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ALASKA DEPARTMENT OF FISH AND GAME
J U N E A U, A L A S K A
MOOSE
STATE OF ALASKA
William A. Egan, Governor
DEPARTMENT OF FISH AND GAME
James W. Brooks, Commissioner
DIVISION OF GAME
Frank Jones, Director
Donald McKnight, Research Chief
RESEARCH CENTER REPORT
By
Robert E. LeResche
Albert W. Franzmann and
Paul D. Arneson
Volume XIII
Project Progress Report
Federal Aid in Wildlife Restoration
Project W-17-4, Jobs l.lR, 1.2R, 1.3R and 1.4R
Persons are free to use material in these reports for educational
or informational purposes. However, since most reports treat only part
of continuing studies, persons intending to use this material in scien-
tific publications should obtain prior permission from the Department of
Fish and Game. In all cases tentative conclusions should be identified
as such in quotation, and due credit would be appreciated.
(Printed November 1973)
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ACKNOWLEDGEMENTS
Many people helped gather data and prepare this report, and we extend
our sincere thanks to the following:
James Davis and Paul LeRoux, who assisted in tagging and relocating
moose and compiling data for the movements and population identity study.
Robert Rausch, who coordinated all the work reported here and sug-
gested directions it might take.
John Oldemeyer, who helped in many ways throughout the studies, and
who was especially helpful with statistical advice.
David Johnson, who was responsible for the field work involved in
the orphan calf survival study. Charles Lucier and his laboratory staff
provided tooth aging, marrow analysis, supply procuring and record keeping
services in a timely and efficient manner.
John Hakala, Bob Richey and Bob Seemel of the u.s. Fish & Wildlife
Service cooperated extensively with the project and provided some unpub-
lished data cited here.
Richard Bishop and John Coady contributed hours of stimulating
discussion of moose ecology.
Drs. James Peek, Peter Lent, Jean Bedard and Ed Telfer made helpful
comments on parts of the manuscript.
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State:
Cooperators:
Project No.:
Job No. :
JOB PROGRESS REPORT (RESEARCH)
Alaska
Alaska Department of Fish and Game, U.S. Bureau of Sport
Fisheries and Wildlife, Kenai National Wildlife Range
W-17-4
l.lR
Project Title: Big Game Investigations
Job Title: Moose Productivity and
Physiology
Period Covered: July 1, 1971 through June 30, 1972
SUMMARY
Natality, mortality and yearling recruitment of moose (Alces alces
gigas) were determined in four 2.6 km 2 (one square mile) pens. Calf
production and survival-to-yearling were 11 percent and 0 percent,
respectively, both lower than in previous years. One pen, stocked with
44 moose in autumn, contained only 14 animals the following spring
because of natural mortality. At the close of the reporting period, the
four pens contained 52 moose, or five per km2 (13 per square mile).
This represents a decline of 31 percent from the pre~eding spring. The
total loss of calves was attributed primarily to the early arrival and
persistence of snow cover.
The available literature in English on the hematology, blood
chemistry, protein polymorphisms and endocrinology of the Cervidae was
compiled and tabulated. A review of blood chemistry of moose and other
Cervidae was published. Since the recurrent stated purpose of these
clinically oriented studies has been evaluation of herd and range condi-
tions, the possible analyses of data in terms of diagnostic patterns are
presented. The available data demonstrate that before such management-
related interpretations can be made, one must consider the effects of
age, sex, season, reproductive condition, and methods of handling.
Given these boundary conditions, experimental studies have demonstrated
effects of nutritional level and disease on blood values in species of
Odocoileus and Alces alces. Serum urea nitrogen levels (BUN) appear to
be directly related to nutritional status in several cervid species.
Measurements of serum albumin and hemoglobin are not sensitive indicators
although they decline with severe deficiencies. A decline in serum
thyroxine with nutritional restriction has been reported in Odocoileus
virginianus. This hormone declines over winter in Alaskan moose. The
metabolic shifts occurring in malnutrition and starvation as reflected
in plasma nonesterified fatty acids, amino acids, ketones, and short-
chain fatty acids await study in the cervids. They offer the possibility
of more sensitive indicators of metabolic status. Protein polymorphisms
have been described for transferrin in Rangifer and Odocoileus, and for
hemoglobin in Odocoileus virginianus. Studies of Alces have shown no
polymorphisms. Fibrinopeptide sequences have suggested that Alces,
Rangifer, and Odocoileus are closely related genera.
i
A selected bibliography on blood chemistry of moose and other
Cervidae is presented.
Weights of 70 moose of known age are presented, as are chemical and
proximate analyses of important forage plants and of rumina from moose
on seral birch and upland willow range. No significant differences were
demonstrated in these analyses of rumina from the two ranges. Analyses
of forage species indicate the need for variety in moose diets. Corre-
lations between protein and ether extract in rumina and fecal pellets
from the same moose were highly significant.
A maximum snow depth of 67 em, with depths exceeding 50 em for
three consecutive months in most habitats, was found to be sufficient
to cause high winter mortality in the Moose Research Center.
A summary of composition, production, rate of growth and shape of
burn, diversity and "edge effect" in the Kenai (1947) burn is presented,
as is a summary of the importance of nonbrowse foods to moose on the
Kenai Peninsula.
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Sunnnary .•
Background.
Objectives. .
Procedures.
Findings ..
Productivity and Mortality
CONTENTS
Blood Chemistry as Indicator of Nutritional Status .
Blood Chemistry of Moose and Other Cervidae: A Selected
Bibliography. . . . . . . • . . . . . . . . • .
Stress Evaluation. . . . . . . . . . . . . . . .
Hair Analyses as Indicators of Nutritional Status.
Weights and Measurements . . • . . . .
Nutritional Analyses of Plants, Rumina and Fecal Pellets
Snow Monitoring. . . . . . . . . .
Browse Production and Utilization.
Food Habits of Tame Moose.
Reconnnendations • •
Literature Cited. .
BACKGROUND
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1
1
2
10
10
29
62
77
77
77
77
77
85
95
.101
.101
The Kenai Moose Research Center was established in 1966-1969 as a
controlled environment within which to study moose-habitat interrelations.
Research at the Center has stressed nutritional aspects of moose growth
and productivity and quality, quantity and availability of plants eaten.
LeResche (1970) and LeResche and Davis (1971) discussed the management
problems considered and the approaches taken at the Hoose Research Center.
OBJECTIVES
To measure natality, mortality and general condition of moose within
four 2.6 km2 (one square mile) enclosures.
To establish baselines by season, age and sex for the following
serological and hematological parameters in moose and to evaluate their
usefulness as indicators of nutritional status in moose:
A. calcium
B. inorganic phosphorus
C. glucose
n. urea nitrogen (BUN)
E. uric acid
F. cholesterol
G. total protein
H. albumin
I. albumin/globulin ratio
J. alpha-1, alpha-2, beta and gamma-globulins
K. bilirubin
L. alkaline phosphatase
1
M. lactic dehydrogenase (LDH)
N. glutamic oxalacetic transaminase (SGOT)
0. hemoglobin
P. hematocrit
Q. white blood cells
R. differential cell count (including segmenters, lymphocytes,
eosinophile, monocytes, basophile)
To estimate browse production and utilization and quantitatively
and qualitatively estimate consumption of all plant material by moose.
To learn changes in rumen protozoa levels in moose on various
winter diets.
To learn nutritional values and digestibilities of the more common
moose forage species of plants.
PROCEDURES
General Description of the Moose Research Center Facility
The Kenai Moose Research Center comprises four 2.6 km 2 (one square
mile) enclosures located in the area of the 1947 burn near Kenai, Alaska.
These enclosures contain representative vegetation of both burned
(regenerative: predominately birch Betula papyrifera and white spruce
Picea glauca) and remnant: (mixed birch-spruce-aspen Populus tremulm:des
stands). Marshland typical of summer range is included as are well-
drained hillocks supporting winter browse species.
The entire area has been type-mapped into 11 vegetation types, and
soil profiles of representative types have been completed. Five five-
acre exclosures are present, at least one within each enclosure.
Twenty-one fenceline traps have been consturcted; 12 within pens
and nine on the outside of the fenceline. Fig. 1 is a generalized map
of the facility showing traps, exclosures, etc.
The log headquarters building sleeps eight, and is accessible by
road during dry seasons. Two-mile-long Coyote Lake provides access by
float or ski plane. The Center may be reached by light plane from
Anchorage in one-half hour.
Populations of moose within the enclosures as of February 1, 19 70,
five months after enclosing Pens 3 and 4, were:
Pen Cows Calves Bulls Total
1 5 0 2 7
2 9 1 2 12
3 7 4 1 12
4 11 5 2 18
32 10 7 49
2
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I 10-4
__r::::]_ Kenai Moose Research Center ~ L.J
I 1-4
b 1-2 1 Headquarters r I I _r/0120-3 PI0-1 1130-1
1-[t ~ere Pen
L......J
I 10-2 [ 3 -I
0
L--1 2-2 [ I 10-3
3 2
_o_Traps
I 0 Exclosures
2-1 3-2
II ,....,
40-4 [ ~ 0 40-3 I One Mile
0 1 4
I b 1 2 ~ 4-2 4-1 ,----, r::::J
Kilometers '-----1
I 40-2
Fig. 1. Map of Moose Research Center.
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I 3
Pens 1 and 2 will be left unmolested in terms of moose numbers,
allowing the populations to increase, decrease, or remain constant as
they will. Pen 3 will be retained at its present population level and
sex structure, as representative of extra·-pen populations in this area.
Pen 4 will be used as an experimental pen, with abnormally high and low
densities of moose present.
Table 1 is a history of major events in construction of the facility
and provides reference as to timing of events leading to the current
description.
Productivity and Mortality
Mortality and natality within pens are measured by daily ground
observations, periodic aerial observations, trapping and use of radio-
tracking devices. General condition is estimated for trapped animals
by methods described below.
Blood
Blood values are determined from serum and whole-blood samples
obtained from trapped and hunter-killed moose and animals immobilized
for marking outside of traps (Job 1.4). Table 2 lists sources of blood
material.
Blood is obtained from live immobilized animals in sterile evacuated
containers by jugular venepuncture. Four or five cc of whole blood are
preserved with EDTH and a thin smear is made; serum is secured by
centrifugation of cooled and clotted blood. Serum is separated into:
1) a NaF tube (1.5-2 cc) for glucose determination; and 2) a 4-5 cc
untreated sample for analyses of other parameters.
Analyses are performed by Alaska Medical Laboratories (Anchorage)
using a Technicon Autoanalyzer St~-12, standard hematological techniques
and electrophoresis.
A review paper was prepared by LeResche, U. S. Seal (Veterans
Hospital, Minneapolis), P. D. Karns (Minnesota Department of Natural
Resources) and Franzmann for the International Symposium on Moose
Ecology, Quebec. It is titled "A review of blood chemistry of moose
and other Cervidae, with emphasis on nutritional assessment", and will
appear in Le Naturaliste Canadien in fall 1973, and in "AZ.ces", the
symposium proceedings. A bibliography of blood studies of the Cervidae
was also prepared.
Stress Evaluation
The influence of stress, particularly that which relates to excit-
ability, on an animal during sampling procedures can influence the
physiologic values obtained (Franzmann, 1972; LeResche et al., 1973b).
Some values are influenced more than others and some may not be affected
at all. The basic objective of this phase of research will be to corre-
late the physiologic values with excitability.
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Table 1. Chronology of establishing the Kenai Moose Research Center.
June 1966: Construction begun.
September-October 1967:
January 196 7:
January 1968:
April 1968:
1968:
April 1969:
June 1969-January 1970:
June-July 1969
August 1969:
October 1969:
January-February 1970:
April 1970:
May 1970:
August 1970:
November 19 70:
March 1971:
May-June 1971:
June 1971:
Browse production estimated in Pens 1 & 2.
Successional plots established and read in
Pens 1 & 2.
Pens 1 & 2 enclosed.
Moose in Pens 1 & 2 collared.
Browse utilization estimated in Pens 1 & 2.
Yearling bull introduced into Pen 1.
Browse utilization estimated in Pens 1 & 2.
Eleven traps constructed in all pens.
Blood collections begun.
Successional plots established and read in
Pens 3 & 4.
Pens 3 & 4 enclosed.
Two male calves introduced into 10-acre
pen in Pen 2.
Numbers of moose in Pens 3 & 4 determined.
Replicate count experiments conducted.
Browse utilization estimated in all four
pens. Plots cleaned of pellets.
Female calf introduced into 10-acre pen in
Pen 2.
Twenty traps complete: 11 inside pens;
nine outside.
Two male yearlings released from 10-acre
pen into Pen 2.
Replicate count experiments conducted.
Pellet-plots counted and cleared in Pen 1.
Browse utilization estimated in Pen 1.
Numbers of moose in pens redetermined and
calves counted.
5
Table 1. (cont'd.)
September-December 1971:
June 1972:
Pen 4 stocked with 44 moose. including
orphaned calves.
Numbers of moose in pens redetermined and
calves counted.
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Table 2. Sources of moose blood for analysis:
NUMBER OF SPECIMENS
Source Serum
Trapping at Moose Research Center:
Pen 1 20
Pen 2 25
Pen 3 15
Pen 4 22
Outside pens 40
Total Moose Research Center 122
Hunts:
1969-70
GMU 15C 32
15B 13
14A 39
14B 14
1970-71
GMU 7 7
15A 28
Total Hunts 133
Tagging:
Bottenintnin Lake (1970) 38
Moose River Flats (1970) 61
Skilak-Tustumena Beach
(1971) 3
Moose River Flats (1971) 60
Total Tagging 162
TOTALS 417
7
June 1969-May 1971.
Whole
Blood Slides
15
16
13
15
26
85
26
6
26
6
2
0
66
23
0
0
0
23
174
5
10
7
12
9
43
49
21
42
9
0
0
121
6
0
0
0
6
170
To determine the excitability stress, heart rate, respiratory rate,
and rectal temperature were recorded from each moose sampled. Each
individual was subjectively judged for excitability prior to and during
handling. The moose is given a score of 1 (not excited), 2 (slightly
excited), 3 (moderately excited), 4 (excited), 5 (highly excited), based
upon observations. The ambient temperature is also recorded.
The physiologic values obtained will be tested to determine the
influence of excitability, if any, upon them. A blood value that can
be used as an excitability state monitor would be very useful, and
would alleviate subjective evaluations.
The utilization of cellular enzyme tests will be studied to deter-
mine their relationship to excitability stress. Cellular enzymes are
released when cells are destroyed during excitability, general stress,
pathologic conditions, and normal cellular aging and destruction (Coles,
1967). An enzyme closely related to excitability stress would also
prove useful in monitoring the relative excitability of an animal.
Hair
Hair analyses to study malnutrition and deficiencies in humans have
been based upon hair shaft diameter and atrophy of hair bulb (Sims, 1968),
hair root protein analysis (Crounse et al., 1970a,b), and trace element
content (Strain et al., 1972). The possibility of utilizing one or more
of these approaches by sampling moose hair was investigated.
For trace element analysis, 30 hair samples taken from moose in May
and June were sent to Cleveland Memorial Hospital for analysis. Samples
will be sent regularly throughout the year to determine if there are
seasonal patterns and, if so, how they relate to blood analyses.
Root-hair protein analysis techniques are under investigation, and
will be incorporated into the sampling procedure if feasible.
Weights and Measurements
Weights and measurements were obtained from trapped immobilized
animals.
Chemical Analysis of Plants and Fecal Pellets
Birch, willow (S'al-ix sp.), lowbush cranberry ( Vacciwium vi tis idaea),
sedges and aspen (hark) were collected for nutritional analysis accord-
ing to plant height, diameter and use-form class. All specimens were
oven-dried at 70° C for 14 days. The following analyses were done by
WARF, Inc.; Madison, Wisconsin:
Elemental:
Protein:
DDM:
JAOAC, 51, 1003 (1968)
AOAC, 16 (1970) 11th ed.
Tilley & Terry, J. Br. Grassland Society, (2) 104
(1963), (Bovine Flora)
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Ash: AOAC, 123 (1970) 11th ed.
Ether extract: AOAC, 128 (1970) 11th ed.
Fiber: AOAC, 129 (1970) 11th ed.
Snow Monitoring
Eight snow plots were established in Pens 1 and 2. One plot was
located in each of the following habitat types: dense hardwoods, thin
hardwoods, sedge meadow, spruce regrowth, birch-spruce regrowth (thin),
birch-spruce regrowth (dense), spruce-Ledum, mature spruce. At approxi-
mately weekly intervals a trench was dug in each plot and thickness and
general structure (e.g.: crystals, powder, ice) of each snow layer were
recorded. The visibility of lowbush cranberry above snow was also
recorded at this time.
Browse Production and Utilization
Browse production and utilization and plant succession are estimated
using methods previously described in detail (Bishop, 1969). A canopy-
cover method after Daubenmire (1959), employing exclosures, is used for
successional measurements and a twig-count method with clipping is used
for production and utilization estimates. During this reporting period
use was estimated in all old plots in Pen 1, the 2.5 ha (10 acre) hold-
ing pen. A paper titled "Distribution and habitats of moose in Alaska"
was prepared for the International Symposium on Moose Ecology by LeResche,
R. H. Bishop and J. W. Coady. It will be published in autumn 1973.
Food Habits of Tame Moose
Tame moose were obtained as calves in October 1969 (two males) and
May 1970 (one female). All were born in spring 1969. They were raised
on natural moose range with calf starter feed (Alaska Mill Feed Co.,
Anchorage) fed supplementally until July 1970. The three were confined
together in a 2.5 ha pen (containing a 1 ha lake) through November 1970.
Summer observations were made within this pen. In November the males
were released into Pen 2 with 14 other moose. "Normal range" winter
observations were made of these animals. "Depleted range" winter
observations were made of the female within the 2.5 ha pen, which had
supported the equivalent of 82 moose per square km during the previous
winter and was supporting 41 moose equivalents per square km during the
winter of observations.
Tame moose were located by radio-telemetry and food intake was
recorded by an observer standing 1-2 m from the moose and recording
species and size of each bite eaten by pencil on an IBM optica] page
reader sheet or by speaking into a tape recorder. Hours of observation
were distributed throughout daylight hours. Bite size in summer was
recorded by estimating number of leaves ingested. Bites were classified
in four categories: less than 5 leaves, 6 to 10 leaves, 11-20 leaves and
more than 20 leaves. Mean number of leaves per bite in each category
were taken as 2.5, 7.5, 15 and 25, respectively. Mean weight of each
9
bite by category was then calculated as 2.5, 7.5, 15 and 25 times the
mean weight of 500 randomly selected leaves from 100 separate plants.
Bite size in winter was recorded by estimating length of stem ingested.
Bites were classified in four categories: less than 3 inches (7. 6 em),
3 to 6 inches (15. 2 em), 6 to 12 inches (30. 5 em) and more than 12
inches. Hean length of stem in each bite category was taken to be 3.8
em, 11.4 em, 22.8 em, and 38.1 em, respectively. Thirty terminal stems
of each of these lengths were then randomly clipped from six plants of
each species considered. Mean weight per bite was calculated using the
mean weight of the appropriate 30 stems. Biomass of food ingested per
day was estimated by applying these weight/bite estimates to estimated
bites/day.
A publication by LeResche and Davis, "Importance of nonbrowse foods
to moose on the Kenai Peninsula, Alaska" was prepared for the Journal
of Wildlife Management (37:2, in press).
Feeding Craters
Feeding crater concentrations were searched for and when found the
habitat type was designated and the area of the concentration measured.
Each crater was then measured in length, width, and depth. Species of
plants eaten and present in each crater and their relative abundance
were recorded. Also, the location of the craters within the plot was
noted to determine if moose used keying characteristics before digging
a crater. Snow profiles were taken at each crater concentration area.
FINDINGS
Productivity and Mortality Within Pens
Table 3 presents complete tagging, breeding, and mortality data for
all moose within the enclosures. Table 4 summarizes these data in terms
of numbers calving and dying. Table 5 calculates natality, yearling
recruitment and change in population size for penned and unpenned popu-
lations, as summarized in Table 6.
On 1 July, 1972, populations within the 4 pens totaled 52 moose (13
per square mile). Components of the populations are tabulated in Table
7, and trends since 1968 are shown in Table 8 and Fig. 2.
These data reflect more stable population levels than might be
predicted from the accompanying severe annual fluctuations in the
"indicator" parameters of calf production, calf mortality and yearljng
recruitment. From 1968 through 1972, known June calf production in the
four MRC pens has varied between 15 and 72 calves per 100 adult females
(11-45% of adult maose) and yearling recruitment has vari.ed from none to
41~45 yearlings per 100 adult females (0-33% of adult moose). Neverthe-
less, absolute numbers have changed only a mean of two pGrcent in the
five years since 196 8. Data collected in the area surrounding the HRC
(Table 5) by various workers follow a similar pattern, with estimated
calf production varying from 21-48 per 100 cows and yearling recruitment
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-------------------Table 3. Histories of individual moose in Kenai Moose Research Center enclosures, July 1971 through June
1972.
PEN 1
Moose II Sex Event Date Age Circumstances
3 F Weight 790 lbs. 31 August 1971 9+ years Trapped
With no calf 13 March 1972 9+ years Observed
Weight 650 lbs. 24 June 1972 10 years Trapped
with no calf
371 F Weight 260 lbs. 31 August 19 71 Calf Roped in trap
Last seen 4 November 1971 Calf Observed
6 F Weight 790 lbs. 24 August 1971 14+ years Trapped
Last seen alive 20 February 1972 14+ years Observed
Found dead 10 May 1972 15 years Helicopter
I-'
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10 F Weight 635 lbs. 11 August 1971 4+ years Trapped
With no calf 6 October 1971 4+ years Observed
Last observed 22 May 1972 5 years Observed
R70-8 F With no calf 23 March 1972 3+ years Helicopter
Last handled 8 June 1972 4 years Trapped
40 F With no calf 23 March 1972 3+ years Helicopter
Last handled 27 June 1972 4 years Trapped
670 F Retagged 8 May 1972 2 years Helicopter
Last observed 25 June 1972 2 years Supercub
61 F Tagged (2 calves 2 September 1971 9+ years Trapped
outside trap)
With no calves 23 March 1972 9+ years Helicopter
Last observed 10 May 1972 10 years Helicopter
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Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, July 1971
through June 1972.
PEN 1
Moose It Sex Event Date Age Circumstances
69 F Tagged 8 May 1972 3 years Helicopter
Last handled 6 Jnne 1972 3 years Trapped
35 M Last handled 2 7 Jnne 1972 4 years Trapped
43 M Weight 1040 lbs. 4 October 1971 4+ years Trapped
Last handled 22 June 1972 5 years Trapped
53 H Tagged 15 July 1971 1+ years Trapped
Last Handled 28 Jnne 1972 2 years Trapped
55 M Died from drug 10 August 1971 1 years Trapped
Weight 500 lbs.
58 M Tagged 31 August 1971 1+ years Trapped
Last observed 20 June 1972 2 years Helicopter
64 ~1 Tagged, weight 620 lbs. 2 April 1972 2+ years Trapped
Last handled 28 June 1972 3 years Trapped
uc F
370UC M After Jnne 3, 1971 these individuals were not again
identified and could be dead or later tagged
1070UC M
4070UC ?
-------------------
-------------------
,....
w
Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, July 1971
through June 1972.
PEN 2
Hoose # Sex Event Date Age Circumstances
1 F Hith one calf 12 August 19 71 8+ years Trapped
With no calf 3 February 1972 8+ years Observed
Weight 700 lbs. 22 June 1972 9 years Trapped
Last trapped 28 June 1972 9 years Trapped
2 (R69-3) F Hith no calf 23 February 1972 7+ years Observed
Last seen (presumed 23 Harch 19 72 7+ years Helicopter
dead)
7(R70-7) F With no calf 27 January 1972 8+ years Observed
With one calf 20 June 1972 9 years Helicopter
Last seen with calf 29 June 19 72 9 years Observed
9 F With no calves 1 February 1972 9+ years Observed
Found dead 8 May 1972 10 years Helicopter
52 F Hith no calf 14 December 1971 4 years Observed
With one calf 20 June 1972 5 years Helicopter
Last handled 29 June 1972 5 years Trapped
R70-2 F Died from drug 15 July 1971 4 years Trapped
R70-4 F Hith 2 calves 5 October 19 71 4+ years Observed
With no calves 17 December 1971 4+ years Observed
Last seen 28 June 1972 5 years Observed
Raquel F IH th one calf 2 July 1971 2 years Observed
Fistula installed 23 August 1971 2 years Trapped
Fistula lost 2 September 1971 2 years Trapped
Height 775 lbs. 24 June 1972 3 years Trapped
!--'
.1:"-
Table 3. (cont't.) Histories of individual moose in Kenai l'foose Research Center enclosures, July 1971
through June 1972.
PEN 2
Moose If Sex Event Date Age Circumstances
62 F Tagged 1 December 1971 19+ years Trapped
Paralyzed from darting, 2 7 January 1972 19+ yesrs Trapped
killed, weight 850 lbs.
63 F Tagged 22 Narch 1972 4+ years Trapped
Found dead 8 May 1972 5 years Helicopter
66 F Died, acute hemorragic 2 April 1972 9+ years Trapped
enteritis, weight
720 lbs,
68 F Tagged 8 May 1972 9 years Helicopter
Found dead, died while 20 June 19 72 9 years Observed
calving
70 F Died, broke neck on 8 May 1972 5 years Helicopter
fence during heli-
copter tagging
76 F Tagged, limps 10 May 1972 9 years Helicopter
With one calf 20 June 19 72 9 years Helicopter
77 F Tagged 10 May 19 72 6 years Helicopter
Weight 605 lbs. 23 June 1972 6 years Trapped
uc F \Hth one calf 20 June 1972 ? Helicopter
36 H Fighting with /f43H 17 October 1971 4+ years Observed
through fence
Last seen 29 June 1972 5 years Observed
-------------------
-------------------Table 3. (cont 1 d.) Histories of individual moose in Kenai Hoose Research Center enclosures, July 1971
through June 1972.
PEN 2
}foose if Sex Event Date Age Ci rcums tan ces
45 M Last seen 28 June 1972 4 years Observed
Richard M Killed being lead by 10 August 1971 2 years Free-ranging
antlers, weight
775 lbs.
Halter H Weight 640 lbs. 23 July 1971 2 years Free-ranging
Died after fistula 30 July 1971 2 years Trapped
surgery
f-' ~~ralter, Jr. M Height 470 lbs. 28 April 1972 1-years Trapped \.J1
( Raquel 1 s cal f) Weight 515 lbs. 24 June 1972 1 years Trapped
54 H Height 355 lbs. 27 July 1971 1 years Trapped
Last seen (presumed 23 February 1972 1+ years Observed
dead)
65 H Tagged 2 April 1972 1+ years Trapped
Last seen 28 June 1972 2 years Observed
73 M Tagged 10 May 1972 3 years Helicopter
Weight 525 lbs. 5 June 1972 3 years Trapped
78 M Tagged 30 June 1972 4 years Trapped
uc M Last seen 30 June 1972 ? Observed
1-'
0\
Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, July 1971
through June 1972.
Noose II Sex Event
R70-2-70UC ?
UC70UC ?
5270UC ?
20 F "Hith no calf, limping
badly
Found dead
27 F Hith no calf
28 F Killed, weight 760 lbs.
38 F Hith no calf
39 F With no calf
Height 535 lbs.
uc F Observed
60 M Height 850 lbs.
Found dead 150 meters
from trap
PEN 2
Date Age Circumstances
After June 3, 1971 these individuals were not again
identified and could be dead, later tagged, or still
uncollared and alive in Pen 2.
Pen 3
12 October 1971 11 years Observed
20 June 1972 12 years Observed from helicopter
20 June 1972 6 years Observed from helicopter
12 August 1971 9 years Dart entered vagina
20 June 1972 18 years Observed from helicopter
20 June 1972 7 years Observed from helicopter
23 June 1972 7 years Trapped
10 May 1972 ? Observed from helicopter
31 August 1971 4 years Trapped
1 September 1971 4 years Observed
-------------------
-------------------Table 3. (con t, d.) Histories of individual moose in Kenai Noose Research Center enclosures, July 1971
through June 1972.
PEN 3
Hoose if Sex Event Date Age Circumstances
2870 F Not bred (rectal 9 Hay 1972 2 years Irrunobilized via helicopter
palpation)
Hith no calf 20 June 1972 2 years Observed from helicopter
2771 F Weight 155 lbs. 27 July 1971 Calf Trapped
Found dead 16 February 1972 Calf Observed
67 F Tagged, could be 2069, 8 April 1972 1+ years Trapped
3970, or UC female
Found dead 9 May 1972 2 years Observed from helicopter
f-72 F Tagged~ could be 2069, 9 l1ay 1972 2 years Immobilized via helicopter -...,J
39 70 ~ or UC female
With no calf 20 June 19 72 2 years Observed from helicopter
75 F Tagged~ could be 2069, 10 }fay 1972 3 years Immobilized via helicopter
3970 ~ or UC female
With no calf 20 June 1972 3 years Observed from helicopter
uc M Observed 20 June 1972 ? Observed from helicopter
PEN 4
22 F Hith no calf 20 June 1972 7 years Observed from helicopter
24 F Found dead 10 May 1972 10 years Observed from helicopter
Table 3. {cant' d.) Histories of individual moose in Kenai Moose Research Center enclosures, July 1971
through June 1972.
PEN 4
~loose if Sex Event Date Age Circumstances
31 F Killed, weight 640 lbs. 25 August 1971 7 years Trapped, did not recover
(R-70-1) from Anactine
34 F Fonnd dead 23 Har 1972 14 years Observed from air
36 F \.Jith no calf 20 June 1972 9 years Observed from helicopter
37 F With no calf 20 June 1972 3 years Observed from helicopter
....
00 R-70-3 F Height 680 lbs, with 11 August 1971 4 years Trapped
calf
With no calf 20 June 1972 5 years Observed from helicopter
A60 F Height 750 lbs. with 10 August 1971 13+ years Trapped
calf
Found dead 9 Hay 1972 14+ years Observed from helicopter
21 H Found dead 26 February 1972 3 years Observed
44 H Found dead 9 Nay 1972 3+ years Observed from helicopter
7 H Trapped 1 September 1971 2 years Trapped
Observed 20 June 1972 3 years Observed from helicopter
-------------------
-------------------
,...,
\.0
Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, July 1971
through June 1972.
PEN 4
Hoose f! Sex Event Date Age Circumstances
uc
2470UC
After 3 June 1971, those individuals were not again
36 70UC identified and could be dead, later tagged, or still
uncollared and alive in pen 4.
R70-l-70UC
R70-3-70UC
4170 M Survived to yearling 27 July 1971 1 years Observed
Last seen 4 February 1972 1 years Observed
57 F Tagged 25 August 1971 1 years Trapped
With no calf 16 June 1972 2 years Observed
59 M Tagged 1 September 1971 l years Trapped
Observed 20 June 1972 2 years Observed from helicopter
118 F Put into Pen 4 16 November 1971 l+ years Trapped
~.Jith no calf 20 June 1972 2 years Observed from helicopter
121 M Put into Pen 4 2 3 November 19 71 1+ years Trapped
Found dead 2 3 March 19 72 1+ years Observed from helicopter
123 F Put in Pen 4, 1 December 1971 ? Trapped
weight 765 lbs.
With l calf 14 June 1972 ? Observed
N
0
Table 3. (cont'd.) Histories of individual moose in Ke;:J.ai Hoose Research Center enclosures, July 1971
through June 1972.
PEN 4
Noose tl Sex Event Date Age Circumstances
126 F Put into Pen 4, 15 December 19 71 17+ years Trapped
weight 790 lbs.
Found dead 8 Harch 1972 17+ years Observed
128 F Broke into Pen 4, 26 January 1972 ? Trapped
With no calf 20 June 1972 ? Observed from helicopter
134 l'l Put into Pen 4 22 March 1972 Calf Trapped ,
weight 360 lbs.
Found dead 23 Harch 1972 Calf Observed
R-72-1 F Tagged, put into pen 4 20 April 1972 10 years Trapped
Found dead 24 !'lay 19 72 10 years Observed
71 F Tagged 9 ~1ay 19 72 3 years Immobilized via helicopter
With no calf 20 June 1972 3 years Observed from helicopter
74 F Tagged 10 Hay 1972 2 years Inunobilized via helicopter
Found dead 3 J U.l"le 1972 2 years Observed
70371 F Tagged, weight 210 lbs. 11 August 1971 Calf Trapped
Found dead 10 May 1972 Calf Observed from helicopter
56 }l Tagged, v.'eight 545 lbs. 11 August 1971 1 years Trapped
Found dead 25 Hay 1972 2 years Observed
96 71 F Tagged and radio-21 September 1971 Calf Trapped
collared, '"'eight 350 lbs.
Found dead 10 }lay 1972 Calf Observed from helicopter
-------------------
-------------------Table 3. (cant' d.) Histories of individual moose in Kenai Hoose Research Center enclosures, July 1971
through June 1972.
