INTERACTIONS OF WHITE-TAILED DEER AND MOOSE ON SYMPATRIC RANGE IN CENTRAL ONTARIO
by
Stephen Kearney, Department of Zoology University of Guelph, Guelph, Ontario
ABSTRACT
The temporal and spatial distributions of white-tailed deer (Odocoileus virginianus) and moose (Alces alces) were studied in relation to habitat on
the 1,200 ha Himsworth Game Preserve in central Ontario. Habitat use was determined by examining systematically arranged plots for deer and moose pellet groups and tracks at 334 locations. Three time periods were in- volved: 15 October, 1973 (leaf fall) to April, 1974; May to June, 1974; and July to August, 1974. Observations of sign were analysed in relation to: 1) 15 habitat types (aspen-white birch-red maple, balsam fir, etc.);
2) habitat factors, including basal area of trees, density of trees, stand height, percent crown cover, percent understory cover, and tree species; and, 3) geographical distribution.
Deer sign was observed at 174 and moose sign at 129 of the 334 locations between October and April, with 83 observations of deer and moose sign being recorded at the same locations. The distributions of both species appeared to be more influenced by individual habitat factors than by habitat types, with moose sign being distributed in relation to the factors important as food and deer to the factors ,important as shelter.
The mixad and coniferous habitat types supported the highest deer densities whereas the highest moose densities were found in the deciduous types.
Sign of deer and moose was recorded at 38 and 31 locations respec- tively in the May to June period. Twenty-eight observations of deer sign but only 12 observations of moose sign were made in the July to August 64 period. The alder habitat type was the only one during spring and summer where sign of both species occurred more than expected, The distribution of deer and moose sign was generally similar to the observed distributions· of habitat factors. The geographical distributions of sign of both cervids indicated little spatial overlap in the summer. - 1 -
Moose (Alces alces Linnaeus) and white-tailed deer (Odocoileus virginianus Zimmerman) are of economic and aesthetic importance over much of their ranges. As a result many aspects of the biology of these two
species have been studied, but little research has been directed toward
their interaction.
Earlier studies had noted declines in moose populations subsequent
to the appearance of deer, the eventual outcome, except for local areas,
being a decline of moose over much of its former southern range (Peterson,
1955; Benson, 1957; Banasiak, 1961; Dodds, 1963). Yet when one reviews
food and habitat studies of moose and deer (Dodds, 1955; Peterson, 1955;
Kelsall, 1963; Telfer, 1965; Behrend, 1966; Prescott, 1968, 1974; Berg,
1971; Peek, 1971), it would appear that little interspecific competition
should occur between the two species except possibly in those areas where
both are present in high densities.
Further interspecific studies were stimulated by the identification
of Parelaphostrongylus tenuis as the causative agent of 'moose sickness'
(Anderson, 1963, 1964), a fatal disease of moose first observed in Minnesota
in 1912 (Thomas and Cahan, 1932), and which was later reported in the
same state (Fenstermacher and Jellison, 1933; Fenstermacher, 1934, 1937;
Fenstermacher and Olsen, 1942; Kurtz et al., 1966), as well as in Maine
(Lamson, 1941; Gilbert, 1974), Nova Scotia (Benson, 1955, 1958; Smith et
al., 1964; Smith and Archibald, 1967), New Brunswick (Smith et al., 1964;
Smith and Archibald, 1967), Quebec (Gilbert, pers. comm,), Ontario (Anderson,
1965), and Manitoba (Lankester, 1974). The reservoir host for!· tenuis
is the white-tailed deer in which a normal host-parasite relationship has
evolved, allowing the parasite to remain enzootic over much of the deer's
eastern range where suitable terrestrial gastropods, which serve as the - 2 - intermediate host, are available (Anderson, 1972).
The primary objective of this study was to investigate the temporal and spatial distribution of white-tailed deer and moose in relation to habitat on sympatric range in central Ontario, and to identify the areas· of overlap. STUDY AREA
The study was conducted in the northern three-quarters of the
Himsworth Game Preserve, an area of approximately 1,200 ha situated 16 km
south of North Bay, Ontario and 0.25 km east of Provincial Highway 11
(Fig. 1).
This area was selected because of: 1) accessibility; 2) well-defined
boundaries consisting of a pipeline right-of-way on the western border,
the Wasi River to the north, the Canadian National Railway tracks to the
east, and a township road to the south; 3) habitat density and interspe~
sion; 4) its natural state, as few people utilized the area and no trapping
or hunting was permitted; and 5) the knowledge that moose and deer oc
cupied this range year round.
The study area and the surrounding region lie completely within the
Precambrian Shield, an area of dense hard rock composed principally of
granite. It is between 240 and 280 m above sea level. The amount of soil
covering the rock varies considerably, and though most of the hills are covered by a stony till, numerous rock outcrops occur (Hoffman~ al.,
1962). The topography is rugged and slopes steep, irregular, and generally
short.
The vegetation present on the Himsworth Game Preserve shares some of
the characteristics of both the deciduous-coniferous and coniferous biomes.
This, along with the rugged topography and the large amount of beaver ac
tivity, resulted in few large continuous stands of any tree species or
associations existing on the area. Conversely there was an unusually
large amount of interspersion and diversity of habitat. Aspen (Populus
tremuloides, P. grandidentata); white birch (Betula papyifera), red maple
(Acer rubrum), and white pine (Pinus strobus), either individually or in - 4 - combination, were the typical upland forest species. Mixed stands con sisting of primarily aspen, white birch, balsam fir (Abies balsamea), and white spruce (Picea glauca) were common on the middle parts of slopes, forming the transition to the balsam fir or white spruce stands which oc curred on the lower levels of slopes. Stands of black spruce (Picea mariana) were found at moist, low-lying sites. Dense swales of alder (Alnus rugosa) within forest types or along the perimeters of openings, also frequently occurred at moist or wet sites. Openings of varying sizes were inter spersed throughout the Preserve. Large areas of inaccessible swamp and bog were present as a result of both the geology and the uncontrolled beaver activity. Moist grassy openings were also present, these often being the result of past beaver ponds. Though few dry upland grassy open ings of any size existed, small patches were present throughout the area CREEKN COMMANDA • I
SCALE 0 10 20 30 KILOMETERS -HIGHWAYS a STUDY AREA SOUND
Figure 1. Geographical location of the Himsworth Game Preserve. MATERIALS AND METHODS
Temporal and Spatial Distributions of Deer and Moose
Sign left by deer and moose was used to determine habitat use. Pel let grouJs and tracks formed the main data base.
