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PREFERENTIAL USE OF \'.1JLLO~v BY :>

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Susitna ,Joint Venture Document Number

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PREFER£NTIAL USE OF WILLOW BY

HOOSE IN INTERIOR ALP.SKA

By

Alaska Col~~rativ2 ~ildlif2 ~~s2arch Unit

University of Al~ska

Fairbanks, Alaska 99701

-~----·-

This work v;as fur1d2d by the Institute of Northern Forestry, U.S. Forest

s~ rvi ce, Fairbanks, P.l c:ska 99701 .

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over t'....:oo consecutive winters at a burn regrohTth a.rea in

Interior Ala.s}·a. Utiliz?tion percent v2ried from 0 to 50.2

t .. erc~nt UV(!T th~ tHO year period • ~. alaxensis and S, planifolia

..,..,~e:re founri to be preferred over So hastata end S, scouleriana •

sp~c1es. Both within-plant and betw~en-plant preferences ~ere

found to be components of within-species preference. Protein

to be unrelated to patterns of between-plant

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f2:=d primarily on curre:nt year shoots of \·lillow (Salix .?.2£_.), aspen

(f~:;:p_u_1_u_s_ tr5_rr~u_l_c;>_i_d~_sj, paper bif~ch (~_r:_tul~ pp_p_,y_ri fera) and balsam poplar

t:(·· •. ~~.mitic:s (Sp:::ncer and Hakala 1964, L::Rt:sche 1974). Of these, willow is

cc!r:sidr:red to b~ the rnost prc:ferrr:d by f>()OSe (Chatelain 1951, tJ,ilke 1969)

P.t:k 1974). The iJ;;portance of Hi.ilows as a winter food for moose has been

;·:ell doc:.K.:::ntc:d in oth:::r areas of North America (Baker et al 1953, Harry

1957, l~nO\\'lton 1958, Pimlott 1961, Berg and Philips 1974).

u i f f i c u l t .Y i n s p e c i e s i de n t i f i c a t i on h a s r e s u l t e d i n mu s t o f the

li:.e::~a~ur'e stating that Salix _spp. is prefen~ed by moose. Betv;een-species

f!r.::-:e:~·enc~s are generally ignored. A few publications, hm-.·ever, have

:ji·•ci..i:·.::nr.ed a species preference by moose. 1·1urie (i961) found that S. plani-

Ht. i·icl~inle:y I'~C:itional Park. t·~ilke (l9E 9) documr:nted that J 1terior Alaska ..

:·oo~e have a decided pr:::ference for~· ~[l_t~~rio.r_, ~· alaxensis, ~- Qlanifolia

snd ~- 9.T .. ~~~c_ul_oides over other availdble willow species.

R-::s::arch concerning within-species sele:ction has likewise been minimal.

L~~~sche (1971) has suggested nonrandom browsing by moose due to the

extreme variances commonly found in random measurements of browse product~on

and utilization. On bitterbrush mule deer range, there is often wide

variation in the degree of utilization among individual plants (Hansen and

Smith 1558). Food habits studies in Western Washington black-tail deer

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jl range have sugg~sted that only 1 percent of available salal plants

~~rown 1~60, Lc.uckhart 1962).

4

The pn:~··=l;t s~_udy w.:s dE:sisr,t:d to e:valuate \'lht:ther bet\-;::en-spe:cies

i-. · ·~-,... ,- · o t""'\ d n' · -~-.a tr..•V It t:- \.. 'i -.... G. J t.. ~ Protein content 2s a possible determinant of

···J·;.. . .-_ .-;--r r (.:. ,.·-:: c; r' -t ...... ~ .. 1'-- ,v_ also inv~stigcted. G3ta ~2re collected in 1977 and 1978.

Sf U~ Y f:,REP.S

Trn~ee study areas n~pn:sentative of v.'illov1ed habitat types

.. :l'c ~-'::1c:ctt:d in a 6,300 ha burn that occ:Jrred in 1971. In addition, one

: r L t•-, c_ tun\' ';... '' !.. • '-' I - .. UJ c rc:a was located in a 70 ytar old mature black spruce stand

to ~~= burn. The size of t:ach study area varied from 50 to 200

~-::. ::.-~ur st:.dy arec:s located ot the \~ickersham Fire Ecology Study Site

20-~11e Elliot Highway, were used in this

: r \' !- c;'.; g;:, ,J. ,· on I V--~1 ~t. • :Jescriptions of t::ach study area are shown in Table 1.

.. -·por-~s *' .:: I 1l 4,.0

Estin~at~s of browse production and utilization for Each spec1es

~~ Each study area was provided by the Shafer twig-count ~~thod (Shafer

~;63). The sa~pling unit chosen was the randomly selected quadrat. The

d:t~rmination of quadrat size and number of replicates (Table l) for each

stLdy area followed the optimization procedures of Cochran (1953) and

Barret (1969). For each willow stem in a quadrat, the data recorded are

sp~cies, numbt::r of twigs browsed and number of twigs available. A stem

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hcS ccnsider~c-d to b::: any t.re:r,c.h that joins \'lith other br~~"'ch::s or r~oots

b~low 40 em. It m~st also be tall~r than 40 em at the point of it's

h~~h~st shoot. Forty em above ground level was chosen rather than ground

level to account for sncw d~pth . . !'.. f.tr::lirr~inar~y Semple take:n during the

1976-17 \·:inter indica~ed tO em to be appcoxiir.ately th'= r:~··an snovi d::pth • . , . . . d t L. ' t .... h . . .... 1 t th r. ~w1g v.·as cons1o::re o ue any sr,oo w1~.- a o1ame~.-er equa o ,e IT'2an

di~r::::~er at the p:;int of brc,wsing (uPB). 1~-::an DPB \·Jas generated by

:r::asuring the DPB of 50 randon:ly selected browsed twigs of each spec1es

in each study area and calculating the m::an. Fifty unbrowsed tvdgs

cc:n~es~·--~r,ding to the same diar:1eter distribution as the DPB sample were

:~i~p~d 1 oven-dried and wsigh~d for mean weight per twig. Each study area

\,c:S ~::·~;ed for prc,duction and utilization during f~ay of 1977 and 1978.