PEN 4
Noose II Sex Event Date Age Circumstances
9771 F Tagged and radio-22 September 1971 Calf Trapped
collared, weight 320
lbs., put into Pen 4
Found dead 14 January 1972 Calf Observed
9871 F Tagged and radio-29 September 1971 Calf Trapped
collared, weight 400
1bs., put into Pen 4
Found dead 15 January 1972 Calf Observed
10671 M Tagged and radio-12 October 1971 Calf Trapped
N collared, weight 350 1-'
lbs., put into Pen 4
Found dead 27 December 1971 Calf Observed
10771 F Tagged and radio-12 October 1971 Calf Trapped
collared, weight 410
1bs., put into Pen 4
Found dead 14 January 1972 Calf Observed
10871 F Tagged and radio-13 October 1971 Calf Trapped
collared, weight 385
lbs., put into pen 4
Found dead 10 December 1972 Calf Observed
10971 F Tagged and radio-20 October 1971 Calf Trapped
collared, \..reight 395
lbs., put into Pen 4
Found dead 15 January 1972 Calf Observed
Table 3. (cant' d.) Histories of individual moose in Kenai }bose Research Center enclosures, July 1971
through June 1972.
PEN 4
Noose /1 Sex Event Date Age Circumstances
11071 H Tagged and radio-21 October 1971 Calf Trapped
collared~ weight 385
lbs., put into Pen 4
Found dead 13 January 1972 Calf Trapped
11471 H Tagged, weight 370 1bs. 2 November 1971 Calf Trapped
Found dead 6 June 1972 Calf Observed
N
N 11571 F Tagged, weight 440 lbs. 3 November 1971 Calf Trapped
put into Pen 4
Found dead 9 May 1972 Calf Observed from helicopter
12071 H Tagged, weight 465 lbs. 23 November 1971 Calf Trapped
Found dead 23 December 1971 Calf Observed
6071 M Tagged, weight 195 lbs. 10 August 1971 Calf Trapped
Found dead 16 June 1972 Calf Observed
uc small F Observed 20 June 1972 ? Observed from helicopter
uc large F Observed 20 June 1972 ? Observed from helicopter
-------------------
-------------------Table 4. Moose natality, mortality, and recruitment in four one-square-mile enclosures.
Net gain (+) or
Adults Died Loss (-) of Adults
(including) (dis counting
Adult Calves Yearlings long experimental
F (M) Calves lost recruited yearlings) manipulation)
PEN 1
June 1972 7 (5) 0 0
June 1971-June 1972 5 2
PEN 2
N 4 0 w June 1972 8 (6)
June 1971-June 1972 8 9
PEN 3
Jtme 1972 7 (1) 0 1
1971-June 1972 2 1 4 3 (1 break in)
June
PEN 4
June 1972 11 (2) 1 0
June 19 71-June 1972 6 11
Table 5. June calf crops and yearling recruitment in Moose Research Center enclosures.
Population (Adult)
% Gain/loss
Calf Crop Yearling Recruitment (excluding (No. 1+
Year Calves/100 F (No. F) Yrlgs/100 F manipulations) years old)
PEN 1
1968 83 (6) 17
1969 0 (4) (no so +17% (6)
breeding bull)
1970 100 (4) No recruitment No change (7)
1971 83 (6) 80 +52% (12)
1972 0 (7) No recruitment No change (12)
X. (not incl. 69-70) 67 37 +17%
N PEN 2 -!:'-
1968 12.5 (8) 25-38
1969 50 (8) 9 +11% (11)
1970 30 (10) No re c rui tmen t -8% (12)
1971 100 (8) 25-38 No change (13)
1972 50 (8) No recruitment No change (14)
-49 12-17 No change X
PEN 3
1969 (August) so (8) 0 No data
1970 29 (7) 38 No change (10)
1971 33 (6) 38 No change (10)
1972 0 (7) No recruitment -20% ( 8)
-28 18 -5% X
-------------------
-------------------
N
V1
Table 5. (cont'd.) June calf crops and yearling recruitment in Moose Research Center enclosures.
Population (Adult)
% Gain/loss
Calf Crop Yearling Recruitment (excluding (No. 1+
Year Calves/100 F (No. F) Yrlgs/100 F mani pu1ations) years old)
PEN 4
1969 (August) 83 (12) 8 No data
1970 45+ (9) 44-56 -21% (14)
1971 75 (8) so +7% (15)
1972 9 (11) No recruitment -7% (14)
X 53+ 26-29 -7%
ALL PENS
1968 43 (14) 21-29
1969 59 (27) 11 +13% (15)
1970 43 (30) 25-29 -10% (39)
1971 72 (29) 41-45 +11% (44)
1972 15 (34) No re c rui tmen t -7% (48) -42 (134) 19-20 +2% (146) X
Unit lSA (Aerial counts by Richey ( unpublished) and LeRoux (unpublished) and LeResche)
(No. F)
1968 47 (1520) 18 (est.) (1520)
1969 48 (438) 7 (est.) (438)
1970 ca: 30 (ca: 500) 23 (496)
1971 21 (657) 14 (166)
1972 28 (106) 5 (248) -39 (ca: 3221) 15 (2868) -3%** X
**Kenai National Moose Range stratified random mile-square quadrant counts (unpublished) indicated
populations north of the Kasilof River of 6700 ± 1410 in winter 1967, 7900 ± 1460 in winter 1971 and
5700 ± 1348 in winter 1972-73. This represents a mean annual decrease of 3 percent for the six years,
disregarding the variations in weather and observers and ignoring the confidence intervals.
Table 6.
Year
1969
1970
1971
1972
Table 7.
Pen 1
Pen 2
Pen 3
Pen 4
All Pens
Pen 1
Pen 2
Pen 3
Pen 4
All pens
Calf production, calf mortality and yearling recruitment in
four 2.6km2 (1 mi.z) enclosures.
Calves Yearlings
Produced Recruited June-June June
(% of Adults) (% of Adults) Calf loss (%) Population
45 10 62
35 20 56 57-58
55 33 6 75
11 0 100 52
Populations within Moose Research Center enclosures as of
20 June 19 72.
Females with
No Calves 1 Calf 2 Calves Yearlings Males Total
7 0 0 0 5 12
4 4 0 0 6 18
7 0 0 0 1 8
10 1 0 0 2 14
28 5 0 0 14 52
SUMMARY
Females Yearlings Calves Males Total --
7 0 0 5 12
8 0 4 6 18
7 0 0 1 8
11 0 1 2 14
33 0 5 14 52
26
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I Table 8. Summary of moose populations within Moose Research Center enclosures,
1968 through June 1972.
I Total
I
Females Yearlings Calves Males Moose
Pen 1 January 1968 6 0 1 0 7
I June 1968 6 1 5 1 13
June 1969 4 2 0 1 7
February 1970 4 2 0 1 7
I June 1970 5 0 4 2 11
June 1971 6 4 5 2 17
June 1972 7 0 0 5 12
I Pen 2 January 1968 8 0 3 1 12
June 1968 8 4 1 1 14
June 1969 11 1 4 1 17
I February 1970 9 0 1 2 12
June 1970 11 0 3 2 16
June 1971 9 3 9 4 25
I June 1972 8 0 4 6 18
Pen 3 August 1969 8 0 4 1 13
I
February 1970 7 0 4 1 12
June 1970 6 3 2 1 12
June 1971 6 2 2 2 12
June 1972 7 0 0 1 8
I Pen 4 August 1969 12 1 10 2 25
February 1970 11 0 5 2 18
I June 1970 9 4-5 4 1 18-19
June 1971 8 4 6 3 21
December 1971 44
June 1972 11 0 1 2 14
I All Pens June-August 1969 35 4 18 5 62
February 1970 31 2 10 6 49
I June 1970 31 7-8 13 6 57-58
June 1971 29 13 22 11 75
June 1972 33 0 5 14 52
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Q)
(/)
80r----------------------------------------,
70
60
g 50
~
.......
0
0
J\
,_ 40 I \
I \
I \
I \
Q)
_Q
E
~ 30 Pen~' ~~/ \
.... , ~-d \
Pen 2 -• 'o---0;::.::> ,6._
--A-........._ ........... ---b
20
.~ P--•--">.~1! ---• ---A
I 0 };"""' Pen~----6.---6. ........ -Pen3----...:•
a I I
Jan 68 Jun 68 Jan 69 Feb 70 Jun 70 Jun 71 Jun72
Fig. 2. Populations within Moose Research Center enclosures:
1968-1972.
28
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varying from 5-23 per 100 cows; but total population changing only three
percent per year since 1967, for a total decline of approximately 15
percent.
Both the }ffiC pens and the surrounding area demonstrate that long-
term population levels may be more stable than annual fluctuations
suggest, and that the effects of an especially "good'' year or an espe-
cially "bad" year may be quickly erased by conditions in succeeding
years. This apparent lack of steady trends likely reflects the relative
short-term instability of a population of near maximum size.
Mortalities recorded within the pens during the reporting period
are listed in Table 9. A disproportionate number of calves are repre-
sented, associated with the deliberate over-stocking of Pen 4 for the
orphan calf survival study (Job 1.2R) and to the severity of the winter
(LeResche and Davis, 1973).
Blood Chemistry as Indicator of Nutritional Status
Specimens of blood analyzed through June 1972 are listed in Table 2.
Preliminary analyses of these data are summarized in the following
publication:
29
Table 9. Mortalities within pens, July 1971-June 1972.
Moose if Sex Age Pen Month -Year Cause
60 M 4 3 September 1971 Found dead 150 meters from trap
he was immobilized in on 8/31/71
20 F 12 3 October 1971-June 1972 Unknown, carcass found
28 F 9 3 August 1971 Killed with immobilizing dart
(intravaginal)
67 F 1+ 3 May 1972 Unknown, carcass found (winter
kill)
21 M 4 4 February 1972 Unknown, carcass found (winter
kill) was weak and in poor
w condition on 2/3/72 0
24 F 12 4 February 1972-May 1972 Unknown, carcass found (winter
kill)
R-70-1 F 7 4 August 1971 Killed with drug
34 F 16 4 February 1972-March 1972 Unknown, carcass found (winter
kill)
70371 F calf 4 December 1971-May 1972 Unknown, carcass fo'lUI.d (winter
kill)
A60 F 15 4 December 1971-May 1972 Unknown, carcass found (winter
kill)
56 M 1 4 February 1972-May 1972 Unknown, carcass found (winter
kill)
-------------------
-------------------
Table 9. (cont'd.) Mortalities within pens, July 1971 -June 1972.
Hoose II Sex Age Pen Month -Year Cause
70171 M calf 4 October 1971-December 1972 Unknown, carcass found (winter
kill)
9671 F calf 4 May 1972 Introduced into pen 4, carcass
found
9771 F calf 4 January 1972 Introduced into pen 4, carcass
found
9871 F calf 4 January 1972 Unknown, carcass found (winter
kill)
w 10671 M calf 4 December 19 71 Unknown, carcass found (winter
I-' kill)
10771 F calf 4 January 1972 Unknown, carcass found (winter
kill)
10871 F calf 4 December 1971 Unknown, carcass found (winter
kill)
10971 F calf 4 January 1972 Unknown, carcass found (winter
kill)
11071 M calf 4 January 1972 Unknown, carcass found (winter
kill)
11471 M calf 4 June 1972 Unknown, carcass found (winter
kill)
Table 9. (cont'd.) Mortalities within pens, July 1971 -June 1972.
Moose If Sex Age Pen Month -Year Cause
11571 F calf 4 Hay 1972 Unknown, carcass found (winter
kill)
12071 M calf 4 December 1971 Unknown, carcass found (winter
kill)
12171 M calf 4 February 19 72 Unknown, carcass found (winter
kill)
126 F advanced 4 March 1972 Unknown, carcass found (winter
kill)
w
N
134 M calf 4 March 1972 Unknown, carcass found (winter
kill), found near trap next day
after trapping, weak and poor.
R 72-1 F advanced 4 May 1972 Introduced into pen and radio-
collared due to pregnancy.
Found carcass one month later.
74 F 1+ 4 May 1972 Immobilized via helicopter,
was weak and poor. Found
carcass three weeks later.
6071 M calf 4 June 1972 Unknown, carcass found (winter
kill)
-------------------
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A REVIEW OF
BLOOD CHEMISTRY OF MOOSE AND OTHER CERVIDAE,
WITH EMPHASIS ON NUTRITIONAL ASSESSMENT
by
Robert E. LeResche
Alaska Department of Fish and Game, Fairbanks, Alaska
Ulysses S. Seal
Veterans Administration Hospital, Minneapolis, Minnesota
Pat D. Karns
Minnesota Department of Natural Resources, Grand Rapids, Minnesota
Albert W. Franzmann
Alaska Department of Fish and Game, Kenai, Alaska
Management of moose (Alces alces) throughout their range histori-
cally has relied upon the crudest assessments of habitat carrying capacity.
These assessments have involved food habits studies (Peek 1973), range
productivity (Seemel 1969) and utilization (Milke 1969, Telfer 1968)
studies, and general population size and trend studies (Spencer and
Chatelain 1953, Spencer and Hakala 1964, Rausch and Bratlie 1965).
Secondary indicators of habitat's carrying capacity, as reflected in
moose population characteristics, have also been relied upon. Popula-
tion composition, natality and recruitment rates have been employed
extensively to indicate populations' relationships with their habitat
(Bishop and Rausch 1973, Pimlott 1959, Simkin 1965).
These secondary indicators in effect summarize the manifestations
of nutritional, behavioral, environmental and genetic forces acting upon
a population. They have been proven inadequate in situations where
delicate control of moose populations is necessary. Such indicators are
of little use as predictors and of even less value in understanding
specific ecological relationships. Therefore, the trend in research
since Peterson (1955) last summarized knowledge of moose has been toward
analyses of individual primary factors rather than of their collective
manifestations. This review concerns methods of measuring nutritional
status of wild moose populations, and presents details of recent studies
of moose blood chemistry as they relate to nutrition.
t1ETHODS OF NUTRITIONAL ASSESSMENT
Nutritional status of individual wild animals can be assessed
grossly or finely, and acutely or chronically. All body tissues reflect
nutritional status and history to a greater or lesser extent, and possi-
bilities for their analyses are as numerous as the tissues themselves.
Ideally, the method for assessing nutritional status of moose populations
should have several characteristics:
33
1. It should be sensitive to subclinical changes in nutritional
status.
2. It should be specific in its indications: i.e. capable of
indicating energy, mineral, protein, etc. balance.
3. It should be able to detect reasonably acute (e.g.: seasonal)
changes in nutritional status.
4. It should involve tissues easily collected from both live (to
allow serial collections from individuals) and dead (to allow
use of hunter-killed and accident-killed specimens) moose and
should be little affected by the stress of collection.
5. It should involve tissues which are easy for untrained personnel
to collect and preserve, and which can be processed by standard
inexpensive methods.
The broad spectrum of methods for nutritional assessment has been
outlined by Jelliffe (1966). Methods we feel are potentially practical
for use on moose include body weight and morphometry (LeResche and Davis
1971), analyses of urine (Blaxter et al. 1966), epithelial tissue
(Squires 1963, Nutr. Rev. 1970), saliva (Murphy and Connell 1970), hair
and nails (Sims 1968, Crounse et al. 1970a,b, Strain et al. 1972, Robson
and El Tahawi 1971), rumen volatile fatty acid (VFA) production (Gasaway
and Coady 1973) and blood constituents.
BLOOD ANALYSES -METHODS AND PATTERNS
Mammalian blood contains liquid (serum) and solid (cellular) frac-
tions. Serology involves analyses of the chemical properties of the
liquid fraction, and hematology involves analyses of the physical
properties of blood cells. Both types of analyses may be related to
nutrition.
Several chemical assays of serum are commonly employed in screening
batteries for human patients (Table 1). These tests have been assembled
to detect and discriminate between disease processes common in human
populations. Interpretation of test results depends upon the magnitude
of a sample's deviation from normal values, and upon the pattern of
results from all tests performed. Table 1 illustrates this concept by
tabulating patterns obtained in infection, malnutrition, tissue necrosis,
and various organ-specific diseases. Differences in hematology also are
associated with these patterns.
The most commonly measured parameters used in serological screening
are those 12 measured by the "SMA-12" Technicon auto-analyzer. Patterns
demonstrated in human and veterinary medicine are the following (cf:
Coles 196 7, Davidsohn and Henry 1969):
Electrolytes:
Calcium: Blood calcium may increase in hyperproteinemia,
34
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-
:....;-.
- -- - ---- -----
Table l. Test pat te ms as socic. ted '.vi th various pathological conditions.
~\ssay
Chc·les tern l
Calcium
I. Bilirubin
Total P!'otein
Albumin
L'ric Acid
BCN
Glucose
LDH
Alkaline
Phosphatase
SGOT
Infection :-!a:..nutri t ion
+
-+
+
See Re• ,:e and Hobbie (l972a,b) and. Searcy (1969).
+ tleT .. lation, -depression.
~ecrosis
+
+
+
+
Disease State
Rapid
Height
:.oss
+
+
+
+
+
Renal
Insufficiency
+
+
+
+
+
+
Hepatic
Disease
+
+
+
-+
+
+
+
--
Pseudohypo-
parathyroidism
+
-
Hypo-
thyroidism
+
+
+
- -
hyperparathyroidism, extreme neoplasia and other abnormal conditions.
Decreased calcium may result from hypoparathyroidism, vitamin D defi-
ciency, acute or chronic renal failure, starvation associated with
hypoproteinemia, parturient paresis (milk fever) and other causes. Thus,
calcium depression during winter months may be related to reduced nutri-
tion and possible hypoproteinemia and decreased vitamin D during this
period. Similarly, depressed Ca may be indicative of range deteriora-
tion in certain instances.
Inorganic Phosphorus: Similarly to Ca, phosphorous is elevated in
hypoparathyroidism, renal failure, and excessive vitamin D. Decreased
inorganic phosphorus often results from simple lack of phosphorus intake,
and as such can reflect dietary deficiencies.
Saccharides:
Glucose: Alterations in blood glucose level may be associated with
pancreatic abnormalities (diabetes or insulin poisoning). Hypoglycemia
may be an indication of extreme starvation, although gluconeogenesis can
maintain blood glucose at high enough levels to sustain life even in
severe starvation if it persists only a short while. For this reason,
uric acid (a product of protein catabolism in gluconeogenesis) is some-
times a more sensitive indicator. Ketosis may be accompanied by hypo-
glycemia, resulting from assorted digestive or nutritional disorders
(including carbohydrate deficiency).
Urea Compounds:
Urea Nitrogen (BUN): BUN level directly reflects dietary protein
intake and protein absorption, and is thus a good indicator of protein
energy balance. In addition, BUN may be pathologically elevated in
cases of excess catabolism of body protein or in renal failure.
Uric Acid: Uric acid is a product of purine metabolism and, as
such, may be increased in starvation for three reasons: 1) increased
tissue protein turnover, 2) decreased renal excretion of the chemical
due to acidosis, and 3) gluconeogenesis, with uric acid as a waste
product.
Sterol:
Cholesterol: Cholesterol level reflects diet, dietary changes and
the state of rumen metabolism. It is elevated when diets high in
saturated fatty acids are consumed. Consequently, starvation conditions
may result in hypocholesterolemia. Pathological conditions that may
alter cholesterol levels include thyroid and hepatic disorders and
nephrosis.
Proteins:
Total Protein: Total protein is a rather insensitive indicator of
nutritional status for it is maintained at near-normal levels except in
extreme distress. In sub-clinical cases of undernutrition, albumin is a
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better indicator. Most commonly, hypoprotememia is the result of trauma
(wound, burns) or renal disease. Occasionally, however, it can result
from gluconeogenesis and, as such, can reflect dietary insufficiencies.
In cases of chronic protein starvation total protein may decrease notice-
ably.
Albumin: Albumin is the smallest of the serum proteins and forms
40-60 percent of the total protein. It is important as a source of
amino acids and for fatty acid transport. Depressed albumin may indi-
cate deficient protein intake or excessive protein breakdown. Decreased
albumin is present in malnutrition and starvation.
Pigment:
Bilirubin: Bilirubin is a pigment liberated in the RE system by
breakdown of hemoglobin. Bilirubin measurement is commonly a test of
liver function, for its elevation is symptomatic of hemolytic diseases.
Other conditions elevating serum bilirubin include jaundice, cardiac
insufficiency and gangrenous pneumonia. Coles (1967) suggests, however,
that bilirubin is elevated only slightly in severe hapatic disease in
bovids, ovids and caprids.
Enzymes:
Alkaline Phosphatase: Alkaline phosphatase hydrolyzes phosphoretic
esters and is concentrated intracellularly in osteoblasts, renal tubules
and the intestinal mucosa. In humans, altered concentrations of this
enzyme are diagnostic for bone abnormalities and liver disease. Levels
normally are elevated during periods of growth and pregnancy, due to
increased osteoblast activity and the enzyme's production in the placenta.
Lowered levels can indicate malnutrition.
Lactic Dehydrogenase (LDH): LDH catalyzes the lactic acid-pyruvic
acid conversion, occurring in most tissues. In humans, elevated LDH
levels are indicative of several pathological conditions (e.g.: pulmonary
or myocardial infarction, hepatitis, cirrhosis, leukemia and carcinomas).
Glutamic oxalacetic transaminase (SGOT): SGOT is elevated in cases
of necrosis involving the tissues rich in the enzyme. Its concentration
is especially high in the heart, liver and skeletal muscle and thus
elevation is a sensitive indicator of myocardial infarction, and hepatic
and muscular necrosis and stress. In cattle, SGOT elevation has been
noted during starvation.
Electrophoretic fractionation of blood proteins is another common
analytic technique using auto-analyzer instrumentation (e.g.: Spinco
Analytrol R). This procedure separates total protein into albumin and
four globulin fractions (alpha-1 , alpha-2 , beta-, and gamma-globulins)
and allows calculation of albumin/globul1n ratio (A/G). Changes in
these values generally are indicative of these conditions:
Alpha globulins: Serum alpha-globulins are elevated during hypo-
proteinemia and acute cellular necrosis (especially alpha2 ).
37
Beta globulins: Beta-globulin levels are elevated in pregnancy and
sometimes during iron-deficient states. They may be depressed during
severe infection and hepatic disorders.
Gamma globulins: The "gamma" fraction defined by electrophoresis
is in reality a complex of proteins with many pathological correlations.
Most generally, gamma-globulins are elevated in many infectious diseases
and organ disorders.
A/G ratio: A/G ratio reflects relative change in albumin and
globulin fractions and is therefore depressed in protein malnutrition
and starvation. It may be elevated in cases of infection or other dis-
orders during periods of adequate protein nutrition.
Hormones:
Thyroxine: Serum thyroxine (T 4 ) level reflects thyroid activity
and thus is correlated directly with fasting metabolic rate and food
intake. This hormone may be analyzed by the method of Nobel and
Barnhart (1969).
Other Enzymes:
Creatine phosphokinase (CPK): This enzyme, similarly to SGOT, is
elevated in tissue breakdown, including muscle injury and vigorous exer-
cise (Rose et al. 1970), when it is released from skeletal muscle, heart
and brain. It is an indicator of physical or mental stress as well as
of trauma and perhaps nutritional stress. This enzyme can be measured
by the method of Nuttall and Wedin (1966).
Changes in serum levels of other enzymes indicate presence of
various disease processes and their possible locations. Use of enzymes
in differential diagnosis depends upon the magnitude of elevation, the
time course, the type of disease process (inflammation, necrosis, trauma,
etc.) and the tissue affected. Identification of the tissue source is
possible by assaying for enzymes of defined localization, Table 2. Such
assays are being developed very rapidly in humans and offer great promise.
Their application to moose will require direct confirmation and estab-
lishment of baseline norms, since variations between species have been
documented (Zimmerman et al. 1965). Another potential use of enzyme
measurements involves tissues as well as serum and may allow detection
of changes in control systems (e.g.: adrenocortical activity). This
might provide a more reliable index than gland weights of the functional
changes in adrenal activity that are postulated to occur in response to
"stress", however produced. At present, the available data on serum
constituents do not provide such a correlation.
Hematology:
The basic battery of hematological values includes packed all cell
volume (PCV), hemoglobin concentration, leucocyte count (WBC), erythrocyte
count (RBC), and differential count of leucocytes. In general, PCV, RBC
and hemoglobin levels reflect oxygen-carrying capacity and homeostasis.
38
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--- - - - --- ------ - --Table 2. Serum enzyme tests -tissue specificity*.
A.
B.
Enzyme
High Specificity
Acid Phosphatase
Alanine transaminase
Arginase
Sorbitol DH
Alcohol DH
5' Nucleotidase
Glutamic DH
Amylase
Lipase
y-glutamyl transpeptidase
Moderate Specificity
Aspartate transaminase (SGOT)
Isocitrate DH
Creatine Phosphokinase (CPK)
C. Low Specificity
Alkaline phosphatase
Lactic dehydrogenase (LDH)
D. Within Cell Specificity
Aspartic transaminase
Alanine transaminase
*Taken from Wilkinson (1970).
Tissue Sources
Prostate, Erythrocytes
Liver
Liver
Liver
Liver
Hepatobiliary tract
Liver
Pancreas, salivary glands
Pancreas
Liver
Liver, heart, skeletal muscle
Liver, heart
Skeletal muscle, heart, brain
Bone, liver, intestinal mucosa, placenta, kidney
All tissues (some isoenzyme specificity)
Mitochondria form, cell sap form
Cell sap only
-
Nutritional deficiencies can produce many anemias and anemia complexes
that are reflected in these values. PCV reflects proportion of erythro-
cytes in whole blood and similarly is correlated with nutritional state.
Interrelationships among all these parameters are too complex to discuss
in detail here.
WBC and differential counts are useful in diagnosing various infec-
tions, intoxications, and neoplasms. Changes in absolute numbers and
relative proportions of the several leukocytic cell types, when considered
with other symptoms, can help differentiate among various disorders.
However, individual variation makes serial bleeding of the same animal
almost essential if small changes are to be detected. No direct nutri-
tionally related changes in WBC or differential counts have been estab-
lished firmly, but such information can prove of value in discovering
pathology resulting from nutritional deprivation.
The patterns described occur in humans and in domestic species.
Many may be expected to apply to other mammals as well. However, direct
confirmation by empirical or experimental data is necessary to draw firm
conclusions regarding patterns in moose. Some possible sources of
artifically altered test results are indicated in Table 3. Some of
these sources of artifact are peculiar to the Technicon SMA 12 systems
which are sensitive to any changes altering the optical properties of
serum, including lipemia (important for many carnivores--but not commonly
seen in cervids), hemolysis, and high bilirubin.
The SMA 12 system, in addition, gives incorrect (low) values for
serum albumin in moose blood, due to dye binding capacities. Therefore,
electrophoresis is used to determine this value.
THE PROBLEM OF STRESS
Severe artifacts may result from the animal's acute physiological
state when the blood is collected. These are related to recent food
intake and to handling stress. The former variables are uncontrollable
in wild animals, and the latter are "controllable" only to the extent
they can be standardized in all animals handled. Usually this extent is
slight, for individuals differ in response to drugs, traps, and handling.
Thus, handling effects have been reported in every study to date (see
bibliography section G). The very substantial effects of one method of
handling an animal to obtain the samples are documented in Table 4. It
is clear that many results depend upon whether the animal was shot,
drugged, or manually restrained. These variables are at least known.
More importantly, the marked effects of restraint 24 hours previously
suggest that not only can short-term serial studies be seriously com-
promised, but that it is possible that levels in immediately obtained
samples from wild animals may reflect the previous several days' activities.
A system to classify individual excitability states based on heart
rate, respiratory rate, and rectal temperature has been developed for
bighorn sheep (Franzmann 1972). Each individual was classified into one
of five classes of excitability (not excited, slightly excited, moderately
40
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---- --- -- ---- ---- --
Table 3. Possible artifactual patterns of test results.
High-Old or Detergent
Recent Protein Lipemic on clot High Contaminated Specimen
~'l.ss ay Heal Neal Serum Serum Bilirubin Convulsions Tubes Hemolysis Dehydration
Cholesterol +(±.) + + +
Calcium + + +
Phosphorous + +
r. Bilirubin +
Total Protein + + + + + +
-"' f-'
Albumin + + + + + +
L'ric Acid + + + +
BID! + + + + +
Glucose + +(±.) + +
LDH + + + + +
Alkaline
Phophatase + + + +
SCOT + + + + +
See Reece and Hobbie (l972a,b) and Searcy (1969).
+ elevation, depression.
Table 4. Acute effects of phencyclidine and promazine administration on
hematology and blood chemistry of eight pregnant, adult does*
(data of Seal, Ozoga, Erickson and Verme (1972a)).
Assay (Units)
Hemoglobin (gm/100 ml)
Red blood cells (106/mm3)
Hematocrit (vol. %)
MCV ()J3)
MCHC (%)
White blood cells (103/mm3)
Fibrinogen (mg/100 ml)
Serum protein (gm/100 ml)
SGOT (K.U.)
CPK (I. U.)
Sodium (mEq/1)
Potassium (mEq/1)
Calcium (mg/100 ml)
Phosphorus (mg/100 ml)
Cholesterol (mg/100 ml)
Restraint
18.6 ± 2.5
13.3 ± 1.7
49 ± 5.7
37 ± 3.6
37 ± 1.0
2120 ± 700
230 ± 35
5.7 ± 0.7
12 8 ± 26
so ± 19
144 ± 6.0
4.3 ± 0.3
9.8 ± 0.5
6.7 ± 1.0
66 ± 7
Post Drugs t p
15.8 ± 2. 7 4.48 .005
11.3 ± 1. 7 3.22 .02
42 ± 6.9 4.02 .01
37 ± 3.1 0.29 NS
37 l_ 0.7 0.36 NS
1500 ± 470 4.47 .005
192 ± 33 10.82 .001
4.9 ± 0.4 5.24 .002
142 l: 32 2.63 .05
101 + 36 5.09 .005
142 :± 3.0 0.64 NS
4.6 :±_ 0.5 1.32 NS
9.7 :t 0.7 1.19 NS
4.9 ± 0.9 4.50 .005
58 ± 5 4.33 .005
*Studies were performed in April 1969 on eight pregnant does. The animals
were guided through runways into restraint boxes and restrained manually.
Blood samples were taken and then each animal was given 1 mg/kg of
phencyclidine and 0.5 mg/kg of promazine by intramuscular injection. A
second set of samples was taken 45 minutes later. The data are recorded
as means i standard deviation. Statistical analysis was by the t-test
for paired variates.
42
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excited, excited, and highly excited). It was concluded that classes of
excitability could be established and would aid in interpreting blood
values. At present, we are accumulating data based on this study to
establish excitability classes for moose.
Evaluating and subsequently classifying stress states in animals
may also be approached through analyses of intracellular enzymes such as
LDH, SGOT and CPK, which escape into the circulatory system when cells
are injured (Coles 1967). Tissue breakdown and subsequent release of
these enzymes has been examined in pathologic conditions in domestic
animals (Blinko and Dye 1958, Whanger et al. 1969). The influence of
handling excitability on SGOT values in bighorn sheep was demonstrated,
but no correlation with degree of excitability was noted (Franzmann and
Thorne 1970). Another potentially useful index of handling is a combina-
tion of CPK and LDH levels which show a different time course of elevation
and decline after handling and tissue damage (Seal et al. 1972a).
Immobilizing drugs have potentially contradictory effects on blood
values. Many of these drugs calm the individuals being handled, decreas-
ing excitability and stress. In addition, however, drugs may have other
more discrete physiological affects due to their pharmacologic action
(cf: Table 4, Harthoorn 1965, Fujita 1970). These must be experimentally
determined for each species studied.
Since obtaining blood from sufficiently large samples of wild
animals is difficult by any means, standardization of procedures is
often extremely difficult. Nevertheless, it is extremely important to
standardize methods as much as possible, and to realize that some
measured values may be more plastic under stress (e.g. LDH, CPK, SGOT,
glucose) than others (e.g.: BUN, uric acid, cholesterol, proteins).
With these relationships in mind, it is possible to use the more plastic
assays as stress indicators, while examining nutritional status using
those values little affected by handling. In any event, it should be
recognized at the outset that even well-documented "norms" for a wild-
life population are not strictly comparable to resting normals for
human or domestic species. Instead, the "normal" values are resting
normals modified by the variable stress of the standardized animal
handling procedure. For this reason, blood studies of wildlife are
concerned with trends, and often require large sample sizes in order to
document differences between populations.
BLOOD STUDIES IN HILD POPULATIONS -STRATEGY AND DESIGN
The multitude of stress-related variables, the difficulty of stan-
dardization, the need for confirmation of relationships for each species,
and the lack of baseline "normal 11 values complicate blood studies of
moose and other cervids. In essence, we may begin at the current level
of knowledge and technology of human blood-nutrition relationships, but
must reconfirm relationships and ascertain normal values, within the
restrictions of hard~to~standardize procedures.
43
. ·---····-·-·-------------------------------,
Until recently, blood studies of moose and other cervids were few
and superficial and only sometimes related to nutrition. A systematic
effort to use blood analyses to evaluate the condition of wild cervids
was first reported by Rosen and Bishoff in 1952. Braend (1962) considered
blood groups in moose, Nadler et al. (1967) studied serum proteins and
transferrins, and Houston (1969) analyzed several serum parameters from
13 moose. Dieterich (1970) reported hematology of six moose and several
other arctic mammals. More thorough nutrition-related studies have been
carried out on other cervidae. Herin (1968) reported 14 blood parameters
for 39 elk (Cervus canadensis). Weber and Bliss (1972) measured 13 para-
meters in wild and captive elk. Kitts et al. (1956) related age and
nutrition to hematological values in black-tailed deer (Odocoileus
hemionus c.). Seal and Erickson (1969) and Seal et al. (1972a,b) have
reported on extensive studies involving nutrition -stress -blood
constituent relationships in white-tailed deer (Odocoileus virginian~g),
These studies have indeed demonstrated that blood assays in these
cervids are subject to more sources of normal variation than are
encountered in human studies. This has increased the complexity of
interpretation substantially and has made the establishment of baseline
normal values difficult. Identified variables include age, sex, repro-
ductive status, season, and method of handling the animal. Evaluation
of an animal's or a herd's condition, a primary motivation for these
studies, has been confounded by these multiple sources of variation
intermingled with other sources of variations in wild animals such as
nutritional history, disease, and parasitism. The advent of experimen-
tal studies on animals of known condition and history (e.g.: Seal's
group on Odocoileus; Alaskan studies on penned moose) now offers the
promise of unraveling these variables and yielding the necessary infor-
mation for interpretation of results from field studies.
