HomeMy WebLinkAboutAPA948ACKNO\JLEIJGEHE1YrS
I would like to take this opportunity to express my
appreciation to those who assisted me during the course of
this study.
Th~ guidance, interest t:~nd critical revie\ .. )f drafts of
this manuscript by my major advisor, Dr. Samuel J. Harbo,
is greatly appreci<:~ted. I would also like to thank my other
committee members, Drs. David R. Klein and Jerry 0. Wolff,
foe their words of encour~gement and critical revi~w of this
m~nuscript. Sincere thanks are ex~cnded to the other
graduate students who expr8ssed interest in this study and
often offered words of encouragement.
A special worJ of gratitude is extended to Drs. John L.
Old.::meyer and Hayn2 R~gel in of the D.:n vee Nild life Reset:~ rch
Center, Kenai Field Station, who.were most helpful in the
design, logistics and data analysis segments of the study.
\
Their experience in .moose habitat evaluation greatly
improved the quality of this study. Thanks is extended to
the U.S. Fish and Wildlife Service for providing needed
employment and logistical support during the course of this
study. The financial and logistical support of the Alaska
Cooperative Wildlife Research Unit, University of Al~skn ~nd
the Institute of Northern Forestry, Fairb~nks is greatly
appreciated. Special t~anks is extended to Dr. L.A. Viereck
iv
TABLE OF CONTENTS
Introductit.")n............................................. 1
Study Areas ...........•. 3
Slikok Lake Study Area. . . . . . . . . . . . . . . . . . . . . 3
Harathon Road Study Area ....•••.•....•••.•. 6
' ) ~l!jicki~rshanl Study Areu .............•.................... 7
t'lethods and Data Analysis. 10
Locatin-g Study Sites ............. . 10
Browse Production and Utilization. 15
Evaluation of Within-species Preference .....•.•.•...... 20
' )
Pellet Group Counts ............••.••••..•..•..•....•... 21
Tracking .•...•••....• . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
-Protein Conteht S~3mpl in::1.· ........................•..... 22
Snow Depth Sampling. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Statistical Approach .............••............•...••.. 24
Discussion of Results .. 26
Bro~/Se Production and Utilization. 26
Browse Species Preference ...•....••...•....•.....•.•..• 32
Within-species Preference .............................. 35
Pellet Group Counts •..••••.••.••.•..••••••..••.•.•.•..• 46
'l'racking ..•....• . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Protein Content ...........•••........•................. 51
Snow De p t h . . . . • • . • • • . . . • . . . . • . . • . • . . . • • • . . . . . . . . . . . . . . . 5 8
vi
vii
Concluding Discussion ...•........•..•.........••..•..•..• 60
BroivSe Production and Utilization ...........•••....••.. 6~J
Protein Content ........................................ 67
Factors Influe~cing Betwe2n-species-Preference ..•....•• 69
F~ctors Influencing Within-species Preference •••••.•... 75
Management Implications ..•..•.............••...••........ 86
hf>pen:J.i:x I ••••••••••••••••••••••••••••••••••••••••••••••• !39
Appe11dix II ... ·'· ......................................... 91
Literature Cited ......................................... 92
LIST CF FIC:UF.E2
Fi;ura 1: Lnc~tinn of the ?likok La~~ stu1y area (L)
~n·3 tl1o t1t:rothnn F.nr:·.J stu:.~y area (G)................... 4
Figure 2: Location of the ~7 i·=!-:-:rshai""\ hur.n stu::'iy 2.rea
(C) ••••••••••••••••••••••••••••••••••••••••••••••••••• ~ 2
Figure 3: Linear regres~ion relating the 1£76-77
winter browsing to availability for the portion of
plant not browsed during the 1975-76 winter. Xl
corresponds to the number of stems per plant not
browsed during the 1975-76 winter and available
during the 1976-77 winter. Yl corresponds to the
nucb~r of steres per plant not brnws~d during the
!~75-76 winter anj browsed during the 197~-77
wintar. The e~uatinn of the r.egr~ssinn line and
the ~ctermin~tion coefficient (r ) are also
shown. The dotted circle indicates that more th3n
one sa~pling point occurs here .••••••• ~ •••••••••••••••• · 42
Figure 4: Linear regression relating the 1976-77
winter browsing to availability for the portinn of
plant br.otr~sed :Juring the 1975-7(:: ~-iinter. X2
corresponds to the nu~ber of steT<s per plant
browsed during the 1975-7~ winter an~ avail9ble
during the 197~-77 winter. Y2 corresponds to the
number of ste~s per plant browsed during the
1975-76 winter and browsed during the 1975-77
w~nter. The equation nf the r.egr~ssinn line and
the determination coefficient (r2 ) ~re alsn
shown. The dotted circle indicates that more than
one sampling point occurs here......................... 4L~
viii
TCJble lt:: 7rac:~ i.n-J .!a tA for shrubs brn•.·JS~.:-: over ::;
r_::!erc·2~t .3t ti1~ ~~i~!-:·:=rsharn stu:~y :~!-::·=··················· 5:3
\
Table 15: ProtGin c~ntent fnr ~ach S)2Ci~s and sit~
at the ~2nai Peninsul~ stu~y ar23s •.•.•••••••.....•..•.
Table 16: Protein content for eac~ s~ecies an6 site
a t t h (~ ~ ·; i ~ h: r:? r sham s t u d y a r 2 a • • • • • • • • • . •••••••••••••••••
T~ble 17: Protein cnntent of the tagged plants at the
t,.; i c k e r sham stu ·:1 y a r e a • • • • • • • • . • • . • . • • • • • • • • • • • • • • • • • • • . 5 7
Table 18: Snow depth measurements for each site at
the Wickersham study area ..•.................•.........
Table 19: Drowse cnnsum9tion an~ pellet grnu~ density
as indices of. moose use at the ~~ick2rsh3::1 stu:iy area •••
Table 2~: Shrub height .:1ata for Sites l and 2 at the
~arathon Foad study area............................... 71
Table 21: Comparison of twig ~ro~uction during t~e
1976 gro~ing season for stems browsed ~uring the 1S75-
76 winter and those not browsed at the Kenai Peninsula
study areas............................................ 79
\
Table 22: Comparison of twiJ 9roductinn during the
1975.growing season fnr stems browse1 during the 1~7~-
76 winter and those not bro~sed at the Wickersham
stu ... :ly ~rea •....•.............•...........•..•..... ".... 81
Table 2~: Cbserve~ distribution of pellet grou?s
fitted against the expected ran6om distribution for
sites 11 and 13 .............................. 0 •••••• Cl • • e. 4
X
)
(
!
2
comm0nly found in random measurements of browse production
anJ utilization. Milke (1969) suggested that the probability
of utilization associated with a browse plant may be relQted
to tlle speci~s composition of brmvse plants in th•: immediate
vicinity. A browse plant of a species with low preference
may show a higher than expected probability of being browsed
if it is surrounded by plant species which are highly
preferred.
The prcsant study was designed to datermine whether
moose show between-species and within-species preferences
among willows in two areas of Alaska with. very different
moos~ population densities. Protein content as a possible
factor influencing preference was also investigated. Dat~
were collected in 1977 and 1978.
Fig 1. L()cation ()f the Slil~ok Lakt? study ar~a (A) and the
Marathon R()ad study area (B) •
\ _,
--------··~--~-·--... ·'
8
Fig. 2. Location of the i'iick•=rsham burn study ur.eu (C).
~~ble 1. Dcscri~tion of sites and sampling quadrats for the three study areas.
Site Description
Dominant
BrO\'ISe
Species
BrovJSe Quadrats
Replicate
Number Size
Pellet
Group Quc.:dr<:~ts
Repl ic<l te
Number Size
------------------------------------------·--~---·------·--·-----------------------------
1 LOVJland; r-larathon R. ,, ~~!:.£!.~Y.!. 20 10 m2 30 lG m2 b.
study are.:.; burned in •' arbusculoides ;) .
1969 .Jnd uncrushed; i? • trciiili rorcr;;-s--------------for.merly bli:ick
spruce climax.
2 Lowland; <·1a r a thon R. s. £~££!.~Y.!. 20 10 m2 30 10 m2
stuay urea; burned in s. arbuscu.loides
1969 and uncrushed; P'. tremufoid'es--
formerly black --·----------
spruce climax.