:":.:::a cunco;:ii-"...c.r:t cts·.::rve:~ion on r:cch stem, the brm·:se history

.:: .~-js..:"'.cs-/abse:nce of br'ov.s~n;) for the 197~-75 and 1975-76 win~ers was

·:::.Jrj-::d during the K1y 1977 s.:::::p1ing period. The browse history of each

s~:::'7l c~·er a three year period was available for analysis. Availability-

~:i 1 i~~~ion gJodness-of-fit statistical t=sting was ~sed to determine

~- . - ~. ' - - .......... .:-c. Y"" ~ .,- -"" • - ., .,, -=~ ·== r'· c: ·-· C:Jit..lai use of st~ms within a species existed.

~~~otive use by moose of each study area was evaluated by fecal

:1·::~ Jrc~up counts (iieff 1968). Optimization procedures of Cochran

(i053) ~~re follow2d for the determination of the most efficient quadrat

si:a a~d replicate number (Table 1). Pellet group data were collected

6t t~~ ~arne time as production and utilization data.

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A series o* 40 p~1rs of willow p~bnts w~re tagged at Study Areas

l and 2 during Octobt:r 1977. One lllf:;;:b~r of e:ach pair had been browsed

tr£ previous \·lint~t~ and th~ other h'2S unbrcwse:d, Othe:r requi r::;rr;::nts for

• I~,:. l.il - pair si..ipulatf~d that th~y beth~~··:·:= sp.::.ci::s end lncat~d l~ss than

10m cpart. Durillg J:pdl 1978, Each ~·1cnt ·.-:as t:>ar •. ln,;d for evide:nce of

browsing during the 19/7-78 winter. If one ~2nber of a pa1r was browsed

both winters and the other unbrowsfd both winters, a sample of 10 or

111::>re cut-rent annual grcwth hd gs was clipped ft~om each plant, oven

dried and analyzed for crude protein content. Statistical tests were

us-::-d to d::;terr.;ine v.·hetr.e:r relative protein content is possibly related

to pr:::fe:r~rrtial use,

r:ach study a rea, a sa:r.ple of current annual growth twigs \·/as

ccl~t:cted fr~om the one or tvm mast iritJ)rtant sp:::cies during February 1978.

Th2 dia~~ter distribution of twigs cli~~~d duplicated the sample

distribution of DPB 1 s coll2ct2d during ~ay 1977. The twig samples were "· ' ovEn driEd and analyzed for crude pro~tin content.

.. .. - -~~- --· ~ - -- .. -------

... L . ., . . :rr··.·,.s::: ·i..lllZctlon -- ,...., _,_ ·- - .. - .. _ ............ - -"··-

~=an trcwse utilization perc~ntag2s varied from 0 to 37.7 percent

the 1976-77 hint2r and from 0.8 to 50.2 percent during the 1977-78

wint~r (Table 2). The results compare favorably with that of Milke (1969)

\-vho fGund browse removal percentages of \villGw in various Interior Alaska

hilltt::r l~anges to vary from 0.1 to 33.8 percent. vlolff (1978) found

uti~ization of~· _SC..Q..!Jj_et~iana at an Int12rior Alaska burn site to vary

fr:ffi 8 to 45 percent. At an Interior Alaska floodplain site, Wolff (1976)

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7 found utilization of wi lJ 0\·ts to be S5 to 56 p~ rcent. Severa 1 wo rh rs

have s.uggested that a r·emoval pc,rcentcge of 50 percent may be optiJT.al.

for a !;.a xi mum sustained yi e 1 d of ha rd1·:ood b r·owse (Spencer and Cha te 1 a in 1QC3

K ~~~ ft in g 1966, Ho 1 ff 19 76) . I - ..) )

Gifferences in rY:;Jval percentages beth·een years was tested b,Y the

chi <quo re m;, thad (Tab 1 e 2) . The chi -square was s i gnifi cant in 5 out of

7 cases and in 4 out of these 5 cases, the difference was attributable

to an appare'ltly higher browsing rate during the 1977-78 winter. l·bose

use of the area as evidenced by the relative a~unt of sign and sightings

.,as definitely higher during this winter .. The reduced biomass of browse

';';oilcblc during the 1977-78 winter as shown in Table 2 my also partially

~r~ws~ s~~cies ?r~f~r~nce5 .. - - --- -- -- ·-·-- -~--...---··------

Hos evaluated by the availability-utili-

:c"cf:Jn chi-square m;,thod after Neu (1974). Illustrated in Table 3 are the J

••su1ts of analysis. In 3 out of 4 cases, the chi-square statistic was

"'sr.ificant. ~- ~axensis •1as utilized IDJre than expected both seasons in

::.dy.,t,rca 3. S. planifolia ~1as utilized r..ore than S, scou1eriana the

··~ '··: ~ ~.,,~son but not the second sea son in Study A rea 2. ~- hastata and

~. ?,1,2_r,_ifu~ •1a re uti 1 i zed 1 ess than expected both seasons in Study Area

: •. It appears that~- alaxensis is most preferred,~- planifo1ia is

~ext and S. hastata and~- 2:£Duleriana a,c least preferred. Other studies

in Interioo Alaska have shown both S. alaxensis and S. planifolia_ to be

i.ighly prefen·ed over other 1vil1ow species (f.lurie 1961, l~ilke 1969),

•;)uld (1977) found for the Colville River area of arctic Alaska that

!: . c l_o,,~sJ.?._ was fTDS t pre fe n·ed wh i 1 e ~- .P}_1I n i fo 1 i a was least preferred

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0f si~ a~ail~ble willow sp~cies. Lack 0f cover in high density

S. F-icrtii~cl~_?- areas was sugg-:sttd as a possible cause of low utilization.

\.. .. . . p ,. .·.1 tr. ~ r. -s ~. ·c. 1 1: s rei r: n:: 11ce s ... 4 ... - .. -- -· .. -.-·- ... -- .. -

\·:i:li~n--spi::cies pre:fcrd1ce of st~~n;s \-.'JS likr:v;ise :::valuatP.d

b C h 1. - s q • . a- .~ p. n1~_ i_ ~--~ c1 d a- r'" ~t.. e r •, ,, P u t 1 9 7 4 ') . by th~ ~\~ila ility-utilization u .... ' - \

TbJ1e 4 illustrates the results of analysis. Due to the low expected

fr'='r~u=nr.ies 1n SOI!le cate:gories, the log-likelihood statistic (G) was

su~stitut~d for the standard chi-square statistic (Sakal and Rohlf

1~f9). Four out of 5 species/study area categories showed significant

G stc:istics. 1n 3 ou: of the 4 speciesistudy area categories, stems

-~ · ;:..~ .. ; . .- •• .:,;...s c."''-·~- •u··!...-..'"'··c:::..d a'ur.:n,, .... .:.. 1°7 11 -75 ar:d 1,q-,,5-76 ,,,;,ntf-.rS ..... _,., :..:-~ -·~) _t..-•1•::: •.\u \.·\\_.,._ - 1 ~ L ... , _,), .. 1 -- n -

•· .. ~~~-:- :.md::rbrov.·sed in proportion to availability. In 1 out of the 4

t~~2g:ries, st~ms bro~sed during the 197~-75 winter and unbrowsed ...

d ... d:.;; F~2 1975-76 h'inter v.:ere b:---::o·.-.sed i'.Ol~e in proportion to availability

""~ r. o:: ~ •• : s "= c, J e n t v.· i n t e r . I n 1 out o f the 4 c a t e q o r 1 e s , s t · 11s u n b ro vis e d

dJri~g the 1974-75 winter and browstd durirg the 1975-76 winter were

:..:ndE:rbt--o\·,·sed in proportion to availability. The p-ttern suggests a

preference by moose for stems browsed yearly and avoidance of those

stems not browsed previously.

S. planifolia plants on Study Ar~a 2 vtere analyzed to determine

wh=th2r pref~rential use was within or betw~en plants or a combinati0·

uf bJth. Only this study area and species m~t the criteria of possessing

int~r·;:,eciate levels nf utilization and a high density of stems recognizable

as plaGts. Two by two contingency table analysis was used to evaluate

vihether preferential use between plants occurred, Table 5 illustrates the

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r:::thc,dology and rfsults. f..s \.;as the cc:se for stems, those plants

brch'Scd the 1975-75 i·.'i nte r tended to be broh·sed more than expected

d~ring the 1976-77 winter •

...... h. l .J:' 1 d b l . 1\l L.. 1n-p ant pct:,f:n:nces were eva uatt:: y regress1on ana ys1s.

Twenty six pl~nts ~~re avbilable for analysis. Each plant was

ccffi~osed of at least one stem browsed during the 1975-76 winter

(category 2) and at 1 east one not br~ov:sed (category 1) The independent

:.:ticbles (X1 and x2) are the number of stems per plant in each category

available as browse during the 1976-77 season. The dependent variables

(Y1 and Y2) are the nurrber of stems per plant in each category broh'sed

dJri 9 ~he 1976-77 season. Results are shown on Fig. l and 2. Both

)·:::-gl~:-ssio'ls are significe:nt (p<:.OOl). Both the slopes and correlation

.::.~iff·icien~s e:re significantly different (p{.OS). Noteworthy is the

·~rg~r s~0pe and much tigher fit observed in the second regression

( f i g . 2 ) v e r s us the f i r s t ( F i g . 1 ) . T he port i on of a p l ant b rows e d "\ .,

c:.~ring the 1975-76 winter tended not only to be browsed at a higher rate

~·~~ f.Jllov:ing winter. but the_ rate was rnore predictable.

Th~ tag~ea plant ~xperiment yielded different results, however.

:~·;t.- cJ· :·v:o cor.tingsncy table analysis ·was again used to evaluate between

;:~~t pr2fer2ntial use (Table 5). The chi-square statistic was not

significant at the .05 level. It is note\'t'orthy that the observed f~quencies

of plc.n~s bro\\'sed both se:asons and plants unbrowsed both seasons are higher

tJ,an t:-.\pected just as in the upper table.

......... ,. ·- ~~ ... ·--·---~~-..,4~~~-··,-·,·· ,. ..... .

-

Illustrated on Table 6 are the rt:sults of protein analysis for

t:;~= study art:as. The range of values was narrow ranging from 5.3 to 6.0

; :l._·.:r,~. f..ll are lo:·Jer' than the 7 percent minimum Gietz et al (1970)

. r·jt:d fol~ good Irule d~er range. Olde:m=yer (1977) reported a mEan

or 6.~ percent for willows on the Kenai Peninsula. Milke (1969)

r.:~ ... ...-~. h values ranging frc•m 5.2 to 7.0 percent for 4 species of Interior

Alas~a willows. His reported value of 7.0 percent for S. alaxensis is

higher than my value of 6.0 percent although both top theii~ respective

lists. Th~ difference may be related to dates of collection of samples.

;~.-:: secsonal changes in protein level has been well documented (Bissell and

~ "" t• r l c~ l .. · ....... tg .. J:>, Tew 1970, Old=~eyer 1977) and apparently is due to differences

'· i ~ :.:Jre and phenol og.Y.

f.. ~.·.t.:il of 6 pairs of tagged plants of~- ol~~ifglia and 5 pairs

•·~ S. s_c~)_uj __ c:r~~~ tagged plants \•;t:re analyzed for crude protein level to

=--~r~i~e ~heth~r proteir content is related to between-plant selection

( ;-:.:;le 7). The paired t-test was used to test for differences in protein

-~··:~::r;~ bst,,·2en the t\'/O browse categories for each species. '\ll percent

c3:::: v:C!s :ransfonned by the arcsin transforrration to allow the use of a

~,,:(6;:-.::tric test (Zar 1974). For each species, the resultant t values were

r1~Jt significant at tne .05 level. Since moose may select within plant as well

~~ 0et~~en plant, the design of the experiment should have taken this

·;rrto account for meaningful results. Due to a higher than expected plant

.. 1 . ( + ' 1 ; ' I (1, I\,. variability, a larger sample size is indicated .

_j _ ..

counts varied ft'om 0 to 52 groups per ha. ,, '.

~T3bl~-S). For the 1976-77 winter, the group counts of Study Area 3

~-~d .:d hi y!.es t, Study At~ea 2 \·:as next a!ld Study Ar.::as l and 4 rC1nked

1~st. For the 1977-78 winter, Area 3 ranked highest, Area l was next

~~d Areas 2 and 4 ranked last (Kruskis-Wallis, p <.OS). A • .L

DE L\..,eEn

~;, :::~- co:~pat'lson shov1ed all a.reas except Area 1 to be the same both

y~al~s (;·~nn-Hhitney, p ( .05). The highest consistent use of willows

in Area 3 ffiay be related to a number of factors. The high density of

11

~~n~~:~·.·s not only provides spatially concentrated forage but cover as well.

1 ~ ~ n' , :... .: gh .:, I ~· t: · i I • I pn:ft:rence for S. alaxensis is nutrition· related rather than

the species preference is an additional factor .

. 0~ cc~sistent use of hrea 4 is to be expected. Since forage plants are

