) and i t s h a b i t a t i s thereby t r a n s i t o r y in na ture ( ~ o b i n i o n 1958) Fire t r a d i t i o n a l l y r e i n i t i a t e s f o r e s t succession, b u t l o ginq has become a s u b s t i t u t e agent f o r reintroducing ea r ly sera1 s t ages bowan 1956).
Some b l a c k - t a i l e d deer populations migrate along dra inages from high s u n e r rznges t o lcwer winter rznges and use much the same rou te year a f t e r y e a r (Cowan 1956).
REfEgENCES CITED
Anderson, C . F. 1959. Nocturnal a c t i v i t i e s of the Columbian black- t a i l e d d e e r , Odocoileus hemionus columbianus Richardson, a f f e c t i n g
IV-P.-2
spotlight c2nsus results in the Oregon Coast Range. M.S. Thesis. Oregon State Univ. , Corvall is. 86 pp.
Cowan, I. McT. 1956. Life and times of the coast black-tailed deer. pp 523-617 in W. P. Taylor, ed. The deer of North Anerica:their history andyanagement. Stackpole Co., Harrisburg and Wildl. Mgmt. Inst., Wash. 668 pp.
Crouch, G. L. 1966. Preferences of black-tailed deer for native forage and Douglas-fir seed1 ings. J. Wi ldl. Manage. 30(3):471-475.
. 1968. Forage availability in relation to browsing of Cougl as-fi r seed1 ings. J. Wi ldl. Manage. 32(3): 542-553.
Dealy, J, E. 1959. The influences of logging practices on Columbian black-tailed deer in the Blue River area of Oregon. M.S. Thesis, - Oregon State Univ., Corvallis. 65 pp.
Hines, W. W. , and C. E. Lands. 1974. Black-tailed deer and Doug1 as-fir regeneration in the coast range of Oregon. pp 121-132 in H. C. Black, ed. Wildlife and forest management in the P a c i f z Northwest. Oregon State Univ., Corvallis.
Lindzey, J. 8. 1944. A study of the Columbian black-tailed deer (Odocoileus hemionus columbianus (Richardson)) and its habits in Oregon. M. S. Thesis. Oreqon State Univ.. Corvallis. 62 pp.
%eh;n, j. M. 1973. pellet counts as a cens& and rang?-use technique for Colwbiar! black-tailed deer. M.S. Thesis, Oregon State Univ., Ccrua>lis. 2 $ p p .
Miller, F. L. 1966. Distribution patterns of black-tailed deer (Odocoileus henionus co:uixbia~us). M.S. Thesis, Oregon State Univ., Corvallis. 90 on. . . r r -
, 1970. Distribution patterns of black-tailed deer (Odocoileus hemionus cslumSianus) in relation to envircnment. J. Mammal. -8-260.
Oreggn Oept. of Fish and Wildlife (n.d.). Guidelines for providing cover for wildlife on forest lands of western Oregon and Washington. Interagency Wildl. Manage. Coord. Guidelines Committee, unpubl. ms.
Reynolds, H. G. 1966a. Use of a ponderosa pine forest in Arizona by deer, elk, and cattle. U.S. Forest Serv. Res. Note RM63. 7 pp. 3
1966b. Use of openings in spruce-fir forests of Arizona by * elk, deer, and cattle. U.S. Forest Serv. Res. Note RM66. 4 pp. d
Robinson, D. J. 1958. Forestry and wildlife relationships on Vancouver Island. For. Chron. 34:31-36.
Taber, A . D. 1960. The black-tailed deer: A review of ecology and management. Int. Union of Conserv. Nat. Resour. Tech. Meet. 8(4):21 pp.
ADDITIONAL REFERENCES
Brown, E. R. 1961. Black-tailed deer of western Washington. Wash. Dept. of Game Biol. Bull. 13. 124 pp.
Leopold, A . S. 1950. Deer in relation to plant succession. Trans. N Am. Wi ldl. Conf. 15: 571-579.
Mitchell, G. E. 1950. Wildlife-forest relationships in the Pacific Northwest region. J. For. 48:26-30.
Rieck, C. A . 1952. Black-tailed deer investigations on a closed area in western Oregon. M.S. Thesis, Oregon State Univ., Corvallis. 74 OD.
Ruf fed Grouse
(Bonasa umbel l u s )
General
r The R u f f e d Grouse (Bonasa umbe l lus ) i s a year - round r e s i d e n t i n dec iduous and mixed d e c i d u o u s / f e r o u s f o r e s t s o f western Washington (Brewer 1980).
Food Requirements
r D i e t c o n s i s t s o f a w ide v a r i e t y o f p l a n t and animal foods. S p r i n g d i e t s i n wes te rn Washington c o n s i s t p r i m a r i l y o f l eaves and f l o w e r s , w h i l e p r i - mary s u m e r f o o d c o n s i s t s o f f r u i t s . (Brewer 1980).
r Buds and t w i g s o f b l a c k cot tonwood, b i r c h , c h e r r y , and b u t t e r c u p leaves a r e i m p o r t a n t w i n t e r food i t e m s i n wes te rn Washington. (Brewer 1980).
0 Brooding c h i c k s depend upon i n s e c t s and o t h e r i n v e r t e b r a t e s f o r p r o t e i n d u r i n g t h e i r f i r s t 15 t o 20 weeks o f l i f e (Bump -- e t a l . 1947).
Cover Requirements
0 I n h a b i t e a r l y t o middle-aged ( 4 0 t o 70 y e a r s ) mixed f o r e s t s , d e c i d u o ~ s and r i p a r i a n communit ies (Edmins te r 1947) below 2000 f t . i n wes te rn Washington, e s p e c i a l l y t h i c k e t s o f b l a c k cot tonwood, v i n e maple, and b i g l e a f maple (Brewer 1980).
0 C o n i f e r s a r e used f o r w i n t e r and escape cover (Edmins te r 1947).
0 Found i n l ow lands and r i v e r bottoms, i n dec iduous t h i c k e t s , ecotones be- tween c l e a r c u t s and f o r e s t s , and i n b rush o f burned o r logged areas (Edmins te r 1947).
Reproduc t i ve Requirements
Orumning:
0 T y p i c a l l y drum f r o m a r e l a t i v e l y l e v e l f a l l e n l o g , rock , o r stump wh ich i s o f s u f f i c i e n t h e i g h t t o a l l o w a v iew o f t h e s u r r o u n d i n g s ( G u l l i o n 1967).
0 Drumming s i t e s t y p i c a l l y a r e i n areas hav ing a h i g h d e n s i t y o f woody stems, e s p e c i a l l y i n t h e shrub and s a p l i n g l a y e r s (Brewer 1980).
0 Optimal drumming h a b i t a t i s p redominan t l y dec iduous f o r e s t w i t h s c a t t e r e d c o n i f e r s , v i n e maple, ma tu re b i g l e a f maple and mature b l a c k cot tonwood; ground cover between 16 i n . and 47 i n . t a l l , p r o v i d i n g 20-50 p e r c e n t v e r - t i c a l o b s c u r i t y , a t l e a s t one o t h e r l o g w i t h i n 32 ft. o f t h e drumming
stage, and p r o x i m i t y t o an edge between matu re f o r e s t and e a r l y suc- c e s s i o n a l s tages ( G u l l i o n 1967, Brewer 1980, Johnsgard 1983).
r H o r i z o n t a l v e g e t a t i o n s t r u c t u r e ( l e a v e s and branches) d i scourages t h e presence o f r u f f e d grouse (Brewer 1980).
r Brood h a b i t a t i s b e s t p r o v i d e d i n l ow land and t r a n s i t i o n a l areas w i t h a dense shrub s t ra tum. Deciduous components o f c l e a r c u t s , edges, and mixed f o r e s t a r e i m p o r t a n t h a b i t a t s f o r t h e i n v e r t e b r a t e foods broods r e q u i r e and p r o v i d e cover f o r c h i c k s (Edmins te r 1947, Brewer 1980).
r Nest s i t e s a r e t y p i c a l l y a t t h e bases o f t r e e s i n open hardwood s tands, bases o f stumps, bushes o r b rush p i l e s (Bump 1947).
r C l e a r i n g s l e s s than f i v e acres i n f o r e s t s o f mixed dec iduous and c o n i - f e r o u s t r e e s may enhance brood h a b i t a t (Edmins te r 1947, Sharp 1963).
I n t e r s p e r s i o n Requirements
r Requi re a h i g h degree of i n t e r s p e r s i o n o f c l e a r c u t , s a p l i n g , p o l e s tage and mature age c lasses i n r i p a r i a n , dec iduous, and mixed c o n i f e r / d e c i d u o u s cover t ypes t o meet seasonal food and cover needs ( E d n i n s t e r 1947, Brewer 1980, Sharp 1963).
Spec ia l C o n s i d e r a t i o n s
r Wioter food and brood h a b i t a t appear t o be c r i t i c a l l i m i t i n g f a c t o r s t o r u f f e d grouse p o p u l a t i o n s i n western Washington (Brewer 1980). I f good brood h a b i t a t i s p r e s e n t , a d u l t s can e a s i l y s u r v i v e i n t h e same con- d i t i o n s (Berner and Gysel 1969).
r Pure c o n i f e r o u s f o r e s t d i scourages r u f f e d grouse use (Brewer 1980) .
r Alder i s n o t a p r e f e r r e d food source i n wes te rn Washington, b u t may be u t i l i z e d by broods and as a food source i n t h e absence o f o t h e r hardwoods (Brewer 1980).
r Logging s l a s h t h a t i f n o t p r o p e r l y t r e a t e d may i n h i b i t movement, espe- c i a l l y o f c h i c k s (Sharp 1963).
REFERENCES CITED
Berner, H. and L. W. Gysel. 1969. H a b i t a t a n a l y s i s and management con- s i d e r a t i o n s f o r r u f f e d grouse f o r a m u l t i p l e use area i n Mich igan. J. W i l d l . Manage. 33(4):769-778.
Brewer, L. 1980. The r u f f e d grouse i n wes te rn Washington. Washington S t a t e Dept. o f Game B i o l o g i c a l B u l l e t i n No. 16.
Bump, G., R. W. Darrow, F. 0. Edminster , and W. F. C r i ssey . 1947. The R u f f e d Grouse. Hol li ng Press, Inc .
Edminster , F. C. 1947. The r u f f e d grouse. The Macmi l l an Company, New York.
G u l l i o n , G. W. 1967. S e l e c t i o n and use o f drumming s i t e s by male r u f f e d grouse. Auk 84(1):87-112.
Johnsgard, P. A. 1983. The grouse o f t h e wor ld . U n i v e r s i t y o f Nebraska Press.
Sharp, W. M. 1963. The e f f e c t s o f h a b i t a t m a n i p u l a t i o n and f o r e s t success ion on r u f f e d grouse. J. W i l d l . Manage. 27(4):664-671.
Washinston Department o f Game. 1982. F i s h and W i l d l i f e Resource S t u d i e s . s u l t a n ~ i v e r P r o j e c t Stage I 1 F i n a l Repor t . Snohomish County PUD No. 1: 217 pp.
APPENDIX A
Word models used i n o r i g i n a l HEP Study ( W D G 1982)
E t i i F E D G F O C j E R i p a r i a n Con?munities
General The r u f f e d g r o u s e ( eonasa umbe l lus ) i n h a b i t s e a r l y s u c c e s s i o n a l
deciduous c o m c n i t i e s . lwo s u b s p e c i e s of t h e r u f f e d g rouse o c c u r i n t h e W i l l a n e t t e V a l l e y - P u g e t Trough Ecoregion: t h e Olympia r u f f e d g r o u s e (B . u. c a s t a n e a ) , which o c c u r s i n t h e Olympic p e n i n s u l a , and t h e Oregon or- - p a c i f i c r u f f e d g r o u s e ( 0 . u . s a b i n i ) , which i s found t h r o u g h o u t c e n t r a l Washington and Oregon ( J a c h a n and S c o t t 1975).
Food Resu i reoen t s Ruffed g r o u s e f e e d on a wide v a r i e t y of p l a n t and animal f o o d s ,
a l t h o u g h ani ,zal f o o d s r a r e l y exceed 5-10: of t h e a d u l t d i e t (Eorn ins te r 1947; Jahnsgard 1 9 7 3 ) . Ec ,z ins te r (1947) r e p o r t e d t h a t between 400 and 500 p l a n t s p e c i e s p r o v i d e food f o r t h e r u f f e d grouse . Bump e t a l . (1947) i d e n t i f i e d 580 k inds of animal f o o d s , predominantly i n s e c t s , e a t e n by r u f f e d g r o u s e i n Kew York.
Cocpos i t ion o f t h e d i e t depends on p l a n t s p e c i e s a v a i l a b i l i t y and t h e seasonal d i s t r i b u t i o n of t h e g r o u s e (Korschgen 1966). D u r i n g w i n t e r , r u f f e d g r o u s e f e e d p r i m a r i l y on buds and twigs of hardwood s h r u b s and t r e e s ( J o h n s g a r d 1973) . Where a s p e n s ( e s p e c i a l l y P o p u l u s t r e ' u lo ides ) a r e a component o f t h e f o r e s t canopy, t h e s t a m i n a t e f l o w e r buas p rov ide t h e c r i t i c a l w i n t e r food r e s o u r c e ( P h i l l i p s 1967 ; G u l l i o n and Svoboda 1972) . I n w e s t e r n Washington, b l a c k cottonwood ( P . t r i c h o c a r p a ) i s t h e only t r e e s p e c i e s t h a t ranks- i n t h e t o p f i v e food i t e m r ( B r e w e r 1978) . Buds and c a t k i n s , l e a f y v e g e t a t i o n , and a v a i l a b l e i n s e c t s form t h e bulk of t h ? s p r i n g d i e t ( Johnsgard 1 9 7 3 ) . S u m e r foods a r e domina ted oy nus?r3us f r u i t s and b e r r i e s and g r e e n v e g e t a t i ~ n , al t ,?ough a c o n s i d e r a b l e anount of i n s e c r s a r e a J s o ~ a t e n ( E d m i n s t e r 1947) . f a l l food i t e m s i n c l u d e numerous n u t s and o t h e r f r u i t s from t r e e s . sh rubs , and h e r b a c e o u s v e g e t a t i o n ( J c h ~ s g a r d l 9 7 3 ) , l e a v e s , s e e d s , buds , and twigs. A d i v e r s i t y o f h a S i t a t s i s n e c e s s a r y t o meet t h e s e a s o n a l food needs of r u f f e d g rouse . I n M i s s o u r i , Korschgen (1966) found t h a t 15% (by volume) of a l l p l a n t foods were f r m h igh canopy t r e e s , 45';; from unders to ry t r e e s , s h r u b s , v i n e s , and b r a m b l e s , and 40% from herbaceous v e g e t a t i o n .
Water Reouirements Kost a r o u s e f o o d s c o n r a i n c o n s i d e r a b l e w a t e r ( Johnsca rd 1 9 7 3 ) and
2
i t i s u n i i k e l y t h a t r u f f e d g r o u s e need a perzanen: cirini.ing w a t e r supp1y. h'nen Grouse a r e found n e a r w a t e r , i t i s r e l a t e d t o t h e i r p r e f e r e n c e f o r :he f o a j o r c o v e r i n t h o s e wet h a b i t a t s r c t h e r t h m a dependence cn f r e e w a t e r ( E a r , i n s t e r 1947) .
Cover Reouirencnts Ruffed g rouse a r e n o t m i g r a t o r y , b u t do i n h a b i t d i f f e r e n t t y p e s o f
h a b i t a t a t d i f f e r e n t t i m e s o f t h e y e a r . They a r e a s s o c i a t e d w i t h deciduous t r e e s , e s p e c i a l l y p o l e - s i z e d o r second-growth hardwoods (Berne r and Gysel 1 9 6 9 ; Muehrcke and K i r k p a t r i c k 1969), t h r o u g h o u t t h e y e a r but c o n i f e r s a r e used f o r w i n t e r and e s c a p e cover ( E d m i n s t e r 1 9 4 7 ; Hasson and Race 1962 c i t e d i n J a c h a n and S c o t t 1975) . Ruf fed g r o u s e i n
t h i s Ecoreg;on a r e t y i i c a l l y i c u n 6 i n lowianos 2nd r i v e r :cti;:i, i r , e c o r o n e s between f o r e s t s anc c i e 2 r i n g s . i n a l o e r t h i c k e t s , and i n b rush t a n g l e s i n burned o r logaed a r e a s (Jackman and S c o t t 1575) . i n wes te rn Wash ing ton , r e d a l d e r (Ainus -- r u b r a ) i s t h e dominant c o v e r s p e c i e s i n g r o u s e h a b i t a t (Brewer 1978).
Ruffed g r o u s e r e q u i r e a h i ~ h d e g r e e o f i n t e r s p e r s i o n o f cover t y p e s o r f o r e s t age c l a s s e s . In New York, Edmins te r (1947) d e s c r i b e d i d e a l g r o u s e c o v e r a s 80-ESP, woods (hardwood t o c o n i f e r r a t i o o f 50:50), 2-4% open l a n d , and t h e remainder i n b r u s h l a n d . I d e a l g rouse h a b i t a t i n Minneso ta i s p r o v i d e d by a m i x t u r e o f f o u r a g e c l a s s e s of a s p e n (P. t r e m u l o i d e s and P . c r a n d i d e n t a t a ) s t a n d s w i t h a common c o r n e r , w i ih e a c h aged b l o c k l e s s G a n 1 0 a c r e s ( 4 h a ) i n s i z e and d i f f e r i n g from a d j a c e n t b l o c k s by. 1 0 t o 1 5 y e a r s ( G u l l i o n 1977) .
S p r i n g h a b i t a t s a r e a s s o c i a t e d w i t h b r e e d i n g and a r e d i s c u s s e d under R ~ z r o d u c t i ve Pequi rements.
Summer g r o u s e h a b i t a t i n Utah i s c h a r a c t e r i z e d by h i g h e r shrub d e n s i t i e s and e f f e c t i v e cover h e i g h t ( 1 7 . 3 i n . , 4 4 . 1 cm) t h a n a r e found on s u r r o u n d i n g a r e a s (Rober tson 1 9 7 6 ) . H a b i t a t used i n t h e a u t u m i s s i m i l a r t o drumming cover ( d i s c u s s e d below) and has l e s s ground cover t h a n h a b i t a t u s e d by a d u l t s i n t h e summer (Berner and Gysel 1969). Opt imal w i n t e r h a b i t a t i s aspen s t a n d s w i t h 5000 t o 8000 s tems p e r a c r e (12 ,500-20 ,00O/ha) with a cancpy c l o s e d abou t 33 f e e t (10 m ) overhead ( G u l l i c n 1377) .
R ~ a r o o u r t i v c R ~ c u i r e c s n t s -. Drumm>ng. H u i i r d S rocse t y p i c a l l y drum from a f a l l e n l o g , a l though
o t h e r o b i e c t s a r e a l s o used. C h a r a c i e r i s t i c s of an a c c e o t a b l e drumminq - - . . ~ ~ ~- - s i t e a r e s u i f i c i e n t he igh t t c a l l o w a view c f t h e s u r r o u n d i n ~ s and a r e 1 a t i v e ; y i e v e l s t o s e (5oag 3 n d S u z a n i k 1 9 6 9 ) . tirunrning s i t e s a r e t y p i c a l l y l o c a t e d i n are;s having a h < a h d e n s i t y o f woody s t e c s ( F , h e r 1963; Boag and Sunanik 1 9 6 9 ; G u l l i o n 1970; Rusch and K e i t h 1971; and R-her t son ? 9 7 6 ) , e s p e c i a l l y i n t h e s h r u b and s z p l i n g l a y e r s . Tree d e n s i t y i s l e s s im2or;ant th?n t n e d e n s i t y o f t h e s f i a l i e r s t e n s (Bcag and S u m n i k 19E9)).
I n A l b e r t a , 6oac and Sunanik (1969) fognd t h a t young w h i t e s p r u c e ( P i c r a o l a u c a ) l e s s ihan 3 .94 i n c h e s (1 dm) doh p rov ided t h e necessa ry -- drumming cover . I n E la ine , Schemni t z (1976) found a g r e a t e r canopy c l o s u r e a t drumming s i t e s (76.7%) t h a n f u r t h e r away from ( 6 4 . 6 3 t h e s i t e s ; i n c o n t r a s t , Gul i ion e r a l . (1962) r e p o r t e d a p r e f e r e n c e i n Ef innescta f o r s t a n i s wi th l e s s t h a n 60% crown c l o s u r e . Optimal drumming t ; a b i t z t i n Minnesota i s i n aspen s t a n d s w i t h 5000 t o 80OG sterrs p e r a c r e (12,SGG-?O,OOO/ha) (Gul l ion 1570) . S t a n d s w i t h stem d e n s i t i e s iower :han 2C00 s tems p e r a c r e ($940/ha) do n o t s u p p o r t d r u n a i n g grouse .
D r u m i n g s i t e s i n Utah were l o c a t e d w i t h i n 4 1 f e e t ( 12 .5 m ) of t h e n e a r e s t open ing (Rober tson 1976). In Elaine, ave rage d i s t a n c e from t h e drumming s i t e s t o a v e g e t a t i v e e d g e was 105 f e e t (32 m) ( r a n g e 0 - 5 0 0 1 , 3-152 m) (Schemni tz 1976). The b e s t g r o u s e drumming h a b i t a t i n w e s t e r n Washington i s found i n a p r e d o m i n a n t l y deciduous f o r e s t w i t h s c a t t e r e d c o n i f e r s and m a t u r e (25+ y e a r s o l d ) b l a c k cottonwood (Brewer 1978). Such s i t e s a r e c h a r a c t e r i z e d by g round c o v e r ( l e s s t h a n 15.7 i n . , 40 cm, t a l l ) , which p r o v i d e s 30-50: h o r i z o n t a l o b s c u r i t y , a t l e a s t one o t h e r
l o o w i t h i n 3 3 f t e i (10 m) o: t h e o r u m i n ~ s:age, a n d p r o x i m i t y t o ar; eoge between mature fc:est an0 e a r l y s u c c e s s i o n a l s t z g e s d a a i n a t e d by shrubs . G u l l i o n and M a r s h a l l (19%) c o n c ~ u o e d t h a t arumming g r o u s e s u r v i v e l o n g e s t i n a un i fo rm f o r e s t t y p e .