3 Upland; Slikok Lake s. Barcloyi 20 10 m2 3" 10 m2
study area; burned in ,. se-s5Tnna u•
1926 and crushed in s. ITarcTa¥I
early 196!:J's; formerly P'. tremufoid c s
\oJh i te S!:)ruc~ climax. 8. P:~ei~If:££~-
4 'Upland; Slikok Luke s. scouleriana 20 10 m2 30 10 m2
study area; timbered IT. e~elEI£££~-a reo adjacent to Site
3 i white spruce/birch
mixture.
---------------------------·--------------------·--------------------------------------
. ~'·')',l.'i' ~~ .• , •• ·-~ •. , :··. ·~ J
Table 1. Description of sites and sampling quadrats for the three study ~re~s
(continued).
Site
5
G
7
8
D•=scr ipt ion
Upl~nd; Slikok Lake
study area; unburned
and crushed in early
l960 1 s; formerly white
Sf>ruce c.lim~x.
Uf>land; Slikok Lake
study area; timbered
area adjacent to Site
5; white spruce/aspen
mixture.
Lowland; Slikok Lake
study area; unburned
ctnd crushed; formerly
black spruce climax.
Lowland; Slikok Lake
study are~; timbered
area ~djacent to Site
7~ black spruce climax.
Dominant
l3rov/Se
Species
S. bebbiuna
s. scouT2rTana
s . a.3rcT5¥r---
p. tr8"muToia e s
S. scouleriana s. n~r.~ra-y-r--s. se-G"sra-n-.:1
s. e:~e~~I~~£.£
Browse Quadrats
R2plicate
Humber Size
20
20
20
20 10 nf
Pellet
Group Quadrats
Ref)lic.Jte
Number
30
30
30
30
Size
10 m2
10 rrf
10 m 2
-------------------------------------------------------------------------------------
'l'.:.ble l. o~~scri;;tion of sites and samplin~J quadrats for the thrf:e study arcns
(t.:ontinued).
Site
10
11
12
D~scription
Upland; Slikok Lake
study are~; burned in
1926 and crusncd in the
early 1960's; formerly
~hite spruce climax.
Upland; Slikok Lake
study area; timbered
area adjacent to Site
9; white spruce climax.
Upland; Wickersham
study area; burned in
1971; formerly black
Si.)ruce climax.
Upland; crushed for
fire break in 1971;
forillerly bl~ck spruce
cl i1nax.
Domin.:lnt
i3rowse
Speci0s .
S. scou.leriana s. sarcTayT ___ _
8. oapyrifcru
~---------
Browse Quadrats
Replicate
Number Size
20
20
20
Pellet
Group •.Juudrats
Rcpl ica te
Number
30
30
20
. 20
Size
250 r.?
I-' w
)
distally from the cut ~nd each twig was placed into a
diameter class. Each diam~ter class was 0.5 rnm wide with
classes ranging from 1.1 -1.5 to 4.6 -5.0 mm. The dpb
sam?le of each species and site in question was also
partitioned into diameter classes. A sample of twigs for
/
each species and site suitable for analysis of protein was
collected possessing either an identical or proportionately
similar diameter distribution as the dpb sample~ The samples
were oven-dried, ground through a 40-mesh screen and
an3lyzed for crude protein by the macrokjeldahl mathod (AOAC
1960). Four replicates per sample were analyzed.
Since the twig quantity was not a problem at the
Wickersham study area, the distribution of twig diameters
was identical to the dpb sample. Three samples instead of
one were usGd for each species and site to permit the
cal~ulation of variance. The samples were analyzed for cruJe
protein level as above except the number of replicates per
sample WdS three. For all study areas, only current annual
growth twigs were collected.
The above methodology represents an attempt to assess
the protein level of browse consumed by moose. For those
spe~ies where second-year growth is eaten in addition to
current annual growth, the results may be misleading. A
similar probl~m was discussed earlier concerning the Shafer
twig-count method. Since the second-year growth twigs
prob~bly contain less protein per unit weight than the
23
' )
' )
current ~nnu~l growth (Cowan et al 1970), the results
obtained may tend to overestimate protein levels of browse
consumed by moose for applicable species.
During 10-11 April 1978, twi3 samples were collected
from the tagged plants ~t Sites 11 and 12 for protein
analysis to determine whether protein level is related to
browse history. Only plant pairs where one plant was browsed
during both the 1976-77 and 1977-78 winters and the other
plant was not browsed both winters were sampled. Thirty or
mGre current annual growth t'.'ligs 3.0 mm in diameter clip!?ed
from each plant constituted a sample. Replication was three
per sample a~d the protein analysis followed the procedure
outlined above.
On 10 March 1978, snow depth was sampled at the four
sites at the Wickersham study area. Fifty me~surements were
taken at each site in a random s~mpling procedure. Since
thaws were rare for the Interior for that winter and
snowfall after 10 March was minimal, the snow depths taken
were assumed to be the deepest moose would face during that
All statistical proc~dures were according to Zar (1974)
and Freese (1962). In situation~ where I suspected the
~
)
25
assumptions of normality were not justified, nonparametric
tests were preferred over their parametric counterparts. For
goodness-of-fit tests, the chi-square statistic was used.
If expected frequen~ies tended to be small ( < 5.0), the
lo3-likelihood statistic (G) was used instead. _If any
expected frequency fell below l.G, the test was not done.
Significance level for all statistical tests were set at the
0.05 level.
)
'l'abl0 ..,
,.) . Browse production and utilization for the Kenai. Peninsul.:t .study nreas
(~ standard error).
--------------------------------------------------------------------------------------
•.rotal 'I'w ig s 'l'o tal 'l'wig s
Available Browsed Product ion Utilization Percent Chi-a:
Site Species Year per ~ ocr .. m2 (kg/h~) (kg /ha) Bro•,.,scd squure
------------------------------------------------------~--------------------------------
1 Bare. 1977 6.09 1. 39 0.56 0~17 72.4 22.6 G.7 2.5 9.2 1.9 14. 4 7b
1978 4.32 1.14 0.21:) 0.03 51.3 17.9 2.4 1.1 4.6 1.6
3ebb. 1977 0.35 lJ. 29 0. Hl :J.08 2.7 2.6 0.8 0.7 23.7 1.6
Arbu. 1977 2.19 1.32 0.65 0.38 50.2 38.8 14.9 11.2 29.7 4.5 123.34b
1978 4.44 1.18 0.26 0.16 102.2 3G.1 5.7 4.2 5.7 3.2
2 Burc. 1977 20.12 3.19 2.97 ~L 74 247.4 55.4 36.5 11. 4 14.7 3.1
1978 20.55 3.87 3.05 0.88 252.8 ()3.2 37.5 13.5 14.8 3. 1
Bebb. 1977 8.18 (:).18 ,, . 03 tJ. 03 1.4 1.6 0.2 r.J. 2 13.9 0.1
1978 ~i. 0 6 0.04 0 0 N.D. N.D. 0
c'\rbu. 1917 1. 51 0.61 0.62 (!). 29 22.2 12.0 9.U 5.4 40.7 7.6
1978 2.41 1.10 0.47 0.31 35.4 21. (J 6.8 5.5 19.3 6.2
3 Scou. 1977 8.91 1. 4~ 5.74 1. 03 204.9 59.1 132.0 39.9 64.4 2.5
Bare. 1977 0.23 0.14 O.OG 0.06 N.D. N.D. . 26. 1 12.4
a Chi-square was used to test for differences in percent utilization between years
for a site und species.
b Signific~nt at the .05 significance level.
t:i.01
30.78b
.I
Table 3. Drowse production and utilization for the Kenai Peninsula study areas
(:t standard error) (continued).