Sampling was carried out on three occasions in the spring and summer of 1974. The first sample was started 10 May and completed 20 May. This
sample was used to derive habitat use for the winter period which ,.,as
considered to be the interval from leaf drop the previous fall (15 October)
through April. The second sample period was from 25 June to 5 July and
the third between 15 August and 26 August. These were used to show habitat use by moose and deer for the late spring - early summer (May-June) and
the mid- and late summer (July-August) periods.
Three hundred and ninety-five "locations" were established in 1973 at
the intersection of grid lines spaced at 200 m intervals in a north-south
direction and 160 m intervals in an east-west direction. Of the 395
locations, 334 were accessible and were located, and marked with a 1.5 m
stake. Trails were flagged between them to facilitate quick relocation.
The remaining 61 unmarked locations were in beaver ponds, swamps, muskeg,
or other inaccessible locations.
Four 2 x 20 m pellet plots were established at each accessible
location where possible. These were arranged systematically to fall along
the transect lines and not overlap (Fig. 2). At 46 of the 334 locations,
it was not possible to delineate all four plots because of adjacent beaver
ponds or other inaccessible sites, resulting in a loss of 53 plots.
Therefore, a total of 1,283 pellet plots was used, with a resulting sam
pling intensity of 0.5 percent of the total accessible area. All pellet
plots were checked in each of the three sample periods. - 7 -
At each accessible location, one 2 x 2 m track plot was made im mediately after completion of the first sample. Placing of these plots was subjective, usually in an opening of sufficient size between pellet
plots one and two (Fig. 2). Track plots were made by removing the litter and duff layers of the forest floor to expose the mineral soil. At a
few places difficulties were encountered due to stoniness, dryness, or
shrub and tree roots at a site, but generally all track plots had a soil
covering which would leave a relatively clear print should a moose or
deer stand on it. The track plots were checked in the second and third
sampling periods.
Observations of animal sightings, bedding sites, trails, tracks, and
pellet groups were made throughout the period May to mid-September to
supplement data from the pellet and track plots. The distance and direc
tion of these observations from the nearest location was recorded.
Habitat Description and Mapping
An intensive survey of the habitat at each location was made twice
during the study, once in late May and again in late July and early August.
Topography of the area around the stake was described as were such features
as terrain, drainage, proximity of water, and transition in forest stands.
Moisture of the site was subjectively classified as wet, very moist, moist,
damp, dry, or very dry. Sample plots for 1) ground (stems less than 1 m in
height); 2) understory (stems 1 to 5 min height); and 3) cover (stems
greater than 5 min height), vegetation were circles with radii of 1, 2,
and 5 m originating at the stake or at the end of one of the pellet plots,
depending on which was subjectively considered as being most representative
of the area (Fig. 2). Two sets of plots were sampled at 20 of the PELLET PLOT # 1 (2 M. x 20 M. l
GROUND VEGETATION SAMPLING PLOT . (1 M. RADIUS> D TRACK PLOT <2 M. x 2 M.) 0
ELLET PLOT # 4 (2 M. x 20 M.)
ELLET PLOT # 2 (2 M. x 20 M.)
UNDERSTORY VEGETATION SAr1PLING PLOT COVER VEGETA Tl ON SAMPLING PLOT ( 2 M. RADIUS> (5 M. RADIUS)
PELLET PLOT # 3 <2 M. x 20 M. l
Figure 2. The arrangement of sample plots at each accessible location. - 9 - locations because of differences in the forest types associated with these locations. These data were considered independently in the analysis.
All cover vegetation with a diameter at breast height (DBH) of 2.5 em or more was recorded by species, diameter, and frequency. Density of all three strata ( the percentage of the sample plot covered by vegetation in that class) was visually estimated and placed in one of the following categories: 1 to 5 percent; 6 to 25 percent; 26 to 50 percent; 51 to 75 percent; and 76 to 100 percent.
A habitat map was produced ·using Ward's cluster analysis method
(Wishart, 1970), aerial photographs, and field observations.
Analysis of Deer and Moose Sign in Relation to Habitat
Observations of deer and moose sign were analysed in. relation to habitat types, habitat factors, and geographical distribution.
The use of habitat types by these two species in October to April,
May to June, and July to August was evaluated by chi-square, in which the distribution of deer or moose observations by habitat types was compared to the observed total distribution of habitat types as determined from locations. Tests resulting in a P:O.OS were considered statistically sig nificant. Values for days of use and numbers of animals by habitat types were obtained for the first sample. These were calculated using a deposition rate of 13 groups per day for deer and moose, and deposition period of 217 days. A Scheffe's multiple range test (Snedecor and Cochran,
1967) was used to determine if there were significant differences in use of habitat types of deer and moose based on the mean density of pellet groups per plot.
Chi-square was used to analyse deer and moose sign in relation to - 10 - individual habitat· factors. Habitat factors analysed were: 1) total basal area; 2) basal area of trees 2.5 to 8.9 em DBH; 3) basal area of trees
9.0 to 26.6 em DBH; 4) total numbers of trees; 5) numbers of trees 2.5 to 8.9 em DBH; 6) numbers of trees 9.0 to 26.6 em DBH; 7) stand height;
8) cover density; 9) understory density; and 10) tree species. Tests resulting in a pSo.05 were considered statistically significant.
The geographical distributions of deer and moose were determined by plotting those locations with sign of either or both species on maps for each of the sample periods. Pellet groups were the only source of
sign in the winter period, whereas in late spring-early summer and mid summer-late summer both pellet groups and tracks on the sample plots and supplementary observations were included. Table 1. Locations established in the major habitat types of the Himsworth Game Preserve.
Habitat No. Percent of Percent of Area Percent of Percent of Type Locations total accessible (ha) total accessible
1 27 6.8 8.1 77.9 6.5 7.6
2 62 15.7 18.6 230.7 19.1 22.5
3 1 0.2 0.3 2.9 0.2 0.3
4 37 9.4 11.1 117.2 9.7 11.4
5 49 12.5 14.7 148.2 12.3 14.4
6 42 10.7 12.6 156.4 12.9 15.2
7 14 3.5 4.2 30.7 2.6 3.0
8 16 4.0 4.8 47.3 4.0 4.5
9 13 3.3 3.9 25.8 2.2 2.5
10 1 0.2 0.3 1.3 0.1 0.1
11 42 10.7 12.6 90.4 7.4 8.8
12 6 1.5 1.8 15.3 1.3 1.5
13 9 2.3 2.7 35.3 2.9 3.4
14 1 0.2 0.3 3.5 0.3 0.3
15 14 3.5 4.2 43.7 3.6 4.2
16 & 17 61 15.4 180.8 14.9 Table 2. Numters of locations within a habitat type on which tree species occurred.