Establishing nutrition-blood parameter correlations in moose popu-
lations involves: (a) determining boundary conditions, or those char-
acteristics of an individual animal that must be known before blood
values may be interpreted, (b) establishing normal values within these
boundary conditions using standardized handling and collecting procedures
and, (c) determining the nature and magnitude of changes wrought in
blood values by known changes in food intake, nutrition, or related
parameters.
Boundary conditions which have been identified in cervid species
are tabulated in Table 5. Those demonstrated to date for moose are
listed in Table 6. Species differences have become apparent in most
measurements. More critical for interpretation of literature data are
the many seasonal changes and the impact of handling methods. Both of
these involve effects of sufficient magnitude to invalidate uncontrolled
comparisons. The effect of time of day (circadian rhythms) has not been
examined for any cervid species, but has been shown to be important in
other mammalian species. Other conditions of possible importance
include posture at time of sampling (standing or recumbent), geographic
location (latitude, elevation, and climate), and perhaps genetically
based intraspecies population differences.
44
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II Table 5. Demonstrated boundary conditions in Cervidae.1 •2
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3 Assay
Hematology
Hemoglobin
Red b load cells
Hematocrit
MCHC
MCV
White blood count
Differential
Chemistry
Cholesterol
Calcium
Phosphorus
Bilirubin
Uric Acid
Total protein
Albumin
Gamma globulin
Glucose
BUN
LDH
SGOT
Alkaline p'tase
CPK
Hormones
Thyroxine
Cortisol
Insulin
Testosterone
Species
+
+
+
0
+
+
+
+
+
+
+
+
+
+
+
+
+
Sex
+
+
+
0
0
0
0
+
0
+
+
Age
0
0
+
+
+
+
+
+
+
+
+
+
+
+
Season
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Repro-
ductive
Status
+
+
+
0
0
+
+
+
+
+
+
+
+
Time
of Day
0
0
0
Handling Fasting
Method or Fed
+
+
+
0
0
+
+
+
+
+
+
+
+
+
+
0
0
0
0
0
+
0
0
+
+
0
0
1 Exclusive of fetal, neonatal, and first one month of life data since many differences
have been documented for these periods in every mammalian species studied including
white-tailed deer and moose.
2 Genera covered include Odocoileus, Alces, Rangifer, and Cervus.
3A "+" indicates significant differences have been reported, while a "0" indicates no
difference, and a "blank" indicates information lacking or inadequate. The initial
condition is that the animals be in good health and nutrition.
45
Table 6. Boundary conditions in Alaskan moose.*
Sex Age Pregnancy Season
Calcium 0 + 0 +
Phosphorus 0 + 0 +
Glucose 0 + + +
BUN + 0 0 +
Uric acid 0 0 + +
Cholesterol + + 0 +
Total protein + + + +
Bilirubin 0 + + +
Alkaline p 'tase 0 + + +
Lactic dehydrogenase 0 + 0 +
SCOT 0 + 0 +
Hemoglobin 0 + +
Hematocrit 0 + +
White blood count 0 + +
*The "O" indicates that no effects of the specified condition were observed
upon the assay. The "+" indicates the occurrence of significant changes
or significant differences and thus that the establishment of norms
requires inclusion of this condition in formulating the data base. The
effects of season are probably complicated by seasonal reproductive and
nutritional factors.
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The only data available concerning the latter conditions are those
gathered in protein polymorphism studies, primarily designed to delineate
subspecies and specific relationships (see bibliography section F). No
polymorphisms in hemoglobin or serum protein patterns have been observed
in several subspecies of moose (Table 7). This is in contrast to rein-
deer (transferrins) and white-tailed deer (transferrins and hemoglobins).
There have been no reports on serum (or tissue) enzyme polymorphisms. In
contrast, the polymorphisms of white-tailed deer hemoglobin are sufficient
to apply to studies of subspecies relationships, selection effects,
population differences, isolates, and geographic clines. In serum, the
fibrinopeptides A (19 amino acids) and B (17 amino acids) are the most
rapidly evolving peptides known at present and, hence, are most likely
to provide information on phylogenetic relationships of closely related
species and genera. Data on the sequences of fibrinopeptides A and B
from moose and six other cervids (Tables 8 and 9) permit inferences con-
cerning phylogenetic relationships among these cervids. Notably, they
indicate the very close affinities of Rangifer, Alces and Odocoileus,
with no sequence differences between the reindeer and moose and only one
difference from the mule deer. Also, the three species of Cervus
exhibited only one sequence difference, in the Sika deer. These peptides
are clearly not useful for species and subspecies studies, but may allow
grouping of genera at the subfamily level.
Seasonal boundary conditions in moose are especially interesting
in view of seasonal changes in energy balance (Gasaway and Coady 1973).
If other conditions, (sex, age, reproductive status, handling method)
are kept constant, alterations in blood values caused by known seasonal
changes in nutrition may be good indicators of blood-nutrition relation-
ships. The relationships are not clear, for climatological stresses and
endocrine relationships (cf: Seal et al. 19 72b) also change seasonally.
Nevertheless results, described below, are encouraging.
BASELINE NORMALS
It is impractical to attempt to develop baseline normal values for
moose similar to those existing for humans and some domestic species.
The number of individuals which must be sampled to meet boundary condi-
tions for all population segments is prohibitive. Nevertheless, most
studies of other wild cervids have presented lumped data with little or
no regard to boundary conditions. Existing lumped data of moose blood
values, including our own unpublished data, are presented in Table 10.
It is evident that moose blood values fall in the same general range as
those reported for other cervids. Little more can be said at this time,
given the great variety of individuals tested and the methods used to
secure samples. However, interesting apparent differences between
groups of moose exist for MCV, cholesterol, glucose, BUN, albumin and
serum enzymes (Table 10). Further analyses af data from tlinnesota and
Alaska, applying the appropriate boundary conditions, may elucidate
these differences if they are real.
47
Table 7. Transferrin* polymorphisms in the Cervidae.
Number Observed Postulated
Location Animals Phenotypes Alleles
S£ecies
Rangifer tarandus (0 bands in homozygotes, 0 bands in heterozygotes)
1. Gahne & Rendel (1961) Sweden 64 6 3
2. Braend (1964a) Norway 132 15 6
3. Braend (1964b) Norway 829 27 8
4. Nadler et al. (1967) Alaska 37 Multiple ?
Odocoileus virginianus (2 bands in homozygotes, 3 bands in heterozygotes)
.f:'-
(XJ
1. (1965) Miller et al. Iowa 150 3 2
2. Seal & Erickson (1969) Minnesota 100 1
3. Seal & Venne (Unpubl.) Michigan 40 3 2
Alces alces (2 bands)
1. Braend (1962) Norway 27 1 1
2. Nadler et al. (1967) Alaska 7 1
3. Nadler (1968) Canada 1 1
4. Seal & Karns (Unpubl.) Minnesota 150 1 1
*All phenotypes had at least two bands on electrophoresis.
-------------------
- - - - --· - - --· -- - ------
Table 8. Fibrinopeptide A sequences.*
Species
Reindeer Ala Asp Gly Ser Asp Pro Ala Gly Gly Glu Phe Leu Ala Glu Gly Gly Gly Val Arg
l'foose
Mule Deer
.j::o-Muntjak Ser Thr
\0
Sika Deer Ser Ser
Red Deer Ser Ser Asp
Elk, American Ser Ser Asp
*Taken from tabulation D-95 of Dayhoff (1972). A dot indicates that the amino acid is identical with that of
the reindeer in that position.
U1
0
-
Table 9. Fibrinopeptide B sequences.*
Species
Reindeer Gly His Leu Ala Asp Tyr Asp Glu Val Glu Asp Asp Arg Ala Lys Leu His Leu Asp Ala Arg
Moose
Nule Deer Asp
Nuntjak Ser Thr
Sika Deer Ser Thr Glu Glu
Red Deer Ser Thr Glu Glu
Elk, American Ser Thr Glu Glu
*Taken from tabulation D96 of Dayhoff (1972). A dot indicates that the amino acid is identical with that of the rein-
deer in that position.
-----· -.. --·----- ----
----------Table 10. Blood data on moose (A!c~s 2~ces ssp.)
Reference
Location
Number Animals
Type
Ages
Sex
Season
Drugs
Hemoglobin (gm/100 ml)
Red blood cells (lo6;mm3)
Packed cell vol. (Vol cells/
100 ml blood)
MCHC (gm/100 ml)
HCV (u3)
Sed rate (mm/hr)
White blood cells (l0 3!mm3 )
Neutrophils (103 /mm3)
Lymphocytes (lo3Jmm3)
Eosinophils (103/mm3)
Cholesterol (mg/100 ml)
Calcium (mg/100 ml)
Phosphorus (mg/100 ml)
Bilirubin (mg/100 ml)
Uric acid (mg/100 ml)
BUN (mg/100 ml)
Glucose (mg/100 ml)
Total protein (gm/100 ml)
Albumin (gm/100 ml)
Gamma globulin (gm/100 ml)
Alk. phosphatase (mU/ml)
LDH (mU/ml)
SGOT (mU /ml)
Thyroxine (T 4 ) (ug/100 ml)
Dieterich (1970)
Alaska
2 (3X)
Captive
4 mos.
M
Surruner
Restraint
X ± S.D.
13.5 ± 0.4
5.25 ± 0.43
38 ± 3
36 ± 2
72 ± 7
27 ± 13
5.87 ~ 1.13 -
2.46 ± 0. 88
3.10 ± 1.11
0.23 ± 0.17
A. a. shir%i
Houston (1969)
Wyoming
13
Wild
lOA, 3Y
4M, 9F
Hinter
~uccinylcholine
X ± S.D.
13 ± 31, 5 ± 1. 82
63 ± 5. 7
6.4 ± 0.9
4.5 ± 1.4
-\--
A . a. andersoni 3
Karns & Seal (Unpubl.)
Ninnesota
ca: 91
Wild
Adult
65M, 26F
Oct. Dec.
Shot
X ± S.E.
13.3 ± 1.3
6.2 ± 1.2
41 ± 5.8
35 -1.4
59 ± 5.6
54 ± 10
9.7 ± 1.3
6.4 ± 1.3
0.5 ± 0.1
0.7 ± 0.1
8.5 ± 3.2
79 ± 8
6.9 ± 0.7
3. 3 ± 0.4
1.6± 0.6
80 ± 11
142 ± 15
59 ± 6
--
LeResche (Unpubl.)
Alaska
ca: 520
Wild, pen
Hi xed
Mixed
- -
Year-round (some serially sampled)
Shot, succinylcholine, M-99, other x =S.D., ± S.E. 0.5
14.9 ± 4.7
40.4 ± 11.8 1.5
37 ± 4.0 0.4
4.19 ± 2.52 0.24
1. 37 ± 0.75 0.16
2.60 ± 0.80 0.16
87 ± 21 0.9
10.6 ± 1.1 0.05
5.6 :! 1.9 0.08
0.4 ± 0.2 0.01
0.6 ± 0.6 0.03
13± 12 0.5
121 ± 54 2.4
6.9 ± 0.9 0.04
3.9 ± 0.7 0.03
0.03
81 ± 71 3.1
339 ± 136 6.0
159 ± 59 2.6
5.4 ± 2.3 0.15
1 Seven animals in July-September; 2 Five animals in October-February; 3 Sex, season and area differences found in this study.
Only data on males here; 4 Sex, age and season differences.
--
-----·-·-··--------------------------------------------------------,
BLOOD-NUTRITION RELATIONSHIPS IN MOOSE
Two major continuing studies of moose blood patterns are underway
in North America: those of LeResche and Franzmann in Alaska and of
Karns and Seal in Minnesota. Preliminary data from both are discussed
here.
The Alaska study was designed to develop practical methods for
assessing population/habitat relationships on different ranges through-
out the state. It has used three primary sources of materials: (1)
moose contained in four 2.6 km2 enclosures at the Kenai Moose Research
Center and trapped, immobilized, bled, weighed and measured periodically;
(2) unenclosed moose immobilized on the Kenai Peninsula for marking,
bled, measured and released and; (3) moose killed by hunters or collected
for rumen VFA studies from several areas of the state.
The Minnesota study has thus far utilized hunter-killed specimens
from two areas of the state, taken at two different times of year. Its
objectives include testing regional differences in nutrition, In addi-
tion blood profiles are being used as indicators of the effects of
parasitism, primarily by Fascioloides magna.
Seasonal Variation
Natural seasonal changes in diet and food intake, coupled with
seasonally varying demands of pregnancy, lactation and breeding, provide
an opportunity to study blood-nutrition relationships on a pseudo-
experimental basis. That is, animals sampled during winter months are
analagous to experimental animals on sub-maintenance diets, and those
sampled during summer are analagous to animals fed high-quality rations.
Seasonal energy budgets of moose calculated from weight change, food
intake and value, rumen VFA production, and theoretical considerations
show profits roughly from Hay through September and deficits from
October through April (Gasaway and Coady 1973 and Figure 1). Although
compounded by changes in photoperiod and resulting endocrine relation-
ships (cf: Silver et al. 1969), pregnancy, prior dietary history and
disease, certain blood values do reflect seasonal changes in energy
balance in predictable ways (Tables 11-12).
Urea nitrogen varies directly in correlation with energy balance
in Alaska moose (Table 11, Figure 1), reaching highest levels during
seasons of positive energy balance. Similarly, October levels in 14
males from northwest Minnesota (8.5mg/100 ml) were significantly (P=.OOS)
higher than December levels (5. 7 mg/100 ml) in seven males. BUN has
been repeatedly correlated with dietary protein and protein utilization
balance in cervids (cf: Houston 1969, Seal et al. 1972b) and appears to
be one of the best correlates, being relatively stable and unaffected
by boundary conditions.
Glucose levels are also correlated with seasonal energy balance in
Alaska. This is somewhat surprising in view of rapid elevation of blood
glucose in handling stress. The fact that trends in glucose levels
follow seasonal energy changes suggests that collection procedures can
52
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-------------------'T! so or ..... ~ ~~ ... ~ ()Q .
r-' .
0 :;o Q)
• Holll > 500 r-'
~Pl ~' Q) ...... __ ·, r-'rt
,~' jll ..... _J (I} 0
' " ;o;";:l ·,~,,.
!5 ~ ...... ~-.....,
g400 ~\: .....
S'O 200 0 (I} 0:·· 0 0 ",-; (I} 0 ro H1 en .
(I} 0 ro
r-' Q) ro n (f) 300 150 rt
::::: ro p.. +-' ' Vl c:r en w f-' Q) 0
0 ~ 200
p..
125 < ll.l _j f-' c ro
(Jl '+-
rt 0 0 100 100 ;:l +-c: c rt
'1 Q) ~ ..... -rt u c ..... ~ 0 0 ::J Q) 0 Pl
f-' (l_ z (JJ May-Jun Jui-Aug Sep-Oct Nov-Feb Mar-Apr rt :::> Pl
rt CD c:
(Jl
-
Table 11. 1 Seasonal variation in selected blood values in adult female Alaskan moose. (Data of LeResche).
n
Calcium (mg/lOOml)
BUN ( mg/lOOml)
Glucose (mg/lOOml)
Bilirubin (mg/lOOml)
Cholesterol (mg/lOOml)
SGOT mU/ml
LDH mU/ml
Total Protein (g/lOOml)
Albumin (g/lOOml)
A/G (ratio)
n
Thyroxine (T 4 ) ug/lOOml
}lay-June
80
10.47 -0.62
(a) 13.0 ± 14.22
(a) 118.6 ± 29.2
0.4 ± 0.23
82.5 ± 21.8
(a) 108.1 ± 36.0
(a) 222.9 ± 88.3
6.9 ± 0.53
4.0 0.47
1. 4 7 ± 0. 38
92
5.8 ± 3.2
July-Aug
30
10.61 ± 0.89
(a) 32.0 ± 7.54
(a)134.9 ± 41.0
0.7 0.34
(a) 97.9 ± 17.2
(a)153.9 ± 44.9
(a)337.7 ± 108.0
(a) 8.1 ± 0.66
4.7 ± 0. 79
1. 54 ± o. 53
53
5.7 ± 1.7
MO:c!THS
Sept-Oct
24
11.28 ~ 0.80
(a) 21.3 ± 11.8
134.4 ± 49.4
0.8 ± 0. 32
(a)l09.4 ± 32.9
158.1 ... 36.7
345.8 ± 81.4
8.2 ± 0.52
5.2 0.52
1.72 -0.34
46
5.2 ± 1.6
Nov-Feb
109
10.8 ±
(a) 6.9 ±
1. 32
4.80
(a)115.4 ± 67.7
0. 39± 0.3
(a) 83.0 ± 13.9
(a)212.1 ± 46.5
(a)389.7 _._ 138.8
(a) 6 . 7 ± 0. 70
(a) 3.8 ± 0.48
1. 35± 0. 30
49
(c) 4.5 ± 1. 4 (b)
Mar-Apr
30
11.1 ... 0.58
(a) 4.0 ± 2. 81 (a)
(a) 90.8 ± 31.6 (a)
0.33± 0.05
86.8 ± 12.5
(a)l21.0 ± 28.7 (a)
(a)291.0 :: 60.8 (a)
6.8 ± 0.48
3.5 ± 0.43
1.09± 0.28
no data
1 All data are given as mean ± standard deviation. The statistical comparisons were made with the "t" test for unpaired variates.
(a) Difference between months significant at P
(b) Difference between months significant at P
(c) Difference between months significant at P
.001
0.025
0.05
---- - ---- -
------ -
-
-------~-----------
Ln
Ln
Table 12. 3 Serum protein electrophoresis of ~tinnesota adult male moose samples. (Data of Karnes and
Seal).
Area N
Northwest
October 16
December 8
Northeast
October 14
1 p ..2. 0.01
2 p ..2. 0.001
Total Protein
gm/100 m1
6.9 ± 0.69
7.3 ± 0.74
6.5 ± 0.65
Albumin
gm/100 ml
3.3 ± 0.39
3.3 ± 0.36
3. 7 ± 0.40 1
Gannna Globulin
gm/100 ml
1. 6 + 0. 56
2.4±0.72 1
1.1 ± 0.37 1
A/G
0.93 ± 0.23
0.84 ± 0.15
1.34 ± 0.26 2
3 All data are given as means ± standard deviation. The statistical comparisons were made with the
"t" test for unpaired variates. The Northeast-October and Northwest-December samples were compared
with the Northwest-October group.
be at least reasonably standardized, however stress-producing they might
be.
Cholesterol was elevated during summer months in Alaska, suggesting
a seasonal increase in dietary fatty acids. No change occurred in blood
cholesterol level between October and December in north¥rest Minnesota.
Cholesterol levels in Minnesota were significantly lower (49-56 mg/100
ml) than in Alaska (82-109 mg/100 ml), suggesting interesting differences
in diet and rumen function (Peek 1973). These differences deserve
further investigations.
Total protein levels in Alaska were elevated in summer and depressed
in winter, probably reflecting changes in dietary protein. The signifi-
cant decline in serum albumin levels between September-October and
November-February further suggests a dietary influence, although no
significant change in A/G ratio occurred. Total protein levels did not
change significantly between October and December in Minnesota (Table 12).
senT and LDH levels in Alaska both peaked during winter months,
suggesting generalized tissue breakdown during this period of negative
energy balance. Annual lows are reached in May and June, when forage
quality suddenly improves. These lows, however, are somewhat surprising,
since near-term pregnancy and early lactation increase physical stress
during this time, The LDH pattern in Minnesota is similar, with DeceniDer
levels significantly lower (204 mU/ml) than October levels (222 mU/ml).
However, SGOT levels in rtinnesota declined significantly from 151 mU/ml
to 85 mU/ml between October and December. Serum concentrations of both
enzymes were higher in Alaska than in Minnesota, perhaps due to differ-
ences in animal handling procedures. These seemingly contradictory
results deserve further exploration.
Serum thyroxine (T4) levels in Alaska (Table 11) followed the
pattern described by Seal's group for white-tail does, which involved a
significant decline in serum thyroxine with reduced nutrition over a
three-year period in experimental animals (Seal et al. 1972b). Their
studies of wild populations also yielded very low values in herds whose
condition is considered poor and whose range is depleted. These studies
also suggested reduced T4 levels were due not only to low nutrition but
also reflected naturally reduced metabolism and food intake in winter.
Our results presented here are the first empirical evidence of such a
mechanism in moose.
Hematological patterns vary seasonally in Minnesota (Table 13),
with a significant decline in erythrocyte numbers and hemoglobin concen-
tration in males from October to December. These changes are consistent
with moderate normocytic anemias, which may result from protein deficiency.
or with parasitic infestation aggravated by normal seasonal decline in
dietary protein. Northwest Minnesota moose do carry a very heavy liver
parasite load. This condition is also reflected in significantly elevated
gamma globulin and depressed albumin levels, relative to those found in
northeast Minnesota (Table 12).
56
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--- ---·----- --'---·---
Table 13. Hematology of adult moose from northern Minnesota (data of Karns and Seal) .
SEX
AREA Hemoglobin (gm/100 ml) RBC (lo6 ;mm 3) Hematocrit (%) MCV ()l3) MCHC (%)
Honth n X ": S.E. n X ± S.E. n X ± S .E. n X::: S .E. n X ± S.E.
~!ALES
NORTHWEST
October 40 13.3 -l.28a 41 6.2 -l.l6c,k 29 41.3 -s.8oe 38 59.2 :': 5.63g 37 34.6 ± l.38h --
2.69e,f,m l.l3h,i December 8 11.1 ± 1. 16a ,b ,j 6 5.0 ± o.ssc,d 30.7 = 5 59.9 :': 4.82 7 36.6 :':
NORTHEAST
lJl October 17 13.5 ± 2.12b 22 6.6 -0.28d,n 15 39.5 ± 5.33f 19 62.3 ± 4.08g 12 33.7 ± 1. 48i ,p
-.J
FEMALES
NORTH\o/EST
October 8 11.6 ± 2.80 7 5.4 ± 0.73k 6 33.5 + 7.40 9 58.5 :': 6.98 9 35.7 :': 2.40
December 10 13.2 ::: l. 75j 9 5.7 ± 0.84 8 36.6 -4.3lm 9 59.8 ± 6.89 9 37.0 ± 2.50
NORTHEAST
October 8 12.3 ± 2.27 7 5. 7 :': 0.85n 5 37.8 ± 3. 77 6 59.2 ± 3.61 7 36.0 :': 1.63P
g, k, n -Difference between means statistically significant at p < 0.05.
j ' m, p -Difference between means statistically significant at p < 0.025.
a, b. c' f, h, i -Difference between means statistically significant at p < 0.005.
d, e -Difference bet'V.'een means statistically significant at p < 0.001.
Physiological Costs of Reproduction
Energy costs of pregnancy and lactation may approach 40-55 percent
of maintenance levels in moose (Gasaway and Coady 1973). These costs
are reflected in weight deficits of 15-20 percent in late July and 6-8
percent in October-December (LeResche and Davis 1971, LeResche unpubl.).
Tables 14 and 15 illustrate how these physiological costs of reproduction
may be reflected in blood values of Alaska moose.
Early-term pregnancy (October through January) resulted in a signi-
ficant increase in glucose and a significant decline in cholesterol
levels, by comparison with non-pregnant adult cows (Table 14). Cows in
late-term pregnancy had slightly elevated glucose levels and greatly
elevated levels of SGOT, alkaline phosphatase and LDH, compared to non-
pregnant cows. These levels of SGOT and LDH suggest greatly accelerated
tissue destruction in late-term, perhaps reflecting the combined effects
of increased fetal demands and very low-nutrition forage. Alternately,
these elevations might indicate fetal or placental injury during handling
or increased cellular turnover at this time. In normal well-nourished
humans, alkaline phosphatase is elevated in late-term pregnancy; whereas,
SGOT is unchanged or slightly depressed.
Glucose elevations in pregnant animals are difficult to explain,
but likely represent accommodation to fetal demands rather than enhanced
energy reserves in pregnancy. Glucose in pregnant cows is depressed
significantly (P<.OOl) in February-April below October-January levels.
This occurs at a time when the fetus is growing most rapidly, and must
represent a decline in this type of energy available for growth. Decreased
cholesterol during early pregnancy might result from lipid uptake by
rapidly-growing fetal myelin structure. Endocrine relationships might
further be involved.
Blood values differ significantly between adult cows with no calves,
those with one calf and those with twins (Table 15). These differences
perhaps reflect some of the nutritional costs of rearing young. Cows
with one calf tested in Table 15 weighed a mean 368 kg, significantly
less than those with no calves (392 kg), in September-December. The
significant increase in BUN levels from cows with no calves to those
with one and those with twins is perplexing. Perhaps a hidden seasonal
bias in the sample is operating, for data from all year are lumped and
progressive calf loss may have resulted in cows with calves being sampled
more heavily in summer months, when BUN levels are high. Alternately,
the values might reflect increased protein catabolism required for
lactation. Relationships are unclear.
Glucose is highest in cows with no calves, significantly lower in
those with one calf, and again significantly lower in those with two
calves. This most likely represents the cost of lactation, which may
involve glycogen stores. SGOT and LDH are significantly elevated in
cows with calves, perhaps signifying, again, greater catabolism of body
tissues in lactating animals. Since lactation ceases in October or
November, however, this year-round elevation is difficult to explain
fully.
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Table 14. Selected blood values in pregnant and non-pregnant adult cow moose, Alaska. (Data of LeResche).
October-January February-AEri 1
Pregnant Not Pregnant Pregnant Not Pregnant
X ± S.D. X± S.D. X± S.D. 1 n X ~ S.D. n n n p
Calcium (mg/lOOml) 26 10.7 ± 0.92 44 10.7 ± 1.1 35 11.3 ± 1.8 42 11.0 ± 0.7 0
BUN (mg/100 ml) 27 5.8 ± 3.7 43 7.8 ± 5.3 35 6.1 ± 4.1 42 5.9 ± 5.1 0
Glucose (mg/lOOml) 27 134.1 ± 92.2 47 107.9 ± 59.5 36 111.4 ± 55.8 43 104.3 ± 42.6 1,4
Bilirubin (mg/lOOml) 27 0.35 ± 0.08 47 0.45 ± 0.35 36 0.35 .._ 0.09 43 0.38 ± 0.19 0
Cholesterol (mg/lOOml) 27 75.7 ± 16.9 47 90.4 ± 13.5 36 82.8 ± 10.6 43 85.2 ± 12.9 1
SGOT (mU/ml) 27 193.2 ± 55.6 46 200.0 ± 46.7 36 224.3 ± 38.8 43 153.1 ± 59.0 3
\J1
\0 Alkaline phosphatase {mU/ml) 27 50.9 ± 26.4 47 49.1 ± 48.1 36 90.7 ± 47.8 43 54.2 ± 45.9 3
LDH (mU/ml) 27 369.3 ± 151.0 47 362.3 ± 112.0 36 413.9 ± 143.7 43 331.9 ± 115.5 3
Total protein (g/lOOml) 12 6.3 ± 0.8 41 6.8 ± 0.7 No data 35 6.8 ± 0.5 0
Albumin (g/lOOml) 12 3.7 ± 0.6 41 3.9 ± 0.6 No data 35 3.6 ± 0.5 0
A/G (ratio) 12 1.49 ± 0.3 41 1.36 ± 0.3 No data 35 1.12 ± 0.3 0
1values for pregnant moose were compared with those for non-pregnant moose during each of the two time periods using the "t" test
for unpaired variates. Probabilities of differences were:
0 -No statistical difference during either time period. (P>.05)
1 -Pregnant vs. non-pregnant difference significant (P<.OOl} during October-January.
2 -Pregnant vs. non-pregnant difference significant (P <. 05) during October-January.
3 -Pregnant vs. non-pregnant difference significant (P<.OOl) during February-April.
4 -Pregnant vs. non -pregnant difference significant (P<.05) during February-April.
Table 15. Selected blood values of adult cow moose leading and not leading calves (year-round values, Alaska).
Data of LeResche.
With No Calf With One Calf With Two Calves
X± S.D. pl X ± S.D. pl X ± S.D. 1 n n n p
Calcium (mg/lOOml) 125 10.5 ± 0.7 50 10.9 ± 0.8 5 10.3 ± 0.9
BUN (mg/lOOml) 129 12.8 ± 11.5 a 49 20.5 ± 12.7 a 5 36.0 ± 33.1 a
Glucose (mg/lOOml) 130 122.6 ± 39.7 51 120.8 ± 38.2 5 114.4 ± 63.8
Bilirubin (mg/lOOml) 130 0.45 ± 0.23 51 0.59 ± 0.28 5 0.82 ± 0.63
Cholesterol (mg/lOOml) 128 86.2 ± 21.6 a,b 50 101.7 ± 26.9 a,c 5 110.4 ± 15.4 b,c
0\
0
SGOT (mU/ml) 128 121.9 ± 42.0 a,b 50 147.7 ± 42.7 a 5 148.4 ± 58.2 b
Alkaline phosphatase (mU/ml) 130 85.0 ± 82.5 a 51 37.2 ± 15.2 a,b 5 86.0 ± 135.9 b
LDH (mU/ml) 128 261.5 ± 99.5 a 50 318.8 ± 88.2 a 5 396.2 ± 121.9 a
Total protein (g/lOOml) 123 7.1 ± 0.7 45 7.6 ± 0.9 5 7.6 ± 0.7
Albumin (g/lOOml) 123 4.1 ± 0.6 46 4.3 ± 0.9 5 4.0 ± 0.2
A/G (ratio) 123 1.41 ± 0.43 46 1.40 ± 0.42 5 1.08 ± 0.11
1 for cows with no calf, the 11 t" test for Values one calf and two calves were compared with one another using
unpaired variates. Probabilities of differences were:
a, b: different at P<.OOl
c: different at P<.Ol
-------------~--~--
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PRESENT APPLICATIONS -NEED FOR FUTURE RESEARCH
It is abundantly clear that the full potential uses of blood
analyses have not been reached. Stress of handling remains a problem,
but results from Alaska indicate that sometimes trends might override
these variables if methods are standardized. The problem of handling
stress is insoluble in wild animals using presently available techniques,
but may be minimized using tame moose and remotely-activated devices
(cf: Farrell et al. 1970). Such devices might make comparisons between
studies feasible in the future.
Seasonal patterns of BUN, glucose, cholesterol, proteins, and enzymes
suggest that these values may be useful as indicators of nutritional
status. Relationships need experimental confirmation, but eventually
such indicators will allow discrimination between ranges of different
qualities. Data are currently being gathered from different ranges in
both Alaska and Minnesota.
The preliminary results of thyroxine assays reported here are very
promising, and endocrine studies should be further pursued. We have
shown that thyroxine levels vary seasonally, but do not know whether
this variation is greater on lower-quality ranges, nor what the relation-
ships between pregnancy, fetal health, and thyroid function are. Studies
with animals at high and low nutritional planes, in either experimental
or natural conditions, are indicated.
The only reported studies on cortisol metabolism are in the rein-
deer by Yousef et al. (1971). TI1ey conclude that winter acclimatization
is accompanied by a decrease in half life and an increase in estimated
secretion rate. No comparable data exist for other cervids. Confirma-
tion of these results would be desirable since their method for measure-
ment of cortisol can be influenced by increases in ketone bodies which
may occur in the winter metabolic state. Simple measurement of serum
cortisol is almost certainly of little value since the levels of binding
proteins are low, the turnover is rapid and levels can fluctuate rapidly.
Measurements of other plasma hormone levels are not presently avail-
able. Seal's group do have data on insulin, glucagon, and TSH levels in
the white-tailed deer which will be published shortly.
An area as yet barely touched is fat metabolism. Cholesterol assays
show some promise, while triglycerides probably do not. The levels of
both are low in cervids. Direct measurements of long-chain fatty acids,
ketone bodies, pyruvate, lactate, acetate, and proprionate, especially
in conjunction with rumen VFA studies, will probably provide a more
direct assessment of the animal's dependence upon its fat reserves for
energy.
Protein assays performed to date have been prom~s~ng, but too crude
to indicate precise metabolic relationships. Measurements of serum and
red blood cell amino acids -especially alanine, glycine, leucine, and
isoleucine -might provide valuable information on the status of protein
used for gluconeogenesis. Creatine-hydroxyproline relationships have
61
yet to be investigated in moose. Albumin-globulin relationships have
successfully detected severe liver parasitism in Minnesota, and these
assays will continue to have such applications.
Serum enzyme patterns have suggested seasonal changes in tissue
destruction rates. Further studies might employ enzymes of higher
tissue specificity in order to explain these patterns more fully.
Thus, the data currently available and the studies in progress with
cervids hold considerable promise of providing the concepts and tools
for evaluating condition of wild cervids and the condition of their
range relative to their needs. It also appears possible to begin unravel-
ing influences of such factors as density and social stress, separately
from nutrition, upon a given population in terms of physiological response.