Site Species Year
Total Twigs
Availab~c
per m
To t a 1 '1"11 i g s
Browses
per m
Production
(kg /ha)
Utilization
(kg./ha)
Percent
Brows~d
Chi-a·
square
---------------------------------------------------------------------------------------
3 I3ebb. 1977 0.90 0.35 0.51 0 .• 20 11. 3 5.5 6.4 3.2 56.7 4.6
4 Scou. 1977 0.32 0.12 9.10 0.08 9.6 5.6 3.1 2.9 31.7 17.2
5 Scou. 1977 0.17 0.10 0.05 ~~.114 N.D. lJ. D. 30.3 11.4 13. 56
1978 0.06 0.04 0.03 0.02 N.D. N.D. 45.4 9.6
Bare. 1977 IL 66 0.36 0.09 0.07 N.D. N.IJ. 13.6 6.1 e. 4 4b
197SJ ~.72 11.32 0.03 0.02 N.D. N.D. 3.5 1.8
3abb. 1977 2.51 0.49 0.54 1L14 36.3 11.9 7.8 3.1 21.5 3.9 15. 79b
1970 4.11 ll.6:?J 0.52 n.o9 52.5 12.5 6.6 1.7 12.5 2.3
,.. Scou. 1977 0.14 0.07 0 N.D. N.D. 0 IJ
1978 0.05 ll. 0 3 !:J.Ol 0.01 N.D. H.D. 10.0 6.3
l3arc. 1977 0.58 0. 24 !L04 fJ. 02 N.D. N.D. 7.0 4.3 8. 59 b
1978 l. 24 0.70 0.02 I.J. (ll N.D. N.D. 1.2 ~L 9
Babb. 1977 0.23 0.09 0.01 J.Ol N.D. N.D. 2.2 1.9
1978 ~.69 !3. 21 0.01 0.01 9.1 3.5 N.D. 7.1 4.7
--------------------------------------------------------------------------------------
achi-square \'las used to test for differ-ences in percent utilization be t\veen years
for a site und S?ecies.
bsignificont <Jt the .05 significance level.
.
,,
~ ~ ,,
t·.
r:· 1\)
())
TQble 3. Browse production and utilization for the Kenai Peninsula stujy areas
(:_t Gtandard error) (continued).
--------------------------------------------------------------------------------------
·rotal •rvJ ig s Total '1'\-iig s
Available Browsed Production Utilization Percent Chi-m
Site Species Year per m2 per m2 (kg/ha) ( kg/ha) Drmvsed square
--------------------------------------------------------------------------------------
9 Oebb. 1977 2.87 8.65 1. 42 0 .• 42 27.8 8.4 13.8 5.2 49.6 6.3
1978 2.79 0.41 0.72 0.15 27.1 6. 4 7.0 2.1 25.9 2.9
Hl Scou. 197i ;).09 0.01 0.04 l1. 0 2 1.3 0. 3 0.6 0.4 47.1 20.6
1978 0.07 0.05 I} (3 ~.9 0.1 f1 0 0 0
Bare. 1977 0.24 l). 20 0.02 0.04 N.D. N.D. 6.3 1.8
a Chi-square was used to test for difference!3 in percant utilization between years
for a site and species.
b Signific~nt at the .05 significance level.
41.92 b
w
0
'i'.:t bl e 4. Browse production and utilization for the Hi.ckersharn burn stuCly arecJ·
{:t_ stund~rd error).
-----------------------------------------------------------------------------·---------
'l'o ta 1 'l'\v ig s 'l'o tit 1 T\·i ig s
Av.::~ilable Browsed Production Utilization Percent Chi-a
Jite Species Year per m2 per m2 ( l~g/hu) (kg/ha) Br ov1scd square
---------------·--------------------------------------------------------------------..... ---
11 Scou. 1977 8.89 1. 93 0.08 "~06 75.6 30.6 0.7 0.6 0.9 0.6 999.36b
1978 7.35 l. 59 1. 22 0.34 69.9 21.0 11.7 4.2 16.7 4.1
12 Scou. 1977 1. 76 0.95 0.28 0.17 14.9 Hl. 8 2.4 1.9 15.9 7.G 0.06
1978 0.13 !1.13 0.02 H.02 1.0 l.C !3. 2 0.2 16.0 0.3
Plan. 1977 6.40 1. 87 1. 71 0.56 67.2 27.3 17.9 8.0 26.6 3. 3 419~4Gb
1978 5.51 l. 56 0.43 0.20 47.6 10.:1 4. 1 2.1 8.6 3.0
13 Alux. 1977 27.10 5.37 10. 2~j 2.18 213.8 69.5 11a.2 32.7 37.7 4.3 13.88 b
1978 14.94 3.08 7.50 1. 91 111.9 34.1 56.4 19.8 50.2 S.G
.f>lan. 1977 12.60 3.94 1. 55 0.68 99.3 36.2 11.2 5.8 12.3 3.2 6.12
1978 11. 37 3. Hl 2.93 0.72 81.3 29.3 21.2 7.1 26.3 G.9
iiust. 1977 8.80 2.67 IL 55 0.23 N.D. N.D. 6.J 2.1 0.01
1978 5.05 l. J9 g,J2 ~). 12 H.D. N.D. 6.2 1.6
14 Scou. 1977 1. 01 0.27 0 0 8.G 3.8 0 0 0 {] 20.5Gb
1978 1. 48 0.18 0.01 0.01 13.9 2.9 ILl 0.1 . ~j. 8 0.8
achi-square was used to test for differences in percent utilization between years
for a site and species.
bsignificant at the .35 significance level.
= w ,.._ _ . ........_ ......... _______ _ J
)
T~blc 7. Two by two contingency tables for within-speci2s
preference at the Ken~i Peninsula study areas.
Sit~ Species
1
2
3
4
5
7
9
s.
Bare.
, .. ., .
arbu.
s.
Bare.
s.
arbu.
s.
scou.
s.
scou.
s.
Bare.
s.
bebb.
s.
scou.
s.
be5b.
s.
scou.
Bro~;sea·
History
Status
of St:ems
6
u
u
B
u
B
u
B
u
B
u
B
u
B
u
B
u
B
u
B
u
No. of Stems
Brov1sed During
19 7 6 - 7 7 ~'Ji n t e r
Obs.
31
16
125
122
34
lv
226 .,
.)
4
1
6
2
41
3
11
0
3 ..•
'J
298
23
Exp.
12.2
48.8
21.7
25.3
58.5
lCD.S
2G.3
17.7
22G.2
8.8
2.9
2.1
3.0
5.0
39.8
4.2
7.1
3.9
1.7
1.3
284.1
37.0
No. of Stems
Not Bro\·/3ed
Durin:J 1976-77
~Hnter
Obs.
23
151
13
32
67
496
12
21
25
7
G
6
3
13
64
8
27
21
22
19
94
28
Exp.
34.8
139. 2
19.3
22.7
133.5
429.5
19.7
13. 3
30.8
1.2
7.1
4.9
6.0
10.0
65.2
6.8
38.9
17.1
23.3
17.7
108.0
14.1
G-b
statistic
17.517
16. I} 4 !·i
133.879
13.378
19.259
1. 395
7.315
o.Gl2
7.424
2.447
19.384
a i3 = Brov1sed durin') the 1975-76 wintG:r. U = Not browsed during
the 1975-76 winter.
b Value of G at the .05 si~nificance level is 3.841
37
'1''-!ble 7. (continued). 'l'\vo by t\·Jo contingency tables for
within-species preference at the Kenai Peninsul~
study areas.
----------------------·--:.---------------------------------------
Sit,~ Species
9 s.
Bare.
s.
bebb.
Browst::a
History
Status
of Stems
B
u
B
u
No. of Stems
Browsed During
1976-77 ~Vinter
Obs. Exp.
4 3.8
4 4.2
78 68.1
1 10.9
No. of Stems
Not Bro'.·JSed
During 1976-77
~Hn ter:
Obs. Ex!?.
8 8.2
9 8.8
22 31.9
15 5.1
G-b
statistic
0.019
32.389
aG =Browsed durin~ the 1975-76 winter. U =Not browsed during
the 1975-76 winter.
bva1ue of Gat the .~5 significance level is 3.841
r
'
constructed for each species and site (Table 8). The
categories used are:
BB = Numb~r of stems browsed during the 1974-75
and 1975-7G winters;
UU = Number of stems unbrowsed during the 1974-75
and 1975-76 winters;
ua = Number of stems unbrowsed durin3 the
1974-75 winter and browsed durin3 the
1975-76 winter and;
BU = Number of stems browsed during the
1974-75 winter and unbrowsed during the
1975-76 winter.
The G statistic was again used due to an occassional low
expected frequency value.
In 4 of 5 species/site combinations, G is significant.
In 3 of the 4 significant combinations, category BB stems
were browsed more than expected during the 1976-77 winter.
Also, in 3 of the 4 combinations, category UU stems were
browsed less than expected during the 1976-77 winter. In 1
of the 4 combinations, category U3 stems were browsed less
than exp2cted, and in 1 of the 4 combinations, category BU
stems were browsed more thQn expected. The pattern suggests
a decided preference by moose for stems browsed yearly.