Habitat No. Type Locations Tree Species
QJ QJ ..c: QJ () () () r-1 !-< ;:J ;:J QJ !-< QJ ..c: -M !-< !-< ..... r-1 ~ ., 4-1 p. p. .~. QJ .0 p. s ., ., p. l:l ,.!
Habitat Types
Seventeen major habitat types were delineated in the Himsworth
Game Preserve. Species cGmposition was the primary factor in identifying these types. The names given to the types represent those tree species which occurred on at least 80 percent of the locations within the type.
Stands in which deciduous tree species predominated covered 534.4 ha and were represented by six major types, four of which occupied mainly upland sites and two which generally occurred at lowland sites. The four upland types included the aspen, white birch, sugar maple, and aspen-white birch-red maple habitat types, and the two lowland types included the alder and the black ash habitat types. Mixed coniferous-deciduous stands were represented by the aspen-white birch-balsam fir habitat type which covered
148.2 ha. Five coniferous types were identified, including the balsam fir, white spruce, black spruce, white pine, and red pine types. The three remaining accessible types included the pipeline type, the open dry field type, and the open wet field type. Two habitat types were in accessible, with one type representing swamps and bogs and the other type open water. Table 1 gives the areas each of the habitat types covered and the number of locations sampled in each type, and table 2 lists the important tree species found in each habitat type.
Deer and Moose Habitat Use
October, 1973 to April, 1974
Deer sign was found in the aspen-white birch-red maple habitat type significantly more than expected in the winter (Table 3). The amount of sign in the open habitat types was significantly less than expected. The - 14 - number of observations of sign in the remaining types was about equivalent to their availabilities. One hundred and forty-four (82.8 percent) of 174 deer observations were in the aspen (17), white birch (33), aspen-white birch-red maple (26), aspen-white birch-balsam fir (27), balsam fir (21), and alder (20) types. These types collectively accounted for 77.7 percent of the total accessible locations.
Moose sign was significantly more numerous than expected in the white birch, aspen-white birch-red maple, and balsam fir types, with 70
(54.3 percent) of 129 observations being recorded in these types which accounted for 42.7 percent of the accessible locations. The aspen-white birch-balsam fir and pipeline habitat types had significantly less sign than expected, and the amount of sign present in the remaining types was as expected. The aspen (13), white birch (28), aspen-white birch-red maple (21), aspen-white birch-balsam fir (14), balsam fir (21), and alder
(15) types contained 86.9 percent of the 129 moose observations (Table 3).
The presence of deer sign indicated that this species showed some preference for sites having a basal area of 20.1 to 30.0 sq m/ha and 764 to 1274 stems/ha of trees 9.0 to 26.6 em DBH, a stand height of 12.4 to
18.4 m, a crown cover of 6 to 50 percent, and an understory cover of 6 to
25 percent. There was significantly less deer sign in openings in the winter period. (Figs. 3 to 11).
ThQ distribution of moOSQ sign indicatod that thio opccico ohowcd some preference for stands having a basal area of 0.1 to 5.0 sq m/ha and
127 to 637 stems/ha of trees 2.5 to 8.9 em DBH, a crown cover of 1 to 5 percent, and an understory cover of 1 to 5 percent. Moose sign was rare in open sites (Figs. 3 to 11).
Most observations of deer and moose sign (at least 65 percent of Table 3. Numbers of Accessible Locations with Deer and Moose Sign by Habitat Types in October-April.
Habitat No. Deer Moose Type Locations* Sign Sign
1 27 8.1)** 17 9.8) 13 10.1)
2 62 18. 6) 33 19. 0) 28 21. 7)
3 1 0.3) 1 0.6) 0 o. 0)
4 37 11.1) 26 14. 9) 21 16.3)
5 49 14. 7) 27 15. 5) 14 ( 10.9)
6 42 12. 6) 21 ( 12.1) 21 ( 16.3)
7 14 4.2) 9 5.2) 3 2.3)
8 16 4.8) 6 3.4) 5 3.9)
9 13 3. 9) 10 5.7) 7 5.4)
10 1 0.3) 1 0.6) 0 o. 0)
11 42 12. 6) 20 11. 5) 15 ( 11. 6)
12 6 1.8) 1 0.6) 0 o. 0)
13 9 2. 7) 0 0.0) 2 1.6)
14 1 0.3) 0 o. 0) 0 0. 0)
15 14 4.2) 2 1.1) 0 0. 0)
Total 334 (100.1) 174 (100. 0) 129 (100.1)
p = 0.001 0.006
Represents the expected distribution on the Himsworth Game Preserve.
** Figures in parentheses expressed as percent of total. - 16 -
the 174 deer observations and 129 moose observations) were recorded at
sites with a basal area of 0.1 to 5.0 sq m/ha and 127 to 1274 stems/ha of
trees 2.5 to 8.9 em DBH, and a basal area of 0.1 to 30.0 sq m/ha and 127
to 1911 stems/ha of trees 9.0 to 26.6 em DBH. Stands with a height gre~t
er than 12.3 m also accounted for a minimum 65 percent of the observations
of sign of both species, as did stands with a crown cover of 1 to 50 per
cent, and an understory cover between 1 and 25 percent (Figs. 3 to 11).
Deer sign occurred at sites having aspen, white birch, red maple,
balsam fir, and/or white pine significantly more than expected. White
birch occurred at 78.7 percent (137) of those locations with deer sign,
but was present at only 67.1 percent of 334 locations. Similarly, aspen,
red maple, and/or balsam fir were present at 69.5 percent (121), 36.8 per
cent (64), and 54.0 percent (94) of the 174 locations with deer sign res
pectively, but were found on only 62.6 percent, 29.9 percent, and 47.0
percent of ·all locations. Sign also indicated some preference by deer for
sites with white pine and white spruce. There was little use of locations with alder. Moose sign occurred significantly more than expected at sites
having aspen, white birch, red maple, and/or white spruce, with these tree
species occurring at 72.1 percent (93), 74.4 percent (96), 39.5 percent
(51), and 35.6 percent (46) of the 129 locations with moose sign. Balsam
fir was present at sites with moose sign to a greater degree than expected, while willow and black ash occurred less frequently than expected (Fig. 12).