~~~ely scatt2red, it 1 s main valuf would be to provide cover.

Cor~parative estimates of use in terms of rroose-days/ha (t~t.D./ha) ('

~~s cc~?~t2d from pellet group data and browse utilization data (kg/ha).

?.::::;.;.;"!.s arc: ~.hown on Table 8. Assuming an adult moose consurr.·2S 5 kg dry

•·-=~,~r,: vf \·:x~dy browse per GGY (Gasawcy and Coady 1974), moose use varied

to 24.28 M.D./ha in the four study areas. If one assumes a daily

.::.:-:~.:.:::ion ~ate of 13 groups/day (Neff 1968, \•Jo.lff 1976), rroose use varied

f·0~ 0 to 4.00 M.D./ha. Results of the two groups do not coincide, particularly

St~dy Ar~a 3. This largely may be a problem related to basic assumptions.

~·i=an consur.-:pt·ion rates vary greatly in the literature, LeResche and Davis

\"t?7l) docu;;;ented values as low as 1.3 kg/day dry weight for r~enai

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n:lose \·;hile Pall;.er (1944) found consuwption rates to be

.;:$ ~.i'gh as 16 kg/day d.ry wE:·ight. Like\'lise, r':pOrted defecation rates

'.·:.~·s s:"eatly d·.:p•:nd~nt both on sex and age structure of the studied

>·= •il5 as \ ... ·211 as fcJJ~e:ge t::aten (Neff 1968). Fronzifionn et al (1976)

. .:·.<. ~>::r.t.:~d a daily de:fe:cation rate for Kt:nai Penir:sula noose at 14.6

~:r fvm3les and 19.6 for males. Unless you kno~ the mean daily

... ~ ... ~ ... ,. v:') ,. ' :"" •.. 1. . ... l. ':- and/or defecation rates of moose in your study area,

,:.,_·;:~_;tation of moose-days of use by either r.tethod is incnnclusive.