S o c i a l i n t e r a c t i o n i s a l s o i ~ p o r t a n t i n t h e s e l e c t i o n of drumming s i t e s by r u f f e d g rouse (Boag 1976).
Nest ing. Most r u f f e d g rouse n e s t s a r e l o c a t e d a t t h e b a s e s of t r e e s i n open hardwood s t a n d s , a l t h o u g h o t h e r s i t e s such a s t h e b a s e s of stumps, under s l a s h , b u s h e s , o r b rush p i l e s a r e commonly used (Edmins te r 1947). Aspen s t a n d s i n Minnesota w i t h stem d e n s i t i e s l e s s t h a n 2000 stems p e r a c r e ( 4 9 4 0 / h e c t a r e ) a r e p r e f e r r e d n e s t i n g c o v e r ; n e s t i n g females make e x t e n s i v e use of emerging aspen l e a v e s ( G u l l i o n 1977) . Nesting h a b i t a t i n New York i s chosen t o p r o v i d e v i s i b i l i t y , p r o t e c t i o n , an escape r o u t e , p r o x i m i t y t o e d g e s , and t o s a t i s f y an a p p a r e n t d e s i r e f o r s u n l i g h t ( i d m i n s t e r 1947). Nearby undergrowth i s u s u a l l y s p a r s e and canopy c l o s u r e i s low (Edmins te r 1947; G u l l i o n 1977).
Broods. I n s e c t s and o t h e r i n v e r t e b r a t e s make up 50-75% o f t h e d i e t of r u m r o u s e c h i c k s d u r i n g t h e i r f i r s t few weeks o f l i f e (Edmins te r 1947; Johnsgard 1973) , d e c r e a s i n g i n importance w i t h age . By l a t e J u l y o r e a r l y August , t h e d i e t of t h e c h i c k s i s s i m i l a r t o t h a t o f t h e a d u l t s , w i t h p l a n t foods compr i s ing t h e bu lk of t h e d i e t .
Brood cover i s t h e most i m p o r t a n t component of r u f f e d g r o u s e h a b i t a t ( E e r n e r and Gysel 1969) and i s t y p i c a l l y p rov ided by lowland s i t u a t i o n s w i t h a dense s h r u b s t r a t u m . Prime brood h a b i t a t i n Minnesota i s found i n r e g e n e r a t i n g aspen s t a n d s w i t h 5000-12,ODDc stems p e r a c r e (12,350-29,63O/ha) (Gul l i o n 1970) . - Eegenera t ing b,ardwood s t a n d s p r o v i d e brood h z b i t a t f o r 7-10 y e a r s f o l l o w i n g c u t t i n g , a f t e r which t h e y become t o o open f o r brood use ( S h a r p 1963; G u l l i o n 1977). Lowland s p e c i e s such a s a l d e r s (Alnus s p p . ) [ P a i x e r 1963; Godfrey 1975) and wi l lows ( S a l i x s p p . ) (Roberrson ?976) a r e u s u a l l y t h e most important woody s p e c c n brood h a b i t a t . A s h i f t i n brood h a b i t a t use was noted i n Michigan from lowlands ( e a r l y s x n e r ) t o uplands and t h e u p l a n d - t r a n s i t i o n zone (rnid- suxrner) (Berner and Gysel 1969) . The t r a n s i t i o n zone between a lowland s o i l and a w e l l - d r a i n e d upland s o i l i s t h e f o c a l p o i n t of brood a c t i v i t y .
Although hens w i t h broods may t r a v e l l o n g d i s t a n c e s t h r o u g h upland a r e a s dur ing t h e f i r s t few weeks a f t e r h a t c h i n g (Godfrey 1975) , t h e y remain i n a r e l a t i v e l y smal l a r e a once on t h e brood range . Chambers and Sharp (1958) r e p o r t e d t h a t t h e c r u i s i n g r a d i u s o f most broods was l e s s t h a n 0.25 m i l e ( 0 . 4 kn). Broods may range o v e r 10-20 a c r e s ( 4 . 0 - 8 . 1 ha) once e s t ~ b l i s h e d on t h e i r summer range ( G u l l i o n unpub. ms.).
Coni fe r s a r e used by broods f o r r o o s t i n g (Godfrey 1975) . Typ ica l b:ood r o a s r i n g s i t e s a r e clumos of t h r e e o r more c o n i f e r s 2-6" ( 5 . 1 - 15 .2 cm) dbh (Hungerford 1951).
~ o e c i a l ' H a b i t a t Reouirements Ruffed a r o u s e r e o u i r e d u s t i n o s i t e s f o r f e a t h e r c a r e and t o keep
e c t o p a r a s i t e ; a t a minimum. o u s t i n g s i t e s a r e s e l e c t e d which p r o v i d e a sandy s u b s t r a t e , exposure t o t h e s u n , p r o x i m i t y t o dense c o v e r , and maximum v i s i b i l i t y t o d e t e c t approach ing p r e d a t o r s (Hein 1970). Dus t ing s i t e s can be p rov ided by a v a r i e t y o f s i t u a t i o n s and a r e p r o b a b l y never l i m i t i n g .
;r:?:soersion Kt.oui r e f i ~ c z t i The i n r e ~ s j e r s i o r , or cover types i s vei,. i m s r t ~ n ; t o r u f f e d g r o u s e
( see Food, Cover , and Reoroouct ive f i equ i recezrs ) . Home ranges of r u f f e d g rouse vary by s e x , a g e , and season . In
Elinnesota, males o c c u p i e d an average home range of 22 a c r e s ( 8 . 9 h a ) fro;, Earch-June , b u t had a reduced range of 1 6 . 5 a c r e s ( 6 . 7 h a ) d u r i n g the drumming s e a s o n ( A r c h i b a l d 1975). I n t h e same s tudy , f e m a l e s occup ied home ranges of 4 0 . 8 a c r e s ( 1 6 . 5 ha) r e l a t e d t o change i n h a b i t a t s from a lowland-upland edge i n e a r l y s p r i n g t o upland s i t e s f o r n e s t i n g . i n the f a l l , j u v e n i l e s a r e more mobile than a d u l t s (Hale and Dorney 1 9 6 3 ) , and j u v e n i l e females a r e more mobile t h a n j u v e n i l e & a l e s ; i n M i n n e s o t a , j u v e n i l e Faales occup ied f a l l ranges averag ing 7.7 a c r e s ( 3 . 1 h a ) compared t o gn average r a n g e of 13 .4 a c r e s ( 5 . 4 ha ) f o r females (Godfrey and Harshal l 1959). Hale and Dorney (1963) found t h a t i n Wisconsin o n e - h a l f t o t w o - t h i r d s of a g r o u s e p o p u l a t i o n remained i n an a r e a 0 . 5 m i l e s ( 0 . 8 Im; 1 2 5 a c r e s , 50 .6 ha) i n d i a z , e t e r throughout t h e year .
Snecia l C c ~ s i d e r a t i o n s - 7ne r u i f e o o r o u s e i s a b i r d of d i s t u r b e d f o r e s t h a b i t a t s ( G u l l i o n <
1577) . Hchever, a l t e r a t i o n of f o r e s t cover over an a r e a g r e a t e r t h a n 10 ac res ( 3 ha) i n s i z e w i l l r e s u l t i n reduced b reed ing d e n s i t i e s ( G u l l i o n 1570). Ex tens ibe a r e a s of a s i n g l e cover t y p e a r e not a s v a l u a b l e t o r u f f e d g r c u s e a s i s t h e i n t e r s p e r s i o n of s e v e r a l h a b i t a t t y p e s .
S p c t - l u h e r i n g ( i d m i n s t e r 1947) , burning (Sharp 1 9 7 0 ) , and t i m b e r carzgenent m y i , ~ p r o v e g rouse h a b i t a t ( G u l l i o n 1977) . Graz ing by l i v e s t o c k m y a d v e r s e l y a f f e c t brood h a b i t a t (Rober tson 1 9 7 6 ) . e x t e n s i v e t i c b e r h a r v e s t i n g may reduce breeding d e n s i t i e s , and l a c k of t i m b e r r tan;S22ent nay a l i o w f o r e s t e d a r e a s t o becocs u n s u i t a b l e f o r r u f f e d g r s z s e (Gul l ior! 1 9 7 7 ) .
k r c ? , i b a l d , ti. L. 1575. Tcm?oral p a t t e r n s of s p r i n g space u s e by r u f f e d grouse . 2 . W i l d l . Ranage. 39(3):472-481.
Eerner , A , , and L . N. C y s e ? . 1969. H a b i t a t e n a l y s i s and management c o n s i d e r a t i o n s f o r r u f f e d grouse f o r a m u l t i p l e use a r e a i n Michigan. J . Wi ld l . Ranage. 33(4):769-778.
6029, I?. A. 1376. I n f l u e n c e of changing grouse d e n s i t y and f o r e s t a t t r i b u ~ f s on t h e occu;ancy of a s e r i e s of p o t e n t i a l t e r r i t o r i e s by male r u f f e d Grouse. Can. J . 2001. 54:1727-1736.
So;;, C . A , , and K . h!. Su,zanik. 1963. C h a r ~ c ~ e r i s t i c s of drumming sites s e i r c t e d bu r u f i e d orovse i n klk.ert;. J . Wi ld l . Kanaoe. ~ ~ ~ < - - 33(3) : 621-628.
b r e ~ e r , L. 1978. P e r s o n a l con;;iunication. h ' i ldl i f e r e s e a r c h b i o l o g i s t . h'ashingt3n Dept. o f Game. Skagi t Wi ld l . Lab. , Nt. Vernon. ( p r e l i m i n a r y r e s u l t s ) .
Bump, G . , R. W, Darrow, F. C. Edminster , and W. F. Cr i s sey . 1947. The r u f f e d g r o u s e : l i f e h i s t o r y . p r o p a g a t i o n , management. New York S t a t e Cons. Dcpt. 915 pp.
Cha-bers, R . E . , and W . M. Sharp. 1958. blovcment and d i s p e r s a l w i t h i n a p o p u l a t i o n of r u f f e d grouse. J. Wildl . !!anage. 22(3):231-239.
i d r . i n s t e r , F . C . i94;. i n e r u f f e o g r o u s e : i t s l i f e s l c r y , ecoiog) ' , and manage-rnt. c i l l r Y 385 F;.
Godfrey, G . A . 1975. home range c h a r a c t e r i s t i c s of r u f f e d g r o u s e broods i n Minneso ta . J. W i l d i . Manage. 39(2):287-296.
Godfrey, G . A , , and W . H . M a r s h a l l . 1969. Brood breakup and d i s p e r s a l of r u f f e d g r o u s e . J . W i l d l . Kanage. 33(3) : 605-620.
Gull ion , G . k'. 1970. F a c t o r s i n f l u e n c i n g r u f f e d g rouse p o p u l a t i o n s . Trans . N . Am. W i l d l . Conf. 35:93-105.
. 1977. F o r e s t m a n i p u l a t i o n f o r r u f f e d grouse . T r a n s . N . AT. Wi d l . Conf. 42: 449-458.
. Ruffed g r o u s e management reccrnnendations. G r a f t o n - B a r r e t t 'di l d l i f e P r o j e c t , Univ. o f Vermont, Burl ing ton . Unpub. ms. 37 pp.
G u l l i o n , G . W . , and W. ti. E:arshall . 1968. S u r v i v a l of r u f f e d g r o u s e i n a b o r ~ a l f i r e s t . The L i v i n g B i r d . 7:117-167.
G u l l i o n , G . W . , and F. J . Svoboda. 1972. T h e b a s i c h a b i t a t r e s o u r c e f o r r u f f e d g r o u s e . P roc . Aspen Syrnp., USOA F o r e s t S e r v i c e t e n . Tech. Rept. N C - 1 : I l ? - l l g .
t u l l i o n , G . W., R. T. King , and W. H . E a r s h a l l . 1962. Kale r u f f e d grouse and 30 y e a r s o f f o r e s t management on t h e Cloquet F o r e s t R e s ~ a r c h C e n t e r , Minneso ta . J. For. 6O(9): 617-622.
Hale , J . B., and R . S . Oorney. 1963. Seasonal movements o f r u f f e d grouse i n h ' i s c o n s i n . J . W i l d l , Ranage. 27 (4 ) : 648-656.
Hein, 0. H . 1970. D u s i - b a t h i n g s i t e s s e l e c t e d by r u f f e d g r o u s e . h'ilzon B u i l . 82 (3 ) :310-314 .
Hungerford, K . E . 1951 . Ruf fpd g rouse p o p u l a t i o n s and c o v e r u s e i n nor the rn I l s h o . T r a n s . N. A m . W i l d l . Conf. 16:216-224.
J a r k x n , S. H., dcd J . M. Scot:. 1975. L i t e r a t u r e r ev ied o f t w e n t y t h r e e s e i e c t e d f o r e s t b i r d s of t h e P a c i f i c t ; o r t h i e s t . USDA F o r e s t S e r v i c e , R r ~ i o n 5 , pp . 153-170.
Johnsg.ard, P . A . 1973. Grouse and q u a i l s of North America. Univ. f iebraska. L inco ln . DD. 253-273.
Korschgen, L . ' J . 1966. i b o d s and n u t r i t i o n of r u f f e d grouse i n EI issour i J . Wi ld i . Pknage. 30(1):35-1DO.
Masson. W . V.. and R . U. !+ace. 1962. I 'pland qame b i r d s . Oregon S t a t e t k ion;. , W i l d l . B u l l . no. 5. 4 4 pp. -
.
Huehrcke, J . P . , and C. M. K i r k p a t r i c k . 1969. Obse rva t ions on ecoloQy and beha\ ; ior o f I n d i a n a r u f f e d g rouse . Proc. Ind. Acad. S c i . 79: 177-165.
Palmer, W . L. 1563. Ruf fed g r o u s e d r u m i n g s i t e s i n n o r t h e r n Mich igan . J . h ' i l d l . P,s:ase. 2 7 ( i ) : E55-6E;.
F h i i l i p s , R. L . 1967. F a l l znd w i n t e r iood h ? b i t s of r u t i e d g r o u s e i n n o r t h e r n U:ah. J . k'i l d l . hianage. 3 l ( 4 ) : 627-823.
Robertson, R. L. 1976 . Ru:fed g rouse h a b i t a t p r e f e r e n c e s and e f f e c t s of l i v e s t o c k g r a z i n g on h a b i t a t u t i l i z a t i o n . H . S . T h e s i s , Utah S t . U n i v . , Logan. 7 1 pp.
Rusch, 0. H . , and L. 0 . K e i t h . 1971. Ruffed g rouse - v e g e t a t i o n r e l a t i o n s h i p s i n c e n t r a l A l b e r t a . J . Wildl . Manage. 35 (3 ) : 417- 429.
Schemnitz, S. 0. 1976. C h a r a c t e r i s t i c s of Maine r u f f e d g r o u s e drumming s i t e s . Res. L i f e S c i . , Univ. Maine, Orono. 23(11):6 pp.
, . C;zrp, k:. E . 1963. T h e efiects o: xs::at manipu!ation and ic:es:
I success5on o n rufieo arouse. 2 . h i l d i . kanage. 27(4 ) :664 -671 . . 1970. The role of f i r e i n r u f f e d g r o u s e hanitar ~ ~ a n a g e m c n t .
Proc. Annu. Tall Timbers Fire icol. Conf. 10:41-61. I;. .
I I . , I I
r a n . W?o sons IXP(Uihl€
AEA & R x m ZIR' LRNO
MI .Y)w Slns L5 a Mhk Kl3AIELr W-iRhDfn son5
CUmAKl SCILS CDBW 9(ML P A M Q . 3 l?LL& 25 ' i u m - M M
I
I ) :
tiLS;TAT SUiTABiLITY INDEX
Ruffed Grouse i n Riparian Cornunit ies
Ecoregion 2410
Winter Food Value (XI) = I 4
Where: I4 = S u i t a b i l i t y index (81) of s tems/acre (winter feeding) .
Cover Value
Because ru f fed grouse require i n t e r s p e r s i o n of severa l f o r e s t age c l a s s e s t o meet t h e i r l i f e r equ i s i t e s , an ove ra l l cover value based on seasonal needs i s not possible.
Reoroductive Values
~ r u i m i n q _ ( X ~ ) = ( I 1 x I,-) 112
h'here: 1 = S I of st:-.s/acre ( d r m ~ e r s ) - 1
i 5 = S I of a v e r z y d i s t ance t o opening.
k s t i n c ( X 4 ) = I j
h'here: I: = S I of s t ens i ac re ( n e s t i n g ) . - Broodins ( X j ) = ( I p x 16) 112
Clnere: I = S I of sterns/acre (broods) . 2
= S I a f s o i i condit ions. : 6
I n t e r s ~ e r s i o n Value ( X 6 ) = i7
Uhere: I7 = S I of average stand s i z e .
The Hab i t a t S u i t a b i l i t y Index i s the lowes t X n value.
Black-capped Chickadee
(Parus a t r i c a p i l l u s )
General
0 Comnon year - round r e s i d e n t i n dec iduous and mixed c o n i f e r / d e c i d u o u s f o r e s t s . ( G a b r i e l s o n and J e w e t t 1940, Brown 1985).
0 A l s o i n h a b i t s l ow land r i p a r i a n areas o f w i l l o w ( S a l i x spp.) and b r u s h a l o n g streams ( L a r r i s o n and Sonnenberg 1968).
-
Food Requi rement s
0 Food supp ly may be most i m p o r t a n t f e a t u r e o f h a b i t a t (WOG 1982)
0 Over 50% o f d i e t i s animal m a t t e r , i n c l u d i n g i n s e c t s and t h e i r eggs, c a t e r - p i l l a r s and moths, p l a n t l i c e , w e e v i l s and s p i d e r s (Pearson 1936) .
0 Vegetab le m a t t e r ea ten i n c l u d e s sma l l seeds, buds, bud s c a l e s (Pearson 1936).
0 C a t e r p i 1 l a r s a r e e s p e c i a l l y i m p o r t a n t f o r young (Sturman l968a) .
0 Forage by g l e a n i n g f o o d f r o m t h e ba rk o f t w i g s , branches, b o l e s , f o l i a g e , f l o w e r s , and f r u i t s o f t r e e s and shrubs (Brewer 1963).
0 D u r i n g b r e e d i n g season, f o r a g e p r e d o m i n a t e l y i n t h e subcanopy ( i n s i d e t h e canopy w i t h l i t t l e o r no 1 i v i n g f o l i a g e ) (Sturman 1968b).
a I n wes te rn Oregon 30 t o 40% o f f a l l and w i n t e r f o r a g i n g o c c u r r e d i n shrubs (Anderson 1970).
0 Most f o r a g i n g i s done w i t h i n 30 f t . o f t h e ground (Brewer 1963).
0 Optimum canopy c l o s u r e s f o r f o r a g i n g occur between 50 and 75% (Schroeder 1982).
0 Optimum h a b i t a t s c o n t a i n o v e r s t o r y t r e e s 49 f t . o r more i n h e i g h t (Schroeder 1982).
Water Requirements
a D r i n k i n g wa te r requ i rements a re met w i t h s u r f a c e wa te r and snow (Odum 1942).
Cover Requirements
r From l a t e f a l l t h r o u g h s p r i n g , b lack-capped ch ickadees excava te n e s t c a v i - t i e s f o r r o o s t i n g (Brewer 1963, Thomas -- e t a1 . 1979).
a P r e f e r dec iduous f o r e s t t y p e s i n wes te rn Washington (Sturman 1968a)
r Excavates c a v i t i e s i n s o f t , decayed wood, occup ies c a v i t i e s made by o t h e r spec ies , and occup ies " n a t u r a l " c a v i t i e s c r e a t e d by decay (Brown 1985).
Reproduc t i ve Requirements
P r imary c a v i t y n e s t e r i n decayed o r s o f t wood (Ddum 1941a; Brewer 1963). A l s o n e s t s i n c a v i t i e s c r e a t e d by o t h e r p r i m a r y excava to rs (Brown 1985).
Optimum h a b i t a t s p r o v i d e 2 snags lac re (Schroeder 1982).
W i l l o w ( S a l i x spp.) and cot tonwood (Populas t r i c h o c a r a ) a r e t h e common n e s t t r e e Y D e c a y e d hardwoods a l s o u s v r d .
Number o f nes t s i t e s i n t h e h a b i t a t does n o t seem t o be a c r i t i c a l f a c t o r i n t e r r i t o r i a l s e l e c t i o n (Sturman 1968a).
Used second-growth a l d e r (A lnus -- r u b r a ) f o r n e s t i n g i n B r i t i s h Columbia ( S m i t h 1967).
Opt imal n e s t t r e e s a r e 9 i n . dbh and 10 f t . o r g r e a t e r i n h e i g h t (Brown 1985).
S p e c i a l C o n s i d e r a t i o n
0 Feed on a v a r i e t y o f i n s e c t s , i n c l u d i n g many regarded as f o r e s t p e s t s (WDG 1982).
REFERENCES CITED
Anderson, S. H. 1970. E c o l o g i c a l r e l a t i o n s h i p s o f b i r d s i n f o r e s t s o f wes te rn Oregon, Ph.D. Thes is , Oregon S t a t e Univ., C o r v a l l i s . 124 pp.
Brewer, R. 1963. E c o l o g i c a l and r e p r o d u c t i v e r e l a t i o n s h i p s o f b lack-capped and C a r o l i n a ch ickadees. Auk 80(1) :9-47.
Brown, E. R. (ed. ) 1985. Management o f w i l d l i f e and f i s h h a b i t a t s i n f o r e s t s o f western Oregon and Washington. P a r t 2. U.S. Dept. A g r i c . , F o r e s t S e r v i c e , P a c i f i c Nor thwest Region. 301 pp.
Gabr ie l son , I. N., and S. G. Jewe t t . 1940. B i r d s o f Oregon. Oregon S t a t e C o l l e g e , C o r v a l l i s . 650 pp.
L a r r i s o n , E. J. and K. G . Sonnenberg. 1968. Washington b i r d s : t h e i r l o c a t i o n and i d e n t i f i c a t i o n . S e a t t l e Audubon Soc., S e a t t l e . 258 pp.
Odum, E. P. 1941a. Annual c y c l e o f t h e black-capped ch ickadee - I. Auk 58:314-333.