There was considerable overlap in the distribution of deer and moose
sign on the study area from October to April, with sign of both species
being found at 83 of the 334 locations. The greatest densities of deer,
based on mean number of pellet groups per plot, were in the northwest
corner of the study area, a region seldom used by moose. The greatest 70
OCTOBER,1973 - APRIL197~ MAY + JUNE,197~ 0 DEER: N= 17~ (F 0. 086) 0 DEER: N= 38 (p= 0. 569) (;Sl fiOOSE:N= 129 (p= 0.026) (;Sl JolOOSE:N= 31 (p= 0.81~)
-10
-20 2 3
JULY + AUGUST, 197~ n DEER: N= 28 (p= 0.039) (;Sl MOOSE:N= 12 (p= 0.926)
4
Figure 3. Distribution of deer and moose sign in relation to the dist-
ribution of basal area of trees 2,5 to 8.9 ern DBH. 35 r- V> z 30 C> s 25 r--- 5: ..... ~ V> 20 ""0 -' < 15 o--~ r--- 1:; 10 g:j 0> w ~ ""u.. 0 n 0 0.1- 10.1- 20.1- > 10.0 20.0 30.0 30.0 BASAL AREA I HECTARE
+20 OCTOBER,1973 - APR!L197~ HAY + JUilE,.l97~ V> z V> DEER: i~= 17~ (p= O.OOll DEER: N= 38 (p= 0.~~1) C> z 0 1J o---- C> IS;! f'()OSE:N= 129 (p= 0.068) IS;! HOOSE:N= 31 (p= 0.796l
JULY + AUGUST,197~ 0 DEER: N= 28 (p= 0.195) IS;! I·IOOSE:N= 12 (p= 0.371)
4 Figure 4. Distribution of deer and moose sign in relation to the
distribution of basal area of trees 9.0 to 26.6 em DBH. V) g 30~i ~ 25 ""~ "'0 20 <>:--' 1- 0 1- 15 s 10 ~
0.1- 10.1- 20.1- 30.1- > 1o.o 20.0 3o.o qo,o qo,o BASAL AREA I HECTARE
+20 OCTOBER,1973 - APRIL197q MAY + JUllE,1974 0 DEER: N= 17q (p= O.OOll 0 DEER: N= 38 (p= 0.070) (:S;I MOOSE:N= 129 (p= 0.036) l:s:l HOOSE:N= 31 (p= 0.335)
'"] -20 2 3
JULY+ AUGUST,1974 0 DEER: N= 28 (p= 0.128) (:S;I MOOSE:N= 12 (p= 0.06qJ
4
Figure 5. Distribution of deer and moose sign in relation to the
distribution of total basal area of trees. 45 1
(I) ""0 37.5
w~ 30 (I) ""C) --' 22.5 ~ t; 15 "'w a!w 7.5 ...=...._ 0 127- 764- 1401- 2038- > 637 1274 1911 2548 2675 STEMS I HECTARE
+20 OCTUBER,1973 - APR!Ll974 MAY + JUNE, 1974 ~en C)"' 0 DEER: N= 174 (p= 0.315> n DEER: N= 38 (p= 0.771) -C) 1-- ~ MOOSE:N= 129 (p= 0.003) 1:SJ HOOSE:N= 31 (p= 0.938)
V>~~ ...... +10 ""V> C)"" ...... C) V>--' C)..: C) 1- ~~ 0 =u ~ffi "'5 >-u=...... fijLL -10 :O.N ~V> = :> "-"'HE -20 2 3
JULY + AUGUSL1974 ODEER: N= 28 (p= 0.127> fSl MOOSE:N= 12 (p= 0,351)
4
Figure 6. Distributio~ of deer and moose sign in relation to the
distribution of density of trees 2.5 to 8.9 em DBH. 35 1
30 5"' Si 25 if
127- 76Q- 1Q01- 637 127Q 1911 2038 smts I HECTARE
+20 OCTOBER,1973 - APR!Ll97Q W\Y + JUNE,197Q oz"' -o"""' 0 DEER: N= 17Q (p= 0. OOll 0 DEER: N= 38 (p= O.QQ7) >-- ""'>-&::; ~ HOOSE:N= 129 (p= 0,092) ~ 1·\00SE:N= 31 (p= 0.723> "'"-'"-'"' +10 ....,o"'"""""' "'_,c..: 0 >- ""~.... ,... 0 "''-'wz ww<=>= ,...... ,"" '-'"' ""'-'-=w ... -10 ~"' "'=u..z .. E: -20 2 3
JULY + AUGUST,197Q 0 DEER: N= 28 (p= O.Q62l ~ NOOSE:N= 12 (p= 0.365)
4
Figur~ 7. Distribution of deer and moose sign in relation to the
distribution of density of trees 9.0 to 26.6 cm'DBH. 2 ~ 30 +20 0 OCTOBER,1973 - APR!L,l974 ;:: 0 DEER: N= 174 (p• 0.002) 25 :> ~ ~IUOSE:N= 129 (p= 0.059) ""'ffi +10 V) 20 _,"'0 I- 15 ""'0 I- i:; z: 10 ~ -10 u.J"' 5 U-"" ~ 0 -20 127- 754- 1401- 2038- 2575- 3312- 537 1274 1911 2548 3185 3822 3950 STEMS I 5ECTARE
MAY + JUN£,1974 JULY + AUGUST, 1974 0 DEER: N= 38 (p• 0.115) 0 DEER: N= 28 (p= 0.175) ~ MOOSE: N= 31 (P• 0. 370) ~ 1100SE:N= 12 (p= 0.535)
3 4
Figure 8. Distribution of deer and moose sign in relation to the
distribution of total density of trees. Cl) 2 1 -o"""'00:: Cl) >-- OCTOBER,1973 - APRIL,1974 z: 30 "">- +20 ;::0 ~::;: 0 DEER: N= 174 (pa 0. OOll 25 "'"'~~ ~ MOOSE:N= 129 (p= 0.072) O
HAY + JUNL1974 JULY + AUGUST, 1971( n DEER: N= 38 (p= O.Ol7J 0 DEER: N= 28 (p= 0.176) ~ liOOSE:N= 31 (p= 0.140) ~ l-lOOSE:N= 12 (p= 0.226)
3 4
Figure 9. Distribution of deer and moose sign in relation to the
distribution of stand height, 35 1
30 "'""0 ~ OCTOBER,1973 - APRIL,1974 25 MAY,1974 - AUGUST,1974 w~ "' 20 0"'__, c< 1- 15 0 1- >-u 10 ffi "'w 5 "'a: "- 76- 51- 26- 6- 1- 100 75 50 25 5 % CROWN COVER
OCTOBER,1973- APRIL,1974 W\Y + JUNL1974 +20 ODEER: N= 174 <•= O.OOOl 0 UEER: N= 38 (p= 0.05ll ES;IMOOSE:N= 129 (p= 0.033) ES:INOOSE:N= 31 (p= 0.584)
+10
-10
-20 2 3 JULY + AUGUST.1974 0 DEER: N= 28 (p= 0.318) IS;l f\UUSE:N= 12 (p= 0.007l
r- I I L L
4
Figure 10. Distribution of deer and moose sign in relation to the
distribution of percent crown cover. 1 "';;::: 60 ;::0
+20 OCTOBER,l973 - APRIL197q flAY + JUNE,l97q ~ DEER: ll= 17q (p= 0. OOll ~ ZV> DEER: N= 38 (p= 0, 937l "'o;;::: -o MOOSE:N= 129 (p= 0.037l . MOOSE:N= 31 (p= 0.770> >--
JULY + AUGUST ,197q 0 DEER: N= 28 (p= 0, 7q0) fSS flJOSE:N= 12 (p= 0.832)
4
Figure 11. Distribution of deer and moose sign in relation to the
distribution of percent understory cover. 70
60 0"' so ~w 40 "'0 -' 5"" 30 I- >-u 20 z w => 0>w 10 ..[f ASPEN WHITE RED ALDER WILLOW BIRCH I·IAPLE
OCTOBER,1973 - APRIL1974 MAY + JUNE, 1974 +20 0 DEER: N= 174 0 DEER: N= 38 "' !S;I MOOSE:N= 129 ~ NOOSE:N= 31 -o"""'0-= I-- ~~ +10 "-'"'"'"-' coo """'0 w "'-'o..: 0 I- :E.O ..... I- 0 "'~ "-'"'ww "'1'r >- w zu..'-'"' -10 w ...,.,a=>"" "'=> ...u..z :c -20 2 3
J~LY + AUGUST, 1974 0 DEER: N= 28 ~ MOOSE: N= 12
L l
4
Figure 12. Occurrence of deer and moose sign in relation to the occur-
renee of individual tree species. 70 1