To ascertain whether selection is occurring within
plant, between plant or both within and between requires a
~~ble B. Avail~bility-utilization chi-syuare tables for evfiluuting
within-s~e:cir.:s fJl'·::ferr:nce ,:_ti: trw ~hcl~~!rsh:>m study .:H.::.::;. Exp•.:ct0d
fruqu~nci.::s and proportions ware b~sad upon the r~lative
quantities of stems avai1~ble from 0u~h cdtegory.
Cdtegory b I.;>efA~cted r· .. o .c
of of Stems
S i tc Species Stem Drowsea
Cbscrv-:d No.c Exp~cted
of Stems Proportion
Browsed Browsed
CJbserveaa
J? r o ~or t ion
Bro·wsed +
95!6 c.r:
G-
statistic
-----------------------------·-·-------·-·---·---------------------·----------·----
11 Scou. BB 2.~ 5 0. 11 0. 28 ~j. 27 9. 6 c;d
uu 9.8 4 U.54 ~:). 2 2 U.25
uo 3. 1 .., 0. 17 ~). l 7 0.22 ..J
DU 3.2 6 0. ] 0 .u 0.33 ,;. '),... u • .c.O
12 :.Jcou. co 8. 1 19 0.1;1 !;i. J 3 0.15 17. 60d
uu 3•L 6 29 u.GfJ iL 50 O.lG
UB 11. 1 7 \1.19 0.12 0.11
BU 4. 2 J 0.0/ ~1 • [) 5 !.1.07
Pl.:!n. BC 2J • 2 42 0.07 0.13 0.05 7 7. 4 3d
0U 21J.G 142 :.}.67 u. 44 0.G7
UB 9. 9 9 n.o3 !L ljJ I;;. 02
BU 4 .. 'J i:J r, C:: 0.41 U.l.:l7 • L. ..; I • ~,".)
aGbs2rved prc)portion bro~vsed .::!: 95 percent confidence intcrvul.
b DB = Stems browse.j during the 1S74-75 an:l 1975-76 vJinters. UU = Stems not
brovi5t.?CJ Juring ti:ic 1974-·;s and 1975-76 winters. UB =Sterns not browsed during
the 1974-15 winter and bro~sed during the 1975-7G wincer. BU =Stems browsed
Jurin·3 tile 1974-75 ;,dnter <111cl not brO\'iS.2d during the~ 1975-7u ·.vinter.
C.lJt:ctuins to stG:m3 br<HJsed during the l'.J./G-77 ~<linter.
dsignificant ~t the .05 significance level.
l
r l 1
~.
I ! •
l
t i. t
i
I
~able 8. n~ailability-utilization chi-squ~re t~bles for ~valuating
within-species preference at the Wickersham sludy area. Expcctuj
froqu.::ncL:s and proportions \•!cr.:! b;)Sf~d UiJOn tn~ relativ·2 yu~ntities
of ste1.1S avi~il.:lblc troi11 c<.1cb Ci)t0-3ory (continued).
-----·------·---------------·----------------------·-----------/--·-----·---------·--·------
Ci!teyory b B;<pected No.0 Ob3ervcd No.c ~xpected
of of Stems o£ Stems Proportion
Site Species Stem Browsed Browsed
13 Alax. 813 6 (1. 3 U7 0.37
uu 40.2 25 0.25
08 48.4 33 ~j. 3 ~J
80 15.1 1 g 0.09
Plan. EB C:J (j iJ
uu 41.7 35 0.65
UB 5.4 Hi 0. 11
BU 1.9 4 fj • (J 4
a Observed proportion browsed ± 95 percent confidence interval.
Observcda
Proi.Jortion
Gro1·1Sed +
95!2. c.r-:-
0.53 0.10
0. 15 O.fl7
u. 20 IJ. OS
0.) 2 ~). G 6
0
0.71
0.28
!} • (:j 8
G-
statistic
d 23.46
7.29
b BB :::Stems brows2d during the 1974-75 and 1975-76 wint-=rs. Ull =Stems tlOt
browsed during the 1974-75 ~nd 1975-76 winters. UD = Sterns not brows2d during
the 1974-75 ~~inter and brov;sed dur inq the 1975-76 winter. BU = Stems· browsed
durin3 the 1974-75 winter and not browsed during the 1975-76 winter.
c Partains to stems browsed during the 1976-77 winter.
d Significant at the .05 significance lev~l.
F i.']. 4. Lin2ar regression rel~ting the 197C-77 winter
:.:JJ:o\·;sing to a·.;:..ilability for thE:: portion of plaut browsed
~'-lrin; tlle 1975-76 winter. X2 corr~sr:.onds to tile: nu;nb::::r of
~·:J.:lilable: durii1~ t~1<2 1976-77 •.vin'.:cr. Y2 corr~sponds to ~he:
. l. f2::Jr':!:3GlOn. lnL: and the jet~rmin~tion cocfficianl ( l-2 ) are
~!so sho~n. ~he jotted circle inJicates that more than one
I
;::.a.TttJlln:J point occurs her:.!.
instead of stems. The chi-square statistic is significant
(p < .001). An inspection of observ2J and expected
fr0qu·2ncies reveals that plants bro\·Jsed durin9 the: 1975-7G
winter sustained proportionately more browsing the following
winter than those unbro0sed. The results from plants tagged
at Sites 11 and 12 showed a similar trend although the
results were not significant at the .OS significan~e level
(T.:ble 10).
46
Mean pellet group counts varied from 0 to 600 groups
per ha for the Kenai study areas (Table 11). For the 1976-77
winter, Site 3 ranked highest, Site 9 was next and the:
remQining 8 sit~s ranked together as the last group
(Kruskis-~vc.l1is, p < .05). For the 1977-78 winter, Si':es 2,9.
and 10 ranked higher than Sites 1,5 and 5 {p < .05). Between
year comparisons showed that, for all sites, the pellet
group density did not change fro~ year to year (.09 < p <
• 7 3) •
For tl1::~ \H;;kersnam study area, the counts were 1mver
than those from the Kenai Peninsula study areas ranging from
2 to 52 groups per ha {Table 12). Site 13 ranked highest;
Site 12 was next and Sites 11 and 14 ranked last for the
1976-77 winter (p < .05). For the 1977-78 winter, Site 13
again ranked highest; Site 11 was next and Sites 12 and 14
ranked last (p < .OS). Between year comparisons showed that
~cibl~ 11. Pellet grou~ counts for the Kenai Peninsula
study aceas.
Sit·~ Yeur
1 1977
1978
2 1977
1973
1977
4 1977
5 1977
1Y7.8
G 1977
1978
7 1971
1977
1977
1978
1977
1978
t·lci.ll1 Nu:i1b0r of P-;.::11"-.:t Groups
por h~ <t St~nd~rd error)
5 ... v (50)
,
:.J
100 ( 7 ~j)
1 .... -> .) _} ( 6 3)
GJ ~j ( 16 3)
1 ~j~j (5G)
-. ..., (33) J .)
{~
f.-i
"
f1
33 (33)
" u
333 ;: 3 3)
1G7 ( (3 4)
33 (56)
100 (56)
--------------------·---------------------------------------
49
\
f
t
l
I
I.
~0blu 12. Pel1ec group counts for th~ Wickersham study ~re2.
Site Ye.:::.r
11 1977
1970
12 1977
1978
13 1977
1978
14 1977
1973
~e~n Number of P2llet Groups
per h~ (~ StDnd~rd Error)
G { 6)
42 ( 7)
2 ~J {9)
14 { 5)
S:J (10)
52 ( 10}.
0
6 { 4)
5o
'1\")t;::l No.
of f'li::lnt.s
Gbse:rved
l\o. of Plants
Cbscrv,~d ho.
o£ !'l a11 ts
SrovJSC::i
E:xpect-<::!::3
Proportion
Br0~s2~-1
Observ-:.::ja
i'rOf·Or tio:-~
Bro·,·iseJ
+ 95~ C.j,.
137.G
144
.U. 5S
G.62
52
liD 131
41.6 52.9
83 5
0.18 G.23
{1.36 0.02
(.08) (.~·32)
acbs~rv~d proportion browsed + 95 percent confidence interv~l.
54
not been adequately accounte::l for. Conclusions should
b~ ~valuated in this light.
The results of protein analysis at the Wickersham study
area are shcrwn in Table 16. The range of values is much
narrower than those found at the Kenai study areas. The
nighest value was 6.9 percent and the lowest was 5.3
perc~nt. Since three samples instead of one were submitted
for orotein d~terillination for each species and site
category, l was able to calculate an estimate of variability
as8ociated with the mean. Each percent value was transformed
by the arcsin transformation to allow the use of parametric
H:~th,">ds of statistics (Zar 1974). ~-lean protein levels ~-;ere
not significantly different between species and site
categories (Nested one-way ANOVA, .20 < p .50).
Results of protein evaluation from the tagged plant
ex 9f! r imen t are shown in Table 17. Six pairs of S; olanifolia --L----~-----
s:H ubs and five p.a irs of ~. ~£~~!:~~!:_~!]_~ shrubs \ve r e S.:lffi!?l ed.
For s. E!:.£!2..!:.f~ll:_~, mean values rang2d from 4.9 to 5.9
' p~rc~nt for plants browsed both the 1976-77 and 1977-78
winters and from 4.8 to 7.2 percent for plants browsed
neither wint-:r. Values for ~-~£~~l~E.!:.~~~ ranged from 6.0
to 5.7 percent for plants browsed both winters and from 5.6
to 5.7 percent for plants browsed neither winter. All
percent data were transformed by the arcsin transformation.
For both species, protein content was not significantly
different between th0 browse categories (paired t-test,
~ablG lG. Protein content for each species and site at th2
Wickersham study area.
Prot~in Content (percent t Standard Error)
.sit·~
.S. scoul-eriana S. alax~nsis
1 5.6 y). 2 N. D.a N.D.
2 5.3 0.2 5.4 0.4 N.D.
"") N.D. N.D. 6.0 0.1 ,J
t. 5. 4 ll.l N.D. N.D.
a D..:t t.:·, not a v .::t i L.iblc:.
.56
CONCLUGING DISCUSSION
Crowse Production and Utiliz~tion
Lrowss production is within the r2nge of that observed
by other workers in Alaskd. Milke (1~69) founj willow
projuction available as browse for ffiOose to ~~erage 2J3.S
kg/ha at his Interior floodplain study area. Wolff (1975)
found for his rip~rian, floodplain study area that willow
production aversgej 31.45 kg/ha for ~n C-year-old st~nd and
99.35 kt;J/11<.:. for. a 15-year-oJJ stand. I found for my rip.Jrian
sit•~ (S1 te 13) thz.t productioii av,:ragGd 325.7 .:.n.J 1S3. 2
kg/ha respectively for the two yedr period. Results of ~11
t
I three studies show that S. olaxensis dominateJ the
p r o c1 u c: t i c) n a t I n t e r i o r r i r-<': r i .:o. n s i t e s i n b~ r m s o f b i o :n a 3 s •
Pro::Juction of 5. scoulerian~ ~t l)ld Interior burned
regrowth and 75-ye~r old mature spruce site w~s found to
av2rage 52.5 anj 21.1 kg/ha respectively (Wolff 1978). These
values co~p&re f~vorably with those I obt~ineJ for a similar
burned rGgrowth site (Site 11) and for a timb~red sitG (Site
14) at the ~-;ic}~e:rshaill study area ('I'able 4).
The upland regrowth ranges ~t the Slikok L3kc study
v.rea app03r to b·~ more productive than th~ir cou,~terp .. .Jrts
at ~i~kcrsham. 1h2 hig~est level of production observed at
the tvickersh::.rn .study C~rea w2s 75.G kg/ha for.§_. ~-~ule.!:~.§..~~:=_
(Table 4). The highest lev2l of production of S. scoulc:riana
~t a compar~ble site at th~ Slikok Lake study area was 2~4.9
62
thc.;t <.t r<::mov.:;l of SC percent mz,y be: optiri1;il for a m.::xiwum
sust~in~d yield of hard~Jod browse (Spencer and Chatelain
lS53, Krofting ~tal l9GG, ~olff lS7G).
Moose appear to browse the upl~nds ranges more
intensively at the Glik~k Lake study area than at
\·;ickc:rs!vnn. 'l'he highe:st utiliz2.tion percent<1g.:.: for the
Slikok L~ke study area was 64.4 p~rcent and the highest
rc~cordec for comp.::rable: uplan::: sit12s ;.lt tlK \dckcrsl1<lm study
dr2~ was 26.5 per~ent. Crete and Au~y (1974) hypothesized
th~t high browsing in~ensiti~s arc rel~ted to high mean dpb
and 11 possess nearly id~ntical densities of ~-!£~~!~!!~~~
twigs (Tables 3 ~nJ 4) but tb2 percent utiliz~tion and rn~an
dpb's ar2 different (Appen~ix I). Percent utilization (or
Sitt:! 3 is (j4.4, for Site 9, 53.4 .:.nd 43.4 ,:;n:J for Site 11,
G.S• :.nd 1G.7. 'HH? mean :Jpb for Sitr::: 3 is 3.6 mm .:;nj the mean
dpb's lumped over th~ two seasons at Sites 9 and 11 are 3.3
an:.~ 3. J. m.-n respectively; trH~ three mee1ns arc significantly
different from c~ch oth0r (ANOV~ ~nd Neum~n-Kuels, p < .05}.
~he diff~renc~s in ms~n dpb ~re r~flected in v~stly
differe:nt estiin.::tt•2S of biom.:,ss pro<'lu(;•.::c1 and utilized. Site
9 shows ct utilization level 1[ to 2C percentage points lower
th~n Site 3 but the utilization level in terms of biom~ss
Eo r S i t c 9 is c: bn u t one-h c:t l f t h.:. r for S i t e J • 'l'h e d i f f <~ r E: n c e
bctwe~n Site 3 ~nj 11 is ev~n more pronounc~d. In spite of
similar numb~rs of twigs dV~il~blc, 2v~il~bil ity in t~rms
)
th~n normal, snow depth p~tterns did not depart
subst~ntially fro~ that st~ted above. Kelsall (1969) cl~ims
that spatially concentrated browse offsets the high
energetic cost of moving through snow. In a winter range of
high snow accumulation, moose would tend to concentrate in
tnose areas where browse is abundant and snow depth is low.
Site 13, a lowland riparian site, had by far the lowest snow
accumulation (T~ble 18), a density of willow twiqs two to
three times higher th~n the upland sites ~nd the highest
utilization levels recorded for the Wickersham study area.
lt is interestin3 to note that Site 13 compares favorably
with the upland Sites 3 an_d 9 in terms of bio;nass utilized,
availability and percentage utilizad. Lower snow
accumulation may give the studied uplan~ ranges at Slikok
Lake greater value as winter range than the studied upland
ran;es at Wickersham.
Possible reasons for th2 lower utilization percent
le~els observed durin3 the 1977-78 winter at the Kenai study
areas are not clear. Setting 0.2 mas the minimum browsing
height may·be one cau3e. That criterion was used during both
winters and w~s based upon measurements from all sites taken
only for the 1976-77 winter. Snowfall accumulation on the
K~nai Peninsula was much greater durin3 the 1977-78 winter
than it was during the 1976-77 winter. (Oldemeycr, personal
cn~munication). Although moose were observed pawing through
snow for browse at the Kenai Peninsula study areas, such
'l'::.d::le 19. ErovJS2 consumptinn und p(!llct group density
as indices of moose us8 at the ~ickccsham
stu-Jy z:tro::a.
fti. D. /haa
(5roNse Consumption
. Da t.:t)
t':. D. /ha
(Pallet Group Data)
·---------·----------------------------------~---·----·-·---
ll
12
1 ....
-·..)
14
1977
1 s~ 7 s
1977
197B
1~77
1978
1S77
1978"
a Monse-d~ys per hectare.
0.14 0.46
? ,, .... _, ~ 3.23
4.0G 1.54
~i. 86 1. 08
2 A ""l r~ -. • L(. 3.85
15.52 4.00
0 0
1;.~2 ~~. 4 6
66
67
{Neff 1968, Wolff 1976), moose use v~ried from a to 4.0
N.D./ha. Results calcul~ted by the two methods do not
correspond well, particularly at Site 13. The lack of
correspondence may be due to errors in the above values used
for consumption and defecation rates. Mean consumption rates
vary 0reatly in the literature. LeResche and Davis (1971)
docuraen ted val Jes as lo~v as l. 3 kg /ha dry \·l2igh t for the
K2n~i Peninsula while Palm~r (1944) found consumption rates
to b2 as high as 16 kg/h~ air dry weight. Likewise, reported
defecation rates vary greatly d2pendent both on the sex and
a~e structure of the population as well as the forage eaten
) (Neff 1968). Franzmann et al (1976) documented a daily
defecation rate for Kenai Peninsula moose of 14.5 for
females and 19.6 for males. Unless the mean daily
consumption ~nd/or defecation rates for moose in a given
are~ are known, comput~tion of moosd days of use by either
method is inconclusiv~.
Protein Content
The ran~2 of valu~s for crud~ protein levels of willows
that I obtained (Tables 15, 15 and 17) were slightly lower
tn3n other reported values for Alaska. Old~meycr et al
(1977) reported a meun value of 6.4 percent for willow on
the Kenai Peninsula. Milke (1969) reported values rangin3
from 5.2 porcent to 7.0 percent for four species of Interior
Alaska willows. His reported value of 7.0 percent for s.
68
~!_~~~Q_~!_~ is higher th:Jn my value of 6.0 percent althou9h
both top their respective lists. Tha differences may be
related to variability in clipping methodology and dates of
sample colle~tion (Bailey 1967}. The observed percent
protein of a twig Jepen~s a great deal upon ho~ fzr from the
terminal bud it was cli~ped. Cowan et al (1979) found that
twigs clipped close to the ter~inal buds had a higher
percent protein than those including a larger portion of the
stem. ~volff (1977} obs2rv2d a similor rel.:ttionship to occur
l,vith twigs of ~· scoult:riana.
The protein content of a twig also varies seasonally
and is related to phenology (Bissell and Strong 1955, Tew
1978). Noteworthy are the significantly higher protein
lev.::ls observed "for th.= tagJed plants of ~· ~£C::~!.~E.~~Q.~ than
those of S. E!.~l2..~f<::!..~~ ('l'able 17}. Th·2 site-specific sumt.Jles
for the two species were not significantly different in
prot~in content. Protein in S. ~£~~!.££~~~~ tended to be
hig~er in the tagged plants than.in the site-specific
samples. A possible explanation of this apparent anomaly
concerns the difference in Jates of sampla collection and
relates to ph2nology. The sitc-s~~cific samples were
collected in early March und the t~gged plants were sampled
in April. Viereck and Little (1972) describe ~· scouleriana
as one of the earliest willows to flower, often forming
catkins prior to snow melt-off. Althoug~ catkins were not
observed on the tagged plant sumples, the timing of the
72
Other ~tudi~s in Alaska h~ve su~y~sted that preference
may b!2 related to height. t1ilk<= (19G9) found thut those
willow species most utilized tenJad to be the tallest
availabl~. Mould (1977) found that seven of eight browse
sp2cies in a riparian habitat in arctic Alaska showed ~
significant positive correlation b2tween the height of plant
and brousing intensity .. Sp2cics which tended to be the
t~llest were browsed the heaviest. Snow depth and the need
for cover were suggested as possible causes for preference.
Although guantitativ2 data on height-use relations are
lacking except for Sites 1 snd 2, l found the above
conclusions on a subjective basis to be true for certain
Sites 3 an1 9, ~· scouleriuna tended to be the
ta.llcst i:lnd ?referred. At Site 13, f·
alax~nsi5 tended to be much taller than S. olanifloia and ----------=-----------
s. ~~~!:_~~~ and wCts the most [.>referred. l\t Site 12, hm.;evcr,
s. scoulcriana tended to be taller but
more preferred. At Sites 1 and 2, the same relationship ~as
Other causes of between-specie3 preference may be
related to nutrition, specifically, protein content. Cow~n
ct al (1970) considered protein content to be an important
indicator of forag2 ~alue. Some studies have documented a
positiva correl~tion between the protein content of a
sp~cies and preference by cattle and sheep {Hobbs et al
1945, H.:1rdison et al 1954, Cook 1959, He.:.dy 19G•i), deQr
74
correl~ted \iith preference (Lon':)hurst et al 19G8, Oh et al
1963).
Some workers maintain that mixing of species in th~
diet is an important aspect of between-species preference
{Longhurst et al 19G3, Cowan et al 1970, Mould 1977). Moose
on h~avily utilized range in the Kenai Peni~sula and
south~entral Alaska was attributed to a need for variety in
the diet (LeHesche u.nd Davis 1973). Oldemey,.?r et al (1977}
postulated that the observ2d decline in diversity of brows0
species on northwestern K2n~i Paninsula winter range over
the past 20 years is at least pu.rtially responsible for the
observed decline in moose popul~tions in recent years.
Proper mixing of species would mu.ximize proportions of
d~sirable nutrients and minimize proportions of undesirable
digestive-inilibitory compounds and to:dns. In addition,
digestive -inhibitory and toxic coumpounds of a species may
)
be buffered by substances found in plants of other species.
Mould (1977) found for the .Colville River ar~a of arctic
Alaska that moose browsed willow and unusu~lly high amounts
' )
of alder {~!_!:!_~ ££.!.~12.£). Pu.p2r birch, asp.=n and cotton;,·mod
were not ~vailable. Alder poss~sses higher levels of protein
than willow while digestibility follows the reverse pattern
(Oldcmeyer et al 1977). Proper mixing of the two species may
possibly optimize the nutrition and digestibility levels in
tne diet. At the Wickersham study area, usa of alder w~s
. ~ -~--. ..__ ___ -~. ---~----· .... --........ ;.. ------·
17.6 percent protein while clippad forage fro~ the same area
containin9 the same pro;?ortion of Sp•::cies had only 6.9
percent protein. The clippin3 procedure attemptad to
duplic~te the browsing of each shrub by deer so as to
minimize the effects of.pl~nt part selection. Bissell found
76
in additional experiments involving captive deer th~t rumen
content protein level differed little from feeJ protein
lev~l. If crude protein lev0ls fall bela~ the 6 to 7 percent
maintain~nce lev8l, rumen functions are thought to be
Lnl?u.ireJ (Dietz l9G5). Preferential browsin9 vmuld permit
moose utilizing my s~uJy area to maintain a diet of 7
percent or hi·:Jher even thoug~ the mean protein levels of t:tll
ffi·2asured spec.i·2S \vere bela·..; 7 p·~rcent (·rable 15 and lG). t·ly
results suggest. that within-species preference is rel~ted
to fa.::tors ott1•2r than i:)rotein level (Table 17). Ho•ti2VC!r,
r2s~lts may h2v2 been jifEsrent if utiliz~tion was
corr~lated with variability of protein content b~twecn stems
rathar than between plctnts. Small sam9le sizes ~ake thQ
results inconclusive.
Other investig~tors h~vc documented that other chemical
components, particularly vol~tile com~ounds, of forage
plants may be important in determining within-species
prcf€:rence. Junif)er utilization by mule de..?r <2.':!'2.£~!!.£.~~
~~~!~~~~) in Utah ;,-:~s found to be negatively correlated \vith
the cs3onti~l oil content of individual shrubs (Smith 1950,
1959). S~gcbrush occurrin3 in the same area was thou~ht to
78
shrubs arc similar in composition to shrubs in an earlier
growth stage; the parcentag~s of phosphorus, protein and
carotene are high and the percentages of lignin and crude
fiber are low. However, Klein (1977) suggested that regrowth
twigs may be less desirable as forage if increased levels
of digestiv2-inhibitory compounds are present. Tables 21 and
illustrate the mean number of twigs available per stem
~uring the 1976-77 winter for two cateanrics of sterns
{brO\·ISi?d and not browsed durinj the 1975-76 winter). Th·:!
~~nn-Whitnay non-parametric test was used to test for
Jifferences in production between categories. In all cases
(e~cept Site 14) where the difference is significant, stems
bro·,;St:!d th'= ?r-<~vious winter tend::=~ to outproducr~ those stems
which were unbrc-H·i5·.:d. Although these! rcsul ts on not
establish cause and effect, stimulDtion of growth by
browsin~ is a possibility. If the probability of a pl~nt or
stem bein3 browsed is related to the number of twigs it
possesses, stimulated regrowth may give the browsed plants
a higher probability of bcinq brn\·;sed tile folln>..Jirlg . .. :.v 1 n .. e r •
Th2 high~r nutritional value of regrowth twigs mdy serve to
increase the prnb3bility still further if preference is
related to nutrition rather th<:!n to dig·::!Stive-inhibitnry
compounds.