The ability to quantitatively describe metabolic status and its endocrine
regulation has increased dramatically in the past five years as the
result of new exquisitely sensitive radio-immunoassay techniques. As a
result, we may soon achieve the precise formulations of these mechanisms
that we need in order to measure ecological and behavioral relationships
of individuals and populations.
ACKNOWLEDGMENTS
Work reported here was supported in part by Federal Aid in Wildlife
Restoration projects W-17-3 (Alaska) and W-117-R (Minnesota) and by USPHS
grant 5 ROL AM-11376 from the National Institutes of Health, NIAMDD. We
are grateful to Dr. J. Peek for his help and review. L. McManus and L.
Ruelle assisted in manuscript preparation.
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on rabbit leukocyte protein, aldolase, and lactate dehydrogenases.
Biochem. Med. 4:336-346.
Marple, D. N., E. D. Aberle, J. C. Forrest, W. H. Blake and M. D. Judge.
1972. Endocrine responses of stress susceptible and stress resis-
tant swine to environmental stressors. J. Anim. Sci. 35:576-579.
Panaretto, B. A. and K. A. Ferguson. 1969. Comparison of the effects
of several stressing agents on the adrenal glands of normal and
hypophysectomized sheep. Aust. J. Agric. Res. 20:115-124.
Rose, L. I., J. E. Bousser and K. H. Cooper. 1970. Serum enzymes after
marathon running. J. Appl. Physiol. 29(3):355-357.
Seal, U. S., J. J, Ozoga, A. W. Erickson and L. J. Verme. 1972. Effects
of immobilization on blood analyses of white-tailed deer. J. Wildl.
Manage. 34:1034-1040.
Todd, J. R. and J. G. Ross. 1968. Biochemical and haematological changes
in the blood of normal sheep following repeated phlebotomy. Br.
Vet. J. 124:353-362.
Wardle, C. S; 1972. The changes in blood glucose in Pleuronectes
platensa following capture from the wild: A stress reaction. J.
Mar. Biol. Assoc. U. K. 52:635-651.
Weltman, A. S., A.M. Sachler and S. B. Sparber. 1966. Edocrine, meta-
bolic and behavioral aspects of isolation stress on female Albino
mice. Aerospace Medicine 37:804-810.
Wilson, W. 0. 1971. Evaluation of stressor agents in domestic animals.
J. Anim. Sci. 32(3):578-583.
74
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H. Analytic Problems
Baer, D. M. and R. B. Krause. 1968. Spurious laboratory values result-
ing from simulated mailing conditions. Am. J. Clin. Pathol. 50:111.
Brittin, G. M., G. Brecher, C. A. Johnson and R. M. Elashoff. 1969.
Stability of blood in commonly used anticoagulants. Am. J. Clin.
Pathol. 52:690.
. , G. Brecher and C. A. Johnson. 1969. Evaluation of the
~----::---::--~
Coulter counter modelS. Am. J. Clin. Pathol. 42:679.
Chien, S., R. J. Dellenback, S. Usami and M. I. Gregersen. 1965. Plasma
trapping in hematocrit determination. Differences among animal
species. Proc. Soc. Exper. Biol. Med. 119:1155.
Cotlove, E., E. K. Harris and G. Z. Williams. 1970. Biological and
analytic components of variation in long-term studies of serum
constituents in normal subjects. III. Physiological and medical
implications. Clin. Chern. 16:1028.
Davies, D. F. 1968. Effects of freezing and thawing serum and plasma
on selected quantitative recoveries. Cryobiology 5:87.
Gibbs, E. L. 1972.
mance part B:
Glass laboratory water stills: Design and perfor-
Performance. In Vitro 8:37.
Gochman, N. and J. M. Schmitz. 1971. Automated determination of uric
acid, with use of a uricase--peroxidase system. Clin. Chern. 17:
1154-1159.
Spate, M. P., M. F. Burks, P. S. Evans and M. E. Tumbleson. 1970. Con-
centrations and activities of bovine serum biochemic consitituents
as a function of storage time and temperature. Clin. Biochem.
3:137.
Hanak, A. and J. Kuo. 1968. The stability of a reconstituted serum for
the assay of fifteen chemical constituents. Clin. Chern. 14:58.
Talstad, I. 1971. Analysis of errors in electronic blood cell counting.
Acta Med. Scand. 190:1.
Tumbleson, M. E. 1969. Modification of the sequential multiple auto-
analyzer (SMA-13/30) for use in animal research studies. Cli.n.
Biochem. 2:35 7.
Williams, G. Z., D. S. Young, M. R. Stein and E. Cotlove. 1970. Biolog-
ical and analytic components of variation in long-term studies of
serum constituents in normal subjects. I. Objectives, subject
selection, laboratory procedures, and estimation of analytic devia-
tion. Clin. Chern. 16:1016.
75
I. Computer Aids
Furlong, N. B. 1972. Computer programs in the biomedical sciences. II.
LABCAL: A computer program for general laboratory calculations.
Tex. Rep. Biol. Med. 30:109.
Gledhill, V. X. and J. D. Mathews. 1970. Computer aided diagnosis: A
learning model. J. Med. 1:249.
Hobbie, R. K. and R. L. Reece. 1972. The use of a computer to suggest
diagnostic possibilities from a battery of blood chemistry values.
Proc. 1972 San Diego Biomed. Symp., pp. 1-9.
Levy, S. W. 19 72. The design and operation of a small computer sys tern
for the clinical chemistry laboratory. Clin. Biochem. 5:146.
Lewi, P. J., D. G. VanRiel and J. Symoens. 1972. The use of computers
in processing data from veterinary field trials. Life Sci. (II)
11:137.
Pemberton, J. R. and P. W. Woodward. 1972. A data handling system for
automated analysis using a programmable calculator. Comp. Biomed.
Res. 5:59.
Reece, R. L. and R. K. Hobbie. 1972. Letters, notes and computers.
The evolution of a laboratory information system. Presented before
Conn. Soc. Pathol., 1/19/72, New Haven, Conn.
values:
57:664.
and 1972. Computer evaluation of chemistry
A reporting and diagnostic aid. Am. J. Clin. Pathol.
Riddick, J. H., Jr., and N. W. Giddings. 1971. Computerized preparation
of average cusum charts for clinical chemistry. Clin. Biochem. 4:156.
--------------~---·• R. Flora and Q. L. VanMeter. 1972. Computerized
preparation of two-way analysis of variance control charts for
clinical chemistry. Clin. Chern. 18:250.
Rowlands, G. J. and R. M. Pocock. 1971. A use of the computer as an
aid in diagnosis of metabolic problems of dairy herds. J. Dairy
Res. 38:353.
VanGelder, G. A. and N. E. Hutton. 1972. Selection and use of labora-
tory computers in veterinary medical research. Am. J. Vet. Res.
33:217.
Warner, H. R., C. M. Olmsted and B. D. Rutherford. 1972. HELP - A
program for medical decision-making. Camp. Biomed. Res. 5:65.
76
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Stress Evaluation
Stress evaluation began in May and data are insufficient for inclu-
sion at this time. Excitability-related parameters were measured from
27 immobilized moose in May and June. Rectal temperatures ranged from
101.1 °F to 108.0°F; heart rates from 52 to 140 beats per minute; and
respiratory rates from 20-88 respirations per minute.
Hair Analyses as Indicators of Nutritional Status
No results are available at this time.
Weights and Measurements
Table 10 lists weights of moose obtained during this reporting
period, additional weights were reported previously by LeResche (1970)
and LeResche and Davis (1971). These data are summarized in Fig. 3.
All available weight and morphometry data from North America are
currently being analyzed for publication in 1974.
Nutritional Analyses of Plants, Rumina, and Fecal Pellets
Results of these analyses are presented in Tables 11-13.
Table 11, although based on few samples from a single site at one
season, suggests the importance of variety to moose. This principle is
discussed at length by LeResche and Davis (1973) and below (Food Habits
of Tame Moose).
Chemical and proximate analyses of rumina of moose from lowland
seral birch ranges and upland willow ranges show no significant differ-
ences between the two groups (Table 12), suggesting that the analyses
performed were inappropriate to detect such differences. Volatile fatty
acid (VFA) production rates might better reflect such nutritional differ-
ences (Gas at-ray and Coady, 19 73).
Table 13 suggests that fresh fecal pellets might prove valuable in
estimating relative levels of nutrients in rumina of animals from differ-
ent areas. Correlations between protein and ether extract in rumina and
pellets were highly significant (P<.OOl), and the correlation of caloric
contents was significant at P<.03>.02.
Snow Monitoring
Snow studies initiated in late winter 1971 (LeResche and Davis,
1971) were continued in the winter of 1971-1972, and snow depths are
listed in Table 14. Snow conditions during 1971-1972 were more severe
than those during the previous winter. A maximum snow depth of 67 em
was recorded, and depths of 50 em or greater were present for three
consecutive months in five of the eight snm.,r plots.
Although 6 7 em of snow is not a great amount v1hen compared with
other parts of Alaska where moose are abundant, it is sufficient to
77
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Table 10. Weights of moose handled at Moose Research Center traps, July 1971 -I June 1972.
Weight I
Date Moose II Pen* Sex Age Remarks Kg lbs
I
15 July 1971 70-2 2 F 4 No calf 298 655
15 July 1971 53(70) 1 M 1 211 465 I 15 July 1971 88 0 F 9 Lactating 357 785
23 July 1971 Walter 2 H 2 Tame 291 640
27 July 1971 5271 2 M calf Died in winter 70 155
27 July 1971 2771 3 F calf Died in winter 70 155 I 27 July 1971 5470 2 M 1 Died in winter 161 355
10 Aug. 1971 43 1 M 4 473 1040
10 Aug. 1971 A60-71 4 H calf Died in winter 89 195 I 10 Aug. 1971 89 0 F 4 Lactating 400 880
10 Aug. 1971 A60 4 F 14 With calf 341 750
10 Aug. 1971 55 1 M 1 228 500 I 11 Aug. 1971 5670 4 M 1 Died in winter 248 545
11 Aug. 1971 70371 4 F calf Died in winter 95 210
11 Aug. 1971 R 70-3 4 F 4 With calf 309 680
11 Aug. 1971 10 1 F 4 Lactating 289 635 I 12 Aug. 1971 1-71 2 F calf Died in winter 84 185
12 Aug. 1971 90-A 0 F 8 With calf 434 955
12 Aug. 1971 90-71 0 F calf 118 260 I 24 Aug. 1971 6 1 F 14+ With calf, died in winter 359 790
12 Aug. 1971 28 3 F 9 No calf 345 760
25 Aug. 1971 R 70-1 4 F 9 With calf 291 640
25 Aug. 1971 R 701-71 4 M calf Died in winter 84 185 I 31 Aug. 1971 3 1 F 9 With calf 359 790
31 Aug. 1971 371 1 F calf Died in winter 118 260
1 Sept. 1971 9171 0 M calf 148 325 I 2 Sept. 1971 61 1 F 9 Lactating 355 780
14 Sept. 1971 93 0 M 3 364 800
19 Sept. 1971 9571 0 F calf 175 385 I 21 Sept. 1971 9671 4 F calf Died in winter 159 350
22 Sept. 1971 9771 4 F calf Died in winter 145 320
29 Sept. 1971 9871 4 F calf Died in winter 182 400
5 Oct. 1971 100 0 F 13 In estrus 430 945 I 7 Oct. 1971 101 0 F 5 In estrus 345 760
7 Oct. 1971 102 0 F 13 In estrus 400 880
8 Oct. 1971 103 0 H 1 268 590 I 11 Oct. 1971 104 0 F 1 284 625
12 Oct. 1971 106 0 F 7 With calf 395 870
12 Oct. 1971 10671 4 M calf Died in winter 159 350 I 12 Oct. 1971 10771 4 F calf Died in winter 186 410
13 Oct. 1971 10871 4 F calf Died in winter 175 385
20 Oct. 1971 10971 4 F calf Died in winter 180 395
20 Oct. 1.971 109 0 F 9 With calf 420 925 I 21 Oct. 1971 11071 4 F calf Died in winter 175 385
27 Oct. 1971 112 0 F 2 291 640
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78 I
------------·-----· I
I Table 10 (cont.)
I Weight
Date Moose If Pen* Sex Age Remarks Kg lbs
I 3 Nov. 1971 11571 4 F calf Died in winter 200 440
2 Nov. 1971 11471 4 M calf Died in winter 168 370
I 4 Nov. 1971 116 0 F 11 261 575
4 Nov. 1971 117 0 F 2 466 1025
17 Nov. 1971 119 0 F 3 311 685
I 23 Nov. 1971 12071 4 M calf Died in winter 211 465
1 Dec. 1971 123 4 F 4 No calf in 1972 348 765
15 Dec. 1971 126 4 F 10 359 790
15 Dec. 1971 125 0 F With calf 373 820
I 27 Jan. 1972 62 2 F 19 386 850
22 Mar. 1972 134 4 M calf Died in few days 164 360
2 Apr. 1972 66 2 F 10 Died in trap. fetus 327 720
I 2 Apr. 1972 64 1 M 1 282 620
25 Apr. 1972 138 0 F 3 Pregnant 355 780
26 Apr. 1972 132 0 F 318 700
28 Apr. 1972 Wally Jr. 2 M calf Supplementally fed 214 470
I 24 June 1972 Wally Jr. 2 M Yrlg. Supplementally fed 234 515
5 June 1972 73 2 M 3 239 525
22 June 1972 1 2 F 8 No calf 318 700
I 23 June 1972 77 2 F 6 No calf 275 605
23 June 1972 39 3 F 7 243 535
24 June 1972 Rachel 2 F 3 No calf 352 775
I 24 June 1972 3 1 F 9 No calf 295 650
28 June 1972 72A 3 F 2 No calf 225 495
1 Aug. 1972 45 2 M 3 384 845
I * Pen "0" indicates moose trapped outside pens.
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co
0
500-------------------------------------------------------------7
(f) 400
E
0
'-
0"
0
c
300
200
•• 0
l 0
0 ~
/ \.; ~~ r',, /'~ .. \ ~./' \ .. / • /• ' o~'\ ' ' , \I ,_, \
e-_ I 1 'I ---o...._ v '
! "-~ I 0 o------u ~\ • : 0 0
6Y' r o \ .. \ '
JL· 'i
0
• 0
•
Age in Months
0
0
0
\
\
\
0\
\
\
0\
0
-• Males N= 18
0
0
---o Females N=63
0
226
Fig. 3. Weights of 81 moose of known age from the northern Kenai Peninsula. Thick solid lines show
seasonal mean weights of males to 48 months age. Dotted lines show seasonal mean weights
of females to 132 months age. Thin solid lines connect sequential weights of individual
animals at various ages, and reflect seasonal fluctuations in weight.
-------------------
- - --- - -- - -- - ---- - - -
Table ll. Nutritional values of plants collected in ::-lay 1971.
Anal-Pro-Ether Carbohy-Digestible1
yses tein Extr. Ash drates Fiber dry matter p Ca K Mg Na Al Ba Fe Sr B Cu Zu Mn Cr
n % % % % % X ~~ % % % % ppm ppm ppm ppm ppm ppm ppm ppm ppm
:c. pap~j~:f"era
Stems 8 8.8 14.6 l.8 49.7 30.8 36.9 0.23 0.38 0.34 0.17 0.027 31 93 78 25 16 9 141 283 <l.S
P. tPe.:rrrLA. l-:;'"Z des
Stems 8 9.1 4.9 2.3 54.3 26.0 39.5 0.22 0.66 0.61 0.15 0.015 16 63 46 53 17 8 80 69 <l.5
Bark 3 3.3 3.1 8.6 59.2 59.2 48.6 0.03 l. 30 0.35 0.18 0.024 36 109 32 109 17 5 152 80 <l. 5
"" >-' -.~ ~)1.-t-I.-s -:. d.aea
'..Thole plants 3 5.4 2.1 2.1 67.0 17.6 50.1 0.09 0.63 0.34 0.17 0.008 148 57 100 10 17 4 14 828 <3.0
Ca.rex CL:;(U2tiZ.is
\·.'hole plants 1 5.4 2.2 3.6 53.0 31.6 0.10 0.15 0.92 0.10 0.010 47 51 547 15 3 2 26 >800 <3
AZ1--::..tB :.::ris;;.::x 3 8.6 1.4 1.3 54.1 24.1 32.5 0.16 0.45 0. 33 0.14 0.010 28 57 56 34 12 7.2 28 98 1.5
~ed'0~!Y.. pc -~ :..:s ::::ne 3 7.6 8.5 2.0 56.0 22.9 43.1 0.12 0.66 0. 38 0.17 0.018 180 81 197 11 26 7.4 26 713 3.0
1 Bovine flora
Table 12. Analyses of rumen contents of moose shot on the Kenai Peninsula
(Units 7 and lSA), November-December 1971.
x sera1 birch x willow
range range
(n=6) (n=8)
Protein (%) 6.1 6.6
Ash (%) 1.5 1.5
Ether Extr. (%) 5.0 2.5
Fiber (%) 43.9 43.9
Carbohydrates (%) 43.6 45.5
Ca1ories/100 g 245 230
DDM (%) 16.3 15.2
p (%) 0.16 0.16
K (%) 0.14 0.14
Ca (%) 0.38 0.49
Mg (%) 0.070 0.070
Na (%) 0.093 0.092
A1 ppm 22 15
Ba ppm 57 53
Fe ppm 225 198
Sr ppm 27 45
B ppm 13 13
Cu ppm 6.9 8.4
Zu ppm 107 118
Mn ppm 233 163
Cr ppm 1.5 1.5±
82
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Table 13. Correlation of nutrient content of fecal pellets with that of
rumens collected from the same animal. November -December
1971.
(n) r p
Protein (5) o. 8835 <.001
Ash (5) 0.3093 >.10
Fat (ether extr.) (5) 0.9828 <.001
Crude fiber (5) 0.4200 >.10
Carbohydrates (5) 0.4200 >.10
Calories (5) 0. 7128 .03> .02
Phosphorus (5) -0.2300 >.10
Potassium (5) 0.2703 >.10
83
Table 14. Snow depth (em) in each of eight habitat types, Kenai Moose Research Center, winter 1971-1972.
November December J anuar;y Februar;y March AEril May
5 12 19 24 3 9 16 23 5 19 25 2 8 20 28 7 14 20 28 5 11 19 27 7
Mature
Hardwoods
(dense) 11 11 9 11 15 19 29 28 33 32 32 47 44 42 40 39 44 42 38 46 44 43 36 Tr.
Mature
Hardwoods
(thin) 13 15 13 15 19 23 33 33 36 32 34 52 51 50 50 50 56 57 51 58 55 56 42 Tr.
Sedge 13 18 12 16 20 25 37 34 44 29 27 43 42 41 38 39 43 44 38 43 44 46 33 Tr.
()) Spruce +--
regrowth 14 18 13 15 20 25 39 36 46 46 45 58 56 57 55 57 60 58 54 62 61 60 47 Tr.
Birch-Spruce
(thin) 17 16 13 15 21 24 37 32 43 43 43 62 58 58 55 51 56 55 51 58 57 56 47 Tr.
Birch-Spruce
(dense) 13 15 11 13 17 24 33 29 43 46 43 55 56 56 50 51 55 53 47 53 52 57 39 Tr.
Spruce-1edum * 9 * 6 10 13 15 19 46 39 39 57 55 57 53 56 61 60 56 58 61 67 51 Tr.
Mature
Spruce 8 8 5 7 9 13 23 23 26 24 24 36 34 34 34 33 34 36 30 36 33 30 26 Tr.
*Not checked.
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cover some important forage plants, including lowbush cranberry (Vaccinium
m:tis-?:daea). Lowbush cranberry may be an important sustaining winter
browse plant in this area (LeResche and Davis, 1973) and its nonavail-
ability could adversely affect wintering moose. Because of the greater
snow depths this past winter Vaccinium was visible for a much shorter
period than during the previous winter. It was no longer visible in
December and remained snow-covered until April or May except in the
dense birch habitat type where it was visible in late March under the
bases of spruce trees. The extensive covering of Vaccinium by snow may
cause greater crater digging activity by moose (see Feeding Craters
section, this report).
Browse Production and Utilization
This project segment returned fully to the Bureau of Sport Fisheries
and Wildlife in May 1971, when John L. Oldemeyer began work. He has
reported his findings in Annual Progress Reports (Oldemeyer, 19 72, 197 3a)
and a review publication (Oldemeyer, 1973b). An abstract of Oldemeyer's
work during this reporting period appears below:
The Kenai Field Station of the Denver Wildlife Research Center
was established in May 1971 with the primary objective to study
moose habitat on the Kenai National Hoose Range particularly
at the Moose Research Center.
A brief history of the Hoose Research Center and moose and
habitat research that has taken place up to this time is
given.
Twenty sites in each of four one-square-mile pens were chosen
for intensive research. All sites in Pens 3 and 4 were
sampled for browse density and frequency of occurrence and
for birch production. Summarized data for the density and
frequency sample are presented; however only preliminary
production analysis has been started.
Several aspects of moose habitat were reviewed in depth in a publi-
cation prepared during this period (LeResche et al., 1973a). Conclusions
relative to the sera! habitat of the Hoose Research Center are reproduced
below:
Fire-mediated habitats are the most ephemeral of Alaskan moose habi-
tats; yet, they support the greatest population explosions and among the
greatest densities of moose in the state. The earliest specific publica-
tions concerning moose management in Alaska (Spencer and Chatelain, 1953;
Leopold and Darling, 1953a and b; Chatelain, 1951, 1952) correctly
stressed the correlation between dense moose populations and forest fires.
The ecological effects of fires in Alaska are extremely complex (cf.
Lutz, 1956; Slaughter et al., 1971; Viereck, in press) and cannot be
considered in detail here. Lutz (1956) presented a diagrammatic repre-
sentation of normal courses of post-fire succession in Alaskan taiga.
Viereck (in press) modified the chart and we reproduce his chart here
85
. ···-··----------------------------
(Fig. 5). The course of succession and the sera! community's value to
moose are determined by a multitude of site (soils, temperature, moisture,
slope and exposure), fire (severity, mode of travel, duration) and mis-
cellaneous (natural reseeding sources, size of openings created) factors.
The general successional courses producing maximum benefits to moose
populations are those involving paper birch-willow-aspen shrub thickets.
Fires have probably always been common in Alaska, but have increased
in frequency and area burned since modern settlement (Hardy and Franks,
1963). Viereck (in press) maintains there are very few Interior forests
that survive burning long enough to reach what can be considered climax
stage. Barney (1971) suggests that 600,000 to 1,000,000 ha burned
annually from 1900-1940. Even with modern fire control, a mean of
approximately 400,000 ha annually have burned between 1940-1969 (Barney,
1971). The general trend in the past three decades has been toward an
increase in numbers of fires (1,138 in the 1940's, 2,583 in the 1950's
and 2,380 in the 1960's in Interior Alaska) but a decrease in the total
area burned (5,018,000 ha in the 1940's, 4,330,000 ha in the 1950's,
2,590,000 ha in the 1960's) and the area burned per fire. Increased
fire control capabilities have thus perhaps decreased the benefits of
civilization to moose.
~haracteristics -Several factors determine the impact a fire-created
successional community will have on moose populations: (1) species com-
position, (2) size of burn and per-area standing biomass and production
of available browse, (3) rate of attaining a) above snow and b) above
moose-reach heights and (4) amount and distribution of mature communities
remaining, diversity of seral communities created and distribution of
mature communities remaining (e. g.: "edge effect" created).
Composition -The species re-invading burned areas in Alaska most
often tend to be the same species present before the fire (Fig. 5).
This results from re-sprouting or growth of root suckers (Spencer and
Hakala, 1964; Viereck, in press). Stands are often nearly monotypic in
shrub species (cf. Table 15), with birch or aspen (on the warmest, driest
sites) predominating. Spruce encroachment usually occurs slowly (Viereck,
in press) in the Interior, but sometimes almost immediately (e.g.: on
some stands on the Kenai Peninsula). Although willows are often among
the first recolonizers, they generally do not achieve great numbers or
production, perhaps because of consistent utilization by moose. Aspen
may be similarly affected. Spencer and Hakala (1964) described how
moose and plant competition had altered the proportion of aspen in the
Kenai (1947) burn. It apparently decreased from 96.5 percent of the
standing shrub biomass in 1952 (Spencer and Chatelain, 1953) to less
than 1 percent in 1967 (Table 15). Wet lowland and permafrost sites are
usually less thoroughly burned (Viereck, in press) and thus may rapidly
recover by vegetative reproduction to a composition similar to that
existing before the fire. Willows, blueberry (Vacciniwn sp.) and dwarf
birch (Betula nana) tend to be more numerous in these stands.
The
upon the
press).
exact composition of species recolonizing a burn may depend
timing of the fire relative to plant phenology (Viereck, in
Willow species produce seeds that are viable for only a matter
86
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-------------------
:::; FIRE
POPLAR BIRCH
BLACK
SPRUCE
DRY-WARM---------------------------------------WET-COLD
FIRE
Fig. 5. Diagrammatic representation of normal post-fire succession in Alaskan taiga (from Viereck, in press.)
Table 15. Characteristics of shrub standing crop and production on Kenai
burn 20 years after the 1947 fire. Modified from Seemel 1969.
Annual
Stems/ Production
Species Community hectare g/hectare
Betula papyrifera Dense Birch 80,046 479,348
II II Medium Birch 47,221 315,987
II II Thin Birch 34,381 249 '316
II II Spruce-birch 19,296 118,118
II II Spruce Regrowth 6, 775 35,226
Populus tremuloides Dense Birch 237 202
II II Medium Birch 339 348
II II Thin Birch 541 8"75
II II Spruce-birch 200 224
" II Spruce Regrowth 168 224
Viburnwn edule Dense Birch 114 101
II II Spruce-birch 89 90
Salix spp. Dense Birch 1,641 9,233
" II Medium Birch 1.196 6,193
II " Thin Birch 692 3,747
II " Spruce-birch 469 2,255
II II Spruce Regrowth 652 2,109
Alnus crisp a Thin Birch 319 718
Betula nana Spruce Regrowth 8, 772 9,008
Totals: Dense Birch 82,038 488,884
Medium Birch 48,756 322,528
Thin Birch 35 '933 254,656
Spruce-hi rch 20,054 120,687
Spruce Regrowth 16,367 46,567
1::18
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of weeks, and various species produce seeds at different times of year.
Further, some species (paper birch, white spruce) produce "bumper" seed
crops only once in several years (Viereck, in press), and the year the
burn occurs might thus affect species composition of the resultin~ seral
community.
There is little detailed information directly relating species com-
position of burn regrowth to subsequent moose population densities.
However, observations suggest that the higher the proportion of first,
willow, and second, birch shrubs produced, the greater the moose densities
that will result during early years of succession.
Production -Biomass production of shrubs can be prodigious follow-
ing burns in Alaska. Seemel's (1969) studies 21 years after the Kenai
(1947) burn indicate annual production of nearly 500 kg/ha in the densest
shrub B. papyrifera stands (Table 15). Standing crop is also large,
with 82,000 shrub stems/ha estimated in the most dense stands. Both
production and standing crop, as well as composition, vary considerably
with individual stand (cf.: Table 15), and thus the proportions of
various communities and variously-productive stands in a burn also
influence moose numbers.
Productivity and quantity of available browse are seldom limiting
factors to moose densities in large burns in Alaska. In the Kenai (1947)
burn of 12 7,600 ha, Seemel (1969) estimated that more than eight moose/
km2 in a 2.6 km2 enclosure used only 18.3 -22.8 percent of the estimated
annual shrub production, depending upon community. Spencer and Chatelain
(195 3) in contrast, had estimated that utilization by only 1. 7 moose per
km 2 on the same burn in 1952 was 45 percent of the annual production of
aspen, then virtually the only shrub present. Since then plant produc-
tion has exceeded moose production in relative terms, even though moose
densities achieved the highest level ever recorded. Spencer and Hakala
(1964) suggested that adequate densities of moose might retard succession
in seral shrub habitats under certain conditions. We feel that these
densities are seldom reached and never persist. Although moose may
alter relative abundance of species in seral communities, we know of no
instance where they have retarded succession over a large burned area.
In the case of the Kenai burn, moose populations appeared to peak at the
very high level of 4-6 moose/km2 in the most productive habitat, at
about the same time Seemel made his estimates. Within four 2.6 km 2
enclosures, populations remained relatively static at 4.6 -5.4 moose/km2
(before calving) over a three-year period, and utilization probably did
not exceed Seemel's estimate (LeResche and Davis, 1971).
Isolated smaller burns serving as winter concentration areas for
moose are sometimes retarded in succession and, on occasion, "over-
utilized" by immigrant moose. Succession on the 1320 ha Kenai Lake burn
(1959) has been effectively retarded by wintering moose, with most shrubs
being browsed to snowline annually. This burn is unusual in that it is
situated in the midst of nearly-mature range that wintered many moose
until the late 1950's, and is about 20 km from the nearest edge of the
larger Kenai burn (1947). Because fires are frequent throughout most of
Alaska, small burns are often near other burns of various ap,es, and are
89
therefore not subjected to intensive use by large concentrations of moose
from a vast area. Furthermore, small isolated burns do not produce wide-
spread moose population explosions sufficient to alter succession. Thus,
in most cases, post-burn succession in large burned areas seems to pro-
ceed at a rate little different from what it would in the absence of
moose. The course of succession may well be altered, however.
Rate of Growth -The sooner recolonizing shrubs achieve a height
that makes them available to moose during winter, the sooner moose popu-
lation explosions occur. Conversely, the more rapidly shrubs mature
into trees, the shorter is the productive life of the burn. Spencer and
Chatelain (1953) estimated that the Kenai burn (1947) produced signifi-
cant browse by 1950, and that by 1953 moose had increased four-fold.
Moose densities peaked on the burn about 1967-1969, and may have
experienced the first stages of a decline in the winter of 1971-1972;
however, they remain very high today.
Rates of growth differ between burns and within large burns, depend-
ing upon species present and site and climatological variables. Within
the Kenai (1947) burn, heights of B. papyrifera plants in adjacent stands
ranged from 1 m to almost 7 min 1970. We do not know whether some sites
were recolonized by paper birch later than others or whether growth rates
alone account for the differences.
Hakala et al. (1971) predicted that browse would "continue to
improve" on areas burned on the Kenai Peninsula (1969) for 23-25 years
after the burn, as has been the case with the 1947 Kenai burn area.
Moose densities could be expected to remain high for this period, and
then decline as happened after a series of fires in the 1880's and 1920's
(Spencer and Hakala, 1964). The limited information available, then,
suggests that the "useful life" of a bum as moose habitat in Alaska is
usually less than 50 years, and that moose densities peak 20-25 years
after the burn.
Shape of Burn, Diversity and "Edge Effect" -The degree of inter-
spersion of communities, or amount of "edge effect", produced by a fire
is very important in determining the fire's effect on moose populations.
Great discontinuity of burning is desirable because it provides, (1)
cover (mature) habitat close to feeding habitat, (2) increased variety
of alternate forage species and (3) staggered maturation rates of
individual stands.
The forest edge ecotone allows moose both to invade a heterogeneous
burn sooner than a large homogeneous burn and to achieve higher year-
round densities. The distance a moose will continually move from cover
into open feeding areas likely varies with season, age, sex, reproductive
status, snow characteristics and a multitude of other factors. It
appears certain, however, that increased mature forest edge will hasten
establishment of moose in a new bum. The rate of moose increase on the
Kenai (194 7) burn, described by Spencer and Chatelain (1953) as 400 per-
cent in five years, was probably near the maximum attainable because of
the tremendous amount of edge created. It is problematical whether
immigration from nearby areas had a significant effect (LeResche, 1973).
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Once the moose population is established, small stands of mature
timber (30-40 m tall birch, white spruce and aspen in the 194 7 Kenai
burn) provide year-round escape cover and winter refugia from deep snow.
In addition, these stands provide alternate food sources that may be of
considerable importance to the maintenance of high moose densities.
LeResche and Davis (1973) showed that Vaccinium V1:tis-1:daea plants were
important to moose in winter and remained available more consistently
in mature stands than in adjacent sera! stands on the Kenai.
Shrub-forest ecotones are not the only important edges created by
fires. Shrub-sedge and shrub-aquatic ecotones are also important at
various times of year. Brush bordering summer feeding bogs, for example,
can be excellent cover habitat for young calves (LeResche, 1966). Dense
shrubs bordering small, drying potholes are preferred habitat for adults
in late fall, when they dig craters in snow to feed on cured sedge leaves.
The optimum amount of edge and/or the optimum size and shape of
individual burned stands in Alaska is difficult to determine because
moose densities depend upon so many variables. The Kenai burn (1947),
however, appears to represent, if not the ideal, at least the most
productive lar~e area of moose habitat known to us. Densities of moose
exceeding 4/km have been achieved over most of the burn (vs densities
of ca 0.08/km2 in similar unburned areas (Spencer and Chatelain, 1953)
through a combination of high forage production, generally mild winters,
abundant alternate foods (especially V. vitis-i.daea), edge effect, and
adjoining upland ranges.
The fire burned 127,600 ha of approximately 260,000 ha, and has
regenerated into several paper birch and birch-spruce communi ties
(Table 15). The entire burn has not been analyzed, but detailed type
maps have been constructed of the reasonably-representative 1024 ha
enclosed by the Kenai Moose Research Center enclosures.