60 "'~ I- :;: 50 ...,<" "' 40 ~__, BALSMl WHITE BLACK WHITE BLACK FIR SPRUCE SPRUCE Pl~E ASH +20 OCTOBER-1973 - APR!Ll974 NAY + JU1JE, 1974 0 DEER: N= 174 0 DlER: N= 38 ~ HOOSE: N= 129 [:S3 HOOSE: N= 31 +10 -10 -20 J 2 3 JULY + AUGUSU974 0 DEER: N= 28 [:S3 NOOSE : N= 12 4 Figure 12. Continued. - 28 - concentration of locations adjacent to one another with moose pellet groups was in the southwest section of the study area, and in comparison to the remainder of the area, relatively little sign of deer was found in this region. Most overlap occurred in a broad diagonal band stretching north east to southwest across the study area. Based on pellet groups, an estimated 38 deer and 21 moose used the Himsworth Game Preserve from October, 1973 to April, 1974. All individuals were assumed to have remained on the area for the total deposition period and possible immigration or emigration was ignored. May to June, 1974 Sign indicated that the alder habitat type was the only type used by both deer and moose more than expected on the basis of its availability. Nine (23.7 percent) of 38 deer observations and 5 (16.1 percent) of 31 moose observations were recorded in this type, which represented only 12.6 percent of all accessible locations. The white birch, balsam fir, and pipeline types collectively represented 35.4 percent of the sample locations and accounted for 55.3 percent (21) of the locations with deer sign and 29.1 percent (9) of the locations with moose sign. On the other hand, the aspen, aspen-white birch-red maple, aspen-white birch-balsam fir, white spruce, black spruce, and white pine types, which made up 46.8 per cent of the total locations sampled, contained 51.5 percent (16) of the locations with moose sign but only 18.4 percent (7) of the locations with deer sign (Table 4). Deer sign was distributed in a significantly different manner from the observed distribution of stand height with observations being more numerous than expected in stands with a crown height of 15.4 to 18.4 m and - 29 - at sites with no trees (Fig. 9). Deer sign was found more commonly than expected at sites with no trees or understory (Fig. 3 to il) and at locations where the total basal area was between 20.1 and 40.0 sq m/ha (Fig. 5). The distribution of sign indicated that deer tended to avoid locations with a basal area of 0.1 to 20.0 sq m/ha (Fig. 4 and 5), a stand height of 12.4 to 15.3 m (Fig. 9), and a crown cover of 1 to 5 per cent or 76 to 100 percent (Fig. 10). Moose sign also occurred more frequently than expected at sites with no trees or understory (Fig. 3 to 11). Distribution of sign indicated some preference in use by moose for locations where the basal area of trees 9.0 to 26.6 em DBH was 0.1 to 10.0 sq m/ha (Fig. 4) and where total basal area was also 0.1 to 10.0 sq m/ha (Fig. 5). Sign was more abundant than ex pected at sites with 127 to 1274 stems/ha (Fig. 8) and with an uneven stand height and a height of 3.1 to 6.1 m (Fig. 9). Sites which offered 0 to 5 percent crown cover were used proportionately more than expected (Fig. 10). Sign of moose was less common than expected at locations where the basal area of trees 9.0 to 26.6 em DBH was 10.1 to 20.0 sq m/ha (Fig. 4), there were more than 2038 stems/ha (Figs. 6 and 8), and stand height ex ceeded 15.4 m (Fig. 9). The association of deer and moose sign with individual tree species changed from winter to May and June. Whereas alder and willow were found at 18.6 percent and 10.2 percent of the 334 locations respectively, 28.9 percent (11) and 15.8 percent (6) of the 38 locations with deer sign had these species. Excluding balsam fir, the remaining tree species were not found at as many of the sites with deer sign as was expected. Sign of moose occurred at locations with alder (observed = 25.8 percent; expected 18.6 percent), balsam fir (observed= 54.8 percent; expected 47.0 percent), Table 4. Numbers of Accessible Locations with Deer and Moose Sign by Habitat Types in May~June. Habitat No. Deer Moose Type Locations* Sign Sign 1 27 8.1)** 1 2.6) 2 6.4) 2 62 ( 18.6) 9 23. 7) 5 16.1) 3 1 0.3) 0 ( 0. 0) 0 0. 0) 4 37 ( 11.1) 2 5.3) 4 12. 9) 5 49 14. 7) 3 7. 9) 5 ( 16.1) 6 42 ( 12.6) 6 ( 15.8) 3 9. 7) 7 14 4.2) 1 2.6) 2 6.4) 8 16 4.8) 0 0. 0) 2 6.4) 9 13 3.9) 0 0. 0) 1 3.3) 10 1 0.3) 0 0.0) 0 0.0) 11 42 12. 6) 9 23. 7) 5 ( 16.1) 12 6 1. 8) 0 0. 0) 0 0. 0) 13 9 2.7) 1 2.6) 1 3.3) 14 1 0.3) 0 0. 0) 0 0.0) 15 14 4.2) 6 ( 15.8) 1 3.3) Total 334 (100.1) 38 (100.0) 31 (100. 0) p = 0.014 0.999 * Represents the expected distribution on the Himsworth Game Preserve. ** Figures in parentheses expressed as percents of total. - 31 - white spruce (observed= 38.7 percent; expected= 29.3 percent), and/or white pine (observed = 12.9 percent; expected = 7.2 percent) to a larger degree than expected. Aspen, white birch, red maple, and willow were not observed at locations with moose sign in proportion to their availabilities, although black ash and black spruce were (Fig. 12). The geographic distributions of deer and moose sign for May to June indicated the two opecies generally used different regions of the Preserve. Deer sign was most numerous in the northwest quarter and along the western side of the area. A number of observations were also made in the southeast section. Moose sign was most numerous in a strip from the centre of the study area southwest to the pipeline. Moose sign was also found in the southeast section but north of the region where most of the deer sign oc curred. Deer and moose sign were found at the same locations on only six occurrences. July to August, 1974 Only 28 observations of deer sign and 12 observations of moose sign were recorded on the plots in the third sample. The presence of sign in dicated that deer used the habitat types as expected based on their avail abilities. Eighteen of 28 observations of sign were from five habitat types, including the aspen (3), white birch (4), balsam fir (4), alder (4), and pipeline (3) types. Eight of 12 observations of sign of moose were from the white birch (3), aspen-white birch-balsam fir (2), and the alder (3) types (Table 5). Deer sign was more abundant than expected at treeless locations during July and August (Fig. 3 to 11). Only 2 of 28 observations of sign were in areas where total basal area exceeded 30.0 sq m/ha (Fig. 5) and - 32 - none was recorded at sites with a basal area of more than 30.0 sq m/ha of trees 9.0 to 26.6 em DBH (Fig. 4). Deer utilized treeless sites and sites with a stand height of 9.3 to 15.3 m more than expected. They did not make extensive use of areas with a stand height exceeding 15.3 m (Fig. 9). Moose seemed to use openings and relatively open forest stands more than expected based on sign in July and August (Fig. 3 to 11). None of 12 observations of sign were recorded at stands where total basal area was greater than 30.0 sq m/ha (Fig. 5) and the basal area of trees 9.0 to 26.6 em DBH was less than 20.0 sq m/ha at 11 of 12 sites with sign (Fig. 4). Similarly