- . 1941b. Annual c y c l e o f t h e black-capped ch ickadee - 11. Auk 58(4):518-535.
- . 1942. Annual c y c l e o f t h e black-capped ch ickadee - 111. Auk 59(4):499-531.
Pearson, T. G., ed. 1936. B i r d s o f America. Garden C i t y Publ . Co., I nc . Garden C i t y . pp. 509-512.
Schroeder, R. L. 1982. H a b i t a t s u i t a b i l i t y i n d e x models: Black-capped c h i c k a - dee. U.S. Dept. I n t . , F i s h W i l d l . Serv. FWSfOBS-82110.37. 12 pp.
Smith, S. M. 1967. Seasonal changes i n t h e s u r v i v a l o f t h e black-capped chickadee. Condor 69 (4 ) :344-359.
Sturrnan, W. A. 1968a. D e s c r i p t i o n and a n a l y s i s o f b r e e d i n g h a b i t a t s o f t h e ch ickadees, Parus a r t r i c a p i l l u s and - P. ru fescens. Eco logy 49(3):418-431.
. 1968b. The f o r a g i n g eco logy o f Parus a t r i c a p i l l u s and P. r u f e s c e n s i n t h e b r e e d i n q season, w i t h compar isons t o o t h e r spec ies o f Parus. Condor
Thomas, J. W., R. G Anderson, C. Maser, and E. L. B u l l . 1979. Snags. pages 60-77. I n J. W. Thomas, ed. W i l d l i f e h a b i t a t s i n managed f o r e s t s - t h e B lue M o u n t a i n s o f Oregon and G h i n g t o n . U. S. Dept. Agr i c . , For . Serv. A g r i c . Handb. 553.
Washington Department o f Game. 1982. F i s h and W i l d l i f e Resource S t u d i e s , S u l t a n R i v e r P r o j e c t Stage I 1 F i n a l Repor t . Snohomish County PUD No. 1 217 pp.
APPENDIX A
Aggrega t ion fo rmu la and s u i t a h i l i t y i n d i c e s o b t a i n e d f rom the USFWS H a b i t a t S u i t a b i l i t y Index Models - b lack-capped ch ickadee (Schroeder 1982).
Black-capped ch ickadee
Percent t r e e canopy c l o s u r e ( V 1 ) Food------c verage h e i g h t o f o v e r s t o r y
I/ t r e e s ( v ~ ) A1 1 t e r r e s t r i a l cover t ypes I '
! i 1 Reproduction-Number of snags 4 t o 10 i n . \ dbh/acre (V3)
L i f e r e q u i s i t e Cover t y p e Equa t ion I
Food A1 1 t e r r e s t r i a l ( V , x v , ) ~ ' ~
Reproduc t ion A1 1 t e r r e s t r i a l " 3
H S I d e t e r m i n a t i o n . The H S I f o r t h e b lack-capped ch ickadee i s equa l t o I t h e l o w e s t l i f e r e q u i s i t e v a l u e .
A P P E N D I X B
W o r d model used i n o r i g i n a l H E P S t u d y (WDG 1 9 8 2 )
June 1978
BLACK-CAPPED CHICKADEE Deciduous Woodlands
General The b lack-capped chickadee (b a t r i c a o i l l u s )
around r e s i d e n t i n t h e deciduous and mixed deciduous i s a common year - f o r e s t s o f t h e
T r a n s i t i o n Zone w i t h i n t h i s Ecoreg ion (Gabr ie l son and J e w e t t 1940). I t a l s o i n h a b i t s l ow land r i p a r i a n areas o f w i l l o w ( S a l i x - spp.) and b rush a l o n g streams ( L a r r i s o n and Sonnenberg 1968).
Food Requirements Over h a l f o f t h e food eaten b y t h e black-capped ch ickadee i s animal
m a t t e r , i n c l u d i n g i n s e c t s and t h e i r eggs, c a t e r p i l l a r s and moths, p l a n t l i c e , w e e v i l s and s p i d e r s (Pearson 1936). Vegetab le m a t t e r ea ten i n c l u d e s sma l l seeds, buds, and bud sca les . C a t e r p i l l a r s a r e an e s p e c i a l l y i m p o r t a n t f o o d i t e m f o r young chickadees (Sturman 1968a). The food supp ly may be t h e most i m p o r t a n t f e a t u r e o f t h e ch ickadee 's env i ronment . Black-capped chickadees may use canopy volume o f hardwood t r e e s as a v i s u a l c l u e t o i n s e c t abundance and b r e e d i n g h a o i t a t s e l e c t i o n .
Black-capped chickadees d u r i n g t h e b r e e d i n g season f o r a g e p re - domina t l y i n t h e sub-canopy (a rea i n s i d e t h e canopy w i t h l i t t l e o r no l i v i n g f o l i a g e ) (Sturman 1968b). I n t h e w i n t e r , t h e y spend most of t h e i r t ime f e e d i n g i n t h e t h i n o u t e r t w i g s o f t r e e s (Smi th 1967). Chickadees f o r a g e by g l e a n i n g food f rom t h e b a r k o f t w i g s , branches, b o l e s , f o l i a g e , f l o w e r s , and f r u i t s o f t r e e s and shrubs (Brewer 1963). I n wes te rn Oregon, 30 t o 40 p e r c e n t o f f a l l and w i n t e r f o r a g i n g occured i n shrubs (Anderson 1970). Most o f t h e w i n t e r f e e d i n g i s done i n deciduous t r e e s (Smi th 1967). Most f o r a g i n g i s done w i t h i n 30 f e e t ( 9 . 1 m ) of t he ground (Brewer 1963).
Water Requirements No s p e c i f i c water requ i rements were found i n t h e l i t e r a t u r e
Cover Requirements L i t t l e i n f o r m a t i o n was found i n t h e l i t e r a t u r e r e g a r d i n g cover
reau i rements o t h e r than r e p r o d u c t i v e needs. Frcm l a t e f a l l t h rough s p r i n g , b lack-capped chickadees use n e s t c a v i t i e s f o r r o o s t i n g (Brewer 1963).
Reproduct ive Requirements Black-canned chickadees a re p r i m a r y c a v i t y n e s t e r s i n decayed o r
s o f t wood (0d;k 1941a; Brewer 1963). ~ ; l l o w and cot tonwood t r e e s (Populas t r i c h o c a r p a ) a r e t h e common n e s t t r e e s . Decayed hardwood t r e e s a r e a l s o used f o r nes t ing . Chickadees i n New York f r e q u e n t l y excavated n e s t s i n we l l -decayed s tubs o r l i m b s o f t r e e s (Odum 1968a j . The number o f n e s t s i t e s i n t h e h a b i t a t does n o t seem t o be a c r i t i c a l f a c t o r i n t e r r i t o r i a l s e l e c t i o n (Sturman 1968a). I n B r i t i s h Columbia, b l a c k - capped ch ickadees used second-growth a l d e r (A lnus r u b r a ) f o r n e s t i n g (Smi th 1967). A second growth mixed f o r e s t (5% deciduou; and 45% c o n i f e r o u s ) t h a t con ta ined many o l d a l d e r s and maples (m macrophyl lum) had an excess o f n e s t s i t e s . Diameters o f t r e e s tubs used f o r n e s t i n g
June 1978
in New York were between 3.5 and 6.0 inches (8.9-15.2 cm) with nest heights ranging from 1 to 40 feet (0.3-12.2 m) (Odum 1941b).
Sturman (1968a) found that the abundance of black-capped chickadees was positively correlated with tree canopy volume. In western Oregon, Oregon white oak ( with tree densities of 120 to 150 per acre (296 dense canopy coverage had higher breeding densities than stands with a low number of trees per acre (56; 138/ha) regardless of the density of canopy coverage. The maximum and minimum breeding densities occurred in stands with the highest and lowest tree densities, respectively.
Special Habitat Requirements No special habitat requirements for black-capped chickadees were
found in the literature.
Interspersion Requirements No specific interspersion requirements were found in the literature.
Special Considerations Black-capped chickadees feed on a wide variety of insects, including
many regarded as forest pests.
REFERENCES CITED
Anderson, S. H. 1970. Ecological relationships of birds in forests of western Oregon. Ph.D. Thesis, Oregon State Univ., Corvallis. - . 124 pp.
Brewer, R. 1963. Ecological and reproductive relationships of black- capped and Carolina chickadees. Auk 80(1):9-47.
Gabrielson, I. N . , and 5. G. Jewett. 1940. Birds of Oregon. Oregon State College, Corvall is. 650 pp.
Larrison, E. J. and K. G. Sonnenberg. 1968. Washington birds: their location and identification. Seattle Audubon Soc. , Seattle. 258 pp.
Odum. E. P. 1941a. Annual cycle of the black-capped chickadee - I. Auk 58: 314-333.
. 1941b. Annual cycle of the black-capped chickadee - 11. Auk 58(4):518-535.
Pearson, T. G., ed. 1936. Birds of America. Garden City Publ. Co., Inc. Garden City. pp. 509-512.
smith,'^. M. 1967. Seasonal changes in the survival of the black- capped chickadee. Condor 69(4):344-359.
Sturman, W. A. 1968a. Description and analysis of breeding habitats of the chickadees, Parus atricapillus and P. rufescens. Ecology 49(3): 418-431.
. 1968b. The foraging ecology of atricapillus and P- rufescens in the breeding season, with comparisons to other species of Parus. Condor 70(4):309-322.
P i l e a t e d Woodpecker
(Dryocopus p i l e a t u s )
General
r A year - round, though uncommon, r e s i d e n t o f dense mature and o l d g rowth c o n i f e r o u s f o r e s t s i n wes te rn Washington (WDG 1982).
Food Requirements
r Feeding h a b i t a t i n c l u d e s areas w i t h h i g h d e n s i t y o f l o g s and snags, dense canopies and t a l l shrub cover ( B u l l and Meslow 1977).
Spend 36 p e r c e n t o f f e e d i n g t i m e f o r a g i n g on logs , 35 p e r c e n t on l i v e t r e e s , and 29 p e r c e n t on snags i n Oregon ( B u l l 1981).
r Animal m a t t e r comprises 75 pe rcen t o f d i e t ( T e r r e s 1980).
r Feed by e x c a v a t i o n and by s c a l i n g b a r k i n search o f i n s e c t s .
r Optimum h a b i t a t occu rs w i t h 10 o r more l o g s 7 i n . o r g r e a t e r d i a m e t e r and/or s t !mps o f t h e same d iamete r and g r e a t e r t h a n 1 f t . i n h e i g h t pe r a c r e (Schroeder 1982).
Water Requirements
r Requi re l a r g e amounts o f water i n c a p t i v i t y wh ich may i n d i c a t e t h e i r water requ i rements i n t h e w i l d (P f i t zenmeyer 1956 c i t e d by Jackman and S c o t t 1976).
Cover Requirements
r Cover requ i rements a r e s i m i l a r t o r e p r o d u c t i v e requ i rements .
r P r e f e r r e d o l d s tands w i t h 70% crown c l o s u r e i n B lue Mounta in of Oregon (Thomas -- e t a1 . 1979).
r Optimum h a b i t a t has 75% o r g r e a t e r canopy c l o s u r e ; s tands w i t h l e s s t h a n 25% canopy c l o s u r e a r e u n s u i t a b l e (Schroeder 1982).
r P r e f e r t i m b e r s tands w i t h sawtimber o f 15 t o 18 i n . dbh i f n e s t s i t e s a r e a v a i l a b l e (Conner -- e t a l . 1975).
Reproduc t i ve Requirements
r Trees s e l e c t e d were a t l e a s t 20 i n . dbh (Jackman 1974) and were a t l e a s t 40 f t. t a l l ( B u l l 1975).
0 P r e f e r t a l l , dead t r e e s w i t h few l i m b s ( S c o t t e t a l . 1977). --
r Optimum h a b i t a t c o n t a i n s 30 o r more t r e e s g r e a t e r t h a n 20 i n . dbh/acre ; h a b i t a t w i t h l e s s t h a n 3 such t r e e s i s u n s u i t a b l e (Schroeder 1982).
0 Optimum h a b i t a t c o n t a i n s snags s u i t a b l e f o r n e s t i n g a t a d e n s i t y o f 6 t o 15 snags p e r 100 acres ( B u l l 1977 and N e i t r o e t a l . 1985). --
S p e c i a l H a b i t a t Requirements
0 U t i l i z e ho les f o r r o o s t i n g year - round (Jackman and S c o t t 1975).
r Fresh r o o s t i n g ho les u s u a l l y excavated i n t h e f a l l (Jackman and S c o t t 1975).
0 Roost h o l e s a re u s u a l l y i n l i v e t r e e s and h i g h e r i n t h e t r e e than n e s t ho les (Hoyt 1957).
I n t e r s p e r s i o n Requirements
0 Home range i s 300 t o 600 acres (Brown 1985).
REFERENCES CITED
Beal, F. E. L. 1911. Food o f t h e woodpeckers o f t h e U n i t e d S t a t e s . U.S. B io . Surv. B u l l . No. 37:164.
Brown, E. R. (ed.) . 1985. Management o f w i l d l i f e and f i s h h a b i t a t s i n f o r e s t s o f western Oregon and Washington. P a r t 1 and 2. U.S. Dept. Agr i c . , F o r e s t Serv i ce , P a c i f i c Region.
B u l l , E. L. 1975. H a b i t a t u t i l i z a t i o n o f t h e p i l e a t e d woodpecker, B lue Mountains, Oregon. M.S. Thes is , Oregon S t a t e Univ., C o r v a l l i s . 58 pp.
B u l l , E. L. 1977. S p e c i a l i z e d h a b i t a t requ i rements o f b i r d s : snag management, o l d growth, and r i p a r i a n h a b i t a t . P a c i f i c Nor thwest F o r e s t and Range Exp. Sta., F o r e s t Serv i ce , U.S. Dept. o f Agr ic . , La Grande, DR. 9 pp.
B u l l , E. L., and E. C. Meslow. 1977. H a b i t a t requ i rements o f t h e p i l e a t e d woodpecker i n n o r t h e a s t e r n Oregon. J. For. 7 5 ( 6 ) : 335-337.
B u l l , E. L. 1981. How woodpeckers s e l e c t and p o r t i o n t h e i r h a b i t a t i n n o r t h e a s t e r n Oregon, Ph.D. D iss . , Univ. Idaho, Moscow. 99 pp.
Conner, R. N., R. G. Hooper, H. S. Crawford , and H. S. Mosby. 1975. Woodpecker n e s t i n g h a b i t a t i n c u t and uncu t woodlands i n V i r g i n i a . J . W i l d l . Manage. 39(1):144-150.
Hoyt , 5. F. 1957. The eco logy o f t h e p i l e a t e d woodpecker. Eco logy 38(2):236-256.
Jackman, 5. M. 1974. Woodpeckers o f t h e P a c i f i c Nor thwes t ; t h e i r cha rac - t e r i s t i c s and t h e i r r o l e i n t h e f o r e s t . M.S. Thes is , Oregon S t a t e Univ. , C o r v a l l i s . 147 pp.
Jackman, S. M., and J. M. S c o t t . 1975. L i t e r a t u r e r e v i e w o f t w e n t y - t h r e e s e l e c t e d f o r e s t b i r d s o f t h e P a c i f i c Nor thwest . Region 6. U.S. F o r e s t Serv i ce . 381 pp.
Mannan, R. W. 1977. Use o f snags by b i r d s , Douglas F i r Region, Western Oregon. M.S. Thes is , Oregon S t a t e Univ., C o r v a l l i s . 114 pp.
N e i t r o , W. A., V . W. B i n k l e y , S. P. Chine, R. W. Mannan, B. G. Marcot , D. T a y l o r and F. F. Wagner. 1985. Snags ( W i l d l i f e Trees) . I n E. R. Brown, ed. Management o f Washington. P a c i f i c Nor thwest Region, F o r e x S e r v i c e , U.S. D e F . A g r i c . 332 pp.
Pf i tzenmeyer , H. T. 1956. L i f e h i s t o r y and behav io r p a t t e r n s o f t h e p i l e a t e d woodpecker r e l a t i v e t o u t i l i t y l i n e s . M.S. Thes is , Pennsy lvan ia S t a t e Un iv . U n i v e r s i t y Park.
Schroeder, R. L. 1982. H a b i t a t s u i t a h i l i t y i n d e x models: P i l e a t e d woodpecker. U.S. Dept. I n t . , F i s h W i l d l . Serv. FWSIOBS-82/10.39. 15 pp.
S c o t t , V . E., K. E. Evans, D. R. P a t t o n , and C. P. Stone. 1977. P i l e a t e d wood pecker . Page 35 i n V. E. S c o t t ed. C a v i t y n e s t i n g b i r d s o f N o r t h American f o r e s t s . USDA F o r e s t S e r v i c e A g r i z Hdbk. No. 511.
Te r res , T. K. 1980. The Audubon S o c i e t y Encyc loped ia o f N o r t h American B i r d s . A l f r e d , A. Knopf, Inc. 1109 pp.
Thomas, J. W., R. Anderson, C. Maser, and E. L. B u l l . 1979. Dead t r e e ( " s n a g " ) requ i rements f o r dependent w i l d l i f e spec ies i n t h e B l u e Mounta ins o f Washington and Oregon. I n J. W. Thomas, ed. W i l d l i f e h a b i t a t i n managed f o r e s t s - The B lue M o u n t a i n s o f Oregon and W a 3 i n g t o n . Pac. Nor thwest f o r . and Range Exp. Stn., U.S. Dept. Agr i c . P o r t l a n d , Oreg. 512 pp.
Washington Department o f Game. 1982. F i s h and W i l d l i f e Resource S t u d i e s , S u l t a n R i v e r P r o j e c t Stage I 1 F i n a l Report. Snohomish County PUD No. 1. 217 pp.
APPENDIX A
Aggrega t ion fo rmu la and s u i t a b i l i t y i n d i c e s o b t a i n e d f rom t h e USFWS H a b i t a t S u i t a b i l i t y Index Models - p i l e a t e d woodpecker (Schroeder 1982).
/ Percent t r e e canopy c l o s u r e (V1)
Number o f t r e e s >20 i n . dbh /ac re ,+2i
Number o f stumps >1 ft. i n h e i g h t A l l F o r e s t e d - Food/Cover/ and >7 i n . d iamete r and/or l o g s Cover Types Reproduc t ion >7 i n . d i a m e t e r p e r 1 a c r e ( V j )
Number o f snags >20 i n . d b h l a c r e \ ( v 6 )
Average dbh o f snags >20 i n . dbh (V7)
112 Western p o r t i o n o f range: Lower o f (V1 x V2 x V3)
Food /cover / rep roduc t ion 1 /2
o r (V6 x V7)
H S I d e t e r m i n a t i o n . The H S I f o r t h e p i l e a t e d woodpecker i s equa l t o t h e l i f e r e q u i s i t e v a l u e o f f o o d / c o v e r / r e p r o d u c t i o n .
Pileated woodpecker
v 2 Percen t t r e e canopy c l o s u r e
v, Number o f t r e e s t u ~ p s > 0 . 3 m ( 1 . 0 f t ) i n h e i g h t and > 18 cm ( 7 i n c h e s ) d i a m e t e r and /o r l o g s > 18 c m ( 7 i n c h e s ) d i a m e t e r / 0 . 4 ha ( 1 . 0 a c r e ) .
v 2 Number o f t r e e s > 51 cm ( 2 0 i n c h e s ) dbh/0 .4 ha ( 1 . 0 a c r e ) .
0 .05 .10 .15 .20 +
v 6 Number o f snags > 5 1 cm ( 2 0 i n c h e s ) dbh /0 .4 ha ( 1 . 0 a c r e ) .
v, Average dbh o f snags > 51 cm ( 2 0 i n c h e s ) dbh.
f iPPENDIX B
Word model used in original HEP S tudy (WDG 1982)
June 1978
PILEATEI WGDPECKER Coni fe rcus F o r e s t
Genera l The p i l e a t e d woodpecker (Dryocopus p i l e a t u s ) i s a yea r -a rcund ,
though uncommon, r e s i d e n t of dense o ld-aged c o n i f e r o u s f o r e s t s i n Eco- r e g i o n 2410.
Food Requirements P i l e a t e d woodoeckers f o r a a e on snaas i n 100+ Year o l d f o r e s t s i n . .
w e s t e r n Oregon ( ~ a h n a n 1977). - snags of"19 i n c h e s 7 4 8 cm) dbh o r l a r g e r were used most f r e q u e n t l y . I n s e c t s comprise most of t h i s woodpecker ' s d i e t , w i t h c a r p e n t e r a n t s (Componotus s p p . ) and t e r m i t e s (w s p p . ) b e i n g t h e most p r e f e r r e d foods (Beal 1911). P i l e a t e d s f e e d on many s p e c i e s o f f r u i t s and nu t s i n t h e f a l l .
Water Requirements P i l e a t e d wood2eckers r e q u i r e l ~ r g e amounts o f wa te r i n c a p t i v i t y
which may i n d i c a t e t h e i r wa te r r equ i rement s i n t h e w i l d (P f i t zenmeyer 1956 c i t e d by Jac!dn and S c o t t 1976). Hoyt (1957) found v e r y few n e s t s l o c a t e d f a r from a s o u r c e o f w a t e r
Cover Reouirements Pi l e a t e d woodoeckers a r e d e ~ e n d e n t on s n a a s f o r f o r a q i n q , r o o s t i n q ,
and n e s t i n g ( ~ c o t t ' e t a l . 1977): In t h e Blue i o u n t a i n s o f ~ T e g o n , -
p i l e a t e d s p r e f e r r e d mature o l d s t a o d s wi th a 70% crown c l o s u r e (Thomas e t a1. i n p r e s s ) . Conner e t a l . (1975) r e p o r t s a p r e f e r e n c e f o r t imber s t a n d s w i t h s a w ~ i m b e r of 1 5 t o 18 inches (38 .1 -45 .7 cm) dbh i f n e s t s i t e s a r e a v a i l a b l e .
R e p r e d u c t i v e Requirements E u l l (1975) r e p o r t s t h a t p i l e a t e d woodpeckers n e s t i n c a v i t i e s o f
panderosa p i n e (Pinus ponderosa) and wes te rn l a r c h ( L a r i x o c c i d e n t a l i s ) . T rees s e l e c t e d had an ave rage dbh of 23 inches (58-r g r e a t e r and were a t l e a s t 40 f e e t (12 m ) t a l l . Jackman (1974) r e p o r t e d t h a t p i l e a t e d s r e q u i r e a n e s t t r e e of a t l e a s t 20 inches (50 cm) dbh. These woodpeckers show a p r e f e r e n c e f o r t a l l , dead t r e e s w i t h few l imbs ( S c o t t e t a l . 1977) .
S p e c i a l H c b i t a t Requirements F i i e a t e d woodpeckers u t i l i z e h o l e s f o r r o o s t i n g yea r -a round (Jackman
and S c o t t 1975). F resh r o o s t i n g ho les a r e u s u a l l y e x c a v a t e d :n t h e f a l l . Bu l l (1975) r e p o r t s t h a t s e v e r a l r o o s t t r e e s a r e used. The ho le used f o r r o o s t i n q i s u s u a l l y i n a l i v e t r e e and f r e q u e n t l y h i g h e r i n t h e t r e e t h a n n e s t h o l e s (Hoyt 1957) .
I n t e r s p e r s i o n Reou i renen t s S t a n d s i z e s of a t l e a s t 300 a c r e s ( 1 2 1 ha) a r e r e q u i r e d t o meet t h e ~
home range requ i rement s of p i l e a t e d koodpeckers i n t h e ~ l u e Mountains of Oreson (Thomas e t a l . i n p r e s s ) . Optimal h a b i t a t shou ld ha\/e a t l e a s t 1 . 2 s n a g s / 2 . 5 a c r e s ( 1 ha ) . Snags should be o v e r 20 i n c h e s (50 cm) dbh.
J u n e 1978
Spec ia l C o n s i d e r a t i o n s P i l e a t e d woodpeckers p r e f e r dense s e c l u d e d t imber and a r e i n t o l e r a n t
o f human d i s t u r b a n c e (Jackman and S c o t t 1975).
REFERENCES CITED
Bea l , F . E . L . 1911. Food of t h e wpodpeckers of t h e Uni t ed S t a t e s . U.S. Bio. Surv . B u l l . No. 37:164.
B u l l , E . L. 1975. H a b i t a t u t i l i z a t i o n of t h e p i l e a t e d woodpecker, Blue Mounta ins , Oregon. M. S. T h e s i s , Oregon S t a t e Univ. , C o r v a l l i s . 58 PP .
Conners, R . N., R. G . Hooper, H. 5. Crawford, and H. S. Mosby. 1975. Woodpecker n e s t i n g h a b i t a t i n c u t and uncut woodlands i n V i r g i n i a . J . W i l d l . Manaae. 3 9 ( l ) : 144-150.
Hoyt, 5. F. 1957. he e;ology of t h e p i l e a t e d woodpecker. Ecology 38(2) : 246-256.
Jackman, 5 . M. 1974. Woodpeckers o f t h e P a c i f i c Nor thwest ; t h e i r c h a r a c t e r i s t i c s and t h e i r r o l e i n t h e f o r e s t . M.S. T h e s i s , Oregon S t a t e U n i v . , C o r v a l l i s . 147 pp.
Jackman, 5. M., and J . M. S c o t t . 1975. L i t e r a t u r e review o f twenty- t h r e e s e l e c t e d f o r e s t b i r d s o f t h e P a c i f i c Northwest . Region 6. U.S. F o r e s t S e r v i c e . 381 pp.
Mannan, R . W. 1977. Use o f snags by b i r d s , Douglas F i r Region, Western Oregon. M. S. T h e s i s , Oregon S t a t e Univ . , C o r v a l l i s . 114 pp.
P f i t zenmeyer , H. T. 1956. L i f e h i s t o r y and behav io r p a t t e r n s of t h e p i l e a t e d woodpecker r e l a t i v e t o u t i l i t y l i n e s . M.S. T h e s i s , Pennsylvania S t a t e Univ . , U n i v e r s i t y Park.
S c o t t , V . E . , K . E . Evans, 0. R . P a t t o n , and C . P. S tone . 1977. P i l e a t e d wcodpecker. Page 35 i n V . E. S c o t t ed. Cavity nes t ' ng b i r d s of North American f o r e s t y USDA F o r e s t T e r v i c e Agric.. Hdbk. NO. 511.
Thonas, J . W . , R . Anderson, C. Maser, and E. L. Bu l l . Dead t r e e ("snag") r e q u i r e m e n t s f o r dependent w i l d l i f e s p e c i e s i n t h e Blue Mountains o f Washington and Oregon. I n J. W. Thomas, ed. W i l d l i f e h a b i t a t i n managed f o r e s t s - The Blue Mountains of Ozcjon and Washington. Pac. Nor thwest For. and Range Exp. S t n . , U.S. Dept. A g r i c . , P o r t l a n d , Oreg. ( i n p r e s s )
P I LEATE5 WOOGPECKER Mfi;FLKLS FCilEST
June 1978
H A B I T A T S U I T A B I L I T Y 1t:DEX
P i l ~ a t e d Uoodpecker in Coniferous Forest
Ecoregion 2410
Food Value ( X I ) = ( I 2 x 13) 1/2
Where: I2 = S u i t a b i l i t y Index (S I ) of dbh of a v a i l a b l e snags
I 3 = SI of number of snags per acre .
Cover Value (X2) ,= ( I 1 x 1 4 ) 1/2
Where: I = S I o f t r e e cancpy c losure . 1
Id = SI of s i ze of timber s tand ( a c r e s ) .
R e p r o d x t i v e Value ( X j ) = ( I 2 x I 3 x I s ) 1 / 3
% e r e : I2 = SI of d b h of ava i lab le snags.
I = SI of number of snags per ac re . 3
I5 = SI of snag height .
Water Value ( X 4 ) = I6
Where: I = SI of d is tance t o water. 6
The Habi ta t S u i t a b i l i t y Index i s the lowest X,, value.
Pine Mar ten
(Mar tes americana)
General
0 The m a r t i n (Mar tes americana) i n h a b i t s mature and o l d - g r o w t h f o r e s t c o r n u n i t i e s t h r o u g h o u t N o r t h America (A1 l e n 1982).
0 The spec ies i s m o s t l y c a r n i v o r o u s , g e n e r a l l y n o c t u r n a l , and a c t i v e th roughou t t h e year ( A l l e n 1982).
Food Requirements
0 Consume a wide v a r i e t y o f f o o d t ypes on an annual b a s i s .
0 M a m a l s are t h e most i m p o r t a n t food i t e m i n w i n t e r . W i n t e r foods incude Doug1 a s ' s q u i r r e l , v o l e s , snowshoe hares and f l y i n g s q u i r r e l (Koeh le r and Hornocker 1977, S o u t i e r r e 1979, Z i e l i n s k i -- e t a l . 1983).
0 Voles a r e u t i l i z e d more t h a n any o t h e r spec ies i n w i n t e r (Weckwerth and Hawley 1962, Koehler and Hornocker 1977, Z i e l i n k s i -- e t a l . 1983).
0 I n v e r t e b r a t e s , b e r r i e s , and p a s s e r i n e b i r d s a r e common f o o d i t e m s s p r i n g th rough f a l l (Weckwork and Hawley 1962, Z i e l i n s k i -- e t a l . 1983).
0 Dense c o n i f e r s on mesic s i t e s a r e t h e ma jo r f o r a g i n g areas used i n t h e w i n t e r (Koeh le r -- e t a l . 1975).
0 Forages i n D o u g l a s - f i r l h e m l o c k f o r e s t s d u r i n g w i n t e r i n Washington ( d e Vos and Guenther 1952).
Water Requirements
0 No i n f o r m a t i o n on wa te r requ i rements was found i n t h e l i t e r a t u r e .
Cover Requirements
I n h a b i t s c o n i f e r o u s f o r e s t s w i t h g r e a t e r t h a n 40% canopy c l o s u r e (Spencer -- e t a l . 1983, Brown 1985).
Found f rom s e a - l e v e l t o t i m b e r l i n e a t a l l t i m e s o f t h e y e a r i n wes te rn Washington (WDG 1975, Maser -- e t a l . 1981).
Use areas above t i m b e r l i n e d u r i n g t h e summer months f o r f o r a g i n g and as t r a v e l r o u t e s (Koeh le r and Hornocker 1977, Spencer 1986).
P r imary h a b i t a t i s mature and o l d g rowth c o n i f e r o u s f o r e s t . S a p l i n g and p o l e s tage c o n i f e r o u s f o r e s t s a r e secondary f o r a g i n g h a b i t a t s (Brown 1985).
r Snags, caves, ground burrows, r o c k p i l e s , s l a s h p i l e s , Douglas ' s q u i r r e l middens ( w i n t e r o n l y ) and stumps a r e f r e q u e n t r e f u g e s i t e s (Master 1980, Spencer -- e t a l . 1983, B u s k i r k 1984).
r Downfa l l a l l o w s mar ten access t o r o d e n t p r e y a c t i v i t y under deep snow (S teven ton and Major 1982).
r Openings (e.g., c l e a r c u t s , meadows, r o a d s ) a r e u s u a l l y avo ided i n w i n t e r (Spencer - e t - a l . 1983), though c r o s s i n g s o f up t o 180 y a r d s i n w i n t e r have been reco rded i n Maine ( S o u t i e r r e 1979).
r Marten r a r e l y use c l e a r - c u t s tands l e s s t h a n 15 y e a r s o l d i n Maine ( S o u t i e r r e 1979).
Reproduc t i ve Requirements
r Reproduc t i ve requ i rements a r e assumed t o be i d e n t i c a l w i t h cover requ i rements ( A l l e n 1982).
r Den i s commonly i n a h o l l o w t r e e o r t r e e c a v i t y , a l t h o u g h any r e f u g e s i t e may be ~ ~ s e d f o r breed ing.
Spec ia l H a h i t a t Requirements
r Home ranges o f males average 640 ac res i n Maine and females average 160 acres i n Montana (S teven ton and M a j o r 1982, Hawley and Newhy 1957).
r Hone range s i z e v a r i e s w i t h c l e a r c u t acreage (S teven ton and Major 1982).
0 P a c i f i c Nor thwest Regiona l F o r e s t S e r v i c e g u i d e l i n e s recommend minimum h a b i t a t s i z e u n i t s of 160 con t iguous acres o f mature o r o l d g rowth f o r e s t (U.S. F o r e s t S e r v i c e 1983).
r Home ranges o f females may o v e r l a p , b u t males defend r i g i d t e r r i t o r i e s .
r Home range boundar ies o f t e n c o i n c i d e w i t h t h e edges o f t o p o g r a p h i c o r v e g e t a t i v e f e a t u r e s , such as l a r g e , open meadows, burns and streams (Hawley and Newby 1957).
r The percen t t r e e canopy c l o s u r e ( s e e V1) and success iona l ( see V3) s tage o f t h e s tand a r e two l i m i t i n g v a r i a b l e s f o r d e t e r m i n i n g t h e s u i t a b i l i t y o f mar ten w i n t e r h a b i t a t ( K o e h l e r and Hornocker 1977, S o u t i e r e 1979, Spencer -- e t a l . 1983).
REFERENCES CITED
A l l e n , A. W. 1982. H a b i t a t s u i t a b i l i t y i n d e x models: marten. U.S. F i s h and W i l d l i f e S e r v i c e , FWS/OBS-82 10.11. 9 pp.
Brown, E. R. ( ed . ) 1985. Management o f w i l d l i f e and f i s h h a b i t a t s i n f o r e s t s o f western Oregon and Washington. P a r t 1. U.S. Dept. Agr ic. , F o r e s t S e r v i c e , P a c i f i c Region. 332 pp.
B u s k i r k , S. W . 1984. Seasonal use o f n e s t i n g s i t e s by mar ten i n s o u t h - c e n t r a l Alaska. J. W i l d l . Manage. 4 8 ( 3 ) : 950-953.
de Vos, A. and S. E . Guenther. 1952. P r e l i m i n a r y l i v e - t r a p p i n g s t u d i e s o f mar- ten . J. W i l d l . Manage. 16(2):207-214.
Hawley, V . D. and F. E. Newby. 1957. Mar ten home ranges and p o p u l a t i o n f l u c - t u a t i o n s . J. Mammal. 3 8 ( 2 ) : 174-184.
Koehler , G. M. and M. G. Hornocker. 1977. F i r e e f f e c t s on mar ten h a b i t a t i n t h e S e l w a y - B i t t e r r o o t Wi lderness. J. W i l d l . Manage. 41:5OO-505.
Koeh le r , G. M., W. R. Moore, and A. R. T a y l o r . 1975. P r e s e r v i n g t h e p i n e mar ten: Management g u i d e l i n e s f o r wes te rn f o r e s t s . West. W i l d l a n d s 2(3):31-36.
Masters , 4. D. 1980. Daytime r e s t i n g s i t e s o f two Ad i rondack p i n e martens. J. Mama1 . 61:157.
Maser, C., B. R. Mate, J. F. F r a n k l i n , and C. T. Dyrness. 1981. N a t u r a l h i s t o r y o f Oregon coast mammals. P a c i f i c Nor thwest Fo r . and Range Exp. S t a t i o n . USDA F o r e s t S e r v i c e Gen. Tech. Rep. PNW-133.
Spencer, W . D., R. H. B a r r e t t and W. J. Z i e l i n s k i . 1983. Mar ten h a b i t a t p r e f e r - ences i n t h e n o r t h e r n S i e r r a Nevada. J. W i l d l . Manage. 47(4):1983.
Spencer, W. D. ph.D. Candidate U n i v e r s i t y o f Ar izona. Persona l Communication w i t h Dave Hays, J u l y 1986.
Steventon, J . D., and J. T. Major . 1982. Mar ten use o f h a b i t a t i n commerc ia l l y c l e a r - c u t f o r e s t . J. Wi ld. Manage. 46(1):175-182.
S o u t i e r r e , E. C. 1979. E f f e c t s o f t i m b e r h a r v e s t on mar ten i n Maine. J. W i l d l . Manage. 43(4):850-860.
U. S. F o r e s t Serv i ce . 1983. Regiona l g u i d e l i n e s f o r i n c o r p o r a t i n g minimum management requ i rements i n f o r e s t p l a n n i n g . Unpubl ished document, Region 6, Por t1 and, Ore.
Washington Department o f Game. 1982. F i s h and W i l d l i f e Resource S t u d i e s . S u l t a n R i v e r P r o j e c t Stage I 1 F i n a l Repor t . Snohomish County PUD No. 1. 217 pp.
Uashington Dept. o f Game. 1975. Mar ten S t a t u s Report . Non-game Data Center , Olympia. 4 pp.
Weckworth, R. P. and V. D. Hawley. 1962. Mar ten f o o d h a b i t s and p o p u l a t i o n f l u c t u a t i o n s i n Montana. J. Wi ld1 . Manage. 26 (1 ) :55-74.
Z i e l i n s k i , W. J., W. D. Spencer and R. H. B a r r e t t . 1983. R e l a t i o n s h i p between food h a b i t s and a c t i v i t y p a t t e r n s o f p i n e mar tens. J. Mamm., 64 (3 ) :387-396.
APPENDIX A
A g g r e g a t i o n f o r m u l a and s u i t a b i l i t y i n d i c e s o b t a i n e d f rom USFWS H a b i t a t S u i t a b i l i t y Mode l s - m a r t e n ( A l l e n 1982).
S u ' t a b i l j t y Index ( S I ) 9~6:hs f o r b , a b i t a t v a r i a k l e s , The r e l a t ; c r z ' ? < ; s between v a r i o c s c o r , d i i i o r s o f ha3i:a: var:;:ies a ~ 3 na>':2: s l j i t ; b i l+ :y f c r t h e T a r t e n a r e g r a p h i c a l l y r e p r e s e ~ r e d i n :h is se : t : cn .
( v 2 ) P e r c e n t o f o r e r s t s r y 1 .o canopy c l o s ~ r e c o n - x
p r i s e d of f i r o r s p r u c e .
r-. .- I n m 0.4
in i
0 . 2 i
( V , ) S ~ c c e s s i a n a l st-,- - . s t a n a . x w
A ) s h r d k - s e e d l i n g - 8 ) p o l e s a p l i n g C ) .I In" .- F 0 . 8 j '0.6 / ' I 0) m a r u r e o r o l d g r o w t h
2 0 . 4 1 /
A B C D
S u c c e s s i o n a l S t a g e
E c u a z ; 3 n s . I n o r d e r t o o b t a i ~ l i f e r e q u i s i t e v a l u e s f a r t h e m a r t e c , t n e S I v a l u e s Tcr a p p r o p r i a t e v a r i a b l e s m d s t b e c o m b i n e d t h r o u g h t h e u s e o f equa- t i o n s . A d : s c a s s i o r : a n d e x p l a n a t i o n o f t h e assumed r e l a t i o n s h i p s b e t w e e n v a r i a b l e s was i n c l u d e d u n d e r M3de; D e s c r i p t i o n , a n d t h e s p e c i f i c e q d a t i c n i n t h i s mode l wss c h o s e n t o m i m i c t h e s e p e r c e i v e d b i o l o g i c a l r e l a t i o n s h i p s a s c : c s e l y a s p c s s i b l e . The s u s g e s t e d e q u a t i o n f o r o b t a i n i n g a w i n t e r c o v e r v a l u e i s p r ~ s e n t e d b e l o w .
( V h ) P e r c e n t o f S - o u n d s u r i s c e c o v e r e d b y d o h n f a l l h * h i c h i s 2 7 . 6 cm ( 3 i n ) i n c i a ~ e t e r .
L i f e r e c u i s i t e C o v e r t y p e E q u a t i o n s
.- L2 m 0 . 4 - + '- 3
-0.2 -
HSI d e t e r m i n a t i o n . S i n c e w i n t e r c o v e r was t h e o n l y l i f e r e q u i s i t e c o n s i d e r e d i n t h i s m s d e l , t h e gS1 e q u a l s t h e w i n t e r c o v e r v a l u e .
-
-
25 5 0 7 5 1 0 0 %
PPPEK3IX B
Word model u s e d i n o r i g i n a l HEP S t u d y ( W D G 1982)
MARTEN
Genera l - The m a r t e n ( N a r t e s amer icana) i s a Smal l c a r n i v o r e i n h a b i t i n g montane and
A-
suba!p ine c o n i f e r f o r e s t s i n t h e P a c i f i c N o r t h w e s t . T h i s member o f t h e m u s t e l i d f a m i l y i s found i n t h e Olympic Moun ta ins , C o a s t Range and Cascade Mounta ins of h 'ash ington and Oregon eas tward t h r o u g h t h e Okanoyan H i g h l a n d s and Sel.kir-k I<oun ta ins (Hagmeier 1956; YOcOm 1974).
Food Requi r m e n t s - Martens e a t a w i d e v a r i e t y o f i t e m s , b u t s m a l l mammals a r e t h e ma in p rey .
t 4 a r ~ e n s p r e y l a r g e l y on m i c r o t i n e s , such as meadow v o l e s ( M i c r o t u s spp.), red -bacLcd v o l e s ( C l e t h r i o n o m y s spp . ) , h e a t h e r v o l e s (Phenacomys i n t e r m e d i u s ) , zr id bog lc- zings (Synaptomy: spp . ) (Cowan and MacKay 1950; K o e h l e r and Hornocker 1977, and o t h e r s ) . T r e e s q u i r r e l s ( T a m i a s c i u r u s spp., G l a n c m s s a b r i n u s ) were i i p o r t ~ n t c o r p o n e n t s o f t h e d i e t i n E ~ h i n g t o n - ( ~ ~ i e w b y i d M o n t a n a ( I . :arshal l 1946; Neckwer th and Hawley 1962) ; t h e i r i m p o r t a n c e appears t o v a r y g r e - t l y f r o m s i t e t o s i t e ( K o e h l e r e t a l . 1975) . Grcund s q u i r r e l s (Spermophi lus s p p . ) a r e t ? k e n d g r i n g t h e summer f rom f o r e s t o p e n i n g s (Cowan and Mackay 1950; t i 6 ) P i k a s (Ochotona p r i n c e s ) a l s o appear i n t h e d i e t (Cowan and I:icK.y 1950; i t r e e t e r a n d B r a u d Deer m i c e (Peromyscus s p p . ) a n d ct,ipt::un),s (Eutamias spp . ) a r e t a k e n o n l y i n l ow numbers (Cowan and MacKay
1977) .
The u t i 1 i z a t i o n o f b i r d s , i n s e c t s , and b e r r i e s v a r i e s s e a s o n a l l y . kcung b i r d s 2 n d eggs a r e t a k e n i n t h e s p r i n g ( L e n s i n k e t a l . 1952; Weckwerth and t l ? r l e y 1962) and g rouse d u r i n g t h e w i n t e r (Newby 1951; Q u i c k 1 9 5 5 ) . I n s e c t s I:dy be h e a v i l y e x p l o i t e d d u r i n g t h e summer i n some a r e a s . C a r p e n t e r a n t s (Cz~:zonotus spp. ) and h o r n e t s (Vespu la spp. ) were found i n o v e r 90% o f t h e s c a t s ~ o l l & i ? ~ o o i n t h e Washington Cascades d u r i n g t h e summer (Newby 195?). B l u e b e r r i e s ! \ f a c c i n i u n - r p ~ . ) and t , a ~ t h o r n b e r ~ - i e s ( C r a t a e y s s p . ) a r e consumed i n t h e ! lc : . r t i t r r , Rock ies d u r i n g l a t e .summer and f a l l M u n e 1961; Weckwerth and l iak i ley 1952; K o e h l e r and Hornocker 1975) .
The m a r t e n forages l a r p e l y on t h e g round . The s p e c i e s a c t i v e l y searches :bc bases o f t r e e s f o r r o d e n t b u r r o w e n t r a n c e s d u r i n g t h e w i n t e r ( K o e h l e r e r 21. 1975) . Elar tens a r e a l s o p a r t l y a r b o r e a l , 2s i n d i c a t e d b y t h e presence c f t r e e s q u i r r e l s and s m a l l b l r d s i n t h e d i e t ( L a r r i s o n and Sonnenberg 1962) . irme s p e c i e s caches c a p t u r e d p r e y under t h e snow o r i n t r e e c a v i t i e s ( M u r i e 1551) .
Dense c o n i f e r s on mes ic s i t e s a r e t h e m a j o r f o r a g i n g a r e a s used by mar tens : r :he w i n t e r ( K o e h l e r e t a : . 1975) . Mes ic s i t e s g e n e r a l l y s u p p o r t h i g h e r i : : ? p c p u l a t i o n s t h a n h y d r i c o r x e r i c s i t e s ( K o e h l e r e t a ) . 1975) . A dense :r:r.C o f c o n i i e r s (canopy c l o s u r e o f more t h a n 30") ~ r o v i d e s b o t h c o v e r and
i r ? c . r#ui;.tir o f t r e e bases f o r i o r a a i n g a c t i v i t i e s ( K o e b l e r e t a l . 1975);
~ ~
A ; ,
:,-,,;:,er, a c o x p l e t e l y c l o s e d canopy may shade o u t t h e v e g e t a t i v e g round c o v e r 11, j i p r c v i d e s food and c o v e r f o r P rey s p e c i e s . I h r t e n s f o r a g e d i n m a t u r e ,pc ::?!as f i r (Pseudotsuga l n e n z i e s i i ) f o r r s t s and m i xed s tands o f Douglas f i r - , wes te rn hemlock (w h ~ t e r o p l y l l a j and w e s t e r n redcedar ( T h u j a
7 i c a t a ) d u r i n g t h e w i n t e r i n Washington, (deVos and Guenther 1952).Most P--_ r l i r : t t r f b r a g i n g a reas were b e l o w 1067 m (3500 f t . ) e l e v a t i o n (Newby 1951) . C e ~ ~ s e s p r u c e ( P i c e a -- s p . ) a n d s u b a l p i n e f ir ( A b i e s l a s i o c a r a ) f o r e s t s more
. t h i : n 100 y e a r s o l d were used f o r w i n t e r f m n d ( K o e h l e r e t a l . 1975) . E l e v a t i o n s v a r i e d from 1341 t o 2377 m (4400 t o 7800 ft.) ( K o e h l e r ;r,i t i c rnocker 1977) . N a r t e n do n o t h u n t i n open ings d u r i n g t h e w i n t e r and z v c i d apen ings more t h a n 100 m (328 f t . ) w i d e ( K o e h l e r e t a l . 1975) .
I S u ~ w e r f o r a g i n g a r e a s f o r m a r t e n s i n c l u d e s mes ic c o n i f e r f o r e s t s and
a d j a c e n t open ings . M a r t e n s s e a r c h f o r v o l e s , b e r r i e s , i n s e c t s , and g round s q u i r r e l s i n open meadows and b u r n s ( K o e h l e r and Hornocker 1977) . The s p e c i e s has been observed h u n t i n g i n r o c k s l i d e s a t o r n e a r t i m b e r l i n e i n B r i t i s h Columbia (Cowan and MacKay 1950) , C o l o r a d o ( S t r e e t e r and Braun 1968) . and I l t 3 h (Hzyward 1952) . M a r t e n s f o r a g e d i n s u b a l p i n e f i r - m o u n t a i n hemlock (1~12 z i e r t e n s i a n a ) f o r e s t s between 1219 and 1676 m (4000 and 5500 m) e l e v a t i o n r!:lrina t h e sumqer i n i r ' ash ing ton (Newby 1.951). S u b a l p i n e f i r - s p r u c e f o r e s t s a t 2743 t o 335; T (9200 t s 11,000 f t . ) e l e v a t i o n were used i n C o l o r a d o (Yeager 2nd Eemington 1956) .
Prey a v a i l a b i l i t y i s p r o b a b l y t h e m a j o r f a c t o r i n f l u e n c i n g m a r t e n d i s t r i b u t i o n ( K o e h l e r e t a l . 1975) . Y a t u r e mes ic f o r e s t s t a n d s s u p p o r t a h i g h v o l e p o p u l a t i o n and show h i g h m a r t e n use . Openings p r o v i d e abundant v o l e s , i r f s e c t s , and b e r r i e s and r e c e i v e h i g h m a r t e n use when these a r e a v a i l a b l e . When 6Lundant b u t c lumped f o o d r e s o u r c e s were p r e s e n t i n A laska , m a r t e n movements wcre rcduced ( L e n s i n k e t a l . 1 9 5 5 ) .
I.!ater Requi r e ~ q e n t s - - .. .
No i n f o r m a t i o n on w a t e r r e q u i r e m e n t s was f o u n d i n t h e l i t e r a t u r e . . . . . . .
Cover Rcqu i re i . ien ts --
D u r i n g t h e w i n t e r , t h e m a r t e n i s mos t a c t i v e i n s tands w i t h more t h a n 30; czncpy c o v e r t o p r o v i d e p r o t e c t i o n f r o ~ i ~ wea the r ( K o e h l e r e t a l . 1975) . T i , ? zpec ies a v o i d s o p e n i c g s ; i f t h e o p e n i n g i s more t h a n 100 m (228 i t . ) w i d e . n , e r t t n s r : i l l n o t c r o s s i t . i i i n t e r den s i t e s a r e l o c a t e d benea th snow under and a round l e a n i n g t r e e s , s tumps, w i n d f a l l s and heavy u n d e r g r o w t h (Newby 1951) . 1 -k r tens b . i l l use f a l l e n l o g s as t r a v e l w a y s .
Kat- tens w i l l use o p e n i n ~ s as w e l l as f o r e s t d u r i n g t h e summer. Areas w i t h o u t overhead c o v e r , such as r e c e n t l y l o g g e d and burned a r e a s , a r e a v o i d e d (deVos 1951; Hawley and t4ewSy 1 9 5 7 ) . Cover i n open ings may be p r o v i d e d b y b r u s h , do:,.ned t r e e s , and herbaceous v e g e t a t i o n ; t h i s l o w l a y e r o f c o v e r i s u s u a l l y b u r i e d by snovl i n t h e w l n t e r .
Ground b u r r o w s , r o c k c r c v i c e s . and t r e e s mav be used by mar tens as r e s t - i n - s < ~ C S . Y a r t e n i n N i n n e i o r a u s u a l l y u t i l i z e d burro!.,s benea th b o u l d e r s a n d I:,;:, r c r h p i l e s , and r o c , c r e v i c e s f o r r e s t i n g ; o n l y 1 o f 13 r e s t S i t e s was l s c a t c d i n a t r e e (l;ech and Facers 1976) . A l l 5 r e s t i n g s i t e s i d e n t i f i e d i n
a fiew York s tudy were p laced i n t r e e s (Masters 19891, Both dtciduous and.conl- / - u s t r e e s were used; t h e i r dbh ranged from 25 t o 55 cm ( , O n , t i ] , , I",),
Rock c r e v i c e s were used f o r cover on a l p i n e rock s l i d e s i n Colorado ( S t r e e t e r and Sraun 1968).
~ e p r o d u c t i v e Requirements -
Snass a r e an impor tan t component of breeding h a b i t a t f o r marten, Tree c a v i t i e s and hollow logs a r e used a s denning s i t e s (deVos 1951). Of 16 marten dens i n Idaho, 1 3 were i n hollow l o g s and 3 were i n hollow stumps (Marshall 1951) . Such snags a r e probably only a v a i l a b l e i n mature c o n i f e r s t a n d s , a l though q u a n t i a t i v e d a t a on denning h a b i t a t a r e l ack ing . Rock p i l e s a r e p o t e n t i a l den s i t e s ; young marten were found beneath one i n Colorado (Reming- ton 1952).
The marten r e q u i r e s geographic i s o l a t i o n t o prevent ove r t r app ing (Koehler e t 21. 1975) . Flarten a r e e a s i l y t rapped and roads may a l low t o o much access t o t h e i r h a b i t a t .
I n t e r s p e r s i o n Requirement
Narten requ;'re mature mesic f o r e s t s t a n d s f o r food and cover dur ing t h e kcinter and f o r den s i t e s i n t h e s u m e r . They forage i n f o r e s t s and openings such a s o l d bu rns , meadows, and rock p i l e s i n t h e sumner. Old-growth f o r e s t with solre i n t e r s p e r s e d openings appears t o r e p r e s e n t opt imal marten h a b i t a t (Neiqby 1951; Koehler e t a1 . 1975) .
Marten popu la t ions may undergo seasonal movements i n some aE.5 (Newby 1951) but no t i n o t h e r s (Haw1 y and Nerib 1957). Home range s i z e s i n Glac e r 5 1 ' - - - - National Park averaged 2 . 4 km (0.94 mi .$ ) f o r males and 0 . 7 km2 (0.27 mi. ) f o r females (Hawley and Newby 1957 . S i z e ranges were 0 . 8 t o 4 . 4 k9 (0.34 t o 1 .69 mi.2) and 0 . 1 t o 1 . 8 km2 ( 0 . 0 3 t o 0.70 mi .2 ) f o r males and females r e s p e c t i v e l y . he home range of r ad io t r acked male marten i n Minnesota was J 10 .5 t o 19.9 k m ( 3 . 9 t o 7 .7 mi . 2 ) i n a r e a ( k c h and Roger 1976). The one 2 f m a l e t r acked i n t h i s s tudy ranged over an a rea of 4 . 3 km (1 .7 ni.2).
Specia l Cons ide ra t ions -
Extens ive logging of high e l e v a t i o n f o r e s t s e l i m i n a t e s marten h a b i t a t (Yeager 1950) . Cu t t i ng o r burning small a r e a s of f o r e s t i n c r e a s e s f o r e s t cormiunity d i v e r s i t y and t h e v a r i e t y and abundance of prey s p e c i e s (Yeager 1950; Koehler e t a l . 1975) . I n t e n s i v e g raz ing on high mountain meadows dec reases herbaceous cover and vo le popu la t ions (Yeager 1950) .
rianagemefit recommendations f o r marten inc lude l e a v i n g l a r g e blocks of r , ? t u r e f o r e s t und i s tu rbed , connec t ing s m a l l e r blocks wi th f o r e s t e d c o r r i d o r s , and n r e s e r v i n ~ a high canopy c l o s u r e by s e l e c t i v e c u t t i n g r a t h e r than c l e a r c u t t i n g (Eoeh lc r e t a l . 1975) .
LITEPATURE CITED
Co..:dn, 1.14. and R.H. MacKaY. 1950. r o o d h a b i t s o f t h e mdr ten , Mar tes amer icana, i n t h e Rocky M o u n t a i n r e g i o n o f Canada. Canad. F i e l d - N a t . GqT-m- 104.
. devos, A. 1951. Recent f i n d i n g s i n f i s h e r and m a r t e n e c o l o g y and management. T r a n s . N. Amer. W i l d l . Conf. 16:498-507.
deVos, A . and S . E. Guen the r . 1952. P r e l i m i n a r y l i v e - t r a p p i n g s t u d i e $ o f n a r t e n . J. W i l d l . Manage. 16(2) :207-214.
H o c j ~ ~ e i e r , E.M. 1956. D i s t r i b u t i o n of m a r t e n and f i s h e r i n N o r t h America. Canad. F i e l d - N a t u r . 70 (4 ) :149-168 .
Habrley. V .D . and F.E. Newby. 1957. M a r t e n home range and p o p u l a t i o n f l u c t u a t i o n s . J . Mammal. 38(2) :174-184.
t l z y ~ a r d , C.L. 1952. A l p i n e b i o t i c comnuni t i e s o f t h e U i n t a Mounta ins , U t a h . E c o l . Konogr. 22:93-120.
k:c;t#ler, G.M. , W.R. [.loore, and A . R . T a y l o r . 1975. P r e s e r v i n g t h e p i n e mar ten: Management g u i d e l i n e s f o r w e s t e r n f o r e s t s . l f es t . W i l d l a n d s 2 (3 ) :31 -36 .
K o e h l e r , G.M. and M.G. Hornocker . 1977. F i r e e f f e c t s on mar ten h a b i t a t i n t h e Se1k;ay-Bi t t e r r o o t W i l d e r n e s s . J . lii I d . Manage. 41 (3) :500-505.
K a r s h a l l , W.T. J . f 4ama l
L e n s i n k , C.J., R.O. Skoog, and J . L . Buck ley . 1955. Foo6 h a b i t s o f m a r t e n in i n t e r i o r A l a s k a and t h e i r s i g n i f i c a n c e . J. Wild.. Hanage..19(3):364-368.
1946. W i n t e r f o o d h a b i t s o f t h e p i n e m a r t e n i n Montana. 2 7 ( 1 ) : 83-84.
K a s t e r s , R.D. J . H a m 6 1
P i n e m a r t e n as a f o r e s t p r o d u c t , J , For , 49(12)r89$&5; A
1980. Dayt ime r e s t i n g s i t e s o f two ad i rondeck p i n e mar tens, 6 1 ( 1 ) : 1 5 7 .
Mech, L.D. and n a r t e n i n 7 ?P.
L .L . Rogers. 1976. S t a t u s , d i s t r i b u t i o n , and movements o f n o r t h e a s t e r n M inneso ta . U,S.D,A, Fo r , Sery, Res, Pap: NC-143,
I b r i e , A. 1961. Som? f o o d h a b i t s o f t h e mar ten, J, M a m e l . 42(4)1516-$21,
l iev~by, Y . E . 1951. E c o l o g y o f t h s m a r t e n i n t h e T w i n Lakes a rea , Chelan County, 1,:ashington. K.S. T h e s i s , Wash. S t a t e Un iy , 38 pp,
. , ..>
? ~ i c k , H . F . 1955. Food h a b i t s o f mar ten (Mar tes amer icana) i n n o r t h e r n F i r i t i s h C o l m b i a . C a n a ~ . F i e l d - k t , m q - : 1 4 4 7 0 7
- . -.E-.1'!3L011, J.D. 1952. Food h a b i t s , g r o w t h , and b e h a v i o r o f t w o c a p t t v e p i n e
: ; a r t e n s J. Manna l . 33 (1 ) :66 -70 .
/" Stree te r , R.G. and C . E . Braun. 1968. Occurrence o f p ine martens, 3 a r t e s americana ( ~ a r n i v o r a ; f j u s t e l i d a e ) i n Colorado a l p i n e areas. Southw. N a t . - - 1 %-*--4~l.
Ueckwerth, R . P . and V.D. Hawley. 1962. Marten food h a b i t s and popu la t i on I f l u c t u a t i o n s i n !.lontant. J. I l i l d . Manage. 26(1):55-74.
, Yeager, L.E. 1950. I m p l i c a t i o n s of some ha rves t and h a b i t a t f a c t o r s on I pine marten management. Trans. N. Am. b l i l d l . Conference 15:313-334.
Yeager, L. E . and J. D. Remington. 1956. S j h t observat ions o f co lorado car tens , 1950-1955. J. Karnnal . 37(4) :521-524.
Yocon, C.F. 1974. S ta tus of marten i n n o r t h e r n C a l i f o r n i a , Oregon, and L l~sh ing ton . C a l i f . F i s h and Game 60:54-57.
ADDITIONAL REFERENCES -
deVos, A. 1952. Ecology and management o f f i s h e r and marten i n Ontar io. Ontar io Dept. Lands For . Tech. B u l l . 90 pp.
, A.T. Cringan, J.K. Reynolds, and H.G. Lumsden. 1959. B i o l o g f c a l i n v e s t i g a t i o n s o f t r a p l i n e s i n no r the rn Ontar io , Ontar io Dept. Lands Tech. B u l l . ( b i i l d l . Serv. 8 ) . 62 pp.
Franc is , G.R. and A . B . Stephenson. 1972. t la r ten ranges and food h a b i t s i n Algonquin P r o v i n c i a l Park, On ta r i o . Ont. Min. Nat. Resour. Rep. ( ~ i l d l . ) No. 91. 53 pp.
Harsha l l , U.H. 1942. The b i o l o g y and management o f the p ine marten Ph.D. Thesis , Univ. f l i c h . , Ann Arbor. 101 pp,
Tr:ining, H. and A. Hensley. 1947. The s t a t u s o f p ine martens i n Ca C a l i f . F i s h and Game 33(3):133-137,
I n Idaho.
l i f o r n i a .
H e r b l a n d - H igh e l e v a t i o n g r a s s l a n d s and meadows
m
Food I
Cover
I
F I E L D MEASUREMENT --
R e l a t i v e p r e y abundance
W i d t h o f o p e n i n g Abundance o f c o v e r
' H A B I T A T - R E Q U I REKECT
Food
L
Cover
I R ~ p r o d u c t i on
.I I n t e r s p e r s i o n
MARTEN
Evergreen F o r e s t
F I E L D M E A S U E E M E N T -. - - .-
Moisture c o n d i t i o n s R e l a t i v e prey abundance Stand age Pe rcen t canopy c l o s u r e P ropor t ion of open a r e a w i t h i n f o r e s t
Stand age Pe rcen t canopy c l o s u r e P ropor t ion of open a r e a w i t h i n f o r e s t
S tand age
P ropor t ion of open a r e a wi th in f o r e s t
Moi=.-turc conditions = vi5u.d a ~ e s s ~ e n 4 of the gcn&L m o w - w r c r e - i m ~ u s i r y eLI,-rL& mil ~ * f * r f i , d plad a r n r n u n i Sype, and - ) D p ~ ~ r a ~ h y (n gut
Evergreen Forest
*rind ayc
&ume> : - M a * % 76
C M ~ P ~ b 5 ~ ~ 1.5 m d e d fo r b V M r ~ y v e r n u d 5 (Koehler and Hornoc-tcr
1477). I -b 'evrr , lcss Shan 100Yo i5 w d e d
E v e r rreo FOG& 3
Doug1 a s ' S q u i r r e l
(Tamiasc iu rus d o u g l a s i i)
General
r P r i m a r i l y a s s o c i a t e d w i t h Douglas f i r f o r e s t s , a l t h o u g h i t occurs i n mixed s tands o f c o n i f e r o u s t r e e s and o f c o n i f e r o u s and dec iduous t r e e s ( B a i l e y 1936, I n g l e s 1965).
r Cone p r o d u c t i o n i s most i m p o r t a n t f a c t o r i n f l u e n c i n g h a b i t a t ( S m i t h 1965 and 1968).
Food Requi rements
r Major food i s seeds o f c o n i f e r s , e s p e c i a l l y Douglas f i r , S i t k a spruce, hemlock, t r u e f i r s and l o d g e p o l e p i n e ( B a i l e y 1936, Smi th 1965 and 1968).
r C o n i f e r p o l l e n , b i r c h and a l d e r c a t k i n s , maple samaras, n u t s , acorns, some s o f t f r u i t s , buds and ba rk a r e a l s o ea ten ( B a i l e y 1936, Smi th 1965).
r Eat t e r m i n a l and l a t e r a l buds o f Douglas f i r , e s p e c i a l l y d u r i n g w i n t e r and s p r i n g ( F i s c h and Dimmock 1978).
r Mushrooms and hypogeous f u n g i eaten ( S m i t h 1965, Maser -- e t a l . 1978, Fogel and Trappe 1978).
r P r e f e r s mature c o n i f e r f o r e s t s w i t h o u t dense u n d e r s t o r y f o r f e e d i n g . O c c a s s i o n a l l y feeds i n deciduous r i p a r i a n zones (Brown 1985).
r Presence o f cone caches i s a good i n d i c a t o r o f Douglas ' s q u i r r e l presence.
Water Requirements
r Most o f t h e water requ i rements a re met by m e t a b o l i c wa te r ( S m i t h 1965 and 1968).
Cover Requirements
r She1 t e r i n c a v i t i e s i n t r e e s and sometimes i n openings between rocks on t a l u s s lopes ( I n g l e s 1965).
r G l o b u l a r n e s t may be b u i l t i n crowns o f t r e e s , and, a t l e a s t i n w i n t e r , t h e s u b t e r r a i n e a n f o o d caches themselves may be used f o r s h e l t e r (WDG 1982).
r Nest , f o r a g e and t r a v e l i n mature c o n i f e r o u s t r e e s (Smi th 1965, 1968).
Reproduc t i ve Requirements
r Occurrence and t i m i n g o f b reed ing and s i z e o f l i t t e r s v a r y d i r e c t l y i n response t o changes i n p r o d u c t i o n o f c o n i f e r seed c rops ( S m i t h 1965 and 1968).
r Nests a r e u s u a l l v i n c a v i t i e s a t l e a s t 20 f t . above around i n t r e e s t h a t a r e a t l e a s t 17 i n . dbh (Brown 1985).
A
Spec ia l H a b i t a t Requirements
0 Body s i z e , j aw muscu la tu re , anatomy, and r e p r o d u c t i v e a c t i v i t y app be adapted t o l i f e i n f o r e s t s where Douglas f i r t r e e s predominate 1965 and 1968).
Near t o ( S m i t h
0 T e r r i t o r i a l t h r o u g h o u t t h e year , t e r r i t o r y s i z e v a r i e s f rom 1 ac re acres (Smi th 1965 and 1968).
S p e c i a l C o n s i d e r a t i o n s
r P a r t i c u l a r l y dependent on cones o f Douglas f i r t r e e s , wh ich a r e r e l a - t i v e l y easy t o open compared t o t h e cones o f such c o n i f e r s as l o d g e p o l e p i n e ( S m i t h 1965, 1968).
REFERENCES CITED
B a i l e y , V. 1936. No. 55.
Brown. E. R. (ed.)
The mdmmals and l i f e zones o f Oregon.
1985. Manaqement o f w i l d l i f e and f i s h hab i
N. h e r . Fauna
t a t s i n f o r e s t s o f wes te rn o r e i o n 'and washington. P a r t 2. U.S. Dept. Agr ic . , F o r e s t S e r v i c e , P a c i f i c Nor thwest Region. 301 pp.
F i s c h , G. G., and E. J. Dimmock 11. 1978. Shoot c l i p p i n g by Douglas ' s q u i r r e l s i n r e g e n e r a t i n g Douglas f i r . J. W i l d l . Manage. 42(2):414-418.
Foge l , R. and J. M. Trappe. 1978. Fungus consumpt ion (mycophagy) by sma l l a n i - mals. Nor thwest S c i . 5 2 ( 1 ) :1-31.
I n g l e s , L. G. 1965. Mammals o f t h e P a c i f i c S t a t e s . S t a n f o r d Univ. p ress , S tan fo rd , C a l i f o r n i a .
Maser, C., J. M. Trappe and R. A. Nussbaum. 1978. Fungal -smal l mammal i n t e r r e - l a t i o n s h i p s w i t h emphasis on Oregon c o n i f e r o u s f o r e s t s . Eco logy ( i n p r e s s ) .
Smith, C. C. 1965. I n t e r s p e c i f i c c o m p e t i t i o n i n t h e genus o f t r e e s q u i r r e l s Tamiasc iurus . Ph.0. t h e s i s , Univ. o f Washington, S e a t t l e .
. 1968. The a d a p t i v e n a t u r e o f s o c i a l o r g a n i z a t i o n i n t h e genus o f t r e e s q u i r r e l s Tamiasc iurus . Eco l . Monogr. 38:31-63.
Thomas, J . W., R. Anderson, C. Maser, and E . L. B u l l . Dead t r e e ( " s n a g " ) requ i rements f o r dependen; w i l d l i f e spec ies i n t h e B l u e Mounta ins of Washington and Oregon. I n : J. W. Thomas, ed. W i l d l i f e h a b i t a t i n managed f o r e s t s - The B lue M o u n t z n s o f Oregon and Washington. Pac. Nor thwest For. and Range Exp. Stn., U.S. Dept. Agr ic . , P o r t l a n d , Oreg. ( i n p r e s s ) .
Washington Department o f Game. 1982. F i s h and W i l d l i f e Resource S t u d i e s , S u l t a n R i v e r P r o j e c t Stage I 1 F i n a l Repor t . Snohomish County PUD No. 1 217 pp.
APPENDIX A
Word m o d e l s u s e d i n o r i g i n a l HEP S t u d y (WDG 1982)
DCYGLAS SQUIRREL Coniferous F o r e s t s
General The Gouglas s q u i r r e l ( T a x i a s c i u r s d o u o l a s i i ) i s p r i m a r i l y a s s o c i a t e d
w i t h D o u g l a s - f i r (Pseudotsug? m e n z i & s i i ) ~ f o r e s t s , a l though i t a l s o occurs i n mixed s t a n d s o f c o n i f e r o u s t r e e s c o n i f e r s u s and deciduous t r e e s ( B a i l e y 1 9 3 6 ; I n g l e s 1965) . The most i m p o r t a n t f a c t o r i n f l u e n c i n g t h e use o f h a b i t a t by Douglas s q u i r r e l s a p p e a r s t o be t h e p roduc t ion of cones by c o n i f e r o u s t r e e s (Smith 1965, 1968) .
Food Requirements Smith (1965, 1968) provided a d e t a i l e d 1 i s t o f food e a t e n o r s t o r e d
by Douglas s q u i r r e l s a n d ' r e d s q u i r r e l s , and concluded t h a t bo th s p e c i e s a r e adap ted t o an e n e r g y - r i c h d i e t . The ma jo r food o f Douglas s q u i r r e l s is s e e d s o f c o n i f e r s , e s p e c i a l l y o f D o u g l a s - f i r , S i t k a sp ruce (P* & h e n s i s ) , henlock (T- s p p . ) , f i r s (Abies - s p p . ) , and lodgepo le p i n e (2- contorts) ( B a i l e y 1936; Smith 1965 , 1968) . Con i fe r p o l l e n , b i r c h ( m a s p p ) and Alde r (Alnus - s p p . ) c a t k i n s , ~ a p l e , (&r s p p . ) samaras , n u t s , a c o r n s , some s o f t f r u i t s , buds and bark a r e a l s o e a t e n ( p a i l e y 1936; Smith !965), a s a r e mushrooms and hypogeous f u n g i (Smith 1965; Maser e t a 1 1979; Fogel and Trappe 1978). During t h e l a t e sumzer and i n f a l l , t h e s q u i r r e l s s t o r e thousands of cones , n u t s and c a t k i n s i n food c a c h e s . These food s t o r e s a r e e x p l o i t e d d ~ r i n g t h e w i n t e r znd , i n t h e e v e n t o f a cone c rop f a i l u r e , d u r i n g t h e f o l l o w i n g su ,m?r (Smith 1965, l ? 6 8 ) .
K a t e r Requirements Free w 6 t e r m y o r may not be 25uneafit i n t h e h a b i t a t of Douglas
s q u i r r e l s . Smith 11955, i 9 6 8 ) s t s t e d t h a t tb,e s q u i r r e l s seldom drank wa te r when i t was p r e s e n t ; however, he no ted t h a t t h e f u n g i e a t e n by t h e s q u i r r e l s a r e an impor tan t source o f preformed w a t e r i n s u m e r , and concluded t h a t most o f t h e w a t e r requirements of t h e s q u i r r e l s a r e met by p r e f o n e d w z t e r .
Cover R e q u i r e r e n t s -- Douglas s q u i r r e l s s h e l t e r i n c a v i t i e s i n t r e e s , and sometimes i n
openings between rocks on t a l u s s l o p e s ( I n g l e s 1965) . Globu la r n e s t s may be b u i l t i n t h e crowns o f t r e e s , a n d , a t l e a s t i n w i n t e r , t h e food caches themselves may be used f o r s h e l t e r .
Smith (1965, 1968) p resen ted d a t a showing a r e l a t i o n s h i p between numbsrs o f c o n i f e r o u s t r e e s over 20 i n c h e s i n c i r cumference and cone c r o p s on s q u i r r e l t e r r i t o r i e s , b u t t h i s r e l a t i o n s h i p was addressed t o food supp ly r a t h e r than d i r e c t l y t o c o v e r . Douglas s q u i r r e l s , hor:ever, a r e tho rough ly a r b o r e a l ; n o t only do t h e y n e s t and f o r a g e i n ma tu re c o n i f e r o u s t r e e s , but a l s o t r z v e l on and i n them whenever p o s s i b l e .
Reoroduct ive R e a u i r e x n t s Smith (1965, 1568) r e p o r t e d t h a t o c c u r r e n c e and t iming of b reed inq
and s i z e o f l i t t e r s va ry d i r e c t l y i n respon:e t o changes i n p r o d u c t i o n - o f c o n i f e r seed c rops a v a i l a b l e t o Doug;as s q u y r r e l s . L i t t e r s a r e produced a t a t ime t h a t would p rov ide mothe r s w i t h a l a r g e food supp ly n e a r t h e end o f l a c t a t i o n . Smith s u g g e s t e d t h a t t h e s q u i r r e l s may be a b l e t o a s s e s s , be fo re b reed ing , t h e s u c c e s s of c o n i f e r p o l l i n a t i o n i n a g iven y e z r , and a l s o how much cached food i s a v a i l a b l e t o s u p p o r t a s p r i n g l i t t e r . ? j e s t s a r e u s u a l l y in c a v i t i e s a t l e a s t 1 5 f e e t (4.5m) above t h e ground i n t r e e s t h a t a r e a t l e a s t 12 i n c h e s (30.5crn) dbh (Thomas e t a l . i n p r e s s ) .
' - h i t ? t R ~ c u i r q e n & Douglas s q u i r r e l body s i z e , jaw m u s c u l a t u r e , anatomy, *nd r e p r o d u c t i v e
a c t i v i t y appear t o be adap ted t o l i f e i n f o r e s t s where Doug las - f i r t r e e s p reZcn ina te ( h i t h (1965, 1968) .
P e o u i r f v n t s Douglas s q u i r r e l s of both s e x e s a r e t e r r i t o r i a l th rouahou t t h e y e a r ;
t e r r i t o r i e s vary i n s i z e from abou t 1 a c r e t o more than 2 a c r e s [Smith 1965, 1963). The d i s t r i b u t i o n o f food and t ime of y e a r r e g u l a t e how t e r r i t o r i e s a r e e s t t b l i s h e d {Smiih 1965, 1 9 6 8 ) .
Soec ia l C o n s i d s r a t i o n s Smith (1565, 1558) sho ied t h a t Douglas s q u i r r e l s and t h e c l o s e l y
r e l z t e d red s q u i r r e l s depend h e a v i l y on t h e 2 v a i l a b l e c r o p of c o n i f e r seeds f o r o v e i - w i n t e r s u r v i v a l and ?cr re;roduc:ion i n t h e f o l l c w i n g s p r i n g . The Douglas s q u i r r e l i s p a r t i c u l a r l y d i p e n d e n t on t h e cones of D o u g l a s - f i r t r e e s , which a r e r e l a t i v e l y e a s y t o open compared t o t h e cones of such c o n i f e r s a s lodgepole p i n e . Smith (1965, 1368) shcwed t h a t a l a c t a t i n g r e d s q u i r r e l e x t r a c t e d ene rgy from D o u g l a s - f i r cones f o u r t imes f a s t e r than she d id f r c v lodge?o le p i n e c o n e s , and t h s t red s q u i r r e l s had s i g n i - f i c a n t l y sa211er l i t t e r s i n y e a r s i n which D o u g l a s - f i r ccnes were n o t a v a i l z b l e dur ing l a c t a t i o n . S i n c e 3oug l?s s q u i r r e l s can e x p l o i t a l t e r n a - t i v e cone s o u r c s s even l e s s e f f e c t i v e l y than can red s q u i r r e l s , t h e i r l i f e h i s t o r y i s a p p a r e n t l y d i r e c t l y d e p e n d ~ n t on the p resence o f Douglas- f i r t r e e s o f s u i t a b l e age and f e c u n d i t y .
Although Douglas s q u i r r e l s f e e d p r i m a r i l y on seeds o f c o n i f e r s , t h e y m y a l s o f eed on t h e t e r ~ n i n a l and l a t e r a l buds o f Douglas f i r , e s p e c i a l l y dur ing w i n t e r o r e a r l y s p r i n g ( F i s i h and Dimock 1978) . The pe rcen tage o f t r e e s a f f e c t e d by shoo t c l i p p i n g by s q u i r r e l s ranged from 1-38% bu t r e s u l t i n g h e i g h t l o s s was found t o be temporary and minimal.
V 1 Average dbh of trees
A - ,- L, - 2,
V2 Stand composition
A-- NO Douglas-fir present 0- - Douglas-fir occurs in small
( < 1 acre) scattered stands C-- Stand composed of 50%
Douglas-fir D-- Pure Douglas-fir stand
Food Value ( X I ) = ( ~ 1 x V 2 ) 1 /2
Where: \rl = Suitability Index (SI) of average dbh of trees.
V2 = SI of stand composition.
Reoroductive Value (X2) = v1
Where: V1 = SI of average dbh of trees.
The Habitat Suitability Index i s the lowest Xn value.
June 1978
REFE?.C:XFS C I T E D
h i l e y , V . l 936 . The m?rmals and l i f e zones o f Oregon. PI. Amer. Fauna No. 55.
F i s c h , G . G . , and E . J . Dinnock 11. 1978. Shoot c l i p p i n g by Douglas s q u i r r e l s i n r e g e n e r a t i n g Douglas f i r . J . Wild1 . Manage. 42 (2 ) :414-418.
Foge l , R. and J. M. Trappe. 1978. Fungus consumption (mycophagy) by smal l a n i m a l s . Northwest S c i . 5 2 ( 1 ) : 1-31.
I n g l e s , L. G. 1965. Marnzals o f t h e P a c i f i c S t a t e s . S t a n f o r d Univ. P r e s s , . S i z n f o r d , C a l i f o r n i a .
I.laser, C . , J . M . Trappe and R. A . ilussSaum. 1978. Fungal-small mama1 i n t e r r e l a t i o n s h i p s wi th emphasis on Oregon c o n i f e r o u s f o r e s t s . Ecology ( i n p r e s s ) .
Smith , C. C. 1905. I n t e r s p e c i f i c c o m p e t i t i o n i n t h e genus of t r e e s q u i r r e l s Tamiasc iu rus . Ph.0. t h e s i s , Univ. o f Uashington, S e a t t l e .
- . 1968. The a d a p t i v e n a t u r e o f s o c i a l o r g a n i z a t i o n i n t h e genus o f t r e e s q u i r r e l s T a n i a s c i u r u s . Ecol . Monogr. 38:31-63.
Thomas, J . W . , R . hnderson, C . Ease r , and E. L . B u l l . Dead t r e c . ("snag") r e ' q u i r e r e n t s f o r dependent w i l d l i f e s p e c i e s i n t h e Blue ,!4ountains o f Washington and Oregon. In J . W . Thomas, ed . W i l d l i f e h a b i t a t i n rnana~ed f o r e s t s - The ~ l u e ~ o u n t a i n s o f Oregon and Washington. Pac. Nor thwest F3r. and Rjnge E x p . S t n . , U S . Dept. A g r i c . , P o r t l a n d , Oreg. ( i n p r e s s ) .
Common Merganser
(Mergus merganser)
General
r The comnon merganser (Mergus merganser) i s found th roughou t t h e y e a r i n - western Washington. Approx imate ly 300 p a i r breed i n wes te rn Washington w h i l e 2,300 i n d i v i d u a l s w i n t e r i n t h e Puget Sound r e g i o n ( B e l l r o s e 1976).
Food Requi rements
r Many k i n d s o f f i s h , e s p e c i a l l y j u v e n i l e salmon and t r o u t (Wood 1986).
r A l s o mussels, f r o g s , smal l e e l s , salamanders, c r a w f i s h , o t h e r s m a l l c rus taceans , leeches, worms, a q u a t i c i n s e c t s and l a r v a e , and t h e stems o f r o o t s of a q u a t i c p l a n t s (Bent 1923, T e r r e s 1980).
r Swims on s u r f a c e search ing v i s u a l l y , t h e n d i v e s and ca tches f i s h by d i r e c t p e r s u i t ; u s u a l l y i n c l e a r , s h a l l o w wa te r 1 t o 6 f t deep ( B e l l r o s e 1976).
r Probes among submerged r o c k s t o f l u s h p r e y (Bent 1923).
Water Requirements
a S p r i n g and summer - use l a k e s and r i v e r s bordered by s u i t a b l e h a b i t a t f o r n e s t i n g ( B e l l r o s e 1976).
r Winter - use s a l t water , a l s o low land l a k e s and r i v e r s t o some e x t e n t (Be1 1 rose 1976).
r Must have c l e a r water . Mated p a i r s w i l l abandon streams t h a t become t u r b i d (Be1 l r o s e 1976).
r The c r e a t i o n o f r e s e r v o i r s has caused an i n c r e a s e i n w i n t e r i n g popu la - t i o n s i n many areas ( B e l l r o s e 1976).
Cover Requirements
r T y p i c a l l y use mature and o l d g rowth dec iduous r i p a r i a n , mixed c o n i f e r , and c o n i f e r o u s f o r e s t s when l o c a t e d a l o n g l a k e s o r st reams (Brown 1985).
Reproduc t i ve Requirements
a P r e f e r n e s t i n g a long m i d - t o - h i g h e l e v a t i o n l a k e s and streams i n wes te rn Washington ( L a r r i s o n and Sonnenberg 1968).
Secondary c a v i t y n e s t e r - p r i m a r i l y n e s t s i n c a v i t y - b e a r i n g t r e e s near water , a l s o n e s t s i n caves, c l i f f s , r o o t t a n g l e s , w e l l h idden l o c a t i o n s on t h e ground, i n n e s t boxes and b u i l d i n g s . P r e f e r u s i n g abandoned p i l e a t e d woodpecker c a v i t i e s (Brown 1985).
Nest h e i g h t ranges f rom 0-200 f t . N e s t i n g may occur as f a r as 575 f t . from water (Be1 1 r o s e 1976).
N e s t i n g occurs between May and m i d - J u l y ( B e l l r o s e 1976).
The same n e s t i n g c a v i t y may be used y e a r a f t e r year .
R i v e r s and l a k e s a r e p r e f e r r e d b r o o d i n g areas (Wood 1986).
I n t e r s p e r s i o n Requirements
0 R i v e r s and lakes must be bordered by s u i t a b l e n e s t i n g h a b i t a t (snags dead down m a t e r i a l ) i n s p r i n g .
0 Home ranges a re approx ima te ly 2-3 r i v e r m i l e s (Brown 1985).
REFERENCES CITED
BEAK. 1975. N o r t h Fork Snoqualmie R i v e r V e g e t a t i o n and W i l d l i f e Study, F i n a l Report on e x i s t i n g c o n d i t i o n s and impacts. Beak C o n s u l t a n t s I n c o r p o r a t e d , P o r t l a n d , Oregon. 93 pp.
B e l l r o s e , F. C. 1976. Ducks, geese, and swans o f N o r t h America. W i l d l i f e Management I n s t i t u t e . 535 pp.
Bent, A. C. 1923. W i l d f o w l . P a r t One, Smi thson ian I n s t i t u t i o n U n i t e d S t a t e s N a t i o n a l Museum B u l l e t i n . 126 pp.
Brown, E. R. (ed.) 1985. Management o f w i l d l i f e and f i s h h a b i t a t s i n f o r e s t s o f western Oregon and Washington. P a r t 2. U.S. Dept. A g r i c . , F o r e s t Serv i ce , P a c i f i c Nor thwest Region. 301 pp.
L a r r i s o n , E . J. and K. G. Sonnenberg. 1968. Washington b i r d s - t h e i r i d e n t i f i c a t i o n and l o c a t i o n . S e a t t l e Audubon S o c i e t y .
T e r r e s , J. K. 1980. The Audubon S o c i e t y Encyc loped ia o f N o r t h American B i r d s . A l f r e d A. Knopf, I nc . 1109 pp.
Washington Department o f Game. 1982. F i s h and W i l d l i f e Resource S t u d i e s , S u l t a n R i v e r P r o j e c t Stage I 1 F i n a l Report . Snohomish County PUD No. 1. 217 pp.
Wood, C. C. 19d6. D i s p e r s i o n o f common merganser (Mergus merganser ) b r e e d i n g i n r e l a t i o n t o t h e a v a i l a b i l i t y o f j u v e n i l e P a c i f i c salmon i n Vancouver I s l a n d streams. Can. J. 2001. 64:756-765.
APPENDIX A
S u i t a b i l i t y i n d i c e s and a g g r e g a t i o n f o r m u l a o b t a i n e d f rom t h e N o r t h Fork Snoqualmie Study ( B E A K 1985).
Water d e p t h ( V l ) e r c e n t o f s t ream o r l a k e w i t h
h i d i n g cover f o r a q u a t i c i n v e r - ~ i v e r / ~ t r e a m \ /OodT t e b r a t e s and v e r t e b r a t e s (V2) Wet1 ands
Cover Age o f a d j a c e n t f o r e s t (V3)
~ a k e l ~ e s e r v o i r ) - Reproduc t ion - Number o f a v a i l a b l e n e s t s i t e s (V4)
The H a b i t a t S u i t a b i l i t y Index i s : V 1 + V2 + V j + Vq
A P P E N D I X B
Word model used i n o r i g i n a l HEP S t u d y (WDG 1982)
Ma1 1 a r d
(Anas p l a t y r h y n c h o s )
General
r The m a l l a r d , (Anas - p l a t y r h y n c h o s ) i s a widespread f r e s h w a t e r w a t e r f o w l spec ies , comnon th roughou t t h e y e a r i n western Washington.
e Approx imate ly 50,000 m a l l a r d s w i n t e r i n t h e Puget Sound reg ion .
Food Requirements
0 P l a n t s compr ise 90 p e r c e n t o f d i e t . P l a n t s eaten i n c l u d e sedges, pondweed, duckweed, and many k i n d s o f seeds ( M a r t i n -- e t a l . 1951, Pehrsson, 1984).
0 J u v e n i l e s l e s s t h a n t h r e e weeks o l d feed p r i m a r i l y on animal m a t t e r . Aqua t i c b e e t l e s , l a r v a e and nymphs o f m a y f l i e s , s t o n e f l i e s , c a d d i s f l i e s , d r a g o n f l i e s , d a m s e l f l i e s , t a d p o l e s , f i s h f r y , and c rus taceans a r e impor- t a n t animal food (Bent 1923, Yocum 1957).
0 Seeds a r e i m p o r t a n t food sources d u r i n g s p r i n g m i g r a t i o n and d u r i n g brood r e a r i n g (Krapu 1981).
0 Food i s p i c k e d up f rom t h e s u r f a c e o r j u s t below t h e s u r f a c e .
Must have open, s h a l l o w water areas f o r f e e d i n g (Johnsgard 1975).
0 G r a i n crops ( i n c l u d i n g co rn , wheat, and b a r l e y ) and p a s t u r e s a r e h e a v i l y u t i l i z e d i n w i n t e r (Yocum 1957).
Water/Cover Requirements
0 Found i n f r e s h w a t e r , p r e f e r s loughs, ponds, marshes, s low-moving streams and r i v e r s and swamps (Johnsgard 1975).
Long nar row s loughs, f l o a t i n g i s l a n d s , and g r a d u a l l y s l o p i n g shore1 i n e s a r e used f o r l o a f i n g ( G i r a r d 1941). Areas w i t h dense v e g e t a t i o n a r e avo ided f o r l o a f i n g (Sowls 1955).
Seasonal wet lands a r e p r e f e r r e d f e e d i n g h a b i t a t by l a y i n g hens (Krapu - e t a l . 1983; Cowardin e t a l . 1985 - --
0 R e s e r v o i r s a r e u t i l i z e d d u r i n g w i n t e r months and d u r i n g m i g r a t i o n p e r i o d s i n wes te rn Washington. (WDG 1982, W a l t e r s 1986).
0 Broods u t i l i z e wet lands hav ing sparse t o dense emergent v e g e t a t i o n and open water. Wetlands w i t h o u t emergent v e g e t a t i o n o r open water a r e u s u a l l y avoided. S h o r e l i n e s ba re o f emergent v e g e t a t i o n a r e seldom used (Berg 1956, Rumble and F l a k e 1983).
r Art i f i c i a 1 i s 1 ands and env i ronments w i t h reduced numbers o f p r e d a t o r s s i g n i f i c a n t l y i n c r e a s e n e s t i n g d e n s i t i e s and h a t c h i n g success (Duebber t and Lokemoen 1980).
Reproduc t i ve Requirements
a N e s t i n g d e n s i t y ( p a i r s / a c r e ) i s h i g h e r i n seasonal we t lands t h a n deep marshes and permanent water areas (Duebber t e t a l . 1983). Seasonal wet1 ands w i t h o u t f i s h p o p u l a t i o n s p r o v i d e E g F - s i z e d i n s e c t s and g r e a t e r o v e r a l l i n s e c t abundance (Pehrsson 1984).
r Deep marshes and permanent wa te r areas a r e used f o r r e a r i n g o f young (Duebbert -- e t a l . 1983). These areas a r e p r e f e r r e d d u r i n g m i g r a t i o n and r e a r i n g , when v e g e t a t i o n and seeds a r e t h e p r i m a r y f o o d source (Pehrsson 1984).
a Beaver impoundments o f t e n c r e a t e s u i t a b l e n e s t i n g h a b i t a t (Beard 1953).
r Nests a r e p l a c e d i n r e l a t i v e l y t a l l herbaceous v e g e t a t i o n i n c l o s e p r o x i - m i t y t o water . V e g e t a t i o n h e i g h t v a r i e s between 8 i n . and 30 i n . i n n e s t i n g areas. Most n e s t s a r e w i t h i n 300 f t . o f wa te r ( B e l l r o s e 1976, Lokemoen e t a ] . 1984). --
a Mature, r e l a t i v e l y dense grasses, rushes, and shrubs a r e p r e f e r r e d f o r n e s t i n g . I n areas w i t h h i g h b r e e d i n g d e n s i t i e s , t h i c k e r , dense shrub v e g e t a t i o n ( n e a r l y 100 p e r c e n t v i s u a l o b s c u r i t y ) i s p r e f e r r e d (Lokemoen e t a l . 1984). --
I n t e r s p e r s i o n Requirements
a Home ranges d u r i n g t h e h r e e d i n g season can be as l a r g e as 700 ac res (Brown 1985).
0 Home ranges o f t e n o v e r l a p (Lokemoen e t a l . 1984), though p a i r s may defend n e s t i n g ponds a g a i n s t o t h e r ma1 1 a r d s T D z b i n 1969).
a A minimum o f t h r e e acres o f n e s t i n g and r e a r i n g we t land h a b i t a t i s needed w i t h i n hen b r e e d i n g home range t o s u p p o r t one m a l l a r d p a i r . Maximum p r o - d u c t i o n w i l l r e s u l t f rom t h e p r o x i m i t y and i n t e r s p e r s i o n o f n e s t i n g and r e a r i n g h a b i t a t (Dzub in 1969).
a Deep marshes shou ld be w i t h i n a 1 m i l e r a d i u s o f s h a l l o w marshes (Jahn and Hunt 1964).
REFERENCES CITED
Beard, E. B. 1953. The impor tance o f beaver i n w a t e r f o w l management a t t h e Senay N a t i o n a l W i d l l i f e Refuge. J. W i l d l . Manage. 17(4):398-436.
B e l l r o s e , F. C. 1976. Ducks, geese, and swans o f N o r t h America. The S tackpo le Co., H a r r i s b u r g , PA., and t h e W i l d l i f e Management I n s t i t u t e , Washington D. C. 535 pp.
Bent, A. C. 1923. Wi ld fow l . Smi thson ian I n s t i t u t i o n U n i t e d S t a t e s N a t i o n a l Museum B u l l e t i n . 126 pp.
Berg, P. F. 1956. A study o f wa te r fow l broods i n e a s t e r n Montana w i t h spe- c i a l r e f e r e n c e t o movements and t h e r e l a t i o n s h i p o f r e s e r v o i r f e n c i n g t o p r o - d u c t i o n . J. W i l d l . Manage. 20(3):253-262.
Brown, E. R. (ed. ) , 1985. Management o f w i l d l i f e and f i s h h a b i t a t s i n f o r e s t s o f wes te rn Oregon and Washington. P a r t 2. U.S. Dept. Agr ic . , F o r e s t Serv i ce , P a c i f i c Region. 332 pp.
Cowardin, L. M., D. S. G i lmer and C. W. S h a i f f e r . 1985. M a l l a r d r e c r u i t m e n t i n t h e a g r i c u l t u r a l env i ronment o f N o r t h Dakota. W i l d l . Monog. 92. 36 pp.
Duebbert , H. F. and J. T. Lokemoen. 1980. H igh duck n e s t i n g success i n a p reda to r - reduced environment. J. W i ld l . Manage. 4 4 ( 2 ) : 428-437.
Duebbert , H. F., J. T. Lokemoen and D. E. Sharp. 1983. Concent ra ted n e s t i n g o f m a l l a r d s and gadwal ls on M i l l e r Lake I s l a n d , N o r t h Dakota. J. W i l d l . Manage. 47(3):729-740.
Ozubin, A. 1969. Comments on c a r r y i n g c a p a c i t y o f sma l l ponds f o r ducks and p o s s i b l e e f f e c t s o f d e n s i t y on m a l l a r d p r o d u c t i o n . pages 239-268. I n R a t t i , J. T., F lake, L. D., and W. A l l a n Wentz, (eds.). Waterfowl eco fogy and management: s e l e c t e d read ings. The W i l d l i f e S o c i e t y . 1982.
G i r a r d , G. L. 1941. The m a l l a r d : I t s management i n wes te rn Montana. J. W i l d l . Manage. 5(3):233-259.
Jahn, L. R., and R. A. Hunt. 1964. Duck and c o o t eco logy and management i n Wisconsin. Wisc. Conserv. Dept. Tech. B u l l . 33. 212 pp.
Johnsgard, P. A. 1975. Waterfowl o f Nor th America. I n d i a n a Univ. Press. B loomington and London. pp. 221-223.
Krapu, G. L. 1981. The r o l e o f n u t r i e n t rese rves i n m a l l a r d r e p r o d u c t i o n . Auk 98: 29-38.
Krapu, G. L., A. T. K l e t t and D. G. Jorde. 1983. The e f f e c t o f v a r i a b l e s p r i n g wa te r c o n d i t i o n s on m a l l a r d r e p r o d u c t i o n . Auk 100:689-698.
Lokemoen, J. T., H. F. Duebbert and D. E. Sharp. 1984. Nest spac ing, h a b i t a t s e l e c t i o n and behav io r o f wa te r fow l on M i l l e r Lake I s l a n d . N o r t h Dakota. J. W i l d l . Manage 48(2):309-321.
M a r t i n , A. C., H. S. Z i m , and A. C. Nelson. 1951. American w i l d l i f e and p l a n t s - a g u i d e t o w i l d l i f e food h a b i t s . Dover Publ . I nc . N.Y.
Pehrsson, 0. 1984. R e l a t i o n s h i p s o f food t o s p a t i a l and tempora l b r e e d i n g s t r a t e g i e s o f m a l l a r d s i n Sweden. J. W i l d l . Manage. 48(2):322-340.
Rumble, M. A. and L. D. F lake . 1983. Management c o n s i d e r a t i o n s t o enhance use o f s t o c k ponds by w a t e r f o w l broods. J. Range Manage. 36 (6 ) :691-694.
Sowls, L. K. 1955. P r a i r i e ducks. The S t a c k p o l e Co., H a r r i s b u r g , Pa., and t h e W i l d l . Manage. I n s t i t . , Wash. D.C. 193 pp.
U.S. F i s h and W i l d l i f e S e r v i c e . 1985. D r a f t h a b i t a t s u i t a b i l i t y i n d e x models - m a l l a r d ( w i n t e r i n g ) . D i v i s i o n o f E c o l o g i c a l Serv i ces . Sacramento, C a l i f o r n i a 5 PP.
Wa l te rs , M. T e r r e s t r i a l b i o l o g i s t , Puget Sound Power and L i g h t Co., Persona l communication w i t h Dave Hays, J u l y 1986.
Washington Dept. o f Game. 1982. F i s h and w i l d l i f e resource s t u d i e s - S u l t a n R i v e r p r o j e c t s t a g e I 1 f i n a l r e p o r t . Snohomish County PUD No. 1. 219 pp.
Yocum, C. F. 1957. Waterfowl and t h e i r food p l a n t s i n Washington. Univ. Wash. Press. S e a t t l e . 272 pp.
APPENDIX A
S u i t a b i l i t y i n d e x o b t a i n e d f rom t h e USFWS d r a f t H a b i t a t S u i t a b i l i t y Index Models - m a l l a r d (USFWS 1985).
Beaver
( C a s t o r canadens is)
General
r A common mammal i n t h e western Washington low lands where numerous watercourses p r o v i d e s u i t a b l e h a b i t a t (Da lques t 1948).
0 F a c t o r s a f f e c t i n g range p o t e n t i a l i n c l u d e topography, s t ream g r a d i e n t , adequate water , abundance o f . f o o d spec ies and l a n d use ( D i c k i n s o n 1971; W i l l i a m s 1961).
Food Requirements
r D i e t i n c l u d e s a wide v a r i e t y o f a q u a t i c and emergent v e g e t a t i o n , f o r b s , f e r n s , shrubs, and t r e e s . P r e f e r r e d spec ies a r e w i l l o w ( S a l i x spp.), cot tonwood (Populus spp.), a l d e r (A lnus spp.), maple (Acer spp.)and ash ( F r a x i n u s spp.)(Denny 1950).
-
0 I n st reams which f l o w west f rom t h e Cascades t o Puget Sound beavers a t e a v a r i e t y o f spec ies o f w i l l o w as t h e i r p r i n c i p a l f o o d ( D a l q u e s t 1948).
0 A l s o e a t o t h e r p l a n t spec ies such as a l d e r , cascara (Rhamnus p u r s h i a n a ) , Douglas f i r (Pseudotsuga m e n z i e s i i ) , r e d h u c k l e b e r r y c c i n i u m - p a r - v i f o l i u m ) and salmonberry (Rubus s p e c t a b i l i s ) i n l e s s e r a m m l q u e s t 194e).
0 Water l i l i e s (Nymphaea spp.) w i t h t h i c k , f l e s h y r h i z o n e s p r o v i d e a good w i n t e r food s o u i n k i n s 1981).
r M a j o r i t y o f f o r a g i n g occurs w i t h i n 300 f t . o f t h e w a t e r ' s edge, a l t h o u g h can f o r a g e w i t h i n 650 f t . o f wa te r ( A l l e n 1982).
r The dbh o f t r e e s shou ld range f rom 1 t o 6 i n . ( A l l e n 19821, b u t p r e f e r t r e e s 3 t o 4 i n . dbh ( B r a d t 1947; Hodgdon and Hunt 1953; Long ley and Moyle 1963; N ixon and E l y 1969).
r Tree and/or shrubs canopy c l o s u r e between 40 and 60% i s an i n d i c a t i o n o f optimum food a v a i l a b i l i t y ( A l l e n 1982).
Water Requirements
r Requ i re permanent supp ly o f wa te r ( A l l e n 1982).
r S t a b l e wa te r l e v e l s a r e o p t i m a l . Large r i v e r s and l a k e s where w a t e r d e p t h and/or f l u c t u a t i o n cannot be c o n t r o l l e d a r e l e s s s u i t a b l e (Murray 1961; Slough and S a d l i e v 1977).
r Stream g r a d i e n t s o f l e s s and 6 p e r c e n t a r e o p t i m a l ( R e t z e r - e t - a l . 1956).
Cover Requirements
r Most d i g bank burrows i n deep and wide wa te r courses ( D a l q u e s t 1948; Kebbe 1978; T h o r i l e y 1978).
0 Dams a r e b u i l t i n s h a l l o w e r l ow g r a d i e n t moun ta in streams (WOG 1982).
r A c t i v e l y e r o d i n g s t ream banks, rocky channels and sandy s o i l s a r e un fa - v o r a b l e h a b i t a t ( R e t z e r -- e t a l . 1956, Henderson 1960).
r On lakes and ponds, l odges a r e f r e q u e n t l y s i t u a t e d i n areas t h a t p r o v i d e s h e l t e r f rom wind, wave and i c e a c t i o n (e.g., c o n v o l u t e d s h o r e l i n e s ) (A1 l e n 1982).
Reproduc t i ve Requirements
r See cover requ i rements .
Spec ia l H a b i t a t Requirements
r A s s o c i a t i o n o f wide v a l l e y s w i t h loamy s o i l s and streams o f l ow g r a d i e n t i s f a v o r a b l e t o beaver occupancy ( H a l l 1960).
r Rapid water l e v e l f l u c t u a t i o n s t h a t exposed o r f l o o d e d r i v e r hank dens a r e d e t r i m e n t a l t o beaver s u r v i v a l ( C l a i r e -- e t a l . 1971).
r Frequent smal l f l u c t u a t i o n s had l i t t l e adverse e f f e c t s ( C l a i r e -- e t a l . 1971).
I n t e r s p e r s i o n Requirements
r I n t e r s p e r s i o n o f food and water i s i m p o r t a n t . C h i e f l i m i t i n g f a c t o r i n beaver c a r r y i n g c a p a c i t y was t h e amount o f a v a i l a b l e food (MacDonald 1956).
0 U s u a l l y t r a v e l up t o 300 f t . f o r food, b u t maximum d i s t a n c e reco rded was 656 f t . ( B r a d t 1936, 1947, Hodgdon and Hunt 1953).
r 90% o f t r e e c u t t i n g o c c u r r e d w i t h i n 100 f t . (30.5 m) o f t h e s t ream ( H a l l 1960).
S p e c i a l C o n s i d e r a t i o n s
r W i l l l i v e i n c l o s e p r o x i m i t y t o man i f h a b i t a t requ i rements a r e s a t i s f i e d (Henderson 1960).
REFERENCES CITED
A l l e n , A. W. 1982. H a b i t a t s u i t a b i l i t y i ndex models: Beaver. U.S. Dept. I n t . , F i s h W i l d l . Serv. FWSJOBS-82/10.30. 20 pp.
B r a d t , G. W. 1938. A s t u d y o f beaver c o l o n i e s i n Mich igan. J. Mammal. 19:139-162.
. 1947. M ich igan beaver management. M ich igan Department Cons., Game Div., 56 pp.
Beard, E. B. 1953. The impor tance o f beaver i n w a t e r f o w l management a t t h e Seney N a t i o n a l W i l d l i f e Refuge. J. W i l d l . Manage. 17:398-436.
Berghofer , C . B. 1961. Movement o f beaver. Proc. 4 1 s t Ann. Conf. W. Assoc. S t a t e Game F i s h Comm. 41:181-184.
C l a i r e , E. W., H. W . S c o t t , and D. E. Sanford. 1971. The p o t e n t i a l impact o f severe water l e v e l f l u c t u a t i o n s on w i l d l i f e resources o f l o w e r Columbia R i v e r . Oregon S t a t e Game Commission, P o r t l a n d . 18 pp.
Da lquest , W. W. 1948. Mammals o f Washington. Un iv . Kansas Pub. Mus. Nat. H i s t . No. 2, 444 pp., 140 f i g s .
Denny, R. N. 1950. Program f o r heaver management and a d m i n i s t r a t i o n i n Colorado. M.S. Thes is . Colo. A&M Co l lege .
D ick inson , N. 0 . 1971. A e r i a l photography as an a i d i n beaver management. N.Y. F i s h & Game J. 18(1) :57-61.
H a l l , J. G. 1960. W i l l o w and aspen i n t h e eco logy o f beaver on Sagehen Creek, C a l i f o r n i a . Eco l . 41(3):484-494.
Henderson F. R. 1960. Beaver i n Kansas. Un iv . Kansas Mus. Nat. H i s t . Misc. Publ . No. 26. Lawrence. 85 pp.
Hodgdon, K. W., and J. H. Hunt. 1953. Beaver management i n Maine. Maine Dept. I n l a n d F i s h e r i e s and Game, Game Div . B u l l . No. 3, 102 pp.
I n g l e s , L. G. 1965. Mammals o f t h e P a c i f i c S t a t e s . S t a n f o r d Un iv . Press. S t a n f o r d , C a l i f o r n i a . 506 pp.
Jenk ins , 5. H. 1975. Food s e l e c t i o n by beavers m u l t i -d imens iona l con t ingency t a b l e a n a l y s i s . Oeco log ia . 21(2):157-173.
Jenk ins , S. H. 1980. A s i z e - d i s t a n c e r e l a t i o n i n food s e l e c t i o n by beavers. Eco logy 61(4):740-746.
,Jenkins, S. H. 1981. Problems, p rog ress , and p rospec ts i n s t u d i e s o f food s e l e c t i o n by beavers. Pages 559-579 i n J. A. Chapman and D. P u r s l e y , eds. Worldwide f u r b e a r e r Conf. Proc., Vol. 1 7
Kebbe, C. 1978. Personal communication. Oregon Dept. o f F i s h and Game. P o r t 1 and, Oregon.
MacDonald, D. 1956. Beaver c a r r y i n g c a p a c i t y o f c e r t a i n mounta in streams i n N o r t h Park, Colorado. M.S. Thes is , Co lorado S t a t e Univ., F o r t C o l l i n s . 136 PP.
N e f f , D. 1957. E c o l o g i c a l e f f e c t s o f beaver h a b i t a t abandonment i n t h e Colorado Rockies. J. W i l d l . Manage. 21:80-84.
Retzer , J. L., H. M. Swope, J. D. Remington, and W. H. R u t h e r f o r d . 1955. S u i t a b i l i t y o f p h y s i c a l f a c t o r s f o r beaver management i n t h e Rocky Mounta ins o f Colorado. Colo. Game and F i s h Dept. Tech. B u l l . No. 2. 33 pp.
R u t h e r f o r d , W. H. 1954. The beaver i n Colorado. Colo. Game, F i s h and Parks Dept., Tech. Publ . No 17. 49 pp.
Thor i l e y , M. 1978. Personal communication. Washington Dept. o f F i s h and W i l d l i f e . Olympia, Wash.
W i l l i a m s , R. M. 1965. Beaver h a b i t a t and management. Idaho W i l d l . Rev. 1 7 ( 4 ) : 3 - 7 .
Washington Department o f Game. 1982. F i s h and W i l d l i f e Resource S t u d i e s , S u l t a n R i v e r P r o j e c t Stage I 1 F i n a l Report . Snohomish County PUD No. 1. 217 pp.
APPENDIX A
S u i t a b i l i t y i n d i c e s and a g g r e g a t i o n f o r m u l a o b t a i n e d f r o m t h e USFWS H a b i t a t S u i t a b i l i t y Index Model - beaver ( A l l e n 1982).
B E A V E R
Water X s t ream g r a d i e n t ( V 7 ) -~verage annual wa te r f l u c t u a t i o n ( V g )
R i v e r / S t r e a m % t r e e canopy c l o s u r e (Vl) % o f t r e e s i n 1 t o 6 i n . ( V 2 )
dbh s i z e c l a s s Win te r Food X o f sh rub crown c l o s u r e ( V 3 )
Average h e i g h t o f sh rub canopy (v4 ) Species c o m p o s i t i o n o f woody
v e g e t a t i o n ( v 5 )
/'water
Average annual w a t e r f l u c t u a t i o n ( v 8 )
/ X t r e e canopy c l o s u r e ( V 1 ) :;;Lands , % o f t r e e s i n 1 t o 6 i n . ( V p )
dbh s i z e c l a s s % o f sh rub crown c l o s u r e ( V 3 )
W i n t e r Food Average h e i g h t o f sh rub canopy ( v 4 ) Species c o m p o s i t i o n o f woody
v e g e t a t i o n ( v 5 ) X o f l a c u s t r i n e s u r f a c e dominated by
y e l l o w and /o r w h i t e w a t e r l i l y ( V 6 )
Lake Water Shore1 i n e development f a c t o r ( v g )
Note: Water p r o v i d e s cover f o r t h e f e e d i n g and r e p r o d u c t i v e a c t i v i t i e s o f t h e beaver.
'J, Pe-cer: o f t r e e s i r 2 . 5 t c 15 2 c r ( 1 t c 6 incnes) 0.1
s i z e c l a s s
B e a v e r
1 2 3 4 (m)
3.3 6.6 9 .9 13.2 ( f t )
v. Average h e i g h t of s h r u b c. spy.
Y, Species csmzzsi:ion o f ~ 3 0 1 ) vejeta+.<on ( t - e e s a?;/or shrubs)
V s Percent o f l a c u r t r i n e sur face da -> r ,a ted by Y E ? ! O * an< '3r w r ~ i i e wa te r l i l y .
5 10 15 20
2 g r a d i e n t
V, Percent stream g r a d i e n t .
U a t e r f l u c t u a t i o n
'J, Average wa te r f l u c - t u a t i o n on annual b a s i s .
A) Small f l u c t u a - t i o n s tb,at have no e f f e c t on b u r r o w o r lodge en t rances .
6) Moderate f l u c - t u a t i o ~ s t h a t a f f e c t bu r row o r l o d c e e n t r a i c e s
C) E x t r m e f l u c t u a - t i o l s 07 wa te r absent d u r l n g P d L l 0 f Yea ' .
1 2 3 4 5
Shore1 i n e development
V, S h o r e ? i n e deve l - opment f a c t o r (see v a r i a b l e d e f i n i t i o n i n F i g u r e 4 ) .
L i f e Cover r e q u i s i t e E q u a t i o n
W i n t e r f o o d Wet1 ands /bog/pond a+b+c 2 . 5
W i n t e r f o o d R i v e r b+c - 1 . 5
W i n t e r f o o d L a k e / R e s e r v o i r - b+c 1 .5 + v 6
whe re : a = woody v e g e t a t i o n v a l u e w i t h i n a c t u a l w e t - l a n d b o u n d c r y . The s u g g e s t e d e q u a t i o n i s :
[ ( V , x v ~ ) ~ ' ~ x v5 ]1 /2 + [ ( V , x v , ) ~ / ~ x V,] 1 / 2
b = woody v e g e t a t i o n v a l u e w i t h i n 100 m ( 3 2 8 f t ) f r o m t h e w a t e r ' s edge . The s u g g e s t e d e q u a t i o n i s :
[(V1 x v2)1'2 x v , ] ~ ' ~ + [ ( V , x v , ) ~ ' ~ x v , ] ~ ' ~
c = woody v e g e t a t i o n v a l u e w i t h i n 100 m ( 3 2 8 f t ) t o 200 m (656 f t ) f r o m t h e w a t e r ' s edge . The s u g g e s t e d e q u a t i o n i s :
0 . 5 [ ( V , x v2) ' " x v,ll'* + [ ( V , x v , ) " ~ x v , ] ~ / ~
K a t e r R i v e r V, o r V,, w h i c h e v e r i s l o w e s t
Wate r L a k e / R e s e r v o i r V, o r V,, w h i c h e v e r i s l o w e s t , i f
l a c u s t r i n e a r e a r 8 ha ( 2 0 a c r e s ) in s u r f a c e a r e a .
V,, i f l a c u s t r i n e a r e a i s < 8 ha ( 2 0 a c r e s )
i n s u r f a c e a r e a .
Wa te r Wet 1 ands /bog/pond v.
H S I d e t e r m i n a t i o n . Based on t h e l i m i t i n g f a c t o r c o n c e p t , t h e H S I i s e q u a l t o t h e l o w e s t l i f e r e q u i s i t e v a l u e o b t a i n e d f o r e i t h e r f o o d o r w a t e r .
P P P E N D I X B
Word model used in original HE? Study (WDG 1982)
June 1978
BEAVER F'parian Comnuni t i e s
C~ner;l , . \ , , The beaver ( C j s t o r canadens i s ) i s a common mammal i n t h e wes te rn
Uash ing ton lowlands where numerous w a t e r c o u r s e s p rov ide s u i t a b l e h a b i t a t ( D a l q u e s t 1948) . I t a l s o occurs th roughou t t h e Wi l l amet t e V a l l e y o f Oregon ( I n g l e s 1965). The fo l lowing f a c t o r s a f f e c t beaver r ange p o t e n t i a l : t opography , s t r eam g r a d i e n t , adequate w a t e r , abundance of food s p e c i e s and l a n d use (Dickinson 1971; Will iams 1961).
Food Resui rements Beavers a r e q e n e r a l i z e d h e r b i v o r e s ( J e n k i n s 1975). T h e i r d i e t
i n c l u d e s a wide v a r i e t y of a q u a t i c and emersen t v e g e t a t i o n , f o r b s , f e r n s , sh rubs and t r e e s . L i s t e d i n o r d e r o f p r e c e r e n c e t h e p r e f e r r e d food s p e c i e s o f beaver i n Colorado a r e ( S a l i x s p p . ) , cottonwood (Po u lus spp ) , s s a n d ash ( F r a x i n u s s ~ h n n y i 9 5 0 ) . I n C a l i f o r n i a , b e a v e r s u t i l i z e d wi l low i n t h e absence o f aspen ( H a l l 1960). R u t h e r f o r d (1954) fouod t h a t , i n Colorado, t h e p r i n c i p a l beav2r food s p e c i e s o f t h e p l a i n s r i v e r b o t t o m h a b i t a t was cottonwood w i t h s p e c i e s o f wi l low a l s o s u p p l y i n g an i n p o r t a n t p a r t of t h e d i e t . A g r i c u l t u r a l c r o p s such a s c o r n and a l f a l f a a r e e a t e n i f a c c e s s i b l e w i t h i n t h e i r c r u i s i n g r a d i u s from wate r . Re tze r e t a l . (1956) found t h a t aspen was t h e most p a l a t a b l e s p e c i e s i n Colorado, a l t h o u g h wi l low was e a t e n i n l a r g e q u a n t i t i e s . Beavers i n h a b i t i n g s t r e a s which f low wes t from t h e Czscades t o t h e Puget Sound a t e a v a r i e t y of s p e c i e s o f wi l low as t k e i r p r i n c i p a l food ( D a l q u e s t 1948) . Othe r b l a n t s p e c i e s such a s a l d e r , c a s c a r a (Rh;nr,us p u r s h i a n a ) , Douglas f i r (Pseudo t suaa m e n z i e s i i ) , red huckl e b e r r y m n ? um p a r v i f o l ium) and s a l n o n b e r r v (Rubus s s e c t z b i l i s ) were e a t e n i n l e s s e r amounts. Beaver showed a G n d e n c v i n ' t h e sumxek t o use both t e r r e s t r i a l and a q u a t i c - i o e c i e s o f herbaceous v e o e t a t i o n . - r .,
E e l g r a s s ( Z o s t e r a mar ina ) , duckweed (Lemna s p p . ) , waterweed (Elodea c a n ; d e i s i s ) , w a t e r l i l y r o o t s (Brasen ia s p x h a r sp ) , c a t t a i l a i l a L l a c k b e r r y (Rubus smdenro&;dabo sp?. , sedge m p . ): b u l r u s h ( S G ; s p p and b u r - r e e d Sparqanlum spp. ) were e a t e n i n Co l - rado (Dennv 1950 The s p r o u t i n q c a p a c i t y of wi l low made i t a d u r a b l e ma ins tay of t h e beaver economy in ~ a l i f o r n i a ; overbrowsed s e c t i o n s r ecovered v i g o r o u s l y and t h e wi l low was u t i l i z e d on a s u s t a i n e d use b a s i s (Hal l 1960).
Water Requirements No d r i n k i n g wa te r requirements were found i n t h e l i t e r a t u r e .
Cover Requirements Most o f t h e beaver s i n t h i s e c o r e g i o n d i g bank burrows r a t h e r than
c o n s t r u c t l o d a e s ( D a l a u e s t 1948; Kebbe 1978; T h o r i l e y 1978). Burrows <
a r e u s u a l l y a s s o c i a t e d w i t h deep w a t e r c o u r s e s . where deep w a t e r i s n o t a v a i l a b l e , a s i n t h e sha l lower mountain s t r e a m s , t h e beaver c o n s t r u c t s dams. The deep pool which forms behind t h e dam p r o v i d e s e s c a p e cover . R ~ t z e r e t a1 ( i 9 5 6 ) found t h a t e r o d i n g , rocky c h a n n e l s were unfavorab le beaver h a b i t a t . Sandy s o i l s may p r e v e n t burrow c o n s t r u c t i o n (Henderson 1960)
/ - June 1978
i II / ( . ReproSuctive Requirements
Cover f o r t h e young i s provided by t h e a d u l t lodge o r burrow (Henderson 1960) K i t s a r e born i n A p r i l and May ( I n g l e s 1965).
Specia l H a b i t a t Requi rements The a s s o c i a t i o n o f wide v a l l e y s w i t h s t reams o f low g r a d i e n t i s
f a v o r a b l e t o b e a v e r occupancy (Hal l 1960). In Colorado, v a l l e y f l o o d p l a i n s t h a t a r e w i d e r t h a n t h e wid th o f t h e channel p rov ide s u i t a b l e beaver h a b i t a t whereas v a l l e y s which a r e o n l y channel wide o r na r rower a r e o n l y marginal ( R e t z e r e t a1 1956). Stream g r a d i e n t s o f l e s s t h a n 6% a r e o p t i c a l . G r a d i e n t s from 7 t o 12% a r e good, and 13 t o 15% are o f q u e s t i o n - a b l e va lue . S t reams wi th g r a d i e n t s g r e a t e r t h a 15% a r e u n s u i t a b l e .
Rapid w a t e r l e v e l f l u c t u a t i o n s t h a t exposed o r f l o o d e d r i v e r bank dens were found t o be d e t r i m e n t a l t o beaver s u r v i v a l ( C l a i r e e t a l . 1971). These f l u c t u a t i o n s a r e most damaging d u r i n g t h e k i t t i n g s e a s o n . F requen t small f l u c t u a t i o n s had l i t t l e adver se e f f e c t s .
I n t e r s p e r s i o n Requirements MacDonald (1956) found t h a t one o f t h e c h i e f l i m i t i n g f a c t o r s i n
beaver c a r r y i n g c a p a c i t y was t h e amount o f a v a i l c b l e food . Most i n v e s t i - o a t o r s have de te rmined t h a t beaver w i l l t r a v e l up t o 300 f e e t (91 .4 m) on :and t o s u i t a b l e food s u p p l i e s ; 656 f e e t (200'm) was t h e maximum obs;rved d i s t a n c e ( B r a d t 1938, 1947; Hodgdon and Hunt 1953). Hal l ( 1 9 6 0 ) r e p o r t e d t h a t 90 p e r c e n t of t r e e c u t t i n g occur red w i t h i n 100 f e e t (30.5 m) o f t h e s t r e a m .
S p e c i a l C o n s i d e r a t i o n s E ~ a v e r s w i l l l i v e i n c l o s e p rox imi tv t o &an i f i t s h a b i t a t r e q u i r e -
ments a r e s a t i s f i e d ( H e ~ d e r s o n 1560). 8ergho;er (1961) s t a t e d t h a t beaver were e x c e l l e n t deve loper s of waterfowl h z b i t a t . Neff (1957) r epor ted t h a t water fowl and muskrat were s i g n i f i c a n t l y more common on beaver occup ied s t r eams . Beaver ponds provided i n t e r s p e r s i o n of food and cover p l a n t s , l o a f i n g s i t e s f o r wa te r fowl , and s h a l l o w w a t e r f o r waterfowl f o r a g i n g (Eeard 1953). Gard (1961) found t h a t b e a v e r ponds produced a h igh s t a n d i n g c rop o f bottom fauna a d a un ique p h y s i c a l h a b i t a t which r e s u l t e d i n g r e a t e r t r o u t p roduc t ion .
REFERENCES C I T E D
Bradt , G . W . 1938. A s t u d y of beaver c o l o n i e s i n Michigan. J. Mammal. 19: 139-162.
. 1947. Michigan beaver management. Michigan Department Cons. , Game Oiv. , 56 pp.
Beard, E . B. 1953. The importance o f beaver i n water fowl management a t t h e Senev Na t iona l W i l d l i f e Refuge. J . Wildl . Manage. 17:398-436.
Berghofer , c . -B . 1961. Movement o f beaver . Proc. 4 1 s t - ~ n n . Conf. W. Assoc. S t a t e Game F i s h Comm. 41:181-184.
C l a i r e , E . W . , H. W. S c o t t , and D. E. Sanford . 1971. The p o t e n t i a l impact of s e v e r e w a t e r l e v e l f l u c t u a t i o n s on w i l d l i f e r e s o u r c e s o f t h e lower Columbia River . Oregon S t a t e Game Commission, P o r t l a n d . l a pp.
June 1978
Da lques t , W. W. 1948. I~lzmnals o f Washington. Univ. of Kansas Pub. Mus. Nat. H i s t . No. 2 , 444 p p . , 140 f i g s .
Denny, R. N . 1950. Program f o r beaver management and a d m i n i s t r a t i o n i n Colorado. M.S. T h e s i s . Colo . A&M Col lege .
Dickinson, N . 8. 1971. A e r i a l photography a s an a i d i n beaver management. N.Y. F i s h & Game J . 18(1) :57-61.
H a l l , J . G . 1960. Willow and aspen i n t h e ecology of beaver on Sagehen Creek, C a l i f o r n i a . Ecol. 41 (3 ) : 484-494.
Henderson, F . R . 1960. Beaver i n Kansas. Univ. Kansas Mus. Nat. Hist. Misc. Publ. No. 26. Lawrence. 85 pp.
Hodgdon, K . W . , and J . H . Hunt. 1953. Beaver management i n Maine. hiaine Dept. In land F i s h e r i e s and Game, Game Oiv. B u i l . No. 3 , 102 pp. -
I n g l e s , L. G. 1965. Mcmmals o f t h e P a c i f i c S t a t e s . S t a n f o r d Univ. P r e s s . S t a n f o r d , C a l i f o r n i a . 506 pp.
J e n k i n s , S . H . 1975. Food s e l e c t i o n by beavers mul t i -d imens iona l con t ingency t a b l e a n a l y s i s . Oeca log ia . 21(2):157-173.
Kebbe, C . 1978. Personal communication. Oregon Cept. of F i s h and Gme. P o r t l a n d , Oregon.
MacDonald, D. 1956. Beaver c a r r y i n g c a c i c i t y of c e r t a i n mountain s t r eams i n North Pa rk , Colorado. M.S. T h e s i s , Colorado S t a t e Univ . , F o r t C o l l i n s . 136 pp.
N e i f , D . 1957. Ecolcgica i e f f e c t s o f bezver h z b i t a t abandonment i n t h e Colorado Rockies. J . Wi ld l . Manage. 21:80-84.
R e t z e r , J . L . , H. M. Swope, J . 0 . R m i n g t o n , and W . H . Ru the r ro rd . 1955. S u i t a b i l i t y o f p h y s i c a l f a c t o r s f o r beaver m a n a ~ e ~ e n t i n t h e Rocky h u n t a i n s of Colorado. Colo. Game and Fish Cept. Tech. B u l l . No. 2. 33 pp.
R u t h e r f o r d , W. H . 1954. The b e a v e r i n Colorado. Colo. Game, F i s h and Parks O e p t . , Tech. Publ . No. 17. 49 pp.
T h o r i l e y , M . 1978. Ps r sona l communication. Washington Dept. o f F i s h and W i l d l i f e . Olympia, Wash.
Wi l l i ams , R. M. 1965. a e a v e r h a b i t a t and mangenent. Idaho Wildl . Rev. 17(4) : 3-7. -
Osprey
(Pandion ha1 i a e t u s )
General
r The osprey (Pandion h a l i a e t u s ) i s a m i g r a t o r y r a p t o r common A p r i l t h r o u g h September i n western Washington.
Food Requirements
r Feed almost e x c l u s i v e l y on f i s h , b o t h f r e s h and s a l t water, u s u a l l y i n t h e 6 t o 10 i n . s i z e c l a s s , a l t h o u g h f o u r t o s i x pound f i s h a r e n o t uncom- mon p r e y (Bend 1937, Hughes 1983).
r No p a r t i c u l a r f i s h spec ies seem t o be p r e f e r r e d , a lmost any f i s h found a t o r near s u r f a c e w i l l be taken.
Water/Cover Requirements
r R e g u l a r l y use l a n d ad jacen t t o l a k e s , ponds and streams t h a t a r e i c e - f r e e by m i d - A p r i l f o r n e s t i n g . Land a d j a c e n t t o r e s e r v o i r s i s a s s o c i a t e d w i t h h i g h e r n e s t i n g d e n s i t i e s t h a n l a n d a d j a c e n t t o f r e e f l o w i n g r i v e r s (Van Daele and Van Daele 1982, Grover 1984).
r U t i l i z a t i o n o f water bod ies appears t o be r e l a t e d t o a v a i l a b l e s u r f a c e area and f i s h p r o d u c t i o n (Van Daele and Van Daele 1982).
Reproduc t i ve Requi rernents
Nest s i t e s a r e n e a r l y always l o c a t e d i n t h e open, a t t h e t o p o f t h e n e s t i n g s t r u c t u r e (Bent 1937).
Bo th a r t i f i c i a l and n a t u r a l nes t s i t e s a r e used. Snags, l i v e c o n i f e r s , and b roken- top t r e e s a re common n a t u r a l nes t s i t e s (Bent 1937, Garber 1972).
P i l i n g s and te lephone p o l e s a r e common a r t i f i c i a l n e s t s i t e s (Ben t 1937).
Nest s i t e s w i t h broad, f l a t , t o p s seem t o be p r e f e r r e d . Nest s i t e s average 3 t o 3.5 f t . i n d iamete r and 1 t o 2 f t . i n d e p t h (Garber 1972).
Nest s i t e s i n t r e e s o r snags a r e 60 t o 160 f t . above ground and u s u a l l y ex tend above t h e s u r r o u n d i n g canopy (Garber 1972).
Nests l o c a t e d on p i l i n g s may be as l i t t l e as a few inches above t h e wa te r (Bent 1937).
Nest s i t e s a r e u s u a l l y l o c a t e d w i t h i n 250 ft. o f a body o f water , a l t h o u g h some ospreys under normal c i r cumstances s e l e c t n e s t i n g s i t e s 2.5 t o 4 m i . from water (Garber 1972, Henny 1977, Seymour and B a n c r o f t 1983).
0 I n d i v i d u a l nes t s i t e s a r e used year a f t e r y e a r by t h e same b r e e d i n g p a i r ( B e r * - 3 3 7 ) .
0 W i l l t o l e r a t e ad jacen t ( a p p r o x i m a t e l y 150 f t . ) osp rey n e s t s (Kush lan and Bass 1983).
S p e c i a l C o n s i d e r a t i o n s
Ospreys n e s t i n g i n areas o f heavy r e c r e a t i o n a l use have exper ienced s i g n i f i c a n t p o p u l a t i o n d e c l i n e s t h r o u g h r e p r o d u c t i v e f a i l u r e (Swenson 1979, Levenson and K o p l i n 1984).
I nc reased d i s t u r b a n c e s ( l o g g i n g , b o a t i n g , e tc . ) o c c u r i n g f rom mid-May t h r o u g h August appear t o have g r e a t e r impacts t o n e s t i n g success t h a n r e l a t i v e l y c o n s t a n t d i s t u r b a n c e s o c c u r i n g p r i o r t o and t h r o u g h o u t t h e n e s t i n g p e r i o d (Levenson and K o p l i n 1984).
Human a c t i v i t v i n t h e immediate v i c i n i t v o f t h e n e s t t r e e can cause n e s t - - abandonment d u r i n g egg l a y i n g and i n c u b a t i o n p e r i o d s (Levenson and K o p l i n 1984).
S p e c i a l l y des igned a r t i f i c i a l n e s t i n g s i t e s may i n c r e a s e n e s t produc- t i v i t y ( t y o u n g l n e s t s i t e ) ( W e s t a l l 1983).
Re-estab l ishment o f b a l d eag le n e s t s near an osp rey n e s t can be p o t e n - t i a l l y d i s r u p t i v e , r e s u l t i n g i n reduced osprey b r e e d i n g success, n e s t f a i l u r e , o r abandonment o f t h e area (Garber 1972, Kushlan and Bass 1983).
P o p u l a t i o n s a r e l i m i t e d by d i s p e r s a l d i s t a n c e s . The p o t e n t i a l f o r osprey t o p ioneer s u i t a b l e h a b i t a t g r e a t e r t h a n 60 m i l e s f rom n e s t i n g con- c e n t r a t i o n s i s l ow (Henny 1983).
REFERENCES CITED
Bent, A. C. 1937. L i f e h i s t o r i e s o f N o r t h American b i r d s o f p rey . Vol. 1. Smi thson ian I n s t i t u t i o n U. S. N a t i o n a l Museum B u l l e t i n #167. U.S. Government P r i n t i n g O f f i c e .
Garber, Dav id P. 1972. Osprey s tudy , Lassen and Plumas Coun t ies , C a l i f o r n i a , 1970-1971. Rep. No. 72-1. C a l i f o r n i a Department o f F i s h and Game. 33 p.
Grover , K. E. 1984. N e s t i n g d i s t r i b u t i o n and r e p r o d u c t i v e s t a t u s o f osp rey a long t h e upper M i s s o u r i R ive r . W i l son B u l l . 96(3):496-498.
Henny, C. J. 1983. D i s t r i b u t i o n and abundance o f n e s t i n g osp rey i n t h e U n i t e d S t a t e s . Page 175-187. I n B i r d , D. M. (ed. ) . B i o l o g y and management o f b a l d eag les and ospreys : proceed ings o f t h e f i r s t i n t e r n a t i o n a l symposium. MacDonald Raptor Research Cent re o f M c G i l l U n i v e r s i t y .
Henny, C. J., M. A. Byrd, J. A. Jacobs, P. D. McLain, M. R. Todd, and B. S. H a l l a . 1977. M i d - A t l a n t i c coas t osprey p o p u l a t i o n s p r e s e n t members, p r o d u c t i v i t y , p o l l u t a n t c o n t a m i n a t i o n , and s t a t u s . J. W i l d l . Manage. 41:254-265.
Hughes, J. 1983. On osprey h a h i t a t and p r o d u c t i v i t y : a t a l e o f two h a b i t a t s . Pages 269-275. I n B i r d , D. M. (ed. ) , B i o l o g y and management o f b a l d eag les and ospreys: proceed ings o f t h e f i r s t i n t e r n a t i o n a l symposium. MacDonald Raptor Research Cen t re o f M c G i l l U n i v e r s i t y .
Kushlan, J . A. and 0. L. Bass, J r . 1983. Decreases i n t h e s o u t h e r n F l o r i d a osprey p o p u l a t i o n , a p o s s i b l e r e s u l t o f food s t r e s s . Page 187-201. I n B i r d , D. M. (ed.) . B i o l o g y and management o f b a l d eag les and ospreys : p r o c e e x n g s o f t h e f i r s t i n t e r n a t i o n a l symposium. MacDonald Raptor Research Cen t re o f M c G i l l U n i v e r s i t y .
Levenson, H., and J. R. K o p l i n . 1984. E f f e c t s o f human a c t i v i t y on produc- t i v i t y o f n e s t i n g ospreys. J. W i l d l . Manage. 48(4):1374-1377.
Seymour, N. R., and R. P. B a n c r o f t . 1983. The s t a t u s and use o f two h a b i t a t s by ospreys i n n o r t h e a s t e r n Nova S c o t i a . Pages 275-281 i n B i r d , D. M. (ed.) . B i o l o g y and management o f b a l d eag les and ospreys: p roceed ings o f t h e f i r s t i n t e r n a t i o n a l symposium. MacDonald Raptor Research Cen t re o f M c G i l l U n i v e r s i t y .
Swenson, J. E. 1979. F a c t o r s a f f e c t i n g s t a t u s and r e p r o d u c t i o n o f ospreys i n Ye l lows tone N a t i o n a l Park. J. W i l d l . Manage. 43(3):595-601.
W e s t a l l , M. A. 1983. An osprey p o p u l a t i o n a ided by n e s t s t r u c t u r e s on San ibe l I s l a n d , F l o r i d a . Pages 287-293. I n B i r d , D. M. (ed.). B i o l o g y and management o f b a l d eag les and ospreys: p roceed ings o f t h e f i r s t i n t e r n a t i o n a l symposium. MacDonald Raptor Research Cen t re o f M c G i l l U n i v e r s i t y .
I U.S. F i s h and W i l d l i f e Serv i ce . 1984. D r a f t h a b i t a t s u i t a b i l i t y i n d e x models - osprey. D i v i s i o n o f E c o l o g i c a l Serv i ces . Sacramento, C a l i f o r n i a . 5 pp.
Van Daele, L. J. and H. Van Daele. 1982. F a c t o r s a f f e c t i n g t h e p r o d u c t i v i t y o f I ospreys n e s t i n g i n w e s t - c e n t r a l Idaho. Condor 84:292-299.
Washington Department o f Game. 1982. F i s h and W i l d l i f e Resource S t u d i e s , D S u l t a n R i v e r P r o j e c t Stage I 1 F i n a l Report . Snohomish County PUD No. 1.
217 pp.
A P P E N D I X A
S u i t a b i l i t y i n d i c e s o b t a i n e d f r o m t h e USFWS d r a f t H a b i t a t S u i t a b i l i t y I ndex Mode l s - o s p r e y (USFWS 1984).