A type map of 254 ha of this mixed habitat, prepared 20 years after
the burn (Fig. 6), was analyzed for sizes of individual stands and amount
of ecotone between each of the 11 communities distinguished (Tables 16-
1}). A "stand" was defined as a contiguous area of one community,
regardless of shape, and ecotone was simply taken as the length of mapped
margin between stands.
The 254 ha contains 624 individual stands ranging in size from .02
ha to 18.4 ha (Table 16). Hean stand size is 0.41 ha, and 86 percent of
all stands are less than 0.5 ha in area; only 8 percent of the stands
are larger than 1 ha, and only 2 percent larger than 5 ha. I~emnan t
mature stands comprise 118 ha, or 46 percent of the area. This mature
forest is extremely segmented, comprising 411 stands distributed through-
out the area.
The large number of stands and their irregular shapes have produced
tremendous amounts of ecotone (Table 17) --112 km in the 2.5 km 2 area.
Mature communities have about 99 km of "edge", sharing 59 km of this
with the various seral shrub communities. Shrub communities themselves
are surrounded by about 103 km of ecotone, bordering on mature forest
and on other seral communities.
91
\0
N
Y I GlUTtON TYP'E MAP' MOOSE INCLOIUIIE N0.4
.. [NAt NATIONAL MOOS[ IIANCE. ALAIIIA ........... _,,
, ... >! .""••r ••••-•'• .-·-..... "' .....
l'i'!"'1 ........... .. ~ .. ~ ............. _ ..
~ •·· o·~~ ... h.
Fig. 6. Vegetation type map of 25~ ha of the 1947 Kenai burn.
---. -
-••c•••••• m .....
LJ u•••
0"""
--- -------- - -------
Table 16. .:;izes of 624 s :ands of ll cor:ununities in 254 ha of the Kenai, Alaska ( 194 7) burn, mapped in 196 7' Kenai ~loose Research
Center, ?en 4.
Total 1\o. Stands of Area
Number Area x area/ Range 1.1-
Co:r:rnuni ty Stands ha stand-ha ha/stand .5ha .5-l.Oha 2.0ha 2 .l-5ha 5-10ha 10ha
Dense Hature Hardwoods 22 43.07 1. 96 .05-12.9 9 5 2 3 2 l
Thin Hature Ha rdvm ods 383 70.69 0.18 .02-18.4 368 7 4 1 1 2
'·[at ure Spruce 6 4.54 0.76 .02-3.9 5 1
Spruce Regro•,nh 10 19.29 1.93 .14-11.6 4 2 2 1 1
Spruce-Birch Regrm.:th 33 17.83 0.54 .02-2.3 22 5 5 1
'-D Spruce-Ledum . ....., 6 5.39 0.90 . 23-2.5 3 1 1 1
Dense Shrub Birch 23 7.30 0. 32 .21-1.3 20 1 2
!-!edium Shrub Birch 40 41.16 1.04 . 03-8.9 24 7 4 2 3
Thin Shrub Birch 77 31.28 0.41 .02-6.2 62 7 5 2 1
Grass 19 1. 82 0.10 .02-0.4 19
Sedge 5 12.05 2.41 .07-10.2 3 1 0 1
Totals 624 254.42 0.41 .02-18.4 539 36 25 ll 8 5
f'ercent of total stands 86~~ 6% 4% 2% 1"' , 1 "/ '"
Table 17. Amount of ecotone (in ~ters) between 11 plant communi ties in 254 ha of the Kenai, Alaska (1947) burn mapped in 196 7'
Kenai ~laos e Research Center , Pen 4 . (Figure 6).
Dense Thin Spruce-Dense Medium Thin
:-:!ature Mature Mature Spruce Birch Spruce Shrub Shrub Shrub
Community Hardwoods Hardwoods Spruce Regrowth Regrowth Led urn Birch Birch Birch Grass Sedge Water
Dense ~Jature Hardvmods 17,581 325 206 356 711 1,728 2,642 51
Thin ~lature Hardwoods 305 2,541 2,795 3,150 22,764 22,967 508 178
~1ature Spruce 76 203 1,016 406 76
Spruce Regrowth 3,455 1,524 305 457 1,524 661 356
'-'> Spruce-Birch Regrowth 203 1,067 4,522 4,472 813 102 .,.. Spruce-Ledum 102 280 1,931
Dense Shrub Birch 3,303 1,016
Medium Shrub Birch 3,303 457 508
Thin Shrub Birch 457 102
Grass 152
Sedge 356
23,600 72 '789 2,407 11' 029 17,785 4,116 9, 755 38,167 37,169 3,099 3,609 508
Total Edge 224,033 D = 112.0 km
2
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This 2.5 km 2 by no means represents an extreme case, but rather is
a fair sampling of the entire 260 km2 area affected by the burn. By
extrapolation, then, the Kenai burn (1947) produced over 60,000 separate
stands, more than 11,000 km of ecotone and 128,000 ha of new shrub
communities. It is not surprising that moose densities achieved such a
high level as a result of this fire.
Moose Use -Seral burn habitats are extremely important wintering ranges
once growth is sufficient to provide available browse above the snow.
Some burns (the Kenai burn [1947], for example) receive an influx of
moose from upland ranges in early winter, and support very dense winter-
ing concentrations. Others (in the upper Little Chena drainage near
Fairbanks, for example) support moose during late summer and fall, but
may be virtually deserted when moose move down into denser forests and
riparian stands in November and December. The most diversified burns
(Kenai 194 7) also support significant densities of moose during calving,
summer and rutting periods and have relatively high resident populations
throughout the year.
Food Habits of Tame Moose
The results of this study, as excerpted from LeResche and Davis
(1973), are presented below.
ABSTRACT
Food intake of three tame moose (Alces alces rrtgas) was observed on
the Kenai Peninsula, Alaska, during summer on normal range and during
winter and spring on normal and depleted range. Plant species and bite
sizes were recorded for 49,308 bites consumed. Food eaten varied between
summer and winter, and moose ate a greater variety of forage than pre-
viously realized. Birch (Betula papyr£fera) leaves comprised 56 percent
(by number of bites) of the summer diet, forbs 25 percent, grasses, sedges
and aquatics 10 percent, and willow (SaUx spp.) 5 percent. Winter diet
on range that had supported average moose densities for the area (15 per
square mile) was 72 percent birch twigs, 21 percent lowbush cranberry
( Vaccin£um vi t-is-idaea) and 6 percent willow and alder (Alnus crispa).
On depleted winter range, stocked for 18 months with abnormally high
moose densities, birch twigs comprised only 22 percent of the diet. The
bulk of bites taken were of lowbush cranberry (51 percent) and foliose
lichens (Pelt-tgera spp.) (23 percent). In Hay, moose consumed 10--30
times as much cranberry and lichen as birch. Availability of understory
forage species during part of the winter is probably an important factor
in supporting the very high moose densities found on this range.
INTRODUCTION
Analyses of moose range commonly stress abundance, production and
use of large woody shrubs (Aldous and Krefting, 1946; Krefting, 1951;
Spencer and Chatelain, 1953; Spencer and Hakala, 1964; Houston, 1968).
This emphasis occurs because browse is the only food available to moose
in winter over much of their range and winter range is classically con-
sidered a limiting factor for moose populations, and because range survey
95
methods depend on the ease by which one can visually estimate browse
conditions (Cole, 1963).
Emphasis on woody shrubs has persisted despite an impressive array
of evidence that moose are extremely catholic in their tastes. Especially
in spring and summer, they consume forbs, grasses, mushrooms, lichens,
and practically all other floral components of their environment. Murie
(1934) reported a great variety of foods taken on Isle Royale. Peterson
(1955) listed more than 100 species and groups of plants taken at least
occasionally by moose. Houston (1968), although stressing that browse
was the most important food class during all seasons in Wyoming, demon-
strated that moose ate significant quantities of grasses, sedges, rushes
and forbs in spring and summer.
This paper reports that moose on heavily utilized range in south-
central Alaska ate considerable proportions of decumbent lowbush cran-
berry (Vaccinium vitis-·tdaea) and lichens during winter, despite an
abundance of available and unused woody browse. Furthermore, only about
65 percent of their food consumption in summer was from erect browse
species.
Summer Foods
Fifty-six percent of more than 28,000 bites taken by the three· tame
moose in July and August consisted of birch leaves. Utilization by
individuals ranged from 49-65 percent birch leaves. Five percent of
bites were of willow species. Moose took all leaves, most bark and
stems up to 2 em diameter from the willow they discovered. Four percent
of bites were leaves and twigs of dwarf birch (Hetula nana). Thus,
during summer, 65 percent of all bites taken were parts of woody browse
plants.
The rema1n1ng one-third of summer forage was primarily forbs (25%),
grasses (3%), sedges (4%) and aquatics (3%). Alder, aspen, lowbush cran-
berry, lichens and mushrooms (mostly Boletus spp.) were taken in trace
amounts.
Moose consumed different nonbrowse species as the season progressed,
apparently according to plant phenology. In late May and June, they ate
newly emergent marsh-growing £orbs, especially cloudberry (Rubus
chamaemorus) and sundew (Drosera rvtundi[ol1:a). Moose gathered in
groups in muskegs to feed on these species in their pre-flower and early
flowering stages. Hoose in boggier areas consumed newly emergent sedges
(Caf'ex spp. ; Eriophorum spp.) and horsetails (1!-'qu/Deturn spp.) during
this period.
Fireweed (EpUoblwn angustifol-Z:um and E. laLij'oliurn) was a favored
food during July. Moose selected plants in the pre-flower stages, and
consumed flowered plants (late July and August) only rarely. Along with
lupine (Lupin us nootkatensis), fireweed contributed most of the forb
biomass consumed during summer. These two species were also the most
abundant £orbs present during this period.
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Lupine was the dominant forb food from the date we first saw moose
consume it (10 July) through late August. Lupine was abundant and the
plants relatively large (20-25 ern in height) for several weeks before
the moose first began to eat the species. During the period of consump-
tion, moose selected almost exclusively plants in the pre-flower state.
Mushrooms, especially Boletus spp., were abundant during the summer
of 1970, and apparently were eaten whenever encountered by moose. Fifty
other nonbrowse plant species consumed in summer are listed in Table 2.
Winter Foods
Winter feeding habits of moose on normal range varied significantly
from those of animals confined to a small area of depleted range. We
did not collect quantitative data from tame moose tmtil February, but
rather observed wild animals during September-January. Plants consumed
were essentially the same as those taken during late winter with one
exception. During this entire period, when snow depth was less than
ca:30 ern, moose dug craters to secure large, still succulent, Carex
plants from boggy areas around marshes and river banks.
Tame moose on normal winter range consumed 72 percent birch stems
in February-May. Most of the remaining plant material eaten was lowbush
cranberry--21 percent by number of bites and by biomass. Lowbush cran-
berry was available throughout the winter, being emergent from the snow
in most habitats on most days. The multitude of dead-fallen burned
spruce poles and the dense lower branches of new growth conifers shielded
lowbush cranberry plants from deepest snow. Mbose commonly dug craters
under these obstructions and often kneeled beside them to feed on low-
bush cranberry plants.
In addition to birch, rnoos.e browsed 6 percent willow and alder, in
approximately equal proportions. A few bites of the large foliose lichen
Pelb:ger•a spp. were consumed on normal range, as were some miscellaneous
£orbs--primarily wintergreen (Pyr•ola secunda) and dwarf dogwood (Cm•nus
canadensis) after snow melt.
The moose on depleted range ate only 23 percent browse during
February-May. Almost all lichens were taken in May when snow cover was
absent from most of the ground surface. Thus, lichens were essentially
a spring food, and not a winter staple. In late April and during Hay,
however, lichens made up more than half the diet, with cranberry compris-
ing almost all other food. Lichens consumed were almost entirely the
large foliose Pel Uger>a spp., which is very abundant ground cover in
many areas of the burn. Other genera (especially Cladmn:a spp.) were
also consumed.
Results of nutritional and elemental analyses of food species
collected in t~y are presented in Table 11.
97
DISCUSSION
Snow is a paramount factor in seasonal distribution and numbers of
mammals in the north (Formozov 1946). Moderate snow conditions on the
northern Kenai Peninsula lowlands contribute significantly to the land's
exceedingly high carrying capacity for moose. The range produces more
than 180 kg (400 pounds) of available browse per acre annually in the
most productive areas (Bishop 1969). While this great biomass of woody
browse is doubtless important, it is only slightly used at current high
moose densities, and similar habitat elsewhere in Alaska supports many
fewer moose on a year-round basis. Other moose populations believed to
approach the 1947 Kenai burn group in year-round density are the Matanuska
Valley herd and the small, recently-established population near Yakutat.
These areas are characterized in most years by snows arriving late in
fall and disappearing or settling early in the spring. Non-woody plants
are thus available to these populations during most months of most years.
In contrast, moose populations on the vast seral ranges of interior
Alaska do not reach densities approaching those on the northern Kenai
Peninsula. Many interior situations present higher quality (predominantly
willow--Milke 1969) browse ranges; however, snow cover is early, persis-
tent, and often deep. The long period during which Interior moose must
eat only woody browse is probably an important factor in limiting moose
densities.
An almost complete loss of calves in much of the 1947 Kenai burn
during the winter of 1971-72 corroborates this thinking. During this
winter, lowbush cranberry was covered by snm.,;r in all habitat types by
mid-December. The snow did not settle or melt appreciably in the succeed-
ing weeks, and by late February all 35 calves in the Moose Research
Center enclosures were dead. Surveys in the vicinity of the Center
showed a calf:cow ratio of .02 (n = 186 cows). Calves died inside and
outside our enclosures with rumens full of birch, and with body weights
decreased 27 -34 percent from September-November weights.
Axiomatically, food quality and digestibility are as important as
quantity and availability in determing ungulate densities (cf: Klein
1970), and quality of browse species is related to successional state
(Cowan et al. 1950). Under the best conditions, woody browse usually
contains less than 10 percent crude protein, 1-7 percent crude fat,
40-60 percent carbohydrates (NFE) and 20-40 percent cellulose (crude
fiber) (cf: Kelsall 1968, Milke 1969, Houston 1968). Kelsall does
report analyses of eight willow species with a mean of more than 20 per-
cent protein, but this is higher than other reports. In protein levels,
at least, the woody parts of all common browse species provide barely
adequate maintenance levels of nutrition as judged from domestic rumi-
nants (Swenson 1970) and other wild cervids (Ullrey et al. 1969, Luick
et al. 1971). Thus, population levels of moose confined to woody browse
for long periods may suffer accordingly.
Depletion or virtual elimination of some browse species--willow,
aspen and highbush cranberry in the Kenai burn--may not only reduce the
proportion of higher-protein forage, but also does reduce variety avail-
able in winter. Variety itself is important to ruminants, for
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digestibility of forage is sometimes altered strikingly by addition of
other material to a one-species diet (eg: Mellenberger et al. 1971).
Nonwoody browse and succulent species were consumed in quantity by
moose when snow cover allowed. The several important nonbrowse species
increased crude nutrient intake as well as providing variety. Peltige~a
lichens contain nearly 20 percent protein and more than 9 percent ash
(Kelsall 1968). Lowbush cranberry, the most important nonwoody winter
food, contains 67 percent carbohydrates and protein levels approximately
equivalent to those in woody browse species.
It is instructive to calculate nutrient contents of several diets
containing various proportions of browse and nonbrowse forage. Moose
eating only birch consume approximately 9 percent protein, 15 percent
fat, 50 percent carbohydrates and 2 percent ash. This diet is only 37
percent digestible, as estimated using bovine flora (WARF Institute).
By substituting 25 percent lowbush cranberry for some birch (thereby
approximating the 75:25 proportions of birch and cranberry eaten on
"normal" Kenai winter range), protein and fat intake decreases somewhat,
but carbohydrate level increases to 54 percent and digestibility is
enhanced, even discounting any synergistic effects.
A 50 percent birch, 25 percent cranberry and 25 percent Pelt-Z:ge~a
diet (approximating the February-May average on "depleted" Kenai winter
range) provides 10.7 percent protein, 3.7 percent ash and 57.7 percent
carbohydrates--increases over a pure birch diet--but a decreased propor-
tion of fat. In any event, biomass of available forage is increased
when nonbrowse plants are available, and this additional plant material
is at least equal to birch browse in nutrition.
We have concluded that the exceedingly high moose densities on the
1947 Kenai burn range result not only from high annual production of
woody browse, but also from the normally moderate snowfall and short
duration of snow cover sufficient to restrict consumption of nonbrowse
species. Seral ranges with good ground cover of lowbush cranberry and
l'el Hge~a lichens will likely have a very high carrying capacity for
moose if snowfall and duration of snow cover permit access to these
plants during at least part of the winter. Nonbrowse plants not only
increase biomass of forage available, but also provide higher protein,
ash, carbohydrate and digestibility levels, as well as enhanced variety.
Feeding Craters
Three concentrations of feeding craters examined in mature hardwood
habitat within the Moose Research Center enclosures contained 84 craters,
or 19 per hectare. One concentration examined in sedge/spruce-Ledum
habitat contained 47 craters, or 58 per hectare. Mean dimensions of
craters measured are listed in Table 18. Locations of craters relative
to surrounding objects are tabulated in Table 19.
Several difficulties were encountered in determining what species
moose ate in the craters and what plants were present. Pawing tore
apart vegetation to the extent that it was uncertain whether it had been
99
Table 18. Dimensions of moose feeding craters in mature hardwood and
sedge/spruce-Ledum habitat types.
Mean (em) Range (em)
n Length Width Depth Length Width Depth
Hardwoods 84 183 102 36 76-381 36-178 22-56
Sedge/spruce-Ledum 47 180 120 20 76-483 28-198 10-31
Table 19. Location of individual moose craters in relation to surround-
ing objects.
in open
near windfall
near birch
near spruce
near aspen
near stumps
near alder
Hardwoods
31%
16%
26%
10%
11%
10%
100
Sedge/
Spruce-Ledum
62%
13%
26%
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fed upon or merely pawed. Granular snow at the base of craters often
caved in, covering the vegetation, and light snowfalls covered many
plants. Because of this, plants eaten from craters in hardwood habitat
types were not recorded. In a few cases craters were not dug deep
enough to reach vegetation (see Table 20).
RECOMMENDATIONS
1. Moose populations in the northern Kenai Peninsula should be
managed on the assumption that they are near maximum levels and slowly
declining. No increase can be sustained by the present habitat, although
some can be expected within 3-5 years in the 1969 burn area.
2. Annual changes in production and recruitment should be inter-
preted primarily in light of long-term trends, with the knowledge that,
in areas of high population densities, these parameters may fluctuate
widely year-to-year.
3. Experimental studies should be performed to confirm suspected
correlations between nutritional status and blood parameters in moose.
These studies should include the evaluation of handling stress and its
influences on blood values obtained.
4. Blood thyroxine levels should be further studied in relation to
metabolic state.
5. All available weight and morphometry information should be con-
solidated and published.
6. The correlations between chemical values of pellets and rumina
of moose on different quality ranges should be further investigated with
a view toward determining whether simple pellet analyses might provide
an index of range quality.
7. Snow depths should be routinely monitored in all habitat types.
Data collected to date should be consolidated with food habits informa-
tion and snow data from Interior Alaska (Coady's) and conclusions drawn
concerning the relative importance of snow conditions to moose in these
two regions.
8. Further data should be gathered concerning feeding craters with
a view toward determining whether or not crater numbers and characteris-
tics are indicative of relative snow-induced stress on moose populations.
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101
Table 20. Plant species present and fed upon in moose feeding craters.
Percent craters in
Percent craters with species present which species was
fed u12on
Plant Species mature hardwood sedge/spruce-Ledum sedge/spruce-Ledum
Vaccinium vitis-idaea 68 34 15
Ledwn palustre 20 36
f-' Rosa acicula:l'is 20
0
N
Lycopodium spp. 8
Betula papyrifera 4
Salix spp. 79 72
Ca:l'ex spp. 70 36
Equisetum spp. trace
No species reached 13 4
-------------------
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----~--~----~ bighorn sheep (Ovis canadensis canadensis) at capture, after
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104
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1969. A note on the blood chemistry of the Shiras moose.
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Klein, D. R. 1970. Food selection by North American deer and their
response to overutilization of preferred plant species. pp. 25-46,
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--------~--~· and~~~-------1953b. Effects of land use on moose
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562.
LeResche, R. E. 1966. Behavior and calf survival in Alaskan moose.
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1970. Moose Report. Ann. Proj. Prog. Rept. Fed. Aid
in Wild!. Rest. W-17-2, Juneau, Ak. 93 pp.
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Naturaliste Canadien (in press).
----~~~~~-· and J. L. Davis. 1971. Moose Research Report.
Proj. Prog. Rept. Fed. Aid Wild!. Rest. W-17-3, Juneau, Ak.
Ann.
156 pp.
---------,---:----. and 1973. The importance of nonbrowse
foods to moose. J. Wild!. Hgmt. 37(3)(In press).
., R. H. Bishop and J. W. Coady. 1973a. Distribution and
----~--.,--------habitats of moose in Alaska. Le Naturaliste Canadien (In press).
---------:------=-'' U. S. Seal, P. D. Karns and A. W. Franzmann. 1973b. A
review of blood chemistry of moose and other Cervidae, with emphasis
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adaptive
calves.
p. 12.
R. G. White and E. Reimers. 1971. Investigations of the
significance of winter growth patterns in female reindeer
Proc. 22nd Alaska Science Conference. Alaska Div. AAAS.
Lutz, H. J. 1956. Ecological effects of forest fires in the interior
of Alaska. U.S.D.A. Tech. Bull. 1133. 121 pp.
Mellenberger, R. W., L. D. Satter, M.A. Millett and A. J. Baker. 1971.
Digestion of aspen, alkali-treated aspen, and aspen bark by goats.
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Milke, G. C.
Alaska.
1969. Some moose-willow relationships in the interior of
Thesis, Univ. Alaska, College.
Miller, W. J., A. D. Haugen and D. J. Roslien. 1965. Natural variation
in the blood proteins of white-tailed deer:Subunit basis. Blood
29:86 7.
Murie, A. 1934. The moose of Isle Royale. Misc. Publ. Mus. Zoo!.,
University of Michigan. No. 25. pp. 1-44.
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saliva to determine the sodium status of cattle and sheep. Aust.
Vet. J. 46:595-598.
Nadler, C. F., C. E. Hughes, K. E. Harris and N. W. Nadler. 1967.
Electrophoresis of the serum proteins and transferrins of Alces
aZces (elk), Rangifer tarandus (reindeer), and Ovis daZli (Dall
sheep) from North America. Comp. Biochem. Physiol. 23:149-157.
1968. Serum protein electrophoresis of Oreamnos americanus
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25:1121-1124.
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thyroxine. Clin. Chern. 15(6) :509-520.
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Buccal smears as an indicator of nutritional
28(10):254-256.
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107
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Silver, H., N. F. Colovos, J. B. Holter and H. H. Hayes.
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PREPARED BY:
Robert E. LeResche
Game Biologist
SUBMITTED BY:
Richard H. Bishop
Regional Research Coordinator
APPROVED BY:
109
State:
Cooperators:
Project No.:
Job No.:
JOB PROGRESS REPORT (RESEARCH)
Alaska
Alaska Department of Fish and Game, U. s. Bureau of
Sport Fisheries and Wildlife--Kenai ·National Moose
Range, and Alaska Cooperative Wildlife Research Unit
W-17-4 Project Title: Big Game Investigations
1.2R Job Title: Moose Behavior
Period Covered: July 1, 1971 through June 30, 1972
SUMMARY
Total mortality of orphaned and unorphaned ~oose calves during the
winter of 1971-72 precluded assessment of differential mortality. Eight
orphaned calves lost a mean of 35 kg from autumn until their deaths in
December or January, or 20 percent of their original body weight. After
apparently being deserted by their mothers in late winter calves aggre-
gated in large groups containing only calves.
Variations in aggregating behavior are related to breeding activities,
mother-young relations, the male social system, the sex ratio of the
population, and external influences of forage, topography and cover.
Aggregations from midwinter through spring are primarily related to the
extrinsic factors of forage location and cover. In addition the solitary
nature of the cow with calf may facilitate the calf's protection from
predation and access to the best forage available to the cow. Aggrega-
tions during the pre-rut, rutting, and post-rutting periods probably can
be attributed to internal behavior patterns associated with breeding and
possibly the establishment and maintenance of social systems, as well as
to environmental factors, including access to high quality forage supplies
in early winter which serves to provide the moose with added sustenance
prior to the severe winter periods where forage accessibility is restricted.
Although the social system among moose populations appears to be
primitive when compared to the highly gregarious systems of wapiti, red
deer and caribou, the differences in aggregation patterns among the
three populations studied here indicate that the species is highly
adaptable to a variety of conditions. The successful adaptation to
different habitats, which range from mountainous, forested terrain where
plant communities are relatively stable, to alpine tundra, and to
undulating boreal forest where successful occupation is primarily
dependent upon the transitory seral stages, must be considered a major
influence upon the social system and the aggregation patterns described
herein.
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Sunnnary . .
Background.
Objectives.
Procedures.
Orphan Calf Survival
Moose Aggregations .
Findings ........ .
CONTENTS
i
1
2
2
2
4
5
5
8
8
Orphan Calf Survival
Moose Aggregations
Results . . . .
Discussion. . . .
Recommendations .
Literature Cited.
12
17
17
BACKGROUND
LeResche and Davis (1971) discussed the importance of moose (Alces
alces) behavior to population dynamics and moose management practices.
Of special note are the importance of the cow-calf bond to calf survival
during the first winter, and the seasonal dynamics of moose aggregations.
Migratory behavior is considered separately under Job 1.4R (LeResche and
Davis, 1971 and this report).
Proper management of moose populations sometimes requires harvest-
ing females as well as bulls. Calves orphaned when their mothers are
taken by hunters may present special management problems. If survival
or physical condition of calves is significantly affected by loss of the
mother, hunting regulations may have to be modified to take this into
consideration.
High mortality of moose calves may occur even when the mother is
present. In an area near Palmer, Alaska, in 1965, over 55 percent
mortality of calves occurred during the first five months after birth
(LeResche, 1966). The greatest mortality occurred within t>vo weeks
after birth, when approximately 26 percent of the calves were lost.
Calf losses during severe winters may also be high. In the Yakutat area
of Alaska, for example, nearly 60 percent of the calves observed in
December 1968 were lost before counts were again made in May 1969 (Rausch,
1971). However, the mortality rates of calves without mothers may be
even higher. Altmann (1958), who did her work in Wyoming, stated that
moose calves without mothers rarely survive the winter. In Alaska
several populations of moose have been established by transplanting
calves which were raised in captivity. Data on mortality of these
calves before release and hunting data on the established populations
are available (Burris, 1971). These calves were released in areas with
no moose or low moose populations and good habitat conditions. The
transplants were successful, but actual mortality rates from the time
of release are not known.
Orphaned calves surviving until spring may be in poorer condition
1
than calves with mothers and therefore be more subject to mortality by
disease, accidents, or predation.
The mother's presence may cease to be a significant factor in calf
survival and condition sometime between fall and spring. If large
enough samples of calves can be obtained at different times, a comparison
of survival rates between samples will be useful for management purposes.
Behavior patterns of moose calves will obviously be altered by loss
of the mother. Social stress may increase due to attempts by orphaned
calves to associate with other moose or even other species. When a calf
loses its mother, the calf may revert to the generalized following
response of its earlier life (Altmann, 1958). It will probably be
rejected by other moose and possibly injured. Altmann (1958) cited an
example of a healthy orphan moose calf being put into an area with a
bison herd. The calf tried to join the herd and was gored by a bison
cow, although there were no obstacles to its escape.
Weaning of calves may be nearly complete by the rutting season, but
calves may depend on their mothers for guidance in feeding. In deep
snow conditions calves may expend more energy without a mother to break
trail for them. Social interactions may cause calves to be driven out
of optimum feeding areas under deep snow conditions.
OBJECTIVES
To gain information on survival rate, physical conditions, and
causes of mortality of moose calves orphaned at various times in the
fall and winter; to compare this information with survival, condition,
and causes of mortality of calves having mothers.
To study certain aspects of the behavior of orphaned calves and
compare to the behavior of calves having mothers.
a) Social interactions
b) Activity patterns and movements
c) Feeding
To examine the seasonal dynamics of moose aggregations on the Kenai
Peninsula and compare them with groupings in other parts of North America,
in reference to habitat and population characteristics.
PROCEDURES
prphan Calf Survival
The study was conducted at the Kenai Moose Research Center on the
Kenai Peninsula. Host of the work was done in Pen number 4. During the
summer of 1971, this pen contained eight adult females, six calves, five
2
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to seven yearlings, and three adult males. In the fall and winter
(1971-72), moose numbers reached a maximum of at least 40 animals due to
introductions of moose from outside the pens. High winter mortality
reduced the Pen 4 population to 13 or 14 moose by June of 1972.
Host observations were made in Pen number 4. One lone calf was
observed in Pen number 3 for several days in February 19 72. Additional
observations of lone calves and calves with cows were made in areas
outside the Moose Research Center.
Between September 21, 1971 and November 23, 1971, eleven calves
were trapped from the area outside the Moose Research Center. These
calves were released in Pen number 4. When the mothers were also
trapped, they were released outside the pens. One calf that was born
in Pen number 4 was orphaned when its mother died from a drug overdose.
This calf was also used in the orphan project.
When a calf was trapped, it was weighed and measured and a blood
sample was taken. All trapped calves were ear tagged and collared for
identification. Seven of the calves were equipped with radio collars.
After being introduced into Pen number 4, calves were frequently
located by one of the following methods: (1) radio-tracking, (2) snow
tracking or accidental sightings from the ground, and (3) aerial
sightings.
When a calf was sighted, its location was plotted on a map. The
map location was often determined by using a hand compass and counting
paces to fencelines or other permanent landmarks.
An attempt was made to determine what vegetation was eaten most
extensively by the calves. The following methods were used to study
feeding: (1) identifying plant after observing calf feeding, (2) locat-
ing calf and backtracking in snow to see what plants were eaten, and
(3) collecting rumen samples from dead calves for later analysis.
Behavior patterns that were observed include:
1. Activity patterns
a. feeding (duration and frequency)
b. resting (duration and frequency)
c. movements (rate, frequency, snow depth, and habitat)
2. Social interactions
a. frequency of contact
b. type of contact
c. description of animal or group contacted
3
Dead calves were thoroughly examined. If the carcass was not
decomposed, it was weighed, measured, and autopsied. The area around
a carcass was investigated for tracks made by the calf or other animals.
In some cases, backtracking in the snow helped to determine rate of
movement and activity prior to death.
Materials collected from dead calves included: (1) femur for
marrow fat determination, (2) mandible for measurement and tooth
eruption, and (3) rumen sample to be analyzed.
Data analysis will include: (1) comparisons including percent fat
in femur marrow and rumen analysis between winter-killed calves and
calves which died of other causes and (2) comparisons of behavior
patterns of orphan calves in Pen 4, lone calves observed outside the
pens, and calves having mothers.
Moose Aggregations
We follow Bergerud and Manuel (1969) in defining an aggregation as
any group of animals which occur within reasonable proximity to each
other. The problem of determining whether these aggregations were
dictated by external factors such as topography or forage supplies, or
whether they are in fact social groups wherein individuals interact
among each other (Etkin, 1964) will be discussed.
Most aggregations discussed in the literature were located by
aerial search using a Piper Supercub, Aeronca Champ, or Bell G3B
helicopter. In addition, summer observations in southwestern Montana
were made from vantage points where moose could be readily located from
the ground due to the open nature of the area. In Minnesota, summer
observations included moose using lakes, streams and ponds, as investi-
gated from a canoe and also by air. All Alaska data were obtained by
aerial search. Information from southwestern Montana was obtained from
1958-60 and from 1965-67, from Alaska between 1970-72, and from Minnesota
between 1967-70. Aggregations have been grouped according to different
periods representative of moose phenological patterns, i.e., calving
period, summer, pre-rut, rut, post-rut, early winter, midwinter, late
winter and spring. These periods seemed to be the most likely times
when group sizes would vary for one reason or another.
Moose were classified according to the following criteria: bulls
were identified by presence of antlers or antler pedicels and lack of
the white vulvar patch (Mitchell, 1970); cows were identified by lack
of antlers or pedicels and presence of the white vulvar patch; calves
were identified on a basis of size (usually there was a mature cow
present to compare sizes with) and the short snout length characteristic
of yonng-of-the-year animals. Because the nature of the terrain precluded
close observation of moose, no classifications were attempted in north-
eastern Minnesota after January 1, when most bulls were either shedding
or had shed antlers. Some individuals were identified according to sex
and age in southwestern Hontana during the winter. Groups were classified
as bulls only, cows only, bulls and cows, ccws and calves, and bull-cow-
calf groups for purposes of analysis. Only groups in which all individuals
4
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were classified were included in the analysis of sex-age groups.
Characteristics of Hoose Populations and Habitat
The Kenai, Alaska, moose population is migratory, and spends summer
months at or above timberline in the Kenai Hountains. Cow:calf ratios
averaged 100:44 from 1968 to 1970 in June (LeResche, 1970), and the bull:
cow ratio averaged 20:100. Hunting seasons are open for bulls with
limited cow permits. Population density is over 3.6 moose per square
mile (Evans et al. , 1966) . Moose are generally readily visible from the
air in this area of Kenai birch (Betula papyrifera var. kenaica), quaking
aspen (Populus tremuloides) , and black spruce (Pice a mariana) , which was
burned over in 1947 (Spencer and Hakala, 1964), and is generally a low
forest.