only two observations of sign were made at sites where there were more than 1911 stems/ha (Fig. 8), and 11 of 12 observations were recorded at locations where there were fewer than 1274 stems/ha of trees 9.0 to 26.6 em DBH (Fig. 7). Deer sign occurred at sites with white spruce and/or willow more than expected, and with white birch, red maple, balsam fir, and/or black spruce less than expected. The number of observations of sign at locations with aspen, alder, black ash, and white pine was as expected. Moose sign occurred at sites with alder and/or willow more than expected but at locations with white birch, red maple, balsam fir, and/or white spruce less than expected (Fig. 12). Apparently moose and deer generally utilized different regions of the Preserve during July and August with no overlap of sign being recorded at any location on the sample plots and only 4 instances of overlap being found with the supplementary observations. Table 5. Numbers of Accessible Locations with Deer and Moose Sign by Habitat Types in July-August. Habitat No. Deer Moose Type Locations* Sign Sign 1 27 ( 8.1) 3 (10. 7) 1 ( 8. 3) 2 62 ( 18. 6) 4 (14.2) 3 (25. 0) 3 1 0.3) 0 ( 0.0) 0 ( 0.0) 4 37 ( 11.1) 2 ( 7.1) 0 ( 0.0) 5 49 ( 14. 7) 2 ( 7 .1) 2 (16. 7) 6 42 12.6) 4 (14.3) 0 ( 0.0) 7 14 ( 4.2) 2 ( 7.1) 0 ( 0.0) 8 16 4.8) 1 ( 3.6) 1 ( 8. 3) 9 13 3. 9) 1 ( 3.6) 1 ( 8. 3) 10 1 0.3) 0 ( 0.0) 0 ( 0.0) 11 42 12. 6) 4 (14. 3) 3 (25. 0) 12 6 l. 8) 1 ( 3.6) 0 ( 0.0) 13 9 2. 7) 1 ( 3.6) 0 0. 0) 14 1 0.3) 0 ( 0.0) 0 ( 0.0) 15 14 4. 2) 3 (10.7) 1 ( 8. 3) Total 334 (100.1) 28 (99. 9) 12 (99. 9) p = 0.930 0.914 * Represents the expected distribution on the Himsworth Game Preserve. ** Figures in parentheses expressed as percents of total. DISCUSSION Habitat Use by Deer and Moose October to April The temporal and spatial distribution of moose and deer in relation to habitat in October to April was difficult to determine. This was be cause habitat use was based on the distributions and densities of pellet groups as observed in early May, and though this indicated an animals' presence in winter, it did not reveal when it was present, nor did it ex clude other areas as possibly having been used. Severinghaus and Cheatum (1956) reported that deer tended to defecate on a "loitering" ground or at their bedding sites, and not necessarily at feeding sites. Des Meules (1962) stated that moose often defecated in or near bedding sites. In northeastern North America, deer often shift from a summer to a winter range at the onset of colder weather with or without snow, and eventually congregate (yard) at sites apparently selected for their shelter rather than food value (Olsen, 1938; Hammerstrom and Blake, 1939; Severinghaus, 1947; Webb, 1948; Carlsen and Farmes, 1957; Gill, 1957; Behrend, 1966). Density of deer within a yard varies with the time of winter, with the greater concentrations occurring as winter progresses (Telfer, 1970), and the severity of the winter, with higher densities being reported in the more severe winters (Severinghaus and Cheatum, 1956). It is known that the area on the Himsworth Game Preserve where the highest densities of deer pellet groups served as a traditional yarding area for deer. Behrend (1966), Verme and Ozoga (1971) and Ozoga and Gysel (1972) have all reported the initiation of winter concentrations of deer to be influenced by regularly increasing levels of windchill in late fall, and ------ - 35 - when windchill was constantly high, the degree of concentration was most influenced by the; deer's "sinking depth" in snow. Conversely, the ending of winter concentrations was influenced by a regularly decreasing level of windchill in early spring. Lower than normal temperatures in November of 1973 (Atmospheric Environment Service, 1975) apparently initiated an early movement of deer to yarding areas. There was little deep snow on the ground for more than one or two days at a time and deer did not move into coniferous shelter for any extended period in early winter. Deer probably did not concen trate in the yard in the study area until February or March. The fact that the best shelter types (considered to include the balsam fir, white spruce, black spruce, and aspen-white birch-balsam fir types) were not used to a greater extent than expected in October to April is misleading. The balsam fir type, based on the density of pellet groups per plot, supplied the greatest number of deer days of use and also supported the highest deer density. The aspen-white birch-balsam fir type ranked third and fifth in these two categories respectively. The white spruce type had the third largest density but was low in terms of days of use because of its relatively small availability. Conversely, the white birch type which supplied the second highest number of deer days of use supported one of the lowest densities. These values were calculated for the entire period of October to April and do not indicate shifts in habitat use as conditions became more severe. In relation to individual factors, deer distribution was influenced by the numbers and basal area of trees 9.0 to 26.6 em DBH, stand height, and crown cover, all of which are important when considering shelter. It appears, therefore, that until December the deciduous forest - 36 - types were used as much as the coniferous shelter types. After December, deer probably used the mixed and coniferous habitat types most of the time, only using the deciduous types when climatic conditions were favourable. It was more difficult to determine habitat use by moose in the winter because of different behaviour than deer. Unlike deer, moose movements are only rarely decreased as a result of severe climatic con ditions in this portion of their range and, therefore, habitat use was not as predictable as it was for deer. Moose do move from summer to winter ranges in Ontario though this movement is not as spectacular as that found in other parts of North America (LeResche, 1974). LeResche (1974) classified moose migrations in Ontario as Type A, which involve short distance movements between two seasonal ranges with little change in elevation; travel over great dis tances would achieve little change in climate or habitat. Peek (1971), des Meules (1964), Berg and Phillips (1972), and Prescott (1974) reported the greatest aggregations of moose occurred in late fall and early winter (immediately after rut). These aggregations decreased as winter progressed. Peek (1971) associated the reduction in group size with the movement of animals to denser cover. Des Meules (1964) reported that as snow depths increased, the numbers of moose in each group decreased. Prescott (1974) cites Houston (1971) who suggested that within the boreal forest biome an evolutionary premium may have been set on small group sizes, since single moose or small groups could more suc cessfully find and utilize scattered winter forage under deep snow con ditions. At this time of year the home range is relatively small (Houston, 1968; Goddard, 1970; Phillips et al., 1973). Moose will use those areas which supply abundant food in winter. - 37 - Berg and Phillips (1972) considered winter habitat to be aspen and tall willow stands. Telfer (1970) noted moose in January appeared to be dis tributed more in relation to food abundance than shelter. However, under severe climatic conditions, moose, like deer, will seek shelter in dense coniferous stands (Prescott, 1974). These conditions are produced most often by excessive snow depth, although snow hardness and density can have a similar effect (Peek, 1971; Coady, 1974). During the present study there may have been aggregation of moose in late fall and early winter after which there was dispersal into smaller groups with overlapping range. There were no extended periods when snow conditions were such that moose would have had to move into dense coniferous shelter. A few localized areas where adjacent plots had higher densities of pellet groups than the surrounding locations suggested moose movements may have been restricted for short periods of time. Open forested types were used predominantly throughout the winter, with hardwoods being most important (c.f. Peek, 1971). The white birch type supplied the greatest number of days of use, with the balsam fir, aspen, and aspen-white birch red maple types being next in importance. The white pine, aspen-white birch-red maple, and aspen types were found to have the highest densities over the winter period. May to June The only habitat type used by both moose and deer to a greater degree than expected during May and June was the alder type. Differences were found in the use of the other types which may have prevented much ecological overlap. Few studies have been done on habitat use by deer in the summer months. Kahn and Mooty (1971), working in Minnesota, stated changes in - 38 - the use of habitat types during the summer corresponded to changes in preference of forage species, and land containing a good distribution of disturbed areas and young deciduous stands provided good summer range for deer. In early spring the importance of openings as areas producing the first green vegetation has been noted for deer (McCaffery and Creed, 1969; Kohn and Mooty, 1971; McCaffery et al., 1974). In the Himsworth study, deer showed a significant preference for the pipeline type but whether this was being used as a feeding area or simply as a travelway is not known. However, the amount of sign in the alder type may indicate animals were using openings as this type often bordered clearings. Deer may pass through the alder type to enter the openings, and it is likely they use the cover provided by the alder. Deer were never seen in the middle of openings but were occasionally observed along the edge. Also, it was not uncommon to find trails through alder. Another possible reason for the heavy use of the alder type may have been related to fawning activities. Such sites would provide excellent cover and had the added advantage of a frequent intermixture uf sllghtly elevated areas supporting mature aspen and white birch. In the present study deer also were found in deciduous types at this time of year. Kohn and Mooty (1971) reported a high use of aspen-dominated stands as revealed by tracks and McCaffery and Creed (1969) noted con sistently higher deer activity in aspen stands as compared to northern hardwoods. At Himsworth, the white birch type received the greatest use of all deciduous types. This was to be expected as this type was more common than the other hardwood types. It is likely the shrub and herbaceous growth within the deciduous types was attractive to deer as such vegetation has been shown to be important in the summer diet (Stegman, 1937; Atwood, - 39 - 1941; Dunkeson, 1955; Stiteler and Shaw, 1966; Skinner and Telfer, 1974). Little is known about use of habitat in summer by moose. It appears moose commonly utilize low-lying and open areas in May and June. Phillips et al. (1973) reported moose in northwestern Minnesota showed a marked increase in the use of low and open willow types in late April and were frequently observed in openings. This led them to conclude there was a distinct difference in the habitat occupied during the summer and fall as compared to that of late winter and early spring. Van Ballenberghe and Peek (1971) suggested moose may spend considerable time in the lowlands adjacent to water, and Peek (1971) noted moose were often in stands ad jacent to waterways which frovided aquatic plants. In the current study moose sign was closely associated with beaver ponds and other waterways. The use of aquatic vegetation by moose was not evaluated but considering the numerous studies which have shown irs importance in the diet (MacMillan, 1953; Peterson, 1955; de Vos, 1956; Peek, 1971; Van Ballenberghe and Peek, 1971; Cobus, 1972) and the large amount present on the Himsworth preserve, it is believed this was an important part of the diet in the summer. There is the possibility that some overlap of deer and moose use did occur at such sites but it was not possible to determine this as these sites were considered inaccessible. The distribution of moose and deer .sign indicated that animals often tended to occupy a specific area. Such behaviour has been commonly re ported for moose. Van Ballenberghe and Peek (1971) found that in summer 2 cows often stayed in an area of less than 2.5 km • Houston (1968) and 2 de Vos (1958) found moose generally had summer ranges of less than 3.2 km . Goddard (1970) reported that in northwestern Ontario moose of both sexes and all ages were quite sedentary in summer. Deer in north-central - 40 - 2 Minnesota had home ranges of 0.7 to 1.9 km between June and August (Kohn and Mooty, 1971). Several studies in the southeastern United States 2 found the home ranges of deer to be generally less than 2.6 km in the summer (Progulske and Baskett, 1958; Thomas et al., 1964; Micheal, 1965; Marchinton, 1966; Marshall and Whittingdon, 1968; Sparrowe and Springer, 1970). July to August There was apparently little change in the use of habitat by deer on the Himsworth area from May to June to July and August. There was a decrease in the use of the alder and white birch habitat types in July and August as compared to May and June, although the alder type continued to be utilized more than expected. Fewer observations of deer sign were recorded in the pipeline habitat type at this time but if the treeless types are considered collectively, their importance did not decrease from May and June. There was little difference in the use of the upland decid uous types (considered to be represented by the aspen, white birch, and aspen-white birch-red maple habitat types) by deer from late spring - early summer to mid- and late summer·, with approximately 30 percent of the ob servations of sign being found at such sites in both samples. Similarly deer tended to use the aspen-white birch-balsam fir type as much in July and August as in May and June. There was a tendency for deer to use stands with a greater number ·of small trees and less total basal area in July and August than in May and June. Behrend (1966), McCaffery and ·creed (1969), and Kohn and Mooty (1971), working in New York, Wisconsin, and Minnesota respectively, also reported no major changes in the use of habitat by deer in summer. In Minnesota it was found the use of upland deciduous areas and upland mixed areas by deer - 41 - remained relatively stable throughout summer and the use of upland coni ferous stands declined somewhat as summer progressed (Kohn and Mooty, 1971). Though a decline in use of openings by deer has been reported to occur in mid-summer, relatively high use of these clearings resumed again in August (McCaffery and Creed, 1969; Kohn and Mooty, 1971). In the current study, moose continued to make extensive use of the alder habitat type and also showed an increase in use of the upland deciduous types in July and August. The distribution of sign indicated that moose avoided the more densely stocked stands and that there was a shift in use from moderately stocked stands in May and June to more sparse ly stocked stands in July and August. There was also an increased use of taller communities in mid- and late summer compared to late spring - early summer. Peek (1971) reported a decrease in movements of moose from 15 August to 7 September. The same situation seems to have occurred in the Himsworth area. Peek (1971) also noted a continued use of lowland sites by moose throughout the summer and that there was also an jncreased use by moose of taller plant communities as summer progressed. Phillips et al. (1973) found the low open habitat types to be used by moose in July and August. In July and August, deer sign was found scattered over much of the area. On Lhe uLhet: hand, moose sign tended to be localized. Such ob- servations can be explained in part by differences in the parent off- spring association of moose and deer. At least three doe - fawn associations and two cow - calf groups were present in the area. Whereas deer can be considered "concealers" (the doe conceals and does not associate with the - 42 - fawn for various intervals of time), moose are "followers" (the cow does not leave the side of the calf for long periods of time) (Stringham, 1974). The fawn for the first couple of months has a much smaller home range than the doe, but the latter must return to the fawn at frequent intervals to allow the fawn(s) to suckle. This would account for the accumulation of deer sign in specific localities in May and June. However, by the age of four months, the fawn will accompany the doe and thus will occupy the same home range as her though its daily activity will not be as great (Micheal, 1965; Byford, 1970). With this movement by the fawn and doe over the entire home range, there would be a reduction in the probability of a buildup of pellet groups in one locality. Moose calves consistently remain with their coes. This cow-calf relationship plus the association of moose with waterways could account for the buildup of moose sign in specific areas. LITERATURE CITED Anderson, R.C. 1963. The incidence, development, and experimental trans mission of Pneumostrongylus- tenuis Dougherty- of- the meninges--of the white-tailed deer (Odocoileus virginianus borealis) in Ontario. Can. J. Zool. 41: 775-792. Anderson, R.C. 1964. Neurologic disease in moose infected experimentally with Pneumostrongylus tenuis from white-tailed deer. Path. Vet. 1: 289-322. Anderson, R.C. 1965. An examination of wild moose exhibiting neurologic signs in Ontario. Can. J. Zool. 43: 635-639. Anderson, R.C. 1972. The ecological relationships of meningeal worm and native cervids in North America. J. Wildl. Dis. 8: 304-310. Atmospheric Environment Service. 1975. Personal communication. Atwood, E.L. 1941. White-tailed deer foods of the United States. J. \olildl. Manage. 5: 314-332. Banasiak, C.F. 1961. Deer in Maine. 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Clustran IA user manual. Computing Laboratory Univ. of St. Andrews, St. Andrews, Scotland. 135 pp. 11 tfi ~f,\ERICAN At. ~oo~ Winnipeg, Manitoba March 1975 88 DELEGATES TO THE 1975 CONFERENCE ATTENDED FROM ALL PARTS OF THE MOOSE RANGE OF NORTH AMERICA