The indivi~uul height of a shrub may influence it's
~hanccs of b~ ing brmvsed. The relationship of plant h.=: igh t
) and between-species preference has alrcajy been discussed.
Bo
'i':~b.12 21. Corn:.x.1rison of t\tig produ.:tion during the 197G
growing se~son s2ason for sterns browsed during Lh2
1~75-76 winter and those not browse~ at the Kanai
Peninsul~ study ~r2~s (continued).
Sit0
6
7
(' :J
lu
i+:~ .:m t..;o •
Br 0'.-iS.: a: of 'l\; ig 5
Sp0cies Cutegory· p2r Stern
Bare. B 2.8B
u 3.95
S_;ou. B 2.76
u 3.33
I3ebb. B 3. 6~J
u 3.20
SC,)i.i. L 2. <i ·.:i
u 1. 67
Bare. c ") •' ""\ ...... ~""'
u ~.~G
Deb!:... D il . c ~1
u 2.8G
.:JC()U. D 4. 2l
u 1. 94
Bare. L 3.42
u 3.15
D·.=bb. .8 S.lS
u 3.31
Bdrc. c '~. 17
u 2.29
Standard
Error
,,
l.lo 52
~L 4';j
0.26
1).51
U.3lr
n.3e
0.26
0.14
l. GO
ti . 19
1. 22
U.37
!).15
J.lS
~L 51
0.50
i). 29
0.51
0.75
1.17
Hann-b
Sample \~h.i.tr1ey
Size Statistic
0 11.0 u
22
38 {u.SGl)
21
25 226.0
19
38 19.0
16
2 16.5
16
5 15.5
5
") c '• .j..,,{. (7.G3l)c
51
12 7~j. 5
13
1 ~1·3 ( 'l .:.. 60 5 )c
16
r 1~.5 0
7
a e = Stems bro\;~~.5 durin~) th .. : 1975-76 ~;inter. U = Ste;n.:; not
bro:.-;s.:::J .Juring the 1975-7G \-linter.
b 'i~i1:.~n s~•;upl<.: sizes ;:J.re gr·-=atr=r than 2-U for tile smaller categori.::s
or gr::ater than 4·J for th2 lurgcr, th.:; norm.J] approxit~hltion was
1.W2.:i (~ar 1974). v~.lu•2S in pur.;ntll•:?sis repres2nt th0 normal
z su~t.istic. Valu2s withnut p;;renth.::!sis r2pr<:S1.?nt th·:' stan-:larci
t•i.:.nn-·,Jbit.Iv.::y statistic (U}.
c SiJnifi-.;.::nt .Jt th8 .05 3ignific.::~nc,.? lev2l.
A similar p~ttern m2y be o~crative conc~rntng within-sp0ci~s
selection. Klein (1977) foull·~= th<:~t snO\·:sho.:: har·~s pr12f(~rrt::::1
t\iigs from the tops of tre:cs and t.Jll shrubs \·;hich norm,=..lly
.::... r 2 o u t o f r e a c h o v c r t •·i i ·;; s f r o in r o o t s u c kt.: r s ;;1 n d 1 o •.-; s h r u b s
of the Sfirr.<:.: species. It w;:;.s sugg!:?St\.:!j th~t the presence of
~igestiv2-inhibitory compounds in the root suck~rs and low
shrubs w~s responsibl~ for ~heir low utilization. In ~r~~s
such as Interior Alss~a wh2rc the influenc~ of hares upon
v2getation is great, such ~n a~ti-herbivorc strdtcgy S8ems
nccc:ssary. If digestive-inhibitory coml?ounds d·2velofH~-::J to
prot2ct shubs fro~ excessive hare browsing affect rnoos2 in
a s i ;n i l c. r v; ~ y , t h t: t a ll :: r s h r u b s o [ .:1 s p e: c.: i 2 s s h o u 1. d b ~-~
j?ref.::rr"'=d by moos'~ o;_>er short·.::r shcubs. During thE· 1977-7f;
wint~r at the Wick2rsh~rn stu~y are~, tall paper birch shrubs
(ov<=r 3m tall) thct hc:,Ci br:~n broi~en c.:Jo\-:n by rn:mse norii12il.ly
h~d only their tops brnws~d. Twigs ~~ailDbl~ at a lower,
raort= conveniE·nt l·..:'Vl.=l \..;er~ ofto:=n unbro.~·sec1. Such, ho;..;-2\'(!f,
;,.·as not co:11monly obseri.r2r} for viillo~;. nos t brn~,s ing of
willa~ seemed to occur ~t a l~vel conv~nient for rnoos0.
Uir~h and willow ffidY 20Ss0ss diffcrant anti-herbivore
str~teyies ~nd, consequently, th~ r~ctors influcn~ing
prefcr~nce for each species muy be different.
Site factors extrinsic to the pl'-lnt may c.;lso be
important in clt:termining -viithin-sp<~cies bro•:sing prcfervnce.
Snow depth w~s observed to Dffect betw2en-speci~s browsing
preft::rcnc•2 ond it mz,y ,"ilso L(~ i.mport,;.nt in d(~termiilin'J
82
·1·aol.;;: 23. Cbserv.;d distr ibutinn of 9~ll..:t 0COUi_iS fitt.::d
~9~inst th2 exp2ct~d r~nso~ distribution for
5 i t 0 s ll c: ;1(; 1 3 .
Frequ2ncy
(grouvs
~it2 Ye~r P0r plo~)
0bs:::rvcd
Disr.ribution
E;{L)·~C t0d
1{<) nd oa1
Distribution
Chi-
ll 1578 " 6 Ll 7.0 0.474
1
")
£.
13 } 977 ,,
Ll
1
2 .,
J
4
1 J
•• .J 1978 ,.
"' 1
2
j
4
7
7
G
7
4
2
1
5 ,...
0
4
2
1
7.4
5.7
5.7
7. 1
4.5
2. Ga
5.5
7.1
4 • c.
-) na "-. ~
0.120
a fr.:::;quen.::y ,:>J te9or L.:s .3 unJ 4 lUillped togeth2r du2 to low
obs..:rvad ~nj 0xp~cted values.
l'ioose in Al.:1ska are prr.~scntly one of thE..· most important
species hunted for both sport and subsistence.
Nonconsumptive use of moose by sightse8rs and photogr~phers
will continue to gain in pro~inence. Nainten~oce of he~lthy,
st~ble populations is desirable for most are~s. Intensive
mc)n,Jg2ment of habitut for a high sustained yi,=dd of moost:
may eventu~lly b~ necess~ry to offset th0 loss of hunting
opportunity caused by l~nj ~i~hdr~w~ls of one form or
~nother. Increased efficicn~y in th0 control of wildfire is
currently reducing future ser~l habitat favor2ble for moose.
Sound hQbit~t mana0emcnt practices nc2d to bQ developed for
the futurt::.
Th·= most com!r:<">II [on.ts of habitat reh~bilit:"tion for
moose as currently practic2d in Alasko are controlled
burning anj mechanical crushing. The Kenai ~ational Noose
H.:mge 2nd the ChugcJCh Nation<il Forest, both located in
southcentral Alaska, are currently the only areas in the
state where habitat rehabilitation is pr~cticcd on ct regular
b~sis. Controlled logging as a man~gement tool will gain in
~rominenc•.= uS th'? ti111b•~r resourc~s of Southcentral cJnc1
Interior Alaska are devalop~d. The maintenance of cJ high
c.mount of ecotone in proportion to the OV(:-raJ 1 acreage
r~habilitcJted is or great importance for all these ~reas
(Eustm.:.n 1974).
86
t~kcn. The observ2d nonran1o~ use pattetn indicates this is
not necessarily so. It is import~nt that plant
characteristics such as nutritional or digestibility factors
\vhich may lead to this. type of use pattern be establishr::a.
It is possible that habitat management practices which favor
some of these ch~racteristics may consequently be develop2d
and· evaludted.
88
AfJ::Ji.:r~Ji~: I. (-::ontinu.;:j) DJ?b an(i m2:.2n t;,vig ·1·t·2ight J:.t.:;
Sitt..::
13
14
f!)r the ti1r2e .:;tuJy .Jr·.=~:;s (_:!: SL::;1J-.:~rj Errt)f)
Dp b ( r.1;n )a
l\lC.lX. 1977 ., G 0. 1 -' .