Prir1-:iple conclusiuns are sur;tnarizt::d 1n the following:

1 ' . .t.' l 0 76 -7 . .... u n n g L.f. e :: - 1 vii nte r, the browse utilization percentages for

;·c:!: sp::cies and stucy ar::a varied from 0 to 37.7 percent and from 0.8

10 50.2 percent during the 1977-78 wintEr.

2~ ,.;n ar.alysis of prefe1~ence of species revesled ~· .alaxensis and S.

:~~.::_r;i_f_oj_i..9_ to be preferred overS. scouler_jana and S. hastata.

3. ~ pr~fe~ence by moose for stems browsed yearly and avoidance of those

;i~~s not browsed previously was documented, The same preference pattern

~~s found to exist b~tween plants of the sam~ species.

4. ~ithin each plant, those stems browsed the 1975-76 winter tended to

'J.;~ browsed more the following winter than those not browsed the 1975-76

h'inter.

5. Crude! protein levels ranged from 5.3 to 6.0 percent. No significant

~'S.JL ''z-~r

o·'

13

,:i -"~dir··«nh=· at the .05 level could be detect~d b:::tween plants browsed u f tf ..... ·;-

\I \\

t~~ ~~/6-77 winter ~nd the 1977-78 winter and those not browsed.

s~/·.:ral applicatio~s of these rr-sults can be r.lade for r:oose

;.:Pc~::2nt in P.laska. Range evaluation studit:s n~ed to take into

c.r·sideration h'ithin-species preference. The 50 percent browsed may

r,~t !:.e i;q~iivalc:nt to the rcn~aining 50 pe:rcb.+: available on the rang:::.

:~,:-jit.at ir11provc:~~~ent work such as controlled burns and mechanical crushing

n2ed to take into account which browse species will grow back and

1n what q~antity.

Additional research along these lines is needed, particularly

:~,a~ing to causes of within-species preference. The role of rumen

j=:~stive in~ibitors as well as other nutritional components besides

... .... : .. ' t • '-lr r ·-~t1C. vn ~ .. -- tf -· .. """"'-" be investigaied. Physical factors such as snow depth

~:r distcr.ce from a t.rail rtay also be imrortant.

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14

f'·!lt·~, T.C., C. Anderson an'~ \\'.T. CnH:tp. J9S3. Food habits study of game

· 1 r ···1d1.,.. :;: ::·~ ~. \)'O. i\1 .1re 17(11) :24-:~1.

.. 1 'T) .. " - 1 ..... l •

J:lf9. Opi ~;.n!T: pJot sr!Jnp1ing 111 cstil:iating

~·! > 1.;, ~.: ::: u t a K :i t h J c f (: n.: n c e t o at h e r h c.: a v i 1 y w i J 1 m .. · e d a rca s . 1\ at . Can .

1Cl :JOl-116.

~.~! · .;•11, H. D. and B. Stn.mg. 1955. T~'le L:rudc: pTotein \'ariations :in the

~ :~_·,;st:> diet of Califc,rnia l~!'::t::r. C:a1if. Fish and Gar.~e 41(~) :145-155.

F.R. 1961. The bJac}:-tai1c·d dt::c:T cf l>c·stern h'ashington. B:iol. Bull.

}. • ~· ; ":, '1• 1 • 'r· C • ' t e ~I f•.,... t r· . .::- C r •• e r- , ' . ~ J• 3 p '1 J, '1 ;\ .• .. -· • J ~' "( ... ~"'"" .. .., L<: L -J' • ul •t'iJ.l ~ J t.~ ... .,.• 1 :· • . • -- -'1 •

"'-.:!:·:;::r,, E.F. 1951. \\'inter range problc~s cf moose in the Susitna

,,. '· 11 '•\' fJ'OC 1 ....... '- •. " \. ..

2nd Ann. A~as1~a Sci. Conf., p. 343-347. •

' , -,; . ·" p.

I··~.::J t.;.R. 1970 .. \.f'ir.al Jlroduct:ion and ft)n1::e q-.Jality. U.S.D.A. Publ. 1\o.

p. ]-9.

A.W., P.D. Arneson and J .L. Oldt:rr.yer. 1976. Daily ,.,inter pellet

(In press)

C:l:c:.,:.-.·ay, W.A. and J .1\'. Coady. 1974. Revie\~' of er.ergy rr:quire:ments and

(\

I::

lt)·, • co

}'::;

. f

~· ,,

' ; . ~

i

\~

\)

:.)

:::J

10

111 ustrated on Table 6 are the r~sul ts of protein analysis for

the study arE:as. The range of values was narn.1\·! ranging from 5.3 to 6.0

~ .. ::,-~..~nt. All are lm·:t::r than the 7 percent minimum Gietz et al (1970)

:F.c::: .:.:r:dc:d for good 1rule deer range. Oldt:rneyer (1977) reported a msan

va1,_.e of 6.4 percent for willov.·s on the Kena·i Peninsl'la. t~ilke (1969)

r-..:~<..r: . .:-d values ranging from 5.2 to 7.0 percent for 4 species of Interior

f...lc:sLa willows. His reported value of 7.0 percent for S. alaxensis is

higher than my value of 6,0 pErcent although both top their respective

lists. The difference may be related to dates of collection of samples.

Th~ seasonal changes ~n protein level has been well documEnted (Bissell and

Tew 1970, Olde~eyer 1977) and apparently is due to differences

. :: (d ~ l:Jre and phenology.

L. :·.t~l of 6 pair·s of tagged p:arts of~· o}.a!~ifg_]~ia and 5 pa1rs

t:.f ~~-· s~cp_uj_e_r_"L.a_f@_ tagged plants \·;~re analyzed for crude protein level to "' ...

::~.·:n:i;.:: hhether proteir content is relat::d to bet\';een-plant select1on

:·•:.:.;1e 7). The paired t-test W3S used to test for differences ·in protein

-~n~~n~ between the two br~wse categori~~ for each species. ~11 pRrcent

::::'ta \vas ~ransfr;r·med by the arcsin t)~ansforrnation to allow the use of a

; .. :(~~:-,2tric test (Zar 1974). For each species, the resultant t values were

r,;Jt significant at tne .05 level. Since moose may select within plant as well

·~ oetwten plant, the design of the experiment should have taken this

.:r~n at.count for meaningful results. Due to a higher than expected plant

. ··, l • , . - •· jl I d:t \.. V3riability, a larger sample size is indicated .

j

I !

I l I I

l i' l c

\ i

,,

11

I·~-:an pellet group counts varied from 0 to 52 groups per ha.

(T3b1e 8)~ For the 1976-77 winter, the group counts of Study Area 3

r ~r.L:d hi gLI?s t, Study Area 2 was next or1d Study Areas l and 4 ranked

:ast. For the 1977-78 winter, Area 3 ranked high~st, Ar2a 1 was next

~nd Areas 2 and 4 ranked last (Kruskis-Wallis, p <.OS). A between

::·: 3r co~·par1son showed all areas except Area i to be the same both

y.::ars (l·~nn-Hhitney, p ( .05). The highest consistent use of willows

in Area 3 ffiay be related to a number of factors. The high density of

hillc;·,•s not only provides spatially concentrated forage but cover as wen.