The southwestern Montana moose population probably moves shorter
distances, usually along a drainage (Knowlton, 1960), and occupies more
densely forested habitats which are not as amenable to aerial observa-
tion as those in the Kenai. Populations may spend all but late winter
in mountainous terrain, at which time they commonly use the narrow willow
(Salix sp.) bottoms along drainages. Cow:calf ratios averaged 100:45
and sex ratios approached equality (Stevens, 1966, 1970). Hunting seasons
are by either-sex permit with cows and bulls taken in similar proportions
(Stevens, 1971).
Population densities range near 0.43 moose per square mile, but
Stevens pointed out that these figures are misleading because an undeter-
mined amount of unsuitable habitat has been included in the estimates.
Generally, these moose are dispersed until winter and then may occur in
winter concentrations at densities of up to 25 moose per square mile.
A typical wintering situation would be several to 25 moose in a willow
bottom of each drainage. It should be mentioned that the winter ranges
of the Jackson Hole (Houston, 1968) and Red Rock Lakes National Wildlife
Refuges (Darn, 1970) are more extensive than commonly occurs through
southwestern Hontana.
The northeastern Minnesota ranges are probably the most densely
vegetated, and least suitable to search from the air. Population
densities were approximately two moose per square mile (Peek, 1971),
the cow:calf ratios averaged 100:47 over a four-year period, and sex
ratios approached equality. These moose appear to be non-mif,ratory
(Van Ballenberghe and Peek, 19 71). No hunting occurred during the
observation periods and human-caused mortality was probably negligible.
It is evident that these moose populations vary considerably in charac-
teristics, and habitat.
FINDINGS
Orphan Calf Survival
Between September 21 and November 23, 1971, 11 calves were trapped
and introduced into Pen 4 without their mothers. A calf which v1as born
5
----------------------------------------------------------------------·-·--·-·--·-
in Pen 4 was orphaned on August 25, 1971, when its mother was acciden-
tally killed. This calf also became part of the orphan project (See
R70171, Table 1).
In November and December 1971, calves were located and observed for
varying lengths of time. Their locations were plotted on a map of Pen 4.
Observations were made of feeding and other activity patterns, and social
interactions.
Between December 10, 1971, and January 15, 1972, nine of the 12
orphan calves in Pen 4 were found dead. The average weight loss for
eight of these calves was 19.8 percent (Table 2). The ninth calf (No.
R70171, Table 1) was born in Pen 4 and weighed 185 pounds on August 25,
1971. When found dead in December 1971, this calf weighed 220 pounds.
The calf probably gained weight after the initial weighing, then lost
weight prior to death.
None of the other three orphan calves in Pen 4 were seen alive
later than December 8, 1971. Two of these calves were found dead during
helicopter tagging in the pens on May 9 and 10, 1972. The final project
calf was found dead on June 6, 1972. It was not possible to determine
the time of death of these three calves.
In Pen 4, four calves with mothers had been observed during the
fall and early winter. None of these calves were seen alive later than
December 19, 1971. Two of the calves had been trapped and marked during
the summer of 1971. Their carcasses were found in May and June 1972.
An unmarked calf carcass was also found in Pen 4 in June 1972.
Several dead calves were found in the other pens and in the area
adjacent to the moose pens during aerial and ground surveys. Many cows
known to have had calves in 1971 were seen in late winter without calves.
No yearlings were seen in the pens during the summer of 1972. Calf
mortality in the moose pens during the winter of 1971-72 was almost
certainly 100 percent.
The 100 percent mortality of calves in Pen 4 was unforeseen and
very few of the early observations were of calves with mothers. There-
fore, it was not possible to compare the behavior of orphan and non-
orphan calves in the same pen during the same time period.
Many dead calves found in the pens and in several other areas were
autopsied, weighed, and measured. Femurs and rumen samples were collected.
Some of the femur marrow values are given in Table 1. Rumen samples
have not yet been analyzed.
Aerial surveys and general observations during the winter of 1971-
72 seemed to indicate that more lone calves were seen than could be
accounted for by the number of hunter-killed cows. It is possible that
calves become weak under severe winter conditions which limit food avail-
ability and hinder movements. Perhaps they are then abandoned by their
mothers. In Pen 3 at the moose pens, a lone calf was tracked and observed
from the ground for several hours between February 10 and February 16,
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I Table 1. Data on project calves.
I Weight Date Weight % Fat-
Moose Date When Found at Condition of Femur
Number Sex Trapped Trapped Dead Death Carcass Marrow
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9671 F 9-21-71 159 kg 5-10-72 Hide and hair
I intact
9771 F 9-22-71 146 kg 1-14-72 118 kg Frozen
9871 F 9-29-71 182 kg 1-15-72 152 kg Legs, ears, stiff
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body soft
10671 M 10-12-71 159 kg 12-27-71 123 kg Partially stiff 9.1
10771 F 10-12-71 186 kg 1-14-72 139 kg Frozen
10871 F 10-13-71 175 kg 12-10-71 142 kg Fresh 8.0
I 10971 F 10-20-71 180 kg 1-15-72 145 kg Frozen
11071 M 10-21-71 175 kg 1-13-72 143 kg Fresh 7.1
11471 M 11-2-71 168 kg 6-6-72 Maggot infested
I 11571 F 11-3-71 200 kg 5-9-72 Maggot infested
12071 M 11-23-71 211 kg 12-23-71 170 kg Odor 8.3
R70171* M 8-25-71 84 kg 12-29-71 100 kg Legs, ears stiff 8.4
2771** F 7-27-71 70 kg 2-16-·72 107 kg Fresh 7.6
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* Born in Pen 4, orphaned August 25, 1971.
I **Born in Pen 3. Later abandoned by mother.
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Table 2. Weight losses of eight calves.
I
Weight when Weight at Weight loss Weight
I Moose trapped kg death kg kg loss
Number Sex (lbs.) (lbs.) (lbs.) (%)
I 10871 F 175 (385) 142 (313) 33 (72) 18.7
12071 M 211 (465) 170 (375) 41 (90) 19.4
10671 M 159 (350) 123 (2 70) 36 (80) 22.9
I 11071 M 175 (385) 143 (315) 32 (70) 18.2
10771 F 186 (410) 139 (305) 48 (lOS) 25.6
9771 F 145 (320) 118 (260) 27 (60) 18.8
I 10971 F 180 (395) 145 (320) 34 (75) 19.0
9871 F 182 (400) 152 (335) 30 (65) 16.2
X kg
I (lbs.) 177 (389) 142 (312) 35 (77) 19.8
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I 7
1972 (see No. 2771, Table 1). No other moose were seen near the calf
during this period. The calf died on February 16, apparently of
malnutrition. The calf was seen with its mother on December 14, 1971,
but had been seen alone as early as January 26, 1972. The mother was
seen again during a helicopter survey of the moose pens on March 23,
1972. She has also been observed and trapped at later dates.
Moose Aggregations
The results are presented below as they will be published in:
Peek, J. M., R. E. LeResche and D. R. Stevens. 1973. Dynamics of
moose aggregations in Alaska, Hinnesota, and Montana. J.
Mammal. (in press).
The other authors' affiliations are: Department of Entomology,
Fisheries and Wildlife, University of Minnesota (St. Paul); and U. S.
National Park Service (Estes Park, Colorado), respectively.
RESULTS
Annual Variation in Group Size
Group sizes reflected the relative density of the three populations,
being highest on the Kenai and lowest in southwestern Hontana (Table 3).
Summer aggregations were smallest, while the post-rutting aggregations
were largest. In addition, the Kenai population tended to aggregate in
March and April. Analysis of variance indicated significant differences
in group sizes between populations and between seasons within populations
at the 90 percent level of probability.
Group sizes varied most on the Kenai, where groups of 1-12 were
observed between July and November, and again in March and April.
Greatest variation in the Minnesota population occurred from October
through early January. Distributions at this time ranged from one to
seven moose. Variations in the Montana population was greatest during
the post-rut-March period. Both variation in group size and average
group size data suggest that the fall and winter periods may be identified
as the periods when these moose populations were most aggregated.
Distribution of Single Individuals
Cows occurred as single individuals more frequently than did bulls.
The exceptions to this occurred during the rut in the Montana and
Minnesota populations and in January and February in the Kenai population
(Table 3). Both sexes were least gregarious in summer in the Montana
and Minnesota populations and in January and February in the Kenai
population. Both sexes were most gregarious during the pre-rut, rut,
and post-rut periods. The percentage of male-female aggregations dropped
rather abruptly after the rutting period in the Kenai population; and
more slowly in the other two populations. The percentage of cows occur-
ring as groups of two or more individuals was higher on the Kenai than
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- - - ----- -- - - - ---- --Table 3. Distribution of aggregations by sex.
Percent Percent Percent
Total Males with Females Other Female Males Females
Number Males Alone Other Males M-F(c) alone (c) Females Calves Seen Seen
Period Groups If % If % II % fl % If % Solitary Alone Alone
:1innesota June-139 58 42 3 2 2 1 75 54 1 1 100 92 96
August
Early Sept. 23 5 22 2 9 7 30 9 39 60 36 56
Late Sept.-27 8 30 1 4 9 33 8 30 1 4 40 44 44
early Oct.
Late Oct.-228 25 11 32 14 59 26 95 42 17 7 81 22 55
November
December 232 30 13 26 11 34 15 130 56 12 5 94 33 74
l1ontana May-June 108 25 23 14 13 3 12 53 49 3 3 100 60 77
July-147 51 35 9 6 8 5 76 52 3 2 100 75 87
August
Late Aug.-89 28 31 12 13 13 15 35 39 1 1 100 53 71
early Sept.
Sept. 16-87 29 33 2 2 22 25 30 34 4 5 57 55 54
Oct. 15
Oct. 16-27 5 19 2 7 5 19 15 56 0 0 71 42 75
November
December 124 19 15 9 7 14 11 76 61 6 5 93 45 79
-----------------------------------------------------------------------------------'
Table 3. (cont' d.) Distribution of aggregations by sex.
Percent Percent Percent
Total l-'lales with Females Other Female Hales Females
Number Hales Alone Other Hales M-F(c) alone (c) Females Calves Seen Seen
Period Groups If % If % 1/ % II % II % Solitary Alone Alone
Kenai Hay-June 317 41 13 9 3 17 5 200 60 so 16 94 61 75
July 267 36 13 24 9 60 19 108 34 39 12 94 30 52
August 324 39 12 22 7 77 24 149 46 37 11 95 28 57
Sept. 1-15 250 21 8 13 5 83 33 86 34 47 19 87 18 40
i-' Sept. 16-100 3 3 2 2 34 34 36 36 25 25 66 9 38 0
Oct. 15
Oct. 16-95 1 1 1 1 42 44 29 31 22 23 65 2 31
November
December 69 4 6 0 0 5 7 44 64 16 23 96 44 68
January 99 17 17 2 2 4 4 65 65 11 11 95 81 81
February 36 3 8 0 0 0 0 29 81 4 11 100 100 88
March 593 30 5 7 1 21 4 414 70 125 21 100 52 74
April 540 54 10 4 1 41 8 314 58 127 24 93 55 65
--- --- --- -- -
------ -
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---------------------------------------
on the other two areas, probably reflecting both the higher density and
the disparate sex ratio of the Kenai population. The data suggest that
the bull may be identified as the more gregarious of the sexes.
Aggregation Patterns for Bulls
The mean group size for Kenai bulls was highest in July and August,
dropped during the pre-rut and rutting periods, increased slightly during
the post-rut period, dropped during the winter and then increased in
March. This was contrasted by the tendency for bulls in the Montana and
Minnesota populations, which tended to be lowest in the summer periods
and during the rut, and highest during the post-rutting and early winter
periods.
Variation in group sizes was greatest in summer in the Kenai popu-
lation. Variation in the Minnesota population was uniformly low until
the post-rut and early winter period. Although classifications ceased
after December, the variation of all groups observed in midwinter
suggested that bull-only group sizes declined during this period. The
Montana populations showed greatest variability in grouping of bulls in
May-June, and during the post-rut and early winter periods. Bulls in
the Minnesota and Montana populations characteristically aggregated more
in late fall than did the Kenai bulls.
Aggregation Patterns for Cows
Differences between aggregating behavior among cows from the three
populations were great. Cows from the Montana population were most
gregarious during the post-rut and December periods, although at no time
was there an apparent pronounced tendency to aggregate. The Kenai cow
population tended to aggregate during the pre-rut, rut, and post-rutting
periods and to disperse during the winter.
Variation in group size was uniformly low in the Montana and
Minnesota populations, and pronounced in the Kenai population. While
variation for bulls appeared greater in the Montana and Minnesota popu-
lations, variation for cows was greater in the Kenai population,
indicating the difference was related to the differences in sex ratio.
Aggregation Patterns for Cow-Calf Groups
There was less tendency for cows with calves to aggregate with
other moose than for cows without calves in the three populations (Table
3). The average group size ranged between 2 and 3 for all three popula-
tions, indicating that the cow-calf or cow-twin calf group was most
frequently observed. Cows with calves in the ~linnesota and Montana
populations did not appear to aggregate with other moose at any time
except during the pre-rut and post-rutting periods. Cows with calves
aggregated more frequently at all times of the year in the Kenai popu-
lation, especially during the migration periods of spring and fall
(LeResche and Davis, 1971).
11
Aggregation Patterns for Cow-Bull Groups
Groups comprised of both sexes were most frequently observed in the
pre-rut, rut, and post-rut periods, but the sexes aggregated more fre-
quently in the Kenai population than in the other two populations.
Aggregations of both sexes in summer commonly included an adult cow and
a yearling bull (yearling bulls may be identified by antler development,
Peek, 1961). Aggregations during the rut were of adult bulls and cows,
probably associated most frequently for breeding purposes. Cows with
calves were associated with bulls in the Hontana and Minnesota popula-
tions from the pre-rutting period on. The Kenai cows with calves also
associated with bulls during summer months and during the migration
period.
Aggregations Above and Below Timberline, Kenai
Flights made in June and July, August, and March on the Kenai
included observations of moose above and below timberline (Table 4).
In all cases but one, group sizes and variation in group size was
greatest above timberline than below. The one exception, cow-calf
groups seen in June below timberline as compared to those seen in July
above timberline, may have been associated with migration activities.
DISCUSSION
The milk group was considered as the basic social unit in moose
(de Vos et al., 1967), and other aggregations appear to be relatively
loose-knit and transitory (Houston, 1968). Interaction among animals
within aggregations during summer and late winter appears to be minimal
most of the time, with individuals often appearing to ignore each other.
The cow-calf group, plus the aggregations associated with breeding
appear to be true social units, as defined by Etkin (1964).
The causes of aggregation have been suggested to be defense against
predators (Lack, 1968; Goss-Custard, 1970), as well as to the pattern of
food availability in terms of food item dispersion and seasonal varia-
tion in abundance (Crook and Gartlan, 1966). Variations in aggregation
size in moose appear to relate to extrinsic factors such as those above,
plus snow depth and hardness, vegetative cover, and intrinsic factors
such as sex of animal, breeding activities, aggressive behavior, family
care, group life (Tinbergen, 1953) and possibly the sex ratio. Popula-
tion density did not appear to influence trends in aggregation changes
through the year, although the largest group sizes and greatest varia-
tion within periods did occur with the densest population.
Influence of Sex Ratio on Aggregation Behavior
Differences between the Kenai population and the other two popula-
tions in aggregating behavior of females may be partially attributable
to the disparate sex ratio on the Kenai as compared with the nearly
equal sex ratios of the other two populations. Cows were definitely
more gregarious on the Kenai than elsewhere. Groups of cows without
12
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- ----- ------------ -
Table 4. Summary of above (AT) and below (BT) timberline aggregations of moose in August, Harch, and June-July, Kenai Peninsula,
Alaska.
Number
AT Number Number Number Number of male Number
or of male of Group of female of Group female of Gr~up
Date BT groups moose x Range groups moose X Range groups moose X Range
August AT(2)1 32 58 1.81 1-8 80 154 1.92 1-12 44 197 4.48 2-11
BT (2) 29 40 1. 38 1-3 63 70 1.11 1-6 17 37 2.18 2-6
Harch AT(2) 18 40 2. 22 1-10 81 130 1.60 1-8 12 66 5.50 2-15
BT (4) 29 29 1.00 389 506 1. 30 1-6 10 30 3.00 2-5
July AT(4) 51 85 1.67 1-7 124 198 1.60 1-8 58 221 3. 81 2-5
June BT(2) 11 13 1.18 1-2 52 73 1.40 1-4 4 10 2.50 2-4
----------------------------------------------------------------------------------------------------------------------------------
1-' Males-females-w 2 Females-calves calves All Grou12s
AT Number Number Number Number Number Number
or of of Gr~up of of Gr~up of of Group
Date BT groups groups X Range groups moose X Range groups moose x Range
August AT(2)1 20 48 2.40 2-6 2 9 4.50 4-5 173 466 2.62 1-12
BT(2) 38 80 2.11 2-4 1 3 3.00 146 230 1.58 1-6
March AT(2) 15 49 3.27 2-13 126 285 2.26 1-15
BT(4) 43 99 2. 30 2-6 477 673 1.41 1-6
July AT(4) 32 73 2.28 2-4 2 7 3.50 3-4 267 584 2.19 1-13
June BT (2) 19 47 2.47 2-7 86 143 1.66 1-7
1 Number of flights. 2 Includes some nol classified according to sex or age.
calves occurred frequently at all seasons except in February and March.
While cows with calves were less social, even in early summer, groups
were observed. This period would be considered that when the cow would
be most protective of the calf (Altmann, 1958). The Kenai bull popula-
tion appeared to associate more readily with cows than the other two
bull populations in question, and less frequently with other bulls, when
ranges in group sizes are considered. This was especially apparent
during the fall and winter periods. The data indicate that the high
percentage of cows and low percentage of bulls in the Kenai population
tends to influence aggregating behavior by causing cows to be more
gregarious and by causing the sexes to mix more frequently during the
year than was observed in the other two populations.
Influence of Predation Upon Aggregation Size
Adult moose in prime condition appear to be relatively immune to
predation, but aged or diseased individuals and calves are vulnerable
(Hech, 1966; LeResche, 1968). In northeastern Minnesota and Ontario,
the incidence of moose remains in wolf (Canis lupus) scats is higher in
summer than in tvin ter (Halverson, 1969; Pimlot t et al. , 1969). This
suggests that predation is most important when aggregation sizes are
lowest, in summer, and also among the cow-calf groups which are the most
solitary. In addition, the late winter-early spring periods of low
aggregation size in northeastern Minnesota are periods when that popula-
tion is probably at its poorest physiological status, following the
winter period, and presumably includes more vulnerable animals than in
early winter or fall. It is therefore possibl£~ that the solitary and
well dispersed nature of the moose population, especially during periods
when individuals would be most vulnerable, may be attributable in part
to the influence of predation.
Undoubtedly the highly aggressive nature of the cow immediately
after parturition, and the tendency to remain with the calf rather than
to conceal it, are predator-induced responses. Tt appears that the
forested terrain which these species occupy during summer months favors
the solitary mode of defense against predation, where vulnerable indivi-
duals remain dispersed and assumedly less easy to locate. Cows with
young calves are not uncommonly found on small islands or in swampy
areas (Rausch, 1959; Peek, 1971) which probabJy favor the maneuverability
of the long-legged moose over its potential predators.
Although wolves are not present in the southwestern :Montana area,
and may not have been a major influence upon moose habitat selection or
mortality because of the abundance of other Cervidae (Cowan, 194 7; Hurie,
1944), the fact that cows are seen in relatively open cover with newborn
calves in this area (Peek, 1961) suggests that the aggressive nature of
the cow would facilitate calf survival if wolves were present.
The tendencies of cows to be aggressive prior to calving, to with-
draw from disturbances, and to defend the calf were factors which Geist
(1963) considered to inhibit development of any greater social organiza-
tion beyond the "milk group" in the Wells Gray Provincial Park, British
Columbia population. In addition, aggressive interaction between cows
14
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has been observed during the rut by Altmann (1959 :423), Geist (1963 :412)
and Peek (1961). This aggressiveness was considered by Geist (1971:124)
to be an antiwolf adaptation. The solitary tendency may be related to
this same influence or possibly to a need to minimize intraspecific
competition for scarce forage sources, or both.
Influence of Breeding Activities on Aggregation Behavior
Aerial searches during the actual breeding perjod would provide
observations of both sexes searching for n~tes, rutting groups, bands
of young individuals, and cows which had mated (Altmann, 1959). The
pre-rutting period is the first time since winter that increases in
aggregation size are readily apparent in the data. Bulls are least apt
to be seen alone during the period, cows without calves tend to associate
with other individuals more, but cows with calves are still solitary.
The tendency is for groups of cows and bulls to decline in size during
the rutting period, suggesting that pre-rutting and post-rutting period
groups are larger and more variable in size than the rutting group.
Actually, it appears that the separation of pre-rutting, rutting, and
post-rutting periods would be artificial, since it may be expected that
groups representing each phenological period may be present at any time
after breeding starts. The possibility exists that pre-and especially
post-rutting groups are aggregating for purposes other than these
associated with breeding.
Aggregations which contain bulls during this period tend to be
largest, so the period of the year when the highest amount of social
interaction occurs is also the period when bulls appear to be most
gregarious. Post-rutting aggregations are especially large. The period
of actual receptivity to breeding in the cow moose is quite short and
most cows are probably bred within a ten-day period centering upon
October l (Edwards and Ritcey, 1958). Spermatogenesis, indicative of
androgen secretion, extends uninterrupted from early September through
November and may have a significant influence upon male behavior until
antler drop in early January (Rausch, 1959; Peek, 1961; Houston, 1968).
It is likely that pre-rutting and post-rutting aggregations serve to
bring males together for purposes of social display. Observations of
bulls in November and December in Minnesota suggest that they are highly
aggressive at that time. Very often, antler polishing and jousting,
plus other activities which may be interpreted as display towards con-
specifics are observed. The possibility exists that dominance heirarchies
among males are established at this time. In summer, males appear less
antagonistic and less gregarious. This is contrasted by the antagonistic
nature of the female which coincides with and explains their solitary
nature. The tentative conclusion which may be drawn is that moose popu-
lations tend to be more patriarchal than matriarchal in nature, since it
appears that the bull is the most social of the two sexes, and appears
to govern what social interaction exists within a population. The bull
tends to seek out other individuals for purposes of social display which
is not the case with the female, whose social interactions center around
breeding and family care. If dominance heirarchies are established or
maintained in late fall and early winter, the purpose may be indirectly
related to reproduction through selection of habitat which facilitates
15
survival and maintenance of condition. Hoose in northeastern Minnesota
tend to disperse into small groups and to occupy the more dense covers
of sparse forage in late winter. Undoubtedly there is a premium in
occupying the cover which provides best forage as well. It is possible
that the more dominant males select prime wintering areas and tend to
exclude younger animals. This exclusion process is probably usually a
passive interaction, wherein the more aggressive animals occupy the best
bedding and feeding sites, causing the subordinate to search for other
areas. In this connection, it is significant that des Meules (1964)
reported that once suitable bedding sites were used up within a yard in
Quebec, moose occupying the area moved out.
Influence of Environment on Aggregation Behavior
The largest group sizes in the Kenai, and in northeastern Minnesota
occurred when moose were occupying the most open cover, whether it be
alpine tundra above timberline, or recently cutover shrub communities.
While both populations were aggregating during the reproductive and
post-reproductive periods, and most certainly the use of open cover
where individuals would be able to observe each other more easily would
be important, the Montana population appeared to aggregate to some
extent in winter while occupying the relatively open cover afforded by
willow bottoms. Moose occupying douglas fir (Pseudotsuga men::vZ:eseii)
in winter tended to be more solitary than those occupying willow bottoms
(Stevens, 1970 and Knowlton, 1960). Dassman and Taber (1956) noted that
deer (Odocoileug hermonus) which inhabit dense cover usually form small
groups or are solitary. Pruitt (1960) found barren-ground caribou
(Rang1:[er tarandus) occurred in smaller bands on taiga winter range than
on tundra summer range. Thus this tendency to aggregate into larger
groups may also have a psychological basis, wherein the larger group
replaces the role of vegetative cover for the individual (Crook, 1970).
Because moose groups are so loose knit, individuals seemingly being
oblivious to one another in late winter, such an explanation does not
completely explain this tendency. In winter the deeper snows on mountain-
sides cause moose to move into the willow bottoms and lower ranges
(Edwards and Ritcey, 1956) and also place the animal in contact with an
abundant and highly palatable forage source. Thus it appears that
topography, snow depth, and forage supplies play a major role in
aggregating moose in mountain areas.
This does not appear to be the case in the relatively even terrain
in boreal forests. Here successful occupation of late winter cover with
sparse forage sources is facilitated by the dispersal of the population
as Houston (1968) and Geist (1971) suggest.
Conclusions
It is concluded that the variations in aggregating behavior are
related to breeding activities, mother-young relations, the male social
system, the sexual composition of the population, and external influences
of forage, topography and cover. Aggregations from midwinter through
spring are primarily related to the extrinsic factors of forage and
cover. In addition the solitary nature of the cow with calf may
16
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facilitate its protection from predation and access to the best forage
available to the cow. Aggregations during the pre-rut, rutting, and
post-rutting periods probably can be attributed to internal behavior
patterns associated with breeding and possibly the establishment and
maintenance of social systems, as well as to environmental factors,
including access to high quality forage supplies in early winter which
serves to provide the moose with added sustenance prior to the severe
winter periods where forage accessibility is restricted.
Although the social system among moose populations appears to be
primitive when compared to the highly gregarious systems of wapiti
(Cervus c:anadensis), red deer (Cervus elaphus) and caribou, the differ-
ences in aggregation patterns among the three populations studied here
indicate that the species is highly adaptable to a variety of conditions.
The successful adaptation to different habitats, which range from
mountainous forested terrain where plant communities are relatively
stable,to alpine tundra, to undulating boreal forest where successful
occupation is primarily dependent upon the transitory seral stages must
be considered a major influence upon the social system and the aggrega-
tion patterns described herein.
RECOMMENDATIONS
1. Further studies should be undertaken to determine the importance
of the cow-calf bond to overwinter calf survival during "normal" winters.
2. Consideration should be given to encouraging or requiring the
harvest of calves when accompanying cows are killed in areas of high
moose populations.
3. Aggregation behavior should be studied in other parts of Alaska
with moose populations having sex ratios less skewed toward females, in
order to determine whether change in aggregation behavior might reflect
important changes in population size or productivity.
LITERATURE CITED
Altmann, M. 1958. Social integration of the moose ealf. Animal
Behaviour. 6(3-4):155-159.
1959. Group dynamics in Wyoming moose during the rutting
season. J. Mammal. 40(3):420-424.
Bergerud, A. T. and F. Manuel. 1969. Aerial census of moose in central
38 (4) :910-916. Newfoundland. J. Wildl. Manage.
Burris, 0. 1971. Unpublished data on moose transplants in Alaska.
Cowan, I. HcT. 194 7.
Parks of Canada.
The timber wolf in the Rocky Mountain National
Can. J. Res. D 25:139-174.
17
Crook, J, H. 1970. The socio-ecology of primates. Pages 103-166 in
J. H. Crook, ed. Social behaviour in birds and mammals. Acad.
Press.
and J. S. Gartlan. 1966. Evolution of primate societies.
Nature. 210:1200-1203.
Dasmann, R. F. and R. D. Taber. 1956. Behaviour of Columbian black-
tailed deer with reference to population ecology. J. Mammal.
37(2) :143-164.
des Meules, P. 1964. The influence of snow on the behaviour of moose.
N.E. Wildl. Conf. Hartford, Conn. 12 pp.
des Vos, A., P. Brokx and V. Geist. 196 7. A review of social behaviour
of the North American cervids during the reproductive period. Am.
}tidl. Nat. 77(2):390-417.
Dorn, R. D. 1970. Moose and cattle food habits in southwest Montana.
J. Wildl. Manage. 34(3):559-564.
Edwards, R. Y. and R. W. Ritcey. 1956. The migrations of a moose herd.
J. Mammal. 37:486-494.
1958. Reproduction in a moose popula-
tion. J. Wildl. Manage. 22:261-268.
Etkin, W. 1964. Cooperation and competition in social behaviour. In
W. Etkin, ed. Social behaviour and organization among vertebrates.
Univ. Chicago Press. 307 pp.
Evans, C. D., w. A. Troyer and C. J. Lensink. 1966. Aerial census of
moose by quadrat sampling units. J. Wildl. Manage. 30:767-776.
Geist, V. 1963. On the behaviour of the North American moose in British
Columbia. Behaviour 20(3-4):377-416.
1971. Mountain sheep; a study in behaviour and evolution.
Univ. Chicago Press. 383 pp.
Goss-Custard, J. D. 1970. Feeding dispersion in some overwintering
wading birds. Pages 3-35 in J. H. Crook, ed. Social behaviour in
birds and mammals. Acad. Press, London.
Halverson, P. B. 1969. Certain seasonal food habits of the timber wolf
in northeastern }linnesota with reference to predation. Unpubl.
Honors Thesis, Macalester College, St. Paul. 28 pp.
Houston, D. B. 1968. The Shiras moose in Jackson Hole, Wyoming. Grand
Teton Natl. Hist. Assoc. Tech. Bull. 1. 110 pp.
Knowlton, F. F. 1960. Food habits, movements, and populations of moose
in the Gravelly Mountains, Montana. J. Wildl. Hanage. 24(2):162-
170.
18
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Lack, D. 1968.
Co., Ltd.
Ecological adaptations for breeding birds.
409 pp.
Methuen and
LeResche, R. E. 1966. Behavior and calf survival in Alaskan moose.
M.S. Thesis. University of Alaska.
1968. Spring-fall calf mortality in an Alaska moose
population. J. Wildl. Manage. 32(4) :953-956.
1970. Moose report. Alaska Dept. Fish and Game Annu.
Proj. Seg. Rept. Vol. XI.
and J. L. Davis. 1971. Moose Research Report. Alaska
Dept. Fish and Game, Annu. Proj. Seg. Rept. Vol. XII.
Mech, L. D. 1966.
Fauna Ser. 7.
The wolves of Isle Royale.
210 pp.
U. S. Natl. Park Serv.
Mitchell, H. B. 1970.
British Columbia.
Rapid aerial sexing of antlerless moose in
J. Wild!, Manage. 34(3):645-646.
Murie, A. 1944. The wolves of Mt. McKinley. U. S. Natl. Park Serv.
Fauna Ser. 5. 238 pp.
Peek, J. M. 1961. Reproduction of moose in southwestern Hontana. M.S.
Thesis, Montana State Univ., Bozeman. 30 pp.
1971. Moose habitat selection and relationships to forest
management in northeastern Minnesota. Univ. Minn. Ph.D. Thesis.
250 pp.
Pimlott, D. H., J. A. Shannon and G. B. Kolenosky. 1969. The ecology
of the timber wolf in Algonquin Prov. Park. Ontario Dept. Lands
and Forests Res. Rept. (Wild!.) 87. 92 pp.
Pruitt, W. 0., Jr. 1960. Behavior of the barren ground caribou. Bio.
Pap. Univ. Alaska no. 3. 44 pp.
Rausch, R. A. 1959. Some aspects of population dynamics of the Railbelt
moose population, Alaska. M.S. Thesis, Univ. Alaska. 81 pp.
1971. Moose Report. Alaska Dept. Fish and Game, Vol. XI,
Proj. Prog. Rep., Fed. Aid Wild!. Rest., Proj. W-17-1.
Spencer, D. L. and J. B. Hakala. 1964. Moose and fire on the Kenai.
Proc. Tall Timbers Fire Ecology Conf. 3:11-33.
Stevens, D. R. 1966.
western Montana.
W-98 R-6. 16 pp.
Moose population and forage inventories in south-
Montana Fish and Game Dept. Job Compl. Rep. Proj.
1970. Winter ecology of moose in the Gallatin Hountains,
Montana. J. Wild!. Manage. 34(1):37-46.
19
1971. Shiras moose. Pages R9-95 in T. W. Mussehl and
F. W. Howell, eds. Game Management in Montana.
Tinbergen, N. 1953. Social behaviour in animals. Methuen, London.
VanBallenberghe, V. and J. M. Peek. 1971. Radio-telemetry studies of
moose in northeastern Minnesota. J. Wildl. Manage. 35(1):63-70.
PREPARED BY:
Robert E. LeResche
Game Biologist
SUBMITTED BY:
Richard H. Bishop
Regional Research Coordinator
APPROVED BY:
Research Chief, Division of Game
20
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State:
Cooperators:
Project No.:
Job No.:
JOB PROGRESS REPORT (RESEARCH)
Alaska
Alaska Department of Fish and Game, U. S. Bureau of
Sport Fisheries and Wildlife (Kenai National Hoose
Range)
W-17-4
1.3R
Project Title: Big Game Investigations
Job Title: Development and Testing
of New Techniques
Period Covered: July 1, 1971 through June 30, 1972
SUMMARY
A one-stage rumen fistulation procedure was not successful on two
tame moose. Mean monthly femur-marrow fat contents of calf moose from
January through May ranged from 11 to 6 percent. Adult femur-marrow fat
levels ranged from 13 to 94 percent (monthly means) on a year-round
basis. A manuscript on precision and accuracy of aerial moose censusing
was submitted for publication during the reporting period.
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Sununary • .