1978 3. 1 fi • 1
f'l::!n. 1977 ' 1 ;:-\ 2 J. ,, .
1978 ") " I' 1 ,_ . J iJ •
Scou. 1977 J. ,,
u 0. ]
1978 ' 1 (l • 1 -' .
He::!n tvli:J
~·i·2 i 9 h t. ( <.J)
1. 88 (\ Cl8 tJ •
0.73 0. () 7
J.. 05 (; . 12
0. 7'u r,
x.; • G7
•'\ !·".J • 84 :J. 16 . ~ 73 C.C7 ..:1 •
90
LITERA~OR8 C[~ED
Allen, R.S. and A.R. Porter. 1954. Effect of add2d 9roun(]
corn on th~ consumption and palatability of stacked
silage. J. Dairy Sci. 37:658.
AOAC. 19GO. OfficiDl methods of analysis of the American
Organization of Agricultural Chemists. Ninth Ed., wash.,
D.C. 832 p.
Bailey, J.A. 19G7. Sam~ling deer browse for crude protein.
J. Wildl. Manuge. 31:437-142.
Baker, T.C., C. Anderson and ~.I. Crump. 1953. Food habits
studies of game animals. Wyo. Wild Lif~ 17(11) :24-31.
B~rret, J.D. and w.~. Guthrie. 19G3.
in estimating browse. J. Wildl.
Optimum plot sampl in·g
Manage. 33:399-403.
Berg, U.E. anu R.L. Philips. 1974. Habitat use by moos~ in
northwestern Hinnesota with reference to other heavily
willo;..J•::d areas. Can. Nat. 101:101-116.
BishofJ, R.iJ. and F~.A. Rau.sch. 1974. Hoose population
fluctuations in Alaska from 1950 to 1972. Can. Nat.
101:559-593.
Bisse.ll, H.D. and H. Stron:J. 1955. 'l'he crude protein
variations in the browse diet of California deer.
California Fish and Game 41{2) :145-155.
Bissell, H.D. 1959.
California Fish
Interpreting chemical analyses of browse.
and Game 45:57-58.
Blaser, R.E., R.C. Hames, II.'l'.
Fontenot and R.H. Engel.
grazing on animal output.
Congress 8:601-606.
Oryant, N.A. Hardison, J.P.
1960. The effect of Selective
Proc. ath Internl. Grassl.
Bo·,,d.;n, D.C., A.E. Anderson and D.E. H-:-din. 1969. Fn~quency
distributions of mule deer fecal group counts.
J. \lildl. H.Jnag.;. 33:895-905.
Bro;.m, E.R. and J.H. Hand-.=ry. 1962. Planting and
fertilization as a possible means of controlling
di.stributions of big game animals. J. For. 60:33-35.
Cnatelain, E.F. 1951. ~inter range problems of moose in
the Susitna Valley. Proc. 2nd Ann. Alaska Sci. Conf.
p. 343-347.
92
Gas2way, W.C. and J.W. Coady. 1974. Review of energy
r~~uirements and rumen fermentation in moose a~d other
ruminants. Nat. Can. 101:227-262.
Hardison, W.A., J.T. Reit, C.M. Martin and P.G. Woolfolk.
1954. D2gree of herbage selection by grazing cattle.
J. Dairy Sci. 37:89-102.
Harry, G.B. 19 57.
Hole, ~vyom ing.
Winter food habits of moose in Jackson
J. 'i'lildl. rlanage. 21:53-·57.
Heady, U.F. 1964. Palatability of herbage and animal
preference. J. Range Manage. 17:76-82.
Hobbs, C.S., \LD. Gullup .~n·:J 3.R. 'I'aylor. 1945. The
composition and apparent digestibility of bluestem
94
grass in the growing stage and in the dry and hay stages
when supplem0nt~d with cottonseed cake. J. Animal
Sci . 4: 3 9 5-4 ,j 2.
J~nsen, C.Il., A.D. Smitb un:1 G.i·l. Scotter. 1972. Guidelines
for grazing sheep on rangel~nds used by big game in
winter. J. Range Manag~. 25:345-352.
Johnson, P.R. and Hartm~n, C.W. 1969. Enviremcntal atlas
of Al~ska. Institute of Arctic Envirementa1 Engineering,
F'~irbanks,. i\l<1ska. 111 !?·
Kelsa~1, J.P. 1969.
deer for snow.
Structural adaptations of moose and
J. Mamilla!. 50:302-310.
Klein, D.R. 1977. Winter food preferences of snowshoe hare
(Lepus americ~nus) in Interior Alaska. Xliith Intl.
con9ress-of"-G'~iae-Biolo':1 is ts. p. 2GG-27 5.
Knowlton, F.F. 1960. Food habits, movements and populations
of moose in the Grav.:lly t·1ountains, r:Iontana. J. ~Vildl.
Manage. 24:162-170.
K r e E t in~ , L • ~·J • , N • H • S t r en 1 u n d an J R • K • S 2 em.~ l • 1 9 G G • 1:: f f e c t
of simulated and natural deer browsing on mountain
maple. J. Wildl. M~nage. 30:431-438.
Laycock, ri.A. and D.A. Price. 197~. Factors influuhcing
forage quality. USDA Misc. Publ. No. 1147. p. 37-47.
Leigh, J.H. 19Gl. The relative palatability of various
varieties of weeping love grass (Eragrostis £~~~~~~).
J. Br. Grassl. Soc. 16:135-140. ----------
P.:; ltuer, I .. J. l94A. f'()o6 rcqu i rer.1en ts of some Alaskan
g ;:uue .:.n imal s. J. f·lc.Hniil.:i l. 2 5: -19-5 4.
Pdck, J.M. 1971. A review of moos~ food hdbits studies
in North America. Nat. Can. 101:195-215.
Pirnlott, D.f:i. 1~61.
North A;nerica.
Th~ ecology and management of moose in
L~ Terre Et Lu Vie No. 2-3:246-265. .
Plice, M.J. 1952.
b 1 1 i v '..! .s to c k .
Sugar vcrs~s intuitive choice of foods
J. R<.<n,.J..:: Hanagc. 5:G9-75.
h:,i::J,;;;.·n, t·LA. 1972. Diff~rt:nc-=s b~L·.i.?en Dougl..:.~s fir g~noty~es
in r~l~tion to browuin~ pref~r~nce by black-tail deer.
Can. J. r .. or. Res. 2:25J-255.
Radwan, M.A. 1975.
Res e .:1 r c h B u 11 • ,
~hat makes deer choosy eaters?
U.SD.i\, Feb. 1975. p. 8-lH.
For•.:?si:ry
R8g0lin, W.L., J.G. ~a~y and O.C.
snowdrifts on mountain shrub
Congress of Gama Biolo9ists.
~~a 11m o . 1 9 7 7 •
communities.
Eff2cts of
XTIIth Intl.
!?· 114-419 •.
Reyno 1~1 s, n. G. .:~nJ i\. rL Sampson.
sprouts as browse for Jeer.
122.
1943. Cnapc:.rrt:~1 crovm
J. ~ildl. Manag~. 7:119-
Sil.:.-fcr, r::.L.
hardwood
1963. The twig-count method for meusurinJ
deer brows~. J. Wild!. Manage. 27:428-437.
Sigm.:.:n, r-1. 1975. ·rhe impoJ~t.:.ncQ of
ov2rwinter mnnse calf survival.
Alaska, Fairbanks. 185 p •
the cow-c~lf bond to
M.S. Thesis, Univ.
.S;nith, ! •. D. 1950. In·~3uirL~s into differential consum()tion
of junipr2r. Utah F.i.sh and Gum.:: Gull. 9:4
Smit~, A.D. 1959. Adcq~~cy of SOille im~ortant browse species
in ov~rwintcrin3 of mule jeer. J. Range Man~ge.
12:8-13.
3pcncer, D.H. and E.F. Chatelain. l s 53.
man<:.19..:;,;·.=nt of
\"li 1 d 1 . Con f .
moose of southcentral
18:539-552.
Pro<Jrcss in th·~
Ali.:ska. Tr. N. l\;n •
Spencer, D.L. and J.B. ilakala. 19G4. Moose and fire on
th2 Ken~i. Pro~. of the 3rd Ann. Tall Timbers Firu
Ecology Conf. 3:11-33.