If t.ht:: high pteft:rence for~· alaxens1s is nutrition· related rather than

r.~titat related, then the species preference is an additional fact0"".·

.0w ~c~sistt::nt use of ~rea 4 is to be expected. Since forage plants are

h~daly scattered, it's m~in val~e would be to provide cover.

COln;Jarative estimates of use in terrr.s of IIDose-days/ha (M.D./ha) "\ \.

~~s cc~?Jt2d from pellet gro~p data and browse utilization data (kg/ha)a

?,:~·~its are shown on Table g. Assuming an adult moose consumes 5 kg dry

~f W)ody browse per day (Gasa~ay ?~d Coady 1974), moose use varied

f~~~.r- 0 ~ . .) 2:1.28 H. D. /ha in the four study areas. If one assumes a daily

.:~.r~~c:ion r·ate of 13 groups/day (Neff 1968, \4olff 1976), rroose us::~ varied

. '

t•\.:X C to 4.00 M.D~/ha. Results of the tvw grouos do not coincide, particularly

St~dy Area 3. This largely may be a problem related to basic assumptions.

H~an consumption rates vary greatly in the literature. l.eResche and Davis

(1971) dc..-.:umentE.d values as low as 1.3 kg/day dry weight for Kenai

t' \

=In

--

12

Iroose \·;hi l e Pa l11.e r ( 1944) found c. on s vr:rpt ion rates to be

"~s·!~igh as 16 kg/day dry \'Ieight. Likev-1ise, r::ported defecation rates

\~·~ s~~atly d~pend2nt both on ~ex and age structure of the ~tudied

:.n.: .:ils as well as fcn~cge eaten (1-~eff 1962). FranZir.ann et al (1976) 0

tk~v~·:,•~nt~~d a daily dtft:cation rate for Kt=nai Peninsula n:CJose at 14.6 0

-; ~ r f t: ma 1 e s a n d l 9 • 6 for rr.a 1 e s . U n 1 e s s yo u k no vi the me a n d a i 1 y

and/or def.::cation rates of JPOOSP. in your study area,

c.::. ·~·utation of l'i1oose-days of use by either nethod is inconclusive.

Pr-in.:iple conclusions are sunr.narized 1n the follm~·ing:

1. ~uring the 1976-77 winter, the brov.•se utilization percentages for

::.c~r: sp':cies and stucy area varied from 0 to 37.7 percent and from 0.8

tG 50.2 ~ercent during the 1977-78 winter.

2. An analysis of preference of spec1es reve3led S. alaxensis and S.

3. r pr0ference by moose for stems browsed yearly and avoidance of those

;. :· · s r.ot browsed pl~eviously was documented. The same preference pattern

~~s found to exist bEtween plants of the same species.

I ~- ~ithin each plant, those stems browsed the 1975-76 winter tended to

t:l~· t.rowsed more the following \';inter than those not brovtsed the 1975-76

h'inter.

5. Crude: protein leve:ls ranged from 5.3 to 6,0 percent. No significant

··-·~ 1...... ~· -•..,~·•~ .. ,_...,_. ·e·~-•" _.,,,.--.~•··-·-······--····~'

I' -jl

13 :'>

cift:r':,:!iCe ot the .05 lev:::l could be detE::ctr::d bct\.;een plants brcv.•sed

u·.e lS/6-/7 winter t~nd the 1977-78 \·.·inter and those not brm·.•sed.

s . .::-;::ral applications of these r,:.sults can be r:1dde for r;oose

;.:nc~·:<·nt in ft.laska. Range evalration stt;diE=s m::ed to take into

c.._r·sidEration within-species preference. The 50 percent bt"'Oh'sed may

!!~t te ;.q~;ivalent to the t~en-:aining 50 pt::rcf::nt available on the range.

such as controlled b~rns and mechanical crushing

~Eed to take into account which browse species will grow back and

1r. \·;hat q~anti ty.

Additional ~~esear~ch along these lines is needed, particularly

~~~6ting to causes of within-species preference. The role of rumen

j~~~stive in~ibitJrs as well as other nutritional components besides

:•~./ .. -;::in r:.:::-ds to be investigated. Physical factors such as snow depth

Jr dist~nce from a trail nay also be i nmo:~tant . .

{1 '· ~ '

.\

-

[

···.l .. · 1 !

' ' \ '

.·

14

, " . - - -. _ . ...,. --

• • ""'J c c· " .l-:.- ·~:.t·J, . . ~ .• :"~TICJCTSOTI and l\'.1~ Cnli::p. 1953. Food habits study of game

· · . 1 ,,. o \' . 1 d I . ,. ,:. : ~· r! s. 11)' • 1 1 .11 e 17(11) :24~:.1.

J:lE9. Opi i;.:lr.; rJ ot smnp1 ing 111 estimating

1. ·.• .- ·,· -~ e . 1 \··1· 1 <ll u · ~ ""e -:: .. · - o c ' 0 ... _ • ..., • • J • , •. a J.d.::. • -'.) • .::-- _ • .:.: ·• .) •

1('1 :}0l-JJ6.

:. : •• ;-. 1 1 l-i 0 'lT d u s .... ,. ,., (l ] 9- ~ -..~ , • ~ .1 J , , , ,· •• ) I) , ._ l. ~ \.,; ! , ~ , • ~ ..- o 1 II e LT''dC fTOtein variations 1n the

1-:::·o,;~.e diet c•f Ca1ifc.r·nia d2er. Calif. F:i sh and Ga::;e 41(2) :145-155.

1 ~"!A p. --1-'-t.

-: :··.c~::-:·n, E.F. 19Sl. \\'inter range probleT!'s c1f moose 1.n the Susitna

\" ·. 11 C\' ' ..... ,.4 .. • ~ T 0 c . : Jl d Ann . A 1 as 1~ a sci . c 0 n f . ' p . 3 4 3 - 3 4 7 .

'

r·s~:., D.R. 1970 •. .\.-:ir.;al .i'roduct5on and fvT<1~E" q'Jality. U.S.D.A. Publ. 1\o.

1~47. p. 1-9.

F':rh:l:r:.Tr~n, A.W., P.D. Arneson and J .L. Oldemyer. 1976. Daily vdnter pellet

(In press)

l·~:. <::. .•• ':::1\' l,..,~ .. , ........ ~.,- '

\\'.A. and J. W. Coady. 1974. Review of ertergy requir£:ments and

J-:lrr!en fern.tmte:t 1or, 1n ~.oosc rmd other TUJrir.ants. ~at. Can. 101:227-262.

f

15

·:~t::st=n, \LO. r~nd J.G. Smith. 1970. Significance of forag.e quality as a

tool in 'dJdlife manage1r1ent. U.S.D.A. Publ. No. 1147. p. 25-31:

l:;:rry; G.B. 1957. Kinter food habits of r.,c,ose in Jackson Hole, Wvo . .. J. KiJdl. ~lanage. 21:53-57.

i:r,md ton, F. F. 1960. Food hab:i ts, moven.ents and popu] ati ons of moose

in the Gravelly Mountains, Montana. J. l\"ildl. t•1anage. 24:162-170.

t:n:fting, L.W., N.H. Stenlund and R.K. Seerr.el. 1966. Effect of simulated

• .

and natural deer browsing on mouptain r:taple. J. \\'ildl. J.1anage. 30:481-

~SS.

L2:.1ckhart, J.B. 1962. Wildlife population ft:ndam<.=ntals. Trans. N. Am. \fildl.

Conf. 27:~33-241.

Le;\e:s.:::he, R.E. and J.L. Da\'is. 1971. 1·~ocse research report. Fedl. Aid

Wildl. Restor. Froj. Rep., Vol. XII W-]7-3. Alaska Dept. of Fish and

Game. Juneau. 88 p.

LeRe.sche, R.E., R.H. Bishop and J. W. Coady. 1974. Distribution and

ha8itats of moose in Alaska. Nat. Can. 101:143-178.

G.C. 191:?9, Some moose-willo"'' relationships in the Interior of --A1:1ska. f'.LS. Tnes_is, Univ. of Alaska, Fairbanks. 79 p.

l·buld, E.D. 1977. Movement <patterns of moose ~n the Colville River area

of Alaska. M.S. Thesis, Univ. of Alaska, Fairbanks. 82 p:

1961~ A Naturalist in Alaska. Devin-Adair Co.J Old Greenwich,

Conn. 302 p.

-

":;

\I

\\ ·;)

~\(·ff 1 D .. J. 1968. The J•e11ct-~.roup count t c·cLnioue '

{)-

for big g~rne trend,

cen$US nnd distribution: A revie~. J. hiJdl. Manage. 32:597-614.

~~.:uJ C.\\'., C.R. ByL·rs <:1nd J.t.L Feek. 1974. :\ tcchniq:Je for r.nalysis of

ut.dj::atio;-:-a.vai1?.bility data. J. h'ildl. :-1cm&ge. 38:541-545.

J"'r,~Jv.:·r, L.J. 1944. Food requirer..ents of sorr:e Alaskan ghme animals. J.

'l..!a::;;~;al. 25:49-54.

16

of r.:0ose food h,~b'i ts studies in ~\orth .:..nerica.

~\at . Can . 1 0 1 : 1 9 5- ~ 1 5 .

1961. The ecology and r::anagt:;;ent of moose in I~Drth iu-::e:rica.

La Terre Et La Vic No. 2-3:~46-265.

~rencer., D.L. and E.F. Chatelain. 1953. I'r0gress in the manager.1ent of moose

~

of soutlJcentral Alaska. Trans. K. ~~- Kildl. Conf. 18:539-552.

~3ic·ncer, D.L. andJ.B. Hakala. 1964. }.;'"'.~·sc a:~cl fircontheKenai. Proc.

3rd .4.J1n. Tall Tirrbers Fire Ecology (onf. ~ f· 11-33.

s~'hal, R.R. and F.J. Rohlf. 1969. !iior.-:_ctry. h'.B. Freemann and Co., San

Francisco, C31jfonria. 776 p.

T{:W, R. K. 1970. Scc::.scnal Yariation in the nutrient content of aspen

foliage. J. i'.'i1dlt. ~·1anagc. 34:475-478.

'\cdf:f, -..J.O. 1976. Utili::ation of ha.TG\\'ood bro1n·se by r.:oose on the Ta:nB..na

flood p1ain of Interior .A1asl:a. U~~;A Research :~otc P~W-267. 7 p.

I

11 I

. 1

i ~

L

d'IP .. I I

f /\Hu j ; LIP SU" i 1l t i 011 ,) l '~ '_,;qj• . ;t r: (.j : . ; ' t ~ : ~I ' I l 1 ( 1 ~ ( . ~ "1 (. : l + 1----.......... -... ----·-----~·---·-- , __ ,._ ...... _ . ...._ .... _____ ,.... -~·

Study Area

1

2

3

4

Jesc(iption

Ridge-top; burned in

l 9 71 ; fo rme r l y c 1 i 111a x

black spruce type

Ridge-top; crushed for

fire break in 1971;

formerly climax black

spruce type

Creek bottom adjacent to

burned area

Ridge-top; adjacent to

s t u d y a rea 1 ; co n t ~ ·o 1

area; 70 year old black ..

spruce stand

....... - .... ., ... _ .. . . .,. ......... _," ·• ·- -------- ···-------·-·---------------------- --"\",_rt-,--~

k:i.!lllclii t

8t·o~·:~ ,_.

Sp(~cies

S. scoul(~riana

Alnus crispa

S. Elan i fo i a

I') subsp. ~_lchra

5. scouleriana

Alnus crispa

S. alaxensis

S . p 1 ani fo l i a

S. hastata

Alnus crispa

S. scouleriana

Alnus crispa

') .... .. - .. ' t """"' t (' i •I ()\•/') ~~ '·< '.JtH.J I 1.1 ;)

i{(;;:; I i ctl Le

i'i Ui!ii) e r Size

20 40 m2

20 40 111 2

30 2 m2

10 250 li12

P l~ , ~ e L G 1·o lJ p C li u (Jr u t s ) .. ~ ~ c ~ '·r. l,t; 1J .1 1 .ol. ....

~i UhiDe r

20

20

20

20

Si'ze ,

- -250 m2

250 Ill 2

2 250 l1l

250 m2

\\

=-"'1:~ ,._.,...,~---.,.~.... - --- ...

. ... .. ' ·' _, ____ ·-·--·-----~~-........_,......._...---____ ,,_,_,.....,.. ... ~ -~!-"~-- •"- _, ----:-""' ____ ~~-,.__--t.r--- -----:---~·-·--·~,. ,.,.._ •. ~ • ·- .. --·-- .... ~·- .. ~ -----

·---.--~--..,........, ---~.----..-------~ ... ..,.--~~--· .. -------··~·~·-- ... --..

'"\ ,,

.•:::

"'

N

~ ·- ~ -~-

_, ·'-···~ ... ~~" '"-

. . . " -' .... ~ - ·• ... -.... -- ·- • • > ~ . . . - . ..... ... .. .......... -·--·~ t-"•'- .........

c'f ~ ;"' ! ')... . ,,.: {. C" i ,., L ., I i '•' ! • ',.

•t I •.; ..... i I 1.1 1 :J~ ~ J ' " t • I J ,.,. •

f\vu.iiub1e B rov1s ecJ Production Utili zution Percent C11 i '":" ' .:_-:_;;

2 2 Study f\xea Species YeFl I" per rn per m (ksJ/ha) ( k ~i/ il i1) !3 !"0\'1 sed square

- . .

\ ". 8.8~: 1.93 0.08 ± 0 .OG 75.6 ± 30.6 0.7 ± 0.6 0.9±0.6 C::• \l

,. 1 s. 1977 999.36

. ' '

·. -•' 't .. ( .

scou1eriana 1978 7. 35 ± 1. 59 1 .22 ± 0. 3<1 69.9±21.0 11.7±4.2 .,G '± 4 .. I • I • I c::>.

2 5. 1977 1 . 76 ± 0. 95 + 14.9± 10.8 2.4±1.9 15.9 ± 7.8 0.061 0.28- 0.17

'" ,,~)

-

scou1eriana 1978 +- + 1.0 ± 1.0 o.2±o.2 16 0.3 0.13-0.13 0.02-0.02

... ,,~ 1977 6 .40+ 1 .87 1 . 71 + 0. 56 6 7 . 2 -t- 27 . 3 17.9+8.0 26 .6 -+ 3. 3 s.

419.46 ""

p 1 ani fa 1 i a 1978 -+ 0.48 + 0. 20 47.6 + 18.0 4.1 + 2.1 8.6 + 3.0

I 5.51_1.56

(' ..::t

..... (J

3 s. 1977 27.10:+" 5.37 10.20 + 2.18 213.8+69.5 -+- 37.7+4.3 i 3 .88 11 0. 2 _, 32.7

alaxensis 1978 14.94 + 3.08 7.50 -+-1.91 Ill . 9 -t- 34 . 1 56 .4 -~"19 .8 50 . 2 -4- 5 . 6 ' (,

s. 1977 12.60-l- 3.94 1.55:!:0.68 90.3-+ 36.2 11.2+5.8 12.3 + 3.2 6.12

p1anifolia 1978 11.37 ± 3.10 2. 98 +- 0. 72 81.3-t-29.8 21.2± 7.1 26.3 + 6.9

s. 1977 a .so± 2.67 o.55+o.23 N/A N/A 6.3-1-2.1 O.OOl,

hastata 1978 s .o5± 1 .39 0.32 + 0.12 N/A N/A 6.2-+ 1.6

- '

\ 1 .01 ± 0.27 8.6-t-3.8 I' 4 s . 1977 0 0 0 20.56

r lf}v -scou1eriana 1978 1 .48 ± 0.18 0.01±0.01 '13.9-+ 2.9 0.1 j: 0.1 0. 8 ·t- 0 .B

N/A- Sample size of DPG's was not sufficient to allow ca1cu1at·ion of biomass.

L ·--.. ~~---

- "~ ' ' --- ~---~--

) I ,___ I

l .. ¥

iaoi~ 3: C:~: -sr,.;"' ,, t.ooi• fu1· <'Viliuating sp·~cies preferenr,e of study are~s 2 and 3. ' J ·-·· ~ .... ,_ ......... , ~< ...... ·-.. - - ---....

·--- -·--·· ·-·---.. ,;<0 --- . - -------------- - .,_.;.

r ,jt

.:.·±.:.

Observed Expected Observed Expect~d Proportion

,;<"'

i ""'

.~ No. Twigs No. Tvli gs Proportion BrO\·Ised +-95% 'Chi-)

.---, Study Area Year Species 8 rov1sed Brow sed Browsed Confidence Level square ·0

*"' " 2 1977 S. pl ani fol 1.§_ 124~~. 34 1362 0.785 0.859 0.020 51 ~65

S. scouler·iana 341 . 66 224 0.215 0.141 0.020 I ... ,~,,-_._ .. ~ .......... -~

,,

1978 S. p 1 ani fo l i a 386.61 380 0.976 0.956 0.023 4.76. S. scouleriana -? 9.39 16 0.024 0.040 0.023

** 3 1977 S. alaxensis 412.14 612 0.558 0.829 0.033 223.99

S. QJanifolia 191 • 86 93 0.260 0.126 0.029 S. has ta ta 134.00 33 0.182 0.045 0.018

** 1978 S. alaxensis 280.09 450 0.476 0.765 0.042 205.22

S. Ql ani fo 1 i a 213.19 119 0.363 0.202 0.040 5. has tata 94.72 19 0. 161 0.032 0.017

** -Significant at the .001 level or less.

I)

L l--::..,. ..,.__....., • ...__ ___ ...,....,.,_._..----~---...,..,---- ~~ ----·----- <~·~~. ,.------- < _ ..... .., ... ~_, ~----~- __ • .., ___ ~~ ~------·-~-----~ -~

( ~:

--~ .,~--------·-~,~~------------.--

--··-......---<. ·-

. 1 . • ' I ' I d l • t; . WiJ; .Jt~ .. i :y-;;'·" .:,.t:.',,,: .~~· 1 udl"L! Lrtti.r:: tn.~ fl(1CI1 specieS ii~'•Li s~...uc~y dreil \COntir.ueo on r;~:xt _,_ ... ---- --.-...... ,_,~-~4--·--.-------------...... -Categotyc Expected 0 (l d ··Onserved '

Study. of LXJ-1eclec.J no? Oos e rved no. d proportion proportion

area Species stem stems browsed stems t)rO\·Jseci browsed browsed G-s ta t i s t i t; .

1 s. 88 1 . 96 5 0.109 0.278 0.265 * 9.603 seoul e ri ana uu 9.75 4 0.542 0.222 0.245b '=l

. Ul3 3.09 3 0.172 0.167 (J "?Q

'' "- L..

13U 3.20 6 0.178 0.333 0.277

I 0.154 b ** I 2 s . BB 8.07 • j 19 0.139 0.328 17.597 -

scouleriana uu 34.57 29 0.596 0.500 0.164 "

UB 11 .14 7 0.192 0.121 0.107

BU 4.23 3 0.073 0.052 0.073

s . BB 21 .19 42 0.065 0.129 0.046b 77.431 ** pl ani folia uu 213.59 142 0.665 0.436 0.069° '··'

UB 9.94 9 0.030 0.028 0~. 022

BU 4.23 3 0.249 0.408 0.068b ;J

* Significant at the .05 level

** Significant at the ~001 level --:?

C.-(_~?·

L ;) ~---------~~~-~~-=-~-~-~---------_:.::"· --- ____ ,_. ·=-···-----~-......,. ___ ....._tiiilllliiMil ............................. ~

c;

-"'

1~\.· ·t .. ~; ·~ 1 ·i;; ~Jt

:J ,;~~ :•

"' t ~

I

I

••

I I I I ~

•

L · . .,

··-···------ ·-·--~--·~-~'-·- - ·----------------· -~...,_

0

~:·

Tab l e 4 : 1\ v a i 1 a o i 1 i t y - u t i"l i z ll i ·i on c n 1 - s q u c1 r e t iJ b i e f o r e a c h s p e c i e s an d s t u c1 y a r e a ( co n t i n u e d) • ·

Categoryc Exp8ctedd (1 d Observed ' · n ') CD

Study of Exrected no. d Ob~ervPd no.d proportion proportion ._:.,

•

a rea Species stem stems browsed sterns browsed browsed browsed r~

!'"' t.- . ',\ ,~l:t~~J&-J_s_t 1 G \::, - - ---··-~ ---::::._~ .. :::::::.;::::~;::.::::::::::·~~-~ ~.--,::~~

3 s . BB 60.32

alaxensis uu 40.21

UB 48.38

BU 15.08

. s. BB 0 -

p1 ani fo 1 i a uu 41 . 69

UB 5.41

BU 1 .90

0

87

25

33

19

0

35

10

4

0.368

0.245

0.295

0.092

0

0.851

0 .110

0.039

0.530

0.152

0.201

0. 116

0

0.714

0.204

0.082

0.097b **-23.462

0.070b

0.078b

0.063

0 7.288

------·------------------------------------·----------------------a Observed proportion browsed 95;. pe-rcent confidence interval

\:-~

b 95 percent confidence interval indicates the observed proportion is significantly different from the expected .

c BB corresponds with stems browsed both seasons. UU corresponds with stems unbrowsed both seasons. UB

corresponds with stems unbrowsed the first season (1974-75 winter) and browsed the second (1975-76 winter) ..

BU corresponds with stems browsed the first season and unbrowsed the second.

d Columns show observed and expecte~ frequencies of stems browsed for the 1976-77 winter.

----~=-..., ~~- ,,~,--....-.._,,_..._ . .,, ___________ ~.,_.------ ..... .,... .._.,__..., " ~'

___ :!.:.....:__"'"''' i!.."> ._,. -- ,..._ _ __ , __ ..,,~, "-""'"- ..,,, __ ~

., '

Table 5: 1\vo by two contj~.ngency tables illustrating between-plant

--·· .. ----------·•*'·'-' -· - ....... ·----~·---

S, p_l ar1_i_fg_l_i __ ~ No. of plants No. of plants rwt

plants ._ ___ -~.!_ St~ browsed during brow:;ed during

P .. l-ea 2: 1976-77 v,ri nte r 1976-77 Hinter ---"---· ........ _ .. ,_ •• ---·---.......,.__, ____ ... - '<•- -~· ----~--- ·------·------- --

- ·------~ ·-----------No. of pl a·nts

br·c\·::r=d during

h ~: . of pi c n t s not

winter

- d f'\1 ' I ... ~~.;. ~- :..n' s · _:~-:. ... _ ;_~~-~-·

:~(... 0f plants

'· .-,. ··st:.d u" -.,Jrt - during

·, ~ns -75 v-d nte r

No. of plants not

t.rowsed during

1975-76 winter

*"'"

Obs.

29

10

27

21

Probability less than 0.001

,..Yp L .• • Obs. Exp.

21 . 59 9. 41

17 .41 15 7.59

0

24.00 13 16.00

24.00 19 16.00

-

-

\)

Chi-

sq Lia re

** 18.766

1 .875

l l \

\ r

0 ·,.

,;) 't O:.::l ..

'• 0

Table 6: Protein content for each species and study area.

f;----~--·-----··-- ---·-··-·---- -·-----------

Study Area Protein content (percent .±. standard error)

S. seoul eri ana S. E_l_a_nj_f.o l i a S. alaxensis

1 5.6 0.2

2 5.3 0.2

3 N.D.

4 5.4 0 .l

a Data not available. o

__ ,. -

5.4

a N . D.

0.4

N.D.

N • D.

N.D.

N • D.

6.0 0.1

N.D.

• '" .•• 1

\\ I

,.,,,

XI Table 7: Protein cont~::nt of the tagged ,plants.

""' -··----..- _ ..... -~------ .. ~-_, ____ _.,.__....., __ ... __ ~-----.. ----·--------

Snt::cies Plant pair Pr·ot~::in content (percent) . ·~ ·- -- ----~ _ .......... - .... -~ ... ,. -t -···---------- .... -- ____.. ... ~ .... ------,,--,--

Plants brov.•sed Plants unbrovJsed

both seasons both seasons

................. ----·~· ... .,.. .. __ -· ·----...... ~ .. ---<' ------

5. 1 5.6 5.2

~:· 1 :.:~ n i f.J l i a ...... - .... _.., --- 2 5.9 7.2

3 5.0 ' 0

4 4.9 5.5

5 5.7 5.3

6 5.8 4.8

s. 7 6.8 6.7 a

~coulsriana ·-. _,_,... ___ 8 6.3 6.5

9 6.5 6 .1

10 6.2 6.6

ll 6.0 5.6

;;

---

c

\\

\

c

-

l J

\

1 ~

1 ; l I i ,,

r r t:

r r

~ i!f·.$-

'~ ---" - ~ ~ c:::::1 L __ :J r::==J c:J ___c::]_=~"c=J. [=:1 ·l['.;;i:J · ""' --··-- · · ·-··

' .. ~11 , ___ , t"' __ .,,.,.,..., ,----, ., . ,..........;. .. ,

Table 8: Pellet group d:ta and comparison of bvo methods for calculating moose use.

No. of pellet fv1. 0 ./ ha a M.D./haa

groups per ha (Forage biomass (Pellet group

Study area Year (+standard error) data) :. data)

-·.~ .. -, •• ";_";;;¥~:

1 1977 6.0 6.0 0.14 0.46

I I I 1978 42.0 7.2 2.34 3.23 ('J

-y ¥

' 2 1977 20.0 8.8 4.06 1 • 54

1978 14.0 5.2 0.86 1 .08

3 1977 50.0 10.4 24.28 3.85

1978 52.0 10.0 15.52 4.00

4 1977 0 0 0

1978 6.0 4.4 0.02 0.46

L a Moose-days per hectare

n Cr

. -';)

;::,

#

"

t;;,

[\. \f? v "'

uo

·I

. -, ·--..,_:

' . . ..... _,_, __ ., _____ ~,_ ...... ....,.,__...._ __ ~,..,-~ ~-·--~--·--~-----·--..----~-~~-----~--.........-~~---~·~-----------.-.......... ._..._ ....... -'~·~''''

........ bf .. 1\t

:r -;:;, :! .

0

~)

' ' --

I

* ~

I

-I I I

~

0 0 l

[j I

!

q i

D

[

c 0 0 D

0

0 ()

10 --.~

\ •

•

• • . ~ • G . ,. . . ..

'\ --+ -f

10 20 30 40

~- c .x.,

Fig. 1. Linear regression relating the 1976-77 winter ~rowsing to availability for the portion of plant not

' browsed during the 1975-76 winter. x1 corresponds to the number ~f stems per plant not bra~sed during the 1975-76 winter and available during the 1976-77 winter. Y1 corresponds

to the number of stems per plant ~o.~ <browsed. during the 1975-7.6 winter and browsed during the 1976-77 winter. The slope (b), intercept (c) and determination coeifficient (r 2 ) are shown

atove. The symbol 0 indice.tes more than one sampling poin+J

occurs here.

-

40

30

20

0

10

• •

10

0

II

20

X 2

30

b-::: 0.80

a = -0.54 r 2= 0.90

40

Fig. 2. T.inear regression relating 1976-77 winter browsing to availability for the portion of plant ._ .....

b~~wsed 6uring the 1975-76 winter. i 2 correspo~ds to the

:;I

I

number of stems per plant browsed during the 1975-76. winter and available during the 1976-77 winter. Y~ corr~sponds to

t::

the nu~ber of stems per plant browsed d~ring the 1975-76 winter and browsed during the 1976-77 winter~ The slope (b),

intercept (c) and determination coeifficient (r2 ) a~e~ sho,..,·n above. (The symbol 0 indi ca-ces more t a:n one sampling point occurs here.)

-

-

;·o.··-1 ' ! . l