Background.
Objectives.
Procedures.
Findings ..•
Reconunendations
Literature Cited.
CONTENTS
BACKGROUND
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2
2
7
Moose (AZces aZces) research and management require methods of
estimating numbers and of handling, marking and following animals. These
techniques necessarily vary with species, location and nature of the
management/research problem. The Moose Research Center, with known
numbers of confined animals, provides a unique test-ground for numbers-
related techniques and for methods and equipment whose effectiveness can
be learned only by relocation of animals.
LeResche and Davis (1971) summarized present knowledge of and needed
future work on techniques of aerial censusing, chemical immobilizing,
radio-tracking, pellet-count censusing and marking moose for movement
studies. LeResche and Lynch (1973) reported on a moose trap developed
at the Moose Research Center and used to capture free-ranging moose in
Alberta.
Van Dyne (1968) reviewed techniques for measuring dietary prefer-
ences and digestibilities of foods selected by large herbivores. Previous
studies at the Moose Research Center have utilized techniques involving
observation of free-ranging animals (LeResche and Davis, 1973 and this
report, Job 1.1), estimating food intake by observing browsed plants
(LeResche and Davis, 1971 and Oldemeyer, 1973), rumen contents analyses
of shot animals (LeResche and Davis, 1971), fecal analyses (this report,
Job 1.1), and in vitro trials. To carry out i~ vivo digestibility trials,
rumen-fistulated animals were needed.
Collection of femurs and analysis of their marrow for fat content
has been done extensively and rather indiscriminately in Alaska in
recent years. Neiland (1970) reported a simplified "dry-weight" method
of estimating fat content in bone marrow, and many femurs have been
analyzed since then. The potentialities and shortcomings of using
marrow fat to indicate nutritional status are reviewed by Cheatum (1949)
and Bischoff (1954). Consolidation of data from various parts of
Alaska was needed to allow at least crude interpretation of moose femur-
marrow fat values.
OBJECTIVES
To develop and/or test techniques for: aerial censusing, pellet-
count censusing, immobilizing, radio-tracking, and marking of moose; to
1
develop a technique for implanting permanent rumen fistulae in moose;
and to establish baselines for interpreting femur-marrow analyses of
moose.
PROCEDURES
Procedures followed were essentially the same as those described by
LeResche and Davis (1971). In addition, bone-marrow data were collected
from several areas of the state and tabulated. Rumen fistulae were
implanted in two tame 14-month-old moose by Dr. R. Herin, Colorado State
Unive.rsity. A 4 em long incision was made in the left abdominal wall
approximately 15 em dorsally from the spine and 5 em posteriorly from
the last rib. The rumen was pulled through the incision (Fig. 1) and
sutured to the incision margin in a circular pattem. The rumen was
incised within the sutures and the base flange of the fistula plug device
(Fig. 2) was folded and inserted into the rumen. The sleeve flange was
then placed over the tube, pulled tight against the abdominal wall
(thereby squeezing the rumen and wall between the two flanges) and the
cable clamp was tightened. The plug was inserted and the animal revived.
The first attempt took two hours, the second, 20 minutes.
FINDINGS
Results of aerial-census evaluation experiments are presently in
manuscript form as: LeResche, R. E. and R. A. Rausch. Accuracy and
precision of aerial moose censusing. (Under review at J. Wildl. Mgmt.).
Radio tracking gear, as described by LeResche and Davis (1971)
proved successful in the orphan calf survival study, reported in Job
1.2R (this report).
One moose (Walter) died from the effects of prolonged chemical
immobilization (M-99) during the first fistulation procedure. The
flanges from the fistula plug assembly implanted in the second moose
(Raquel) caused tissue necrosis, and the plug was rejected in approxi-
mately ten days. The fistula itself healed satisfactorily, but the
hole closed completely about 18 months after the surgery.
Fat levels of femur-marrows of 106 calves and 60 adults are pre-
sented in Table 1 and Fig. 3. Hare data should be consolidated and
analyzed (eg: those from Coady, 1973) before conclusions are drawn.
No further findings concerning technique development and testing
are available at this time.
RECOMMENDATIONS
1. The one step rumen fistulation procedure is not indicated for
moose. Two-stage procedures should be tested.
2
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Figure 1. Rumen being pulled through incision in abdominal wall.
3
Plug inserts here
...,___ ___ Sleeve Flange
----Hose Clamp
Rumen Fistula Device
Figure 2. Schematic drawing of rumen fistula device tested
at Moose Research Center.
4
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I Table 1. Fat levels (expressed as percentage dry weight--cf: Neiland,
1970) of moose femur marrow collected in various months of the
year. Hales and females are lumped, "calves" are animals from
I birth through the following Hay 31, and "adults" are all older
animals. The tabulation includes only those animals killed by
man or other predators and not those obviously suffering from
I malnutrition or disease. The levels therefore approximate
"normal" levels for the time and place collected.
I Calves Adults
(% dry
Month n x weight) range n x range
I June 2 22 2-41 1 45
I July None 3 82 81-85
August None 2 86 75-93
I September None 2 94 94-94
I October None 1 29 (rutting bull)
November None 1 91
I December None 1 85
January 3 11 7-37 7 67 35-90
I February 26 10 6-28 7 77 57-92
I March 35 8 6-25 14 75 27-92
April 33 8 5-16 3 71 63-76
I May 2 6 3-9 4 13 8-20
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100-
80-
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Adult
20 "'"Co lves
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{See table I )
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26
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2 A c c t I I I c
0 ~~---~·-----L-'-----L-·----~·~----~·------~·------~·------~·------~----~------~----~ I I I I
Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr May
Month
Figure 3. Percent dry weight of femur marrow in adult (A) and calf (C) moose during months of the
year. Letters (A or C) indicate mean, vertical lines show ranges and numbers above lines
indicate sample sizes.
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2. Femur marrow fat analyses done in the future should be consoli-
dated and similarly summarized, to provide more valid guidelines for
interpretation. Fat values for marrow of animals suspected to have
suffered from malnutrition should be separately summarized. Marrow
analyses on calves of the year are uniformly low and not valuable for
determining comparative nutritional status.
LITERATURE CITED
Bischoff, A. I. 1954. Limitations of the bone marrot.r technique in
determining malnutrition in deer. Proc. Western Assoc. State Game
and Fish Commissioners, Las Vegas, Nevada. 34:205-210.
Cheatum, E. L. 1949. Bone marrow as an index of malnutrition in deer.
New York State Conservationist. 3(5):19-22.
Coady, J. W. 1973. Moose Report. Fed. Aid Wildl. Rest. Annu. Proj.
Seg. Rep. Juneau, Alaska. (in press).
LeResche, R. E. and J. L. Davis. 1971. Moose Research Report. Fed.
Aid Wild!. Rest. Annu. Proj. Seg. Rep. W-17-3. Juneau, Alaska.
156 pp.
----~---------· and 1973. Importance of nonbrowse foods
to moose on the Kenai Peninsula, Alaska. J. Wildl. Hanage. 37:3
(in press).
---------------· and G. M. Lynch. 1973. A trap for free-ranging moose.
J. Wildl. Hanage. 37(1):87-89.
Neiland, K. A. 1970.
caribou femurs.
Weight of dried marrow as indicator of fat in
J. Wildl. Manage. 34(4):904-907.
Oldemeyer, J. A. 1973. Quality of forage plants moose eat. Le Nat.
Can. (in press).
Van Dyne, G. M. 1968. Heasuring quantity and quality of the diet of
large herbivores. Pages 54-94 in F. B. Golley and H. K. Buechner,
eds. A practical guide to the study of the productivity of large
herbivores. IBP Handbook No. 1. (Blackwell, Edinburgh).
PREPARED BY :
Robert E. LeResche
Game Biologist
SUBMITTED BY:
Richard H. Bishop
Regional Research Coordinator
APPROVED BY:
c~ Di rttot=:iOilOf Gallle
Research Chief, Division of Game
7
State:
Cooperators:
Project No.:
Job No.:
JOB PROGRESS REPORT (RESEARCH)
Alaska
Alaska Department of Fish and Game; U. S. Bureau of
Sport Fisheries and Wildlife; Kenai National Moose
Range
W-17-4
1.4R
Project Title: Big Game Investigations
Job Title: Kenai Peninsula Moose
Population Identity
Study
Period Covered: July 1, 1971 through June 30, 1972
SUMMARY
Thirty-eight moose were marked and released outside the Moose
Research Center, bringing the total number marked to 321. Two hundred
fifty-one resightings of collared moose occurred, bringing the total to
664. An analysis of migrations and population mixing of moose on the
Kenai Peninsula and a review of moose migrations in North America were
published.
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Summary ..
Background.
Objectives.
Procedures.
Findings ..
Recommendations
Literature Cited.
CONTENTS
BACKGROUND
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33
33
Moose (Alces alces) in the lowland areas of the northern Kenai
Peninsula receive considerable hunting pressure in the few restricted
areas where access exists. In late fall, moose herds in these areas
characteristically have a low proportion of bulls, and trophy-size bulls
are extremely rare. Although lowland areas contain a higher proportion
of calves within the herd, calf production in some years is lower than
anticipated (eg: 33 calves:lOO cows in November 1970). Most of the
area in question is sera! birch (Betula papyrifera) range remnant from
the 194 7 burn, and birch browse is in great abundance. However,
substantial numbers of moose have died during severe winters in the area.
Population estimates by personnel of the Kenai National Moose Range
suggest substantial numbers of moose (7900 ± 1400 minimum north of the
Kasilof River in early 1971), but concern has been expressed regarding
the numbers and welfare of the "lowland" moose, especially in relation
to hunting pressure.
The moose traditionally using climax willow (Salix sp.) ranges in
foothills and mountains, but wintering on the lowlpnd areas, receive
little hunting pressure. These groups characteristically exhibit a high
bull:cow ratio and a low proportion of calves.
With the formalization of moose management plans for the Kenai and
the designation of certain areas as trophy, foot-hunting and maximum
sustained yield hunting areas, delineation of these various groups, their
interactions, their seasonal movements, and their calving and breeding
sites, has become imperative. Further, the proposed classification of
more than one million acres of the area as wilderness, as well as the
possibility of a limited access road bisecting part of the area, require
specific knowledge of the migrations of these moose. Descriptions of
populations and their movements would 1) allow harvesting of desired
portions of specified moose herds and prevent harvesting of trophy-class
bulls while they are away from trophy-management areas (and often
antlerless), 2) prevent unnecessary restriction of activities (eg: by
wilderness designation) in areas of key winter range, where habitat
manipulation might someday become necessary, 3) contraindicate develop-
ment of small areas seasonally crucial to large numbers of moose (eg:
during calving, rutting, or wintering) and 4) provide valid data relative
to possible obstructions presented by future proposed highways and other
projects,
1
The literature contains few major studies of moose migrations and/
or movements, and the studies that have been undertaken have shown that
such movements vary with the population studied. Goddard (1970) reported
an Ontario study similar to ours. His recoveries were few (59 of 328
marked moose) but he documented movement from summer to winter ranges
(done previously by Edwards and Ritcey, 1956; Kraft, 1964; and Houston,
1968) and suggested there was no net movement into heavily hunted areas.
Phillips and Berg (1971), with many relocations (2,000) of few (27)
radioed Minnesota moose, recorded individual horne ranges of 2-10 square
miles, winter confinement to less than 100 acres, average daily movement
of 0.60 miles, identical mean daily movements of cows and bulls, and 0.5-
21 mile movements from winter to summer ranges. VanBallenberghe and
Peek (1971) also radio-tracked moose in Minnesota. They showed summer
localization, winter confinement by snow, adjacent winter and summer
ranges of an individual, and a rapid 12-rnile movement by a rutting bull.
Mercer and Kitchen (1968) described dispersal of moose introduced onto
the Labrador Peninsula. LeResche (1968) and LeResche and Davis (this
report; Job 1.2R) reported localization of parturient females and their
new calves, and LeResche (1970) suggested internal triggering as a
factor in moose migrations. Bishop (1970) reported that a Tanana Flats
(Alaska) calf-tagging study suggested that both resident and migratory
individuals were present in these lowlands in spring. Didrickson (pers.
comm.) reported adult moose tagged in the Matanuska Valley (Alaska)
moved nearly 60 miles on occasion.
OBJECTIVES
To identify populations and key habitat areas and to learn seasonal
patterns of movement by moose on the Kenai Peninsula.
PROCEDURES
Table 1 lists moose marked during the reporting period. Except
where noted, moose were captured in fenceline traps (LeResche and Lynch,
1973), marked and released. Thirty-eight reconnaissance flights made
during the period are listed in Table 2.
FINDINGS
Resightings of marked moose are listed in Table 3. LeResche (1972)
prepared a summarj and analysis of resightings through June 1971. No
further analysis is available at this time.
A review prepared as part of this job is reproduced below. It 't·lill
be published in Le Naturaliste Canadien in autumn, 1973.
2
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---------------------------------·--··--------·--
Table 1. Sex, age, location, and markings of moose tagged on the Kenai
Peninsula from July 1, 1971 through June 30, 1972.
Males Females Sex? Calves Total
Moose Research Center 5* 30** 0 2*** 37
Miscellaneous**** 1 1
Total 38
* Two males marked with quad-colored collars; three with solid blue;
all had silver Saflag in left ear.
** Twenty-two females marked with white collars; eight collared with
quad-colored collars; all had silver Saflag in right ear.
*** One male calf tagged with silver Saflag in left ear; one female
calf tagged with silver Saflag in both ears.
****This cow was mired down in the mud near Hidden Lake. Tagged when
rescued.
3
Table 2. Reconnaissance flights by Alaska Department of Fish and Game search-
ing for collared moose July 1, 1971 through June 20, 1972.
Date
1 Sept 71
14 Sept 71
1 Sept 71
23 Sept 71
8 Nov 71
9 Nov 71
8 Nov 71
8 Nov 71
12 Nov 71
3 Jan 72
3 Jan 72
3 Jan 72
3 Jan 72
18 Jan 72
18 Jan 72
18 Jan 72
11 Feb 72
11 Feb 72
3 March 72
3 March 72
3 March 72
3 March 72
29 March 72
29 March 72
29 March 72
29 March 72
7 April 72
7 April 72
7 April 72
7 April 72
7 April 72
19 April 72
25 April 72
15 May 72
15 May 72
14 June 72
14 June 72
14 June 72
Area
Mystery Creek
Mountains between Big Indian Creek
and Thurman Creek
Moose River Flats
Mystery Creek
Swan Lake Canoe System and vicinity
Moose Lake vicinity
North of MRC
North side Kenai River west of
Skilak Lake
East of Bear Lake (GMU 15A)
Slikok-Funny River Road
Tustumena-Skilak Benchland
Bottenintnin Lake-Skilak outlet
Mystery Creek Basin
Swan Lake Canoe System-MRC area
MRF
Skilak Loop-Skilak Lake outlet
Mountains north of Thurman Creek
Sterling Hwy Mile 60-Mile 100
Sterling Hwy Mile 100-Mile 75
Swanson River Road-MRC area
Eastern MRF area and Hystery Creek
Skilak Loop
Tributaries on west side of
Resurrection Creek
Chickaloon River and Big India Creek
Skilak Loop
Mile 62-90 Sterling Highway
Slikok Lake
Moose Creek (Tustumena Lake)
Funny River-Killey River
Skilak Loop
Sterling Highway Mile 75-90
Mountains north of Thurman Creek
Skilak outlet area
Moose River Flats
Bottenintnin Lake
Swan Lake Road-MRC area
Moose River Flats
Tustumena-Skilak Benchland
Total
*Code: Tagged at Moose River Flats: MRF
Tagged at Moose Research Center: MRC
Tagged at Bottenintnin Lake: BL
Tagged at Tustumena Benchland: TB
Part of markings lost so exact
tagging site unknown: Unk
4
Collared Moose Located*
6 MRF, 1 Unk
16 MRF
1 MRF
1 BL, 5 MRF
1 Unk, 1 BL, 6 MRC, 3 MRF
1 MRF, 1 MRC, 1 Unk
2 MRC, 1 Unk
1 Unk
1 BL, 4 MRF, 2 MRC, 1 Unk
0
0
1 MRF, 3 Unk
0
1 MRC, 1 Unk
0
1 MRF, 1 BL, 1 Unk
2 MRF
2 Unk
0
1 MRC
0
1 MRC, 2 MRF
1 MRF
6 MRF
1 BL
0
0
1 TB
0
1 MRF, 1 Unk
2 Unk
1 MRF
1 Unk
1 MRC, 26 MRF, 3 Unk
2 Unk
0
3 MRF
0
123
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-------------------
Table 3. Resightings of collared moose occurring from July 1, 1971 through June 30, 1972.
Tagging
Location
Ski1ak-Tustumena
Benchland
Mystery Creek
1968
Moose River
Flats
Moose Pens
"Bot." Lake
Not Properly
Identified
July Aug Sept
71 71 71
a 9 ~a 9
3 1 1
2 3 5 11 20
1 1
1 3 2
Oct
71
a 9
Nov
71
d 9
2 2
Dec
71
0 9
1
Jan
72
a 9
6
2 5 30 1 1 4
12 4
4 2 3 5
I
1 5 2 2 2 8
Feb
72
0 9
2
Mar
72
a 9
Apr
72
d 9
1
1
5 2 12 1 3
4 2
2 1 1 2
3
June
72
May
72
a 9 a 9
2 1 2
29 5
3 1 3
1
3 1
Total
1
24
141
31
27
27
251
Moose Migrations in North America
by
Robert E. LeResche
Alaska Department of Fish and Game
Fairbanks, Alaska
U.S .A.
Movements of moose (Alces alces ssp.) have long fascinated North
American naturalists and scientists, but have been poorly understood
until recently. Some early naturalists believed changes in abundance
were actually mass migrations to and from large areas (eg. Merrill 1916,
Heape 1931). Others, perhaps correctly, believed that moose they
observed remained life-long in one very small area (Seton 1909, 1929,
Bailey 1930). Still others, unable to discern a pattern in recorded
observations, concluded that moose were nomadic, their movements being
without pattern during most of the year. (Heape 1931). Some observers
recognized patterns of movement, and saw cause-and-effect relationships
between movements and snow, rainfall, food quantity, food quality or
even feces-tainted range (Heape 1931, Hosley 1949, Peterson 1955). Only
recently have more definitive studies in British Columbia (Edwards and
Ritcey 1956), Montana (Knowlton 1960), Minnesota (Berg 1971, Van
Ballenberghe and Peek 1971), Wyoming (Houston 1968), Ontario (Goddard
1970), and Alaska (LeResche 1972) revealed the true complexity of the
migration phenomenon in moose.
Moose populations in North America vary from the extremes of seden-
tary to truly migratory. Every degree of migratory behavior in between
the extremes seems represented on the continent. Differences in
behavioral patterns are certainly correlated with habitat and terrain
occupied, but may also be related to taxonomic form, annual variations
in weather, and population structure. The central questions with regard
to moose migrations are these:
1. How is migratory behavior, or lack of it, related to habitat,
terrain and taxonomy?
2. What are the immediate mediating causes of migratory behavior;
snow, forage, internal stimuli?
3. What role does tradition play in movement patterns of individuals
and populations?
4. What is the adaptive value of migratory behavior where it exists,
ru1d why is it absent from some populations?
This review considers migration in North American moose in light of
these questions. Migratory behavior is defined as movement between
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separate seasonal home ranges, regardless of distance traveled. Local
movements (within a home range) and dispersals into new areas and newly-
created habitat are also discussed.
LOCAL MOVEMENTS
The common denominator of an individual's use of space is its "home
range", considered in this review to be the area in which the individual
accomplishes its normal activities during a given period of time. "Home
range" and related concepts of localized activity have been defined and
redefined as often as any terms in animal ecology and behavior (Burt
1940, Godfrey 1954, Kaufman 1962, Jewell 1966, Mohr and Stumpf 1966).
My present definition ignores the fine points but allows comparisons
between reported moose studies. Local movements occur within the home
range. Home range may shift seasonally, and individuals may occupy more
than one home range in a year. Territories, or defended areas, have
never been reported for moose (Geist 1963) with the possible exception
of cows with calves (Altmann 1958, LeResche 1966).
Size of Home Ranges
Seasonal home ranges of moose are consistently small throughout
North America. Casual observations and formal studies all have concluded
that, regardless of how far moose habitually move between seasons, home
range during a given season seldom exceeds 5-10 km2 • This is true in
areas where moose migrate and where they are primarily sedentary; and in
all habitats studied.
Peterson (1955) felt that moose in Ontario (A. a. americana) frequently
passed a lifetime in an area of 32-800 km2 (a radius of 2-10 mi 2 ). DeVos
(1956) reported summer home range was about 6.0 km2 for a cow with a
calf occupying lowlands in Ontario, and Goddard (1970) characterized
more in similar Ontario habitat as having "relatively small" winter and
summer home ranges.
Telemetry studies in Hinnesota habitat characterized by low relief
and mixed deciduous and deciduous-coniferous forests have confirmed very
small home ranges in both winter and summer (A. a. andersoni). Berg
(1971) and Van Ballenberghe and Peek (1971) reported summer home ranges
averaging 4.6-14.3 km2 (the latter made 75 percent of their relocations
within 2.6 km2) and winter home ranges averaging 2.0 km2. Van Ballenberghe
and Peek (1971) observed an adult female that remained in a 2.4 ha area
for 25 days in January and February. They concluded that "winter home
ranges consisted of a series of (very small) high use areas connected by
wanderings of various distances".
Moose (A. a. shit'aE£) in more mountainous areas of the western
United States, similarly, have small home ranges. Houston (1968) and
Knowlton (1960) studied seasonally migratory populations in Wyoming and
Montana, respectively. In W~oming, 24 of 25 moose studied had summer
home ranges less than 3.8 km , and 35 of 39 had winter ranges this small.
7
In Montana, summer home ranges varied between 2 and 8 km 2 . Stevens
(1970) also felt that moose he studied in another area of Montana were
"relatively sedentary" in winter.
A lees alces g-igas in Alaska have seasonal home ranges similar in
size to other subspecies. Murie (1944) described an adult cow that
spent most of her time throughout the year in 8 km2 near a road. LeResche
(1966) estimated sunnner home ranges of 4-5 km2 for females with calves
in one migratory population. Another migratory population had summer-
fall above timberline home ranges of less than 32 km2, and an apparently
resident lowland population occupied home ranges year-round of less than
20 km 2 (LeResche 1972 and unpubl. data).
Sex, Age and Home Range Size
Collectively, studies to date indicate that calves and their attend-
ing cows have smaller home ranges for a short time following parturition
than do other moose, and that yearlings have larger home ranges than
other moose. Home ranges during rut seem to vary with the sex structure
of the population.
Summer home ranges of cows with calves in various areas are
sun~arized in Table 1. Altmann (1958), McMillan (1954), LeResche (1966),
LeResche and Davis (1971), and Berg (1971) documented extremely small
home ranges of cows and newborn calves in Wyoming, Alaska and Minnesota.
These home ranges seem to increase beginning when the calf is one to
three weeks old (Berg 1971, LeResche and Davis 1971), until summer,
summer-fall and winter home ranges are equivalent to those of cows and
bulls. Houston (1968) and Saunders and Williamson (1972) could show no
differences between size of seasonal home ranges of cows and bulls in
Wyoming and Ontario. McMillan (1954), Peek (1962) and Berg (1971) found
no differences between eventual summer home ranges of cows with calves
and cows without calves. LeResche (1966) did note such a difference,
but this was likely due to the fact that cows without calves began their
fall migration during his observations. Knowlton (1960) suggested from
scant evidence that bulls' summer home ranges were about four times as
large as those of cows with calves.
Knowlton (1960), Houston (1968), Berg (1971) and LeResche and Davis
(1971) showed that, during rut, bulls' movements on summer-fall range
became erratic, and home range increased accordingly. This change is
more properly considered a transition into another area of activity
(i.e.: "rutting home range") than a true change in summer home range.
Rutting home range of females may be a function of their abundance in
the population relative to bulls (Houston, 1968). This idea is
substantiated by my observations in areas of Alaska with very few bulls
(circa 10-20 bulls:lOO cows). Cows in these areas become much more
active and mobile during rut than those described by Houston in Wyoming,
where bull:cow proportions approached 81:100.
Data on yearling home ranges are limited and contradictory, but do
suggest they may be larger than those of older moose. Seton (1909)
believed young bulls wandered erratically over a wider area than mature
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Table 1. Home ranges of moose cows with calves. Modified from Berg,
1971.
Home Range
Time Interval Area (km2) Reference
Montana 6 July-23 September 2.2 Knowlton 1960
Wyoming 6 June-15 September 5.1 McMillan 1954
Ontario 15 August-31 October 6.0 DeVos 1956
Minnesota 11 June-22 August 5.9 Van Ballenberghe
Peek 1971
Minnesota 15 May-1 October 15.4 Berg 1971
Minnesota 15 May-1 October 16.9 Berg 1971
Alaska 6 June-31 October 8.4 LeResche 1966
9
&
animals. Houston (1968) maintained yearling moose in Wyoming had no
"established" home ranges and moved further and more erratically than
older animals. Pimlott (1959) and Simkin (1965) showed that, in
Newfoundland and Ontario, yearlings were more vulnerable to hunting than
other moose, and suggested larger home ranges might be a contributing
factor. However, Saunders and Williamson (1972), in Ontario, found no
significant difference related to age in linear movements of tagged
moose.
Especially in light of the contrast between individual constancy of
home ranges in adult moose and the species' tendency to disperse and
explore (below), it seems logical that young moose should have larger
home ranges (or broader local movements) than older animals. This
remains to be confirmed by more precise studies, however.
Tradition and Home Ranges
Moose in all areas of North America tend to return to the same
individual home ranges year after year. Since Seton (1909) spoke of
moose's "judgement founded on their memory of the country", study after
study has confirmed traditional use of the same areas by the same
individuals, and suggested that patterns persist over several generations.
DeVos (1956) observed a tagged calf returning as a yearling to the
area of its birth after a winter's absence. Knowlton (1960) noted that
the same concentration areas were used each winter in mountainous areas
of Montana. Geist (1963) recognized that individual moose returned each
year to their "accustomed summer range" in British Columbia.
Houston (1968) confirmed strong individual tradition among Wyoming
moose. His study included both Type B and Type A populations, and
demonstrated that marked individuals from the Type A group returned year
after year to permanent summer home ranges. He believed that the Type B
group, similarly, had permanent summer ranges. Specific winter home
ranges were used annually by members of the Type A population. Type B
animals were apparently less attached to specific winter ranges, although
still traditional to a great extent. Twenty of 33 (61%) marked animals
returned to established home ranges for at least two consecutive years.
Five others (15%) returned only in alternate winters, and the remaining
24 percent had not returned within one or two winters. Houston felt
that the severity of winters influenced the return of the Type B group.
Berg (1971) conclusively demonstrated that radio-marked moose in Hinnesota
lowlands returned to their established sununer-fall and winter home ranges
for at least two consecutive years. Each individual (males and females
were included) migrated annually between two established home ranges.
Studies in Alaska since 1963 have confirmed strong traditionality
in many areas by following marked calves and adults (Bishop 1969a, 1969b,
LeResche and Davis 1971, LeResche 1972). This traditional use is
demonstrated both by populations (Fig. 1, 2) and by individuals (Fig. 3).
Figure 2 shows seasonal locations over a 2 1/2 year period of collared
moose from groups marked at various times of year in different areas of
the Kenai Peninsula (Fig. 1). The figure includes moose from three
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-··--··--·---··-·-----------------·--·· ·-·-·-I
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,...~---·-.....,
/ Bench land ) . '--. ---. ....,_, . ,.,
KENAI PENINSULA
Fig. 1. Locations of seasonal ranges on the Kenai Peninsula, Alaska.
11
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G.
oosrAv£0 •r ·D"~>•••r c-. r.:)< •••• A.
~Mooot A.
(;!IDSiollt' lMP
C)<e ... l MIO
oLE~~~~_L~~
.J»>· MAR-MAV· .Ill.· KP• P¥Jol•
FUI -~ IIUO OCT D£C
M()(7/h
25 08SEAVED _,, c.
o\T:
,.
50 b. OOS.£11vt:D
OBSER\1£0 .. T:
~Moo .. Rloer
c:J194? .....
•r·D~··"'' Loop
&1'/.lJ•'lH '"'"
L::)(oNII loUt.
c=, ........
I<IJD6.~~ehl-
d.
o._ .... ~,.,..,..,._-, ... :":7;"~."":-: ... ~>tr"· ..A.::.IIk-=-· -'::w~,:-.""":N(N=-,u ...,... JuN *10 OCT ore
Monlll
Fig. 2. Bimonthly locations ot groups of moose tagged
at four locations on the Kenai Peninsula,
Alaska: (a) marked at Mystery Creek in
October, (b) marked at Skilak Loop in March,
(c) marked at Moose River in May and June,
(d) tagged at 1947 burn in all months.
(From LeResche 1972).
12
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0
t
0
Moose ·
Research
Center
10
Skilak Loop
5 Miles
I
8 Kilometers
6-f . --~'
6 0 10 I T 7 \~ Mystery Creek
67 Kenai River
Fig. 3. Migration patterns of two male and two female moose
over a two-year period on the Kenai Peninsula, Alaska.
"T" is tagging site, digits are monthly resighted
(!=January, 12=December), "cc" indicates two calves
present. (From LeResche 1972).
13
migratory groups (a, b, c) and a predominately resident population (d).
Each group concentrated annually in the area they were marked. At other
times of the year, some groups were widely dispersed. Groups a, b and c
represent animals with traditional autumn (rutting), winter and early
summer (calving) ranges, respectively.
Individuals of each group had fixed spring, summer and autumn ranges
(Fig. 3). These respective ranges were occupied at rather constant time
intervals for the period of the study (LeResche 1972).
MOVEMENTS BETWEEN SEASONAL HOME RANGES
"Migration" is most commonly taken to mean regular annual movements
of some distance, involving return to at least one common area during
each annual cycle. The many regular shifts between seasonal home ranges
reported for moose populations in North America are true migrations in
this sense. Much confusion has arisen because not all populations are
migratory, and because migratory and resident groups not infrequently
occupy common areas during part of the annual cycle, and sometimes during
rut. The interactions between migratory populations (or population
segments) and resident populations (or segments) lead to seasonal aggre-
gations and segregations, and therefore affect habitat relationships and
population dynamics. Any consideration of the significance of migrations
to moose populations must consider these factors.
All seasonal movement patterns reported for moose in North America
represent one of three general types (Fig. 4): Type A; short distance
movements between two seasonal ranges with little change in elevation;
Type B; medium to long distance movements between two seasonal ranges
with significant differences in elevation between high summer-fall
ranges and lower winter ranges, and; Type C; medium to long distance
movements between three distinct seasonal ranges with significant differ-
ences in elevation between high summer-fall ranges and lower winter and
spring ranges. These types do not represent absolutely distinct patterns,
but rather form a continuum, (Table 2). These patterns differ in timing
and regularity, distance traveled, change in elevation involved, and
number of seasonal home ranges visited annually. All represent seasonal
oscillations between at least two areas or habitats, however.
Timing and Regularity of Movements
Timing varies between individual populations, and probably reflects
differences in general seasonal climatic changes and habitats. In
addition, timing varies year-to-year in some populations, perhaps
reflecting annual climatic differences.
Localized (Type A) populations in northwest Minnesota moved 2-6 km
from summer-fall to winter ranges in late November, and all returned
within a 10-day period in April in one spring (Berg 1971). In similar
habitat in northeast Minnesota, moose occupy a series of high-use winter
areas beginning in January, and wander short distances between these
areas until May-June, when they become localized on summer ranges
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0
r-<t > w
_J
w
Transition 1------1----+-----2----
t----3 ----+-----1 ---+---2---
1-Summer/Fall
Home Range
2-Winter Home Range
3-Spring Home Range
MAY-JUN JUL-AUG SEP-OCT NOV·OEC JAN·FEB MAR-APR
TYPE A
TYPE 8
TYPE C
Fig. 4. Types of moose migration patterns reported from North America.
Broken line in Type A representation indicates periods of move-
ment. "Transition" period in Type B may be longer or shorter
than shown, and represents winter home ran.ge habitat.
15
Table 2: Summary of seasonal movements of moose in North America. Because methods of study and reporting results vary, designation of migration
type is necessarily arbitrary. Designations are made to help structure consideration of the many separate situations represented.
Location (study)
N.E. Minnesota
(Van Ballenberghe
and Peek, 1971)
N.W. Minnesota
(Berg, 1971)
Wyoming
~-' (Houston, 1968)
"'
British Columbia
(Edwards & Ritcey,
1956)
British Columbia
(Edwards & Ritcey,
1956)
Summer/Fall range
(elevation-m)
coniferous-deciduous
mosaic (500-700)
open willow
floodplain willow
(2200-2300)
subalpine forests
(2400)
lowland burns
(500)
Tanana Flats, Alaska lowland mosaic
(Bishop, 1969) (200-250)
Kenai, Alaska lowland mosaic
(LeResche, 1973) (75-100)
------
Distance
between
(km)
local
(2-6km)
2-lOkm
8-16km
local
?
local
0-16k!"_
Winter range
(elevation-m)
Distance
between
(km)
Spring (calving)
range
(elevatfon-m)
coniferous-deciduous local
mosaic ( 500-700) ( 2-6km)
mature willm-1/
hardwoods
floodplain willow
(2200)
subalpine forests
(2400)
lm.;land burns
(500)
lo~{land burns
(200-250)
lowland burns
(75-100)
2-lOkm
8-16km
?
?
local
?
local same as summer-
(0-16km) fall
-- -----
Distance
between
(km)
Migration
type Remarks
A
A
A
A
A
A
A
Local shifts in winter
home range depending
upon weather
Definite, rapid but
short-range seasonal
shifts-no vertical
component
Share winter-spring
range with larger
Type B group (below)
Both groups represent
small resident segments
of a much larger Type B
population
See above
Share spring-summer &
some winter range with
TvPe C population
Share spring & some
winter range with Type
C population {below)
- -----
---- ------
Table 2: Continued
Location (study)
Northwest Territories
(Barry, 1961)
Arctic North Slope,
Alaska (Alaska Dept.
Fish & Game, 1973)
British Columbia
(Edwards & Ritcey,
1956)
Montana (Knowlton,
1960)
Kenai , Alaska
(LeResche, Unpubl.)
(benchland residents)
Wyoming (Harry, 1957;
Houston 1968)
Summer/Fall range
(elevation-M)
Distance
between
(krn)
Winter range
(eleva ti on-m)
tundra
( o:!:.)
to 170km riparian willow
?
riparian willow &
tundra (100-200)
0-75km
subalpine forests 20-60km
(2400-2500)
subalpine meadoi?S & 2-7kn
upland willoHs ( 3000)
alpine tundra & 2-20km
upland willm-1s
1000+
upland forest 15-40km
{3300)
riparian willm-r
( 200-600)
lowland burns
(500-700)
riparian willow
(2000)
riparian willm-1
( 500-800)
floodplain willow
(2200-2300)
-
Distance
between
(km)
to 170km
0-75km
20-60km
l-7km
2-20km
15-40km
--
Spring (calving)
range
(elevation-m)
-
Distance
between
(km)
-- ---
Migration
type Remarks
A-B
A-B
B
B
Move along rivers long
distances, with little
change in elevation
Move along rivers long
distances, with little
change in elevation;
some cross crest of
Brooks Range through
high passes
Share winter & spring-
summer ranges with two
Type A populations
(above)
Very short movement;
but great elevational
difference
B Share spring-summer
range with much larger
Type C group (below)
B Type A in some years,
depending upon snow.
Shares winter range
with smaller Type A
group (above)
Table 2: Continued
Distance Distance Sprin!! (calving} Distance
Summer/Fall range between Winter range between range between Migration
Location (study) (elevation-m) (km) (elevation-m) (km) (elevation-:r.J) (km) type Remarks
Kenai , Alaska upland •~i llows 30-40km lowland burns 20+km lowland bogs 13-20km c Share ranges with
(LeResche, 1973) (600-750) (80-100) (80±_) groups of Types A &
A-B (above)
Tanana Flats, Alaska upland willows 30-SOkm lowland burns 20+km lowland bogs 15km c Share ranges with Type
(Bishop, 1969) (1000) {200-500) ( 200-300) A group (above)
---- ----------- - -
--
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(Van Ballenberghe and Peek 1971). Wyoming moose, representing both
Type A and Type B groups, move onto winter ranges from mid-December
to mid-January; then return to spring-summer ranges from late March to
mid-April (Houston 1968). Type B groups have left this range by mid-
June. More northerly groups with Type A movements (Kenai. Alaska;
LeResche 1972 and Tanana Flats, Alaska, Bishop, 1969) follow roughly
the same schedule in their local movements.
Timing varies considerably annually and regionally among Type B and
Type C groups. Moose in mountainous British Columbia move rapidly from
late winter range to summer range in April (Edwards and Ritcey 1956).
In one year, the migration was complete by mid-April. In other years it
was still in progress in May, a lull in movement (attributed to cold
weather) having occurred in the interim. In contrast, movement from
upland summer ranges to lower wintering areas is normally a gradual
process in this population, occurring from mid-September until abrupt
upward movement begins again in April. Progression of the migration
varied between years, perhaps due to differences in snow accumulation.
In Montana, moose from a Type B group first arrived at lowland
winter concentrations between mid-December and mid-January, and continued
to drift slowly onto winter ranges as late as March (Knowlton 1960).
Return to summer range is more rapid, occurring mostly in May and June.
Two Type C populations in Alaska return to summer-fall ranges later
than moose in Montana and British Columbia, and remain on them for a
shorter time. Movement to upland summer-fall ranges is gradual, in
contrast to that in Montana and B. C., and is not complete until rut,
in late September and October. Conversely, movement from upland ranges
to winter ranges is sudden, and may occur as early as mid-November.
This, too, contrasts with the slow winter-long descents described in
Montana and B. C.
l1ovements in Interior Alaska (Tanana Flats) are similar in all
respects to those at Kenai, except that ascent to summer-fall areas may
not be detectable until as late as early September. In both locations
in Alaska, timing of the precipitous fall downward migration varies
year-to-year, apparently depending upon snowfall. In at least one
instance at Kenai, major downward movement was delayed until January.
Environmental and Physical Distances between Seasonal Ranges
Distances between seasonal ranges can be considered as horizontal
distances, elevational distances, or environmental distances. To be
adaptive, movements must involve a change in environment, or travel an
"ecological distance" from one environment to another. As suggested by
Edwards and Ritcey (1956), a change in elevation accomplished by a
relatively short horizontal movement may result in environmental changes
similar to those resulting from extremely long latitudinal migrations.
In regions where juxtaposition of disparate habitats is extreme, only
short horizontal movements may be necessary to realize the maximum
environmental changes practicable. In vast flat areas of homogeneous
habitat, no reasonable migration can achieve a significant environmental
change.
19
Seasonal migrations reported in North America involve horizontal
movements of from one to more than 170 km, and include examples of all
distances in between. No distinct relationship--direct or inverse--is
evident between elevational shift and horizontal movement. Rather, the
horizontal and elevational distances traveled are linked to environmen-
tal differences achieved at each location studied.
The shortest horizontal and elevational distances traveled occur,
by definition, in Type A populations. In low-relief Minnesota habitats,
distances between summer and winter home ranges are only 0-10 km,
although definite and regular seasonal movements do occur (Van Ballenberghe
and Peek 1971, Berg 1971). Of the 11 winter home ranges documented by
Berg, three were adjacent to, and two were included within summer-fall
home ranges of the same individuals. No spectacular advantages accrued
to the moose that moved these short distances, for habitat was similar
(though not equivalent) on both ends of the movements.
Type A movements in other areas, although covering only short
horizontal and elevational distances, may involve large ecological
changes. Resident moose both at Kenai and at Tanana Flats, Alaska,
redistribute seasonally on lowland ranges, moving only a few kilometers.
In so doing, however, they move from open bogs to mature hardwood
forests, dense sera! shrub ranges, or riparian willow habitats (LeResche
et al., this volume).
The two northernmost populations listed in Table 2 (Northwest
Territories and Arctic North Slope, Alaska) both spend summer on tundra
and intertidal ranges near the Arctic Ocean. Both groups undergo medium
to long distance horizontal movements along river courses. These move-
ments have small elevational components, but involve major ecological
change, from exposed areas where survival in winter might be difficult
(Miller et al., 1972), to riparian willow habitat which offers more
available forage in winter and protection from weather. These movement
patterns are considered as intermediate between Types A and B.
Type B movements, which involve a change in elevation and nearly
always a rather major ecological change, may involve as little as a 2 km
horizontal movement (Table 2). These movements are characteristic in
mountainous terrain, and typically proceed between high subalpine or
upland forests (summer-fall range) and low sera! shrub or riparian
habitat (winter range). The longest horizontal movements, in British
Columbia and Wyoming, are 60 km and 40 km, and also involve the greatest
change in elevation (Edwards and Ritcey 1956, Harry 1957, Houston 1968).
The British Columbia population moves from subalpine forests 2400-2500 m
elevation to lowland burns 500-700 m elevation. Migratory moose in
Jackson Hole, Wyoming, move from upland forests as high as 3300 m to
flood plain willow communities 2200-2300 m elevation. Similar ecological
movements are accomplished with shorter horizontal and elevational move-
ments by moose in Montana and Alaska (Knowlton 1960, LeResche unpubl.).
In the Gravelly Mountains, Montana, moose move between subalpine meadows
and upland willows at 3000 m and riparian willow about 1000 m lower,
while traversing only 2-7 km horizontally in the steep terrain. In the
Kenai Peninsula "benchland'', a group of moose moves from subalpine
20
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willows and alpine tundra at 1000 m elevation to riparian willow
communities at 500-800 m, traversing only 2-20 km horizontally. This
group sometimes oscillates between the habitats (depending upon whether
bare ridges exist above timberline) several times in winter, in a manner
similar to that described by McDowell and May (1942) in Montana.
Both Type C populations in Alaska move elevationally 500-800 m and
horizontally 30-50 km between summer-fall and winter ranges; and 0-300 m
and 20 km between winter and spring ranges. These movements are impor-
tant environmentally in terms of habitat (Table 2) and climatological
variables (Coady, this volume).
Traditional Routes of Travel
Because specific home ranges are used repeatedly by individuals, we
might expect routes of travel to be similarly traditional. The little
data available suggest this is the case. Knowlton (1960) described the
following of drainage patterns and Houston (1968) diagrammed routes
repeatedly used which also followed topographic features. Edwards and
Ritcey (1956) believed that migrations retained the same essential
elements year after year.
Berg (1971) and LeResche (1972) present evidence that individual
moose repeatedly use the same routes of travel between seasonal ranges.
In Minnesota, Berg radio-tracked a cow and a bull and found that each
followed the same routes during spring and fall movements. LeResche's
information from Alaska is partially included in Figure 3, which illus-
trates movements of three individuals over 2 1/2 years. The individual
moose were observed often enough during seasonal movements to suggest
that routes they followed were at least very similar from one year to
the next.
Segregations, Aggregations and Seasonal Movements
Seasonal movements of moose. alter the spatial relationships between
individuals and groups, as well as redistributing moose in relation to
their habitats. When animals of different ages, sexes, and reproductive
conditions move differentially, migrations cause a reordering of group
makeup. When migratory and "resident11 groups converge or diverge,
aggregations and segregations occur. The social and demographic changes
resulting from such interactions are often overlooked when causes of
migrations are sought.
Population segments may differ in timing and extent of seasonal
movements, or in whether they migrate at all. Hany studies have shown
that 11 migratory" and "resident 11 populations may share the same ranges
at any time of the year. Houston (1968) studied a ';resident" (i.e.
remaining in his study area year-round) Type A group and a "migratory"
(i.e. visiting the study area only in winter) Type B group in Wyoming.
These moose were aggregated during most winters, but extent of ingress
by the Type B group was related to snow conditions. Type B moose were
about t~.,rice as numerous as resident individuals. Edwards and Ri tcey
(1956) described migrations of a vast majority of moose they studied in
21
-----------------------------------····----·· ------------·-·-·
British Columbia, but noted "a few" moose were year-ronnd residents on
this group's summer and winter ranges. A similar situation is present
in the Tanana Flats, Alaska (Bishop 1969b), where one Type A group
remains in lowlands all year (but redistributes seasonally), and a
larger Type C group moves between various upland ranges and the lowlands.
Both groups aggregate on lowland spring calving ranges. Another small
segment of the population remains in upland areas throughout the year,
redistributing seasonally and mixing with Type C groups in autumn.
"Resident" and migratory groups interact in a complex manner on the
Kenai Peninsula (Fig. 2). The "resident" (Type A) lowland group inter-
mingles with the Type C migratory groups on winter and spring areas.
The Type C groups, in addition, congregate with upland "residents"
(Type B) in summer and fall, on upland ranges. Probably 40-50 percent
of the moose involved undertake Type C movements, about 40 percent are
Type A lowland "residents", and the remaining 10-20 percent have Type B
movement patterns.
Segregations and aggregations assume greater social importance when
they involve distinct age and/or sex classes of the population. Definite
segregations by sex, age, and reproductive status have not been reported
in Type A populations although redistribution of bulls during rut and of
yearlings has been suggested above. Seasonal movements in these popula-
tions seem to involve all classes of animals.
Social segregation is common in some Type B and Type C moose popu-
lations. Peterson (1955) noted that cows, calves and yearlings tended
to move to upland ranges later in the spring than bulls in Montana and
British Columbia. Moose summering in upland ranges in British Columbia
had higher twinning rates, and perhaps higher pregnancy rates, than
those resident in the lowlands (Edwards and Ritcey 1958). This suggests
a differential movement pattern between females of different reproductive
capacities.
Figure 5 illustrates progression of season~l movements by bulls and
cows of a Type C population studied on the Kenai Peninsula, Alaska
(LeResche 1972). Bulls leave lowland spring ranges beginning in May and
June, and all reach upland ranges by early July. Cows without calves
are delayed in comparison to bulls, but precede cows with calves to
upland ranges. Many cows with calves do not leave lowland home ranges
until August, and many never travel as far into the hills as bulls.
Rutting occurs after the bulls drift slightly downward, to elevations
where cows are concentrated. Sexual segregation reoccurs after rut,
when many cows begin downward movement to winter home ranges and most
bulls return to the highest upland ranges. Full intermixing is not
reestablished until all animals congregate on winter ranges, usually by
January.
During spring, summer and early winter, this migratory population
is largely segregated from the Type A group of almost equal size, which
remains in the lowlands. There are significant differences in sex, age
and reproductive success between the Type C and Type A groups. Bull:cow
ratio may approach 60-80 in the Type C groups; whereas, it is 20-40 in
22
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-------------------
N
Lio.l
750
~
f:: 1 ~ 500 c:: I .CJ ......... I ...,.._
~ ~ I
Q_) /~ ' ........... ' lw 274
c:: I \) 250 n) I ·~ ~ --d
0 ._.J_A_Ns:!!MJI-~M-A-R~-~MA~Y -~11133Z!JJ u .... c=_i-~-~~ SEP-1 NOv-
FEB APR JUN AUG OCT DEC
Month
Fig. 5. Mean bimonthly elevations where bulls and cows were sighted in a study of
a Type C moose population on the Kenai Peninsula, Alaska. (From LeResche
1972).
the "resident" group. Type C animals are significantly older (LeResche
1970), and calf:cow ratios are significantly lower in the Type C groups.
These differences result from the more intense hunting that occurs on the
lowlands, but may also reflect changes in movement patterns with age.
Type C groups on the Kenai segregate from ether Type C groups as
well as from lowland residents. Because individuals return annually to
traditional summer-fall ranges, in specific upland drainages, breeding
groups tend to retain separate identities year after year. Many of
these groups contain only 100-500 moose, and are composed of the same
individuals each autumn. During other times of the year, individuals
may be far apart (LeResche 1972).
Mediating Factors in Seasonal Movements
Proximate factors that initiate or halt migrations may be climato-
logical or botanical and may involve "internal clock" mechanisms. Almost
all descriptions of seasonal movements by moose have noted correlations
with one climatological or habitat-related factor or another. Such
presumptive data show that primary mediating factors differ between
situations. There is recent evidence that internal time cues are also
involved.
Weather, and particularly snow conditions, are the most often
reported mediating factor in moose migrations. Formozov (1946) and
Nasimovitch (1955) have reviewed the influence of snow on distribution
and movements of mammals, including moose. Snow depth and characteris-
tics definitely affect local movements and habitat selection by moose
throughout North America, and cause "yarding" in many instances (Seton
1909, Heape 1931, Peterson 1955, Edwards 1956, Harry 1957, Van Ballenberghe
and Peek 1971). Snow-moose relationships vary according to many factors,
which are discussed in depth by Coady in this volume. In general, snow
causes or restricts movements by altering the energy cost of traveling
or living in a habitat, and by altering the availability of forage.
"Rigor of the winter" alters location of winter range of a Type B
moose group in Wyoming (Houston 1968). The migratory group only wintered
in Houston's lowland study area during more severe winters. Knowlton
(1960) believed that winter snow depths in Montana initiate movements
downward from upland summer ranges. Edwards and Ritcey (1956) concluded
that temperature is the "basic factor" mediating Type B movements in
British Columbia, but that snow depth alters the effects of temperature.
Spring movements halt when temperature drops in this region, and resume
with a rise in temperature. However, no upward movement occurs during
warm periods in winter because snow depths prevent it.
Peek (1962) felt that precipitation ~ffects summer distribution of
moose in Montana, where greater use of upl-and habitats occurs in dry
summers.
Abundance and quality of available forage change with snow condi-
tions and with habitat. Hosley (1949: 15) concluded that "depth of snow
and the resulting effect on food availability .... determine the locations
24
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of moose in winter." Stevens (1970) described yarding of Montana moose
due to snow conditions and movements from the yards after available
forage was exhausted.
Springtime movements from winter ranges may be mediated by disappear-
ance of snow or by phenology of growing plants. Probably both factors
are important. The abrupt shift from downward drift to rapid upward
migration in spring described for Type B populations in British Columbia
(Edwards and Ritcey 1956) and Montana (Knowlton 1960) suggests a sudden
release by disappearance of snow rather than a movement initiated by new
plant growth. Knowlton, nevertheless, concludes that "greening up" of
forage initiates spring movements. In contrast, Type C Kenai Peninsula
groups move from winter range onto spring concentration areas in April
and May and then disperse slowly to upland summer-fall ranges beginning
in June. Most snow disappears from upland ranges in May, so this move-
ment is more likely related to plant phenology.
It is impossible to know whether seasonal movements would occur
without proximate climatological and phenological stimuli. However,
there is some evidence of internally mediated migratory behavior in
moose. Sexual stimuli cause movements of bulls preceding and during
rut. Houston (1968) postulated that the extent of movements of cows
might be similarly affected, being perhaps Ha function of the density
of breeding males."
My studies of moose enclosed in four 1 mi 2 (2.6 km2 ) pens near
Kenai, Alaska, indicate that these moose have wigratory tendencies con-
sistent with actual movements of unenclosed moose. Most individuals
were born outside the pens and now are enclosed on the Kenai lowlands
near the interface between a major spring range (where Type A and Type C
populations aggregate), and an extensive lowland wintering area (Fig. 1).
Animals enclosed in the pens are from both Type A and Type C populations.
The movements of unenclosed moose are described above and by LeResche
(1972). Sightings of enclosed moose along fencelines over a 34 month
period were analyzed for directional tendency (i.e.: a moose sighted
along the north fenceline of a pen had a northward tendency, etc.).
Frequency of sightings of moose along north, east, south and west fence-
lines differed significantly by time of year (Table 3), and directional
tendency was consistent with known migrations of unenclosed moose.
During seasons of most significant migrations by unenclosed moose
(e.g.: movement to calving concentration areas in March and April and
migration to upland summer and rutting areas in July-August), locations
of enclosed moose differed most from those predicted by x2 expectations
under a null hypothesis of no seasonal change in distribution (Table 3).
During February-April, within pen locations were weighted north and west
(the direction of movement from wintering to calving areas) to contribute
44 percent to the x2 value. In July and August, when unenclosed moose
were migrating east and south to upland summer rutting ranges, enclosed
moose were seen more often than expected along east and south fencelines.
This deviation from expected values contributed 27 percent to the total
x2 value. Similarly, a westerly trend within the pen contributed 18 per-
cent to x2 during the months (November -January) when unenclosed moose
were gradually moving westward from rutting to wintering areas.
25
Table 3. Proportions of 757 observations of moose along enclosure fence-
lines by season and compass direction. Kenai MOose Research
Center. May 1969 -March 1972.
Fence line
Season North East South
Nov. -Jan. .21 .18 .43
Feb. -Apr. . 36 .20 .24
May -June .15 .33 .31
July -Aug. .07 .38 .49
Sept. -Oct. .18 .33 .43
Total Observations
n.b. Proportions of animals along each fence line
by season (P .001; x2=70.43, 12df).
26
West
.18
.20
.11
.06
.07
Number of
Observations
112
80
150
175
240
757
differ significantly
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During pre-calving, calving and post calving periods (May -June)
and during pre-rut and rut (September -October) unenclosed moose are
essentially resident in calving and rutting areas. During these periods
enclosed moose differed but little from expected distribution patterns,
thereby suggesting no migratory tendency (Table 4).
Enclosed moose are, of course, influenced by the external stimuli
of day length, temperature, plant phenology, etc. Nevertheless, the
directional tendencies they exhibit are appropriate only in light of
patterns (probably learned) cued by internal timing mechanisms. For
example, in autumn the penned moose are already present on winter range,
and are not affected by snow or vegetation on upland ranges. Still,
their directional tendency is in the direction necessary to move from
upland ranges to winter ranges.
DISPERSALS
There can be little doubt that moose have evolved as an opportunis-
tic species, adapted to dispersing into subclimax habitats periodically
created by fires and other disturbances (Geist 1971). This ability is
manifest in prehistoric and more recent extensions of moose distribution
throughout North America (cf: Anderson 1924, Peterson 1955, Mercer and
Kitchen 1968, LeResche et al. this volume, Krefting this volume, Dodds
this volume, Kelsall and Telfer this volume). It is further suggested
by the countless instances of increases and declines in areas of sub-
climax forests (eg. Leopold and Darling 1953, Peterson 1955, Lutz 1960,
Spencer and Hakala 1964).
In fact, actual extensions of range have sometimes been more apparent
than real in recent centuries (cf: Lutz 1960, Barry 1961, LeResche et al.
this volume). Instead, supposed "dispersals" have in reality been only
increases in previously low density populations. Nevertheless, true
range extensions certainly have occurred, sometimes with the aid of
transplants, in Western and Southeastern Alaska (Alaska Department of
Fish and Game 1973, LeResche et al. this volume) and parts of Canada.
It is perhaps a moot point what proportion of large increases in
moose numbers is due to immigration and what proportion results from
increased productivity of resident moose. However, some recent studies
indicate there is at least a short period of delay before apparent
"immigration" occurs into an area of vacated habitat. The basis for
this delay is likely the traditionality of movement patterns discussed
here.
Goddard (1970) concluded from an eight-year study of marked moose
in Ontario that there was no net movement of moose into heavily hunted
areas. He felt that sustained harvest from the heavily hunted area
depended upon enhanced production rather than immigration. We observed
a similar pattern during early years of state management of moose in
Alaska. Intensive hunting along road systems (especially in the
Matanuska Valley and near Kasilof, Kenai Peninsula) significantly
decreased roadside resident (Type A) populations. Late winter densities
27
Table 4. Seasonal directional tendencies of enclosed and unenclosed
moose. Kenai Peninsula, Alaska. 1969-72. "Statistical
expectation" is the expected chi-square value with null
hypothesis of no directional variation by season, and relative
movement is included only when observed frequently exceeds
expected by more than 20 percent.
Season
Nov. -Jan.
Feb. -Apr.
May -June
July -Aug.
Sept. -Oct.
Enclosed Moose
Movement
Relative to
Statistical
Expectation
West
North/West
None
South/East
East
Percent
Contribution
to x2
18%
44%
4%
27%
7%
28
Activity of
Unenclosed Moose
Slow westward
migration to
wintering areas.
Northwestward
migration to
calving areas.
Occupation of
calving areas
Eastward migration
to summer and
rutting areas
Rutting
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remained high due to an influx of Type C moose, but in summer and autumn,
when Type C groups were away, populations were depressed from previous
levels. There was no evidence of a significant influx of outside moose
into the Type A populations. Both these examples suggest a delay in
filling hunter-created niches in Type A populations.
The rate at which newly-created habitat is colonized by moose is
poorly known. Spencer and Chatelain (1953) and Spencer and Hakala (1964)
document increase in moose numbers on the 128,000 ha Kenai (1947) burn
area. Spencer and Chatelain (1953) believe that increase was probably
about fourfold from 1950-1953, and it occurred "largely as a result of
diversion of moose from other wintering areas". Spencer and Hakala
(1964) estimate an increase from 2,000 moose in 1949 to 4,500 in 1955,
which represents an increment of 9 to 21 percent annually. Whichever
of these estimated increases is most accurate, it is entirely possible
that reproduction by previously resident moose on the areas in question
accounted for the entire increase in numbers. Neither publication
reports a concurrent decrease of moose numbers in nearby areas, from
which they feel immigrants originated.
The best information on moose dispersals comes from Labrador (Mercer
and Kitchen 1968). Animals introduced in southeast Labrador in 1953
have since expanded into an area of about 4,400 km2, at a rate of about
10 km per year. Simultaneously, moose occurring naturally in southwest
Labrador spread northeast at the rate of 13 -24 km per year. Rate of
dispersal depends on quality and quantity of habitat available, of
course, as well as on intensity of other limiting factors, such as
disease, hunting and climate. It is interesting to note, however, that
true dispersals (as opposed to increases in density) are often relatively
slow.
THE CENTRAL QUESTIONS
How is migratory behavior related to habitat, ter.rain and taxon~
No precise relationships are evident, for often several general
movement patterns are present in a single population of moose occurring
in one habitat. In general, however, the most complex movement patterns
occur in areas with the greatest compression of disparate habitats and
climates. This situation almost always involves hilly or mountainous
terrain, where different climates and biota are very close to one another.
Only local movements (Type A) have been reported from the flattest
terrain studied, in Minnesota and Ontario. An exception occurs in the
arctic, where long migrations occur in relatively flat areas. Here,
however, summer habitats are probably absolutely uninhabitable in winter.
In Ontario and Minnesota travel over great distances would achieve little
change in climate or habitat. This pattern is confused, however, by
resident (Type A) populations occurring in almost all areas studied--
including those areas where more complex movement patterns (Types B & C)
occur.
29
There is no evidence that any of the four subspecies of North
American moose has consistently different migratory behavior than any
other.
What are the innnediate mediating causes of migratory behavior?
Seasonal movements are mediated by snow conditions, available
forage and a combination of both. There is some evidence for internally
mediated migratory tendency. Autumn movements from summer-fall to
winter ranges are most often attributed to snow accumulation; whereas,
reverse spring movements are associated with disappearance of snow and
"greening up" of forage. Weather changes during migration periods some-
times alter the speed and progress of movements.
What role does tradition play in migratory patterns?
Tradition is strong in movement patterns of individual moose. Move-
ment patterns of populations change over time, but similarly contain
strong elements of tradition. Specific home ranges and specific routes
of travel are used on an annual basis by individuals in all areas studied.
Overall annual movement patterns of populations and population segments
change little year-to-year during periods when habitat relationships
remain relatively stable. The extent and timing of movements vary with
climatological and phenological events, but the pattern remains constant.
The rates and mechanisms of establishing migration patterns in
newly-created habitats are obscure. It is probable that several years
pass before traditional patterns are changed and new patterns established.
Young moose and rutting bulls move most erratically. These animals
might thus be the pioneers of new movement patterns.
What is the adaptive value of migratory behavior?
The result of migrations is alteration of individuals' environments.
Movements change physical (climatic), biotic (forage and interspecific)
and social (intraspecific) relationships. It is within these relation-
ships that we must search for the ultimate benefits accrued from migratory
behavior. Relationships changing seasonally are responsible for annual
movement patterns; those changing over decades or centuries are ultimately
responsible for dispersals. In the final analysis, species seek the
environmental relationship in which they have the greatest reproductive
potential. Movement patterns will thus be selected for if they have
reproductive value (MacArthur 1972). Basic to reproduction is survival,
and basic to survival is energy balance. Movement patterns will thus be
selected for if they place individuals in a favorable energy balance (cf:
West 1960, Cox 1961, 1968). The influences of forage quantity and
quality and climatic variables on energy balance of moose are extremely
complicated but are now at least partially understood (Gasaway and Coady,
this volume, Oldemeyer, this volume, Coady, this volume). In the grossest
terms, the energy advantages achieved by a moose migrating from deep
snow to less deep snow, from non-available forage to available forage,
or from less nutritious forage to more nutritious forage are intuitively
obvious. We can expect more thorough accounting of these relationships
30
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once energy costs and benefits of seasonal environments are measured
precisely.
Social factors are even less well understood than physical and
extraspecific biotic factors of migrations. Their consideration is
clouded by the fact that movements achieving overwhelmingly beneficial
climatic or forage relationships might result in decreasing social
benefits. The converse--social benefits being important enough to over-
ride unfavorable climatic or biotic relationships--might also occur.
Social benefits are presently difficult to measure, but may be sought
in aggregations and segregations that occur as the result of movement
patterns. The most generally apparent aggregations are breeding aggre-
gations and winter concentrations; the most obvious segregations are
post--parturition segregations of cows with calves and segregation of
populations during breeding.
Aggregation for breeding is necessary to reproduction, and all move-
ment patterns must result in this one aggregation. The larger "herd",
which might be intermixed on calving or wintering areas, often splits
into many localized rutting populations. These populations are segre-
gated from one another during breeding. This results in significant
pre-rut movements by some segregated population segments. Several para-
meters, such as summer distribution of population segments and bull:cow
proportions, influence these "gametic migrations," but in all cases
their adaptiveness is obvious. Less obvious are the benefits derived
from segregation of small populations year after year in traditional
breeding ranges. Perhaps such distributions insure best use of "spotty"
habitats (upland riparian willow stands, for example). In addition,
regular return to traditional breeding grounds by the same individuals
insures contact during the rutting season, and perhaps allows efficient
social patterns to develop and persist,
In some reported cases, upland Type A populations aggregate with
Type B or Type C populations, which have migrated from the lowlands, for
rut. These groups might interact for only the late summer-fall period
each year. In contrast, lowland Type A populations are generally isolated
during rut, in lowland areas that Type B and C groups have left. Perhaps
this segregation preserves genetic tendencies for both localized (Type A)
and long-distance (Types B & C) migratory behavior in the same large
area. Localized behavior may be adaptive only when a substantial portion
of a population leaves during part of the year. It may be especially
adaptive to reproducing females; whereas, the benefits of migration might
be greatest to other individuals. Thus, both types of behavior are
retained in the same area.
It is difficult to ascribe social adaptiveness to movements that
result in aggregations on winter range; probably selection pressure for
these patterns originates largely in climatic and forage advantages.
Nevertheless, aggregations do occur in late winter (cf: Peek et al. ms)
in large expanses of good habitat where clumping is probably not required
by snow or forage conditions. Social advantages of these groupings
might perhaps be sought in predator-prey relationships or socialization
of short yearlings before their rejection by cows.
31
Summer segregation of some migratory populations by sex and repro-
ductive status might have social advantages. Part of the adaptiveness
of this pattern certainly derives from localization of cows in relatively
"predator-safe" habitats at a time when travel is di.fficult for new
calves; contrasted with the forage (energy balance) advantages of upland
habitats to animals unencumbered with young. Perhaps social advantage
also derives from minimizing contact between young calves and moose other
than their dams, at the time when their first socialization occurs.
MANAGEMENT CONSIDERATIONS
Moose migration patterns have significant influence on designation
of special-use lands (eg. wilderness areas, recreational areas, highway
routes) and development of hunting seasons and limits. Annual movements
are especi.ally important when they involve travel from inaccessible
areas to roadside areas. Segregation of localized traditional breeding
groups and the slowness of dispersal into social niches further complicates
hunter management. The root of most migration-related management problems
lies in management of land areas rather than moose populations. Their
solutions lie in learning the nature and timing of movement patterns and
the geographical origins of the groups and population segments involved.
Hanagement problems are unique to each situation, but several
general examples will illustrate their nature. Traditionality of
seasonal home ranges and slowness of dispersal into vacated home ranges
greatly influence hunter management along road systems where there are
"resident" populations and in upland drainages where the same individuals
breed repeatedly. In such situations, these groups must be considered
as isolated populations rather than as portions of larger reservoirs of
animals, and must be harvested only within their limited annual increments.
When lands are designated for special uses--to protect certain
moose "herds" or to provide for trophy hunting of large bulls, for
example--it is essential to understand seasonal movement patterns of
all segments of the group in question. Without such understanding,
critical habitats (eg. winter range, calving areas) outside the desig-
nated area might be destroyed, thereby affecting any moose from the
"protected" area that might use them seasonally. Similarly, "trophy 11
areas near "maximum sustained yield" areas are futile if trophy animals
migrate to the latter areas during hunting seasons. Conversely, maximum
sustained harvest management will fail in situations where moose are not
present in accessible areas when hunting is allowed. Benefits from
intensive habitat manipulation for production of moose to be harvested
may be minimal if lands important to the same moose at other times of
the year are not also appropriately manipulated or protected.
Finally, knowledge of moose movements is critical to planning com-
patible human developments. Construction of a single highway or pipeline
barrier across a major migration route could affect moose over tens of
thousands of square kilometers. Misuse of a very small critical seasonal
range for only a month each year could result in the disruption of
thousands of moose over a vast area. These possibilities are not
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hypothetical, but very real in some areas (c:f: LeResche 1972). Where
migrations do occur, managers must realize that free movement is of
major importance to healthy moose populations.
RECOMMEND AT IONS
1. No more moose should be tagged on the northern Kenai Peninsula
other than at the Moose Research Center traps.
2. No further reconnaissance flights should be made for the sole
purpose of locating marked moose, but sightings of marked animals seen
incidentally to other flights should be recorded.
3. Recommendations of LeResche and Davis (1971) should be followed.
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Bailey, V. 1930.
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34
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---------------·• R. H. Bishop and J. W. Coady. 1973. Distribution and
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Con£. (abstract).
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sparrow in relation to migration.
PREPARED BY:
Robert E. LeResche
Game Biologist
SUBMITTED BY:
Richard H. Bishop
Regional Research Coordinator
37
1971. Radiotelemetry studies of
J. Wild!. Manage. 35(1):63-71.
in the energy balance of the tree
Auk 77:306-329.
APPROVED BY: