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Growth in length and weight of northern brown bears: differences between

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Article in Canadian Journal of Zoology · February 2011

DOI: 10.1139/z88-145

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 Growth in length and weight of northern brown bears: differences between sexes
and populations
MICHAEL
C. S. KINGSLEY
Department of Fisheries and Oceans, 501 University Crescent, Winnipeg, Man., Canada R3T 2N6
JOHNA. NAGY
Alberta Environmental Centre, Vegreville, Alta., Canada TOB 4L0
AND

V. REYNOLDS
HARRY
Alaska Department of Fish and Game, Fairbanks, AK 99701, U. S.A.
Can. J. Zool. Downloaded from www.nrcresearchpress.com by Renmin University of China on 05/28/13

Received January 19, 1987

KINGSLEY, M. C. S., NAGY,J. A., and REYNOLDS, H. V. 1988. Growth in length and weight of northern brown bears: differ-
ences between sexes and populations. Can. J. Zool. 66: 98 1- 986.
Growth curves were fitted to data on age, length, and spring weight for individuals from three populations of the brown bear,
Ursus arctos, in northern Canada and northwest Alaska. Females reached 90% of asymptotic length before sexual maturity
and before the age of first production. Their weight remained approximately in proportion to the cube of their length. Males
reached 90% of asymptotic length 0.7 to 1.7 years later than females, and had asymptotic lengths 10- 15% greater. Males
continued their growth in weight even longer, and reached asymptotic weights 80- 100% greater than females. Variation
between these populations was small compared with the total range of variation in the species.

KINGSLEY, M. C. S., NAGY,J. A., et REYNOLDS, H. V. 1988. Growth in length and weight of northern brown bears: differ-
ences between sexes and populations. Can. J. Zool. 66 : 98 1-986.
Des courbes de croissance ont kte ajustees a des donnkes sur l'ige, la longueur et la masse au printemps chez trois
populations d'Ours bruns (Ursus arctos) dans le nord du Canada et le nord-ouest de ]'Alaska. Les femelles atteignent 90% de
la longueur a l'asymptote avant la maturitC sexuelle et avant l'ige de la premikre reproduction; leur masse reste a peu prks
For personal use only.

proportionnelle au cube de leur longueur. Les miles atteignent 90% de leur longueur 0,7 a 1,7 ans plus tard que les femelles et
leur longueur a l'asymptote est de 10 a 15% plus grande. Les miles continuent leur croissance en masse encore plus longtemps
et leur masse a l'asymptote est de 80 a 100% plus ClevCe que celle des femelles. La variation entre ces trois populations est
faible comparativement a la variabilitk totale qui existe chez I'espkce.
[Traduit par la revue]

Introduction
The brown bear, Ursus arctos, now ranges over north-
western mainland North America, including Alaska, the
Yukon Territory, the northern and western Northwest Terri-
tories, British Columbia, western Alberta, and parts of the
northwestern United States; its former range extended further
south and east (Rausch 1963). The species is dimorphic; this
article quantifies size differences between the sexes in three
northern North American populations by comparing growth
curves fitted to spring weight and total length. Fitting a growth
curve, or "size -age curve" (Fitzhugh 1975), summarizes
size-at-age data in a small number of parameters, and eases
comparison of different populations (e.g ., Gros 1980) or of the
same population at different times (Kingsley 1979; Innes et al.
1981)o; seasons (Kingsley et al. 19837 wood et al. 1962) or
under different stresses (Fendley and Brisbin 1977; Zach and
Mayoh 1984).
three populations lived at the limit the range
of the species. Two lived in northern Canada: one on the
Tuktoyaktuk Peninsula and Richards Island on the Arctic coast
of the northwestern District of Mackenzie (Nagy et al. 1983b)
and the other in the Arctic Mountains of the northern Yukon
Territory (Nagy et al. 1 9 8 3 ~ ) The
. third inhabited the Brooks
Range in northwestern Alaska (Reynolds 1980, 1981).
Methods
Study areas
All three study areas were at or near the Arctic coast of North
America. Their locations and topography are briefly described below;
Printed in Canada / Imprime au Canada
more detail is given in Nagy et al. (1983a, 1983b) and Reynolds
(1980, 1981).
The northwest Mackenzie study area included the Tuktoyaktuk
Peninsula and Richards Island, which consist of low-lying open
Arctic tundra on fluvial and deltaic deposits (Mackay 1963). A
southerly extension towards Inuvik included some higher ground,
with open boreal forest and forest -tundra transition.
The other two study areas were mountainous, and more varied in
topography and elevation. The northern Yukon study area lay
between the Trail and Blow rivers in the British Mountains; its
northern limit was 30 km from the Arctic coast. It drained generally
northward via the Trail, Babbage, and Blow rivers, but the extreme
southern part drained south into the Old Crow River. The third study
area lay on the north slope of the western Brooks Range of northwest
Alaska, east of the Kokolik River and drained also by the Utukok
River.
Field methods
The northern Yukon study was carried out from 1972 through 1974,
that in northwest Mackenzie from 1974 through 1978, and that in
northwest Alaska from 1977 through 1980. Bears were caught after
being darted with phencyclidine hydrochloride, alone or combined
with promazine hydrochloride or acepromazine maleate, from
helicopters. Each bear was measured along the spine from nose to tip
of tail, to the nearest centimetre, and weighed with a dial scale
suspended from either the helicopter or a tripod, to the nearest kilo,
gram in the Alaska study and to the nearest 5 lbs (2.27 kg) in the
Canadian ones. A premolar tooth was extracted and subsequently sec-
tioned for age estimation.
Northern brown bears vary greatly in weight between spring and
fall. As this weight variation differs between the sexes (Kingsley et al.
1983), we used only data from bears caught in May and June. This
spring period was also selected by Glenn (1980). In all three studies,
 982 CAN. 1. ZOOL. VOL. 66, 1988

240- MACKENZIE MALES 2 4 0 N YUKON MALES 2401 ALASKA MALES

120
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90

L = L(1-ex ( -k i~-Fi)ll
L = 188
60 k = 0.416

I
0 5
R =
I0
- 1 .03

Rge ( y e a r )
15
I
20
I
25
I
0 5 I0
Rge (year1
15
-
20 25 0 5 10
Rge (year)
15 20
I
25

1 1 1
240 MACKENZIE FEMALES N YUKON FEMALES 240 ALASKA FEMALES
For personal use only.

I I I 1 I
0 5 I0 15 20 25 0 5 10 15 20 25 0 5 10 15 20 25
Rge (year) Rge ( y e a r ) Rge ( y e a r )

FIG. 1. The relationship of length in spring to age for brown bears (Ursus arctos) from northwest District of Mackenzie, northern Yukon, and
northwest Alaska. Growth is quickly completed in both sexes and disparity of final length is small.

some data points were obtained from bears that had been caught and
measured in a previous year or years.
Fitting growth curves
All bears were assigned ages of (n 0.3) years (n is an integer).
Growth curves (von Bertalanffy 1938) were of the forms
and
where l(a) = length (cm) and w(a) = weight (kg) at age a (years);
L = asymptotic (final) length (cm) and W = asymptotic weight (kg);
k = growth-rate constant (year-'); and A = fitting constant (extra-
polated age at zero size) (years).
The growth curves were fitted by the Gauss - Newton algorithm of
the procedure NLIN of the statistical analysis system SAS@ (SAS
Institute Inc. 1982). The data points were not weighted.
Derived parameters and scaling weight on length
The maximum rates of growth (kglyear or cmlyear) and the ages at
which 90% of final size was reached were calculated from the param-
eters of the fitted curves. T o scale weight on length, a corpulence
index (CI; kg/m3) was calculated by dividing the predicted weight at
age (kg) by the cube of the predicted length (m).
Results
The von Bertalanffy curves fitted the data well (Figs. 1
and 2) and examination of residuals did not indicate that any
+ other standard curve should have been used. The asymptotes
were estimated with good precision, especially those for
length, but there were larger errors in k and A .
All populations had similar patterns of growth in weight
(Table 1). Parameter values for the fitted curves were within
one standard error, with the single exception of the female
weight asymptote. Northern Yukon females had an asymptotic
weight about 15%less than those from the other two popula-
tions. Northwest Mackenzie bears, of both sexes, had the
highest k , values and the highest maximum growth rates.
The populations differed more in their patterns of growth in
length (Table 2). Northwest Alaskan males had a greater
asymptotic length than Canadian males, but grew more slowly:
k, was 30% less and maximum growth rate 27% less than the
for Canadian bears, and these bean did not reach 90%
length 6-5 years, compared with 4.5 -4.8
Yean for Canadian bears. Northwest Alaska females had
greater asymptotic length and slower growth rates than their
Canadian counterparts, but the differences were small.
 KINGSLEY ET AL.

I
MACKENZIE MALES N YUKON MALES 240] ALASKA MALES

200
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A 1 - 1 .38

0 5
I
I0
Age ( y e a r 1
I
15 00
4
Age ( y e a r I
20 25
0
0
I
5 I0
1

Age ( y e a r 1
15 20 25
1

I MACKENZIE FEMALES 240

1
N YUKON FEMALES ALASKA FEMALES
For personal use only.

o! I I
0 5 I0 15 20 25 0 5 10 15 20 25
Age ( y e a r 1 Age (year) Age (year1

FIG.2. The relationship of weight in spring to age for brown bears (Ursus arctos) from northwest District of Mackenzie, northern Yukon, and
northwest Alaska. Growth of males is prolonged and disparity of final weight is large.

Northern Yukon females were asymptotically shorter, as well
as lighter, than those of the other populations.
The standard deviations about the fitted lines were about
10 cm in length for all populations, and about 12 kg in weight
for all females; male weights were more variable, especially
for those from northern Yukon.
Corpulence indices for females showed little variation with
age (Table 3). Northwest Mackenzie and northwest Alaskan
females ranged between about 19 and 21 kg/m3; Yukon
females started lower at 17 kg/m3. Mackenzie and Yukon
males became increasingly corpulent with age, reaching values
of 28 kg/m3; Alaskan males reached values of only about 24
kg/m3 and, because of their slow growth in leng,th, did not
reach those values until late in life.
Discussion
Growth curves conveniently summarize size-at-age data in a
small number of parameters but, because they do, they must be
interpreted with caution (Knight 1968). They cannot track tem-
porary variations in growth rate: the equations used here model
the rate of growth as changing smoothly with time. They can-
not be extrapolated beyond the data; in particular, the fitting
parameter A (extrapolated age at zero size) cannot be regarded
as estimating the point in time when growth started, nor the
backward extrapolation of the curve as portraying prenatal or
neonatal growth.
We did not follow the growth of cohorts through time, but
analysed the cross-sectional relationship of size and age within
each population averaged over a period of 3 to 5 years. There-
fore, the fitted curves are valid in population terms rather than
for individuals or cohorts.
The length standard deviation about the fitted curves was
about 10 cm or 5 -6% of the asymptotic length; coefficients of
variation of linear measurements within mammal populations
are seldom less than 3.5 -4 % (e.g ., Rausch 1963; Thomas and
Everson 1982; Bryden et al. 1984; Wiig and Andersen 1986).
In an Alaska Peninsula population of U. arctos Glenn (1980)
found that "all body dimensions exhibited a high degree of
variation beyond that attributable to age," but he presented no
quantitative summaries of variation, only examples. Besides
the variation inherent in wild populations, part of the scatter in
the present study may have been due to the inability of the von
Bertalanffy curves to follow the growth pattern exactly, and
part to measurement error.
Female growth and maturity
Maturity and cessation of growth are associated in female
mammals with the start of reproduction; Laws (1956) asso-
ciated reproductive maturity (in marine mammals; no similar
 984 CAN. J . ZOOL. VOL. 66, 1988

TABLE1. Parameters of growth curves w (a) = W (1 -exp(-~,(u-A,)))~ fitted to data on age and spring weight of brown bears
in the northwest District of Mackenzie, N.W.T., northern Yukon Territory, and northwest Alaska

Max. growth Age (years) No. of SD

W (kg) k (year-') A (years) rate (kglyear) at w = 0.9 W points (kg)*
Males
Northwest Mackenzie 195 (3.2) 0.281 (1 1.8) - 1.38 (32.7) 24.4 (10.1) 10.6 41 16.9
Northern Yukon 188 (8.5) 0.245 (36.2) - 1.04(191.3) 20.5 (29.6) 12.7 24 30.9
Northwest Alaska 187 (5.4) 0.248 (19.9) -1.39 (70.5) 20.6 (15.8) 12.2 44 22.4
Females
Northwest Mackenzie 108 (3.7) 0.410 (16.1) - 1.06 (50.8) 19.7 (14.1) 7.2 45 14.7
Northern Yukon 93 (4.9) 0.371 (24.7) - 1.14 (79.5) 15.3 (21.1) 7.9 24 10.3
Can. J. Zool. Downloaded from www.nrcresearchpress.com by Renmin University of China on 05/28/13

Northwest Alaska 105 (2.8) 0.380 (12.4) - 1.13 (40.8) 17.7 (1 1.0) 7.7 59 12.9
NOTE:Values in parentheses are asymptotic error coefficients of variation in percent.
*Root mean squared deviation of points about fitted line.

2. Parameters of growth curves 1 (a) = L(1 -exp(-kl(a-A,))) fitted to data on age and length of brown bears captured in
TABLE
spring in the northwest District of Mackenzie, N.W.T., northern Yukon Territory, and northwest Alaska

Max. growth Age (years) No. of SD

L (cm) k (year-') A (years) rate (cmlyear) at 1 = 0.9L points (cm)*
Males
Northwest Mackenzie 188 (1.5) 0.416 (13.6) - 1.03 (34.5) 78.2 (12.8) 4.5 42 9.6
Northern Yukon 188 (1.9) 0.375 (43.3) - 1.31 (130.6) 70.5 (42.1) 4.8 31 13.3
Northwest Alaska 198 (1.5) 0.283 (9.9) - 1.63 (19.0) 56.0 (8.8) 6.5 40 9.1
Females
Northwest Mackenzie 170 (1.4) 0.481 (12.5) -0.96 (26.1) 81.8 (1 1.7) 3.8 49 10.6
For personal use only.

Northern Yukon 163 (2.0) 0.479 (33.0) -0.82 (109.5) 78.1 (34.3) 4.0 25 10.4
Northwest Alaska 172 (1.3) 0.375 (10.9) -1.26 (18.2) 64.5 (10.1) 4.8 47 9.7
NOTE: Values in parentheses are asymptotic error coefficients o f variation in percent.
*Root mean squared deviation of points about fitted line.

study seems to have been published for wild terrestrial
mammals) with the attainment of 87 -90 % of final length.
Females in the populations studied here reached 90% of
asymptotic length at ages of 3.8 -4.8 years. Although 4.5- and
5.5-year-old females were seen with attending males, indicat-
ing probable oestrus, ages at which females first bore cubs
averaged 6 - 8 years. The Alaskan bears grew in length more
slowly than the Canadian ones, having lower values for kl,
lower maximum absolute growth rates, and greater 90% ages;
the Alaskan females were estimated to have the greater mean
age at first production, 8 years compared with the Canadian
value of 6 years. It is not clear why the Alaskan bears should
have grown more slowly in length; growth in weight was simi-
lar in all three populations.
Females took 2.9-3.9 years longer to reach 90% of
asymptotic spring weight than to reach 90% of asymptotic
length (Tables 1 and 2). Female Alaska Peninsula brown bears
took 2 years longer to reach 90% of weight than to reach 90%
of length (Glenn 1980). But if growth were isometric, weight
would be proportional to the cube of any length, and any frac-
tion of final weight would necessarily be reached later than the
same fraction of final length. The cubic exponent in the von
Bertalanffy weight curve is based on such a cubic relationship,
so the k values for length can be directly compared with those
for weight; the relation of growth rates in length and weight
can also be studied using a corpulence index.
Even when this correction was applied, female growth in
spring weight was still slower than growth in length, but only
by a small amount. Values of k for weight were 77 - 101% of
those for length (Tables 1 and 2). Corpulence indices of
females did not change much with age (Table 3), rising from
juvenile values of 17-19 kg/m3 to mature values of
2 1 -22 kg/m3. Female brown bears measured in spring on the
Alaska Peninsula had asymptotic length of approximately
205 cm and weight of 190 kg (measured from data plots in
+
Glenn (1 980) ) for a CI of 22 kg/m3. Three 6 -year-old female
brown bears captured in April, May, or June in Jasper National
Park, Alberta, had a mean CI of 22.4 kg/m3 (data from Appen-
dix 2 of Russell et al. (1979)); the mean of 14 CI values for
spring-caught female brown bears (ages unknown) from the
southern Yukon was 18.5 kg/m3 (data from Appendix B of
Pearson ( 1975)) .
Growth in length of female bears was substantially complete
before first reproduction, and growth in weight was nearly iso-
metric with length. Bears of different ages and populations
showed similar relations of weight to length.
Male growth and sexual dimorphism
Sexual dimorphism in mature body size is universal among
the Nearctic bears. In these populations of the brown bear,
males grew to larger sizes than females in both weight and
length, but not by having higher absolute rates of growth. The
maximum rates of growth in length, calculated from the fitted
curves, were less for males than for females; in weight, they
were greater. Males got bigger by growing for longer. Males
did not reach 90% of asymptotic length until 0.7- 1.7 years
later than females, and kl values for males were 75 - 86 % of
those for females. Length dimorphism was 18-26 cm, i.e.,
11 - 15% of female length.
Attainment of large male size by prolonged growth was
more pronounced for weight than for length. For males, k for
weight was 65 -87% of k for length and growth in weight
 KINGSLEY ET AL.

TABLE3. Length at age, weight at age, and corpulence indices (CI) for brown bears of northwest
Mackenzie, northern Yukon, and northwest Alaska predicted from fitted growth curves

Males Females

Length Weight CI Length Weight CI

Age (years) (cm) (kg) (kg/m3) (cm) (kg) &dm3)

Northwest Mackenzie
1
2
3
4
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5
6
8
10
15
20
Northern Yukon
1
2
3
4
5
6
8
10
15
20
For personal use only.

Northwest Alaska
1
2
3
4
5
6
8
10
15
20
NOTE: Corpulence index is equal to weight (kg) divided by the cube of the length (rn)

among males continued longer than growth in length: they
took 5.7 -7.9 years longer to reach 90% of asymptotic spring
weight than to reach 90% of asymptotic length. Growth in
weight among males continued longer than in females, k , for
males being 65 -69 % of the female value.
Other studies on bears have produced similar results. From
plots of spring weight against age for the brown bears of the
Alaska Peninsula (Glenn 1980), it appeared that male weight
had still not reached its maximum value by age 10 years,
although female weight levelled off at about 5.5 years.
Pennsylvania black bears (Ursus americanus), for which Alt
(1980) plotted good, seasonally separated data, also showed
long-continued growth in weight of males, as did New York
black bears (Sauer 1975). Among the polar bears (Ursus mari-
timus) of the Beaufort Sea, males continued to grow in girth
for longer than females (Kingsley 1979).
The result of prolonged weight growth appeared in the
values of the corpulence index and sexual dimorphism at
maturity. The corpulence index for males continued to increase
with age and in the oldest Canadian males reached values of
28 kg/m3, in the Alaskan, 24 kg/m3. Dimorphism in weight, at
78 - 102%, was greater than that in length; a cubic relationship
applied to length dimorphism of 11 - 15% predicts only
37 -52 % weight dimorphism.
The breeding behaviour of bears is not well known, and their
degree of polygyny has not yet been measured directly
(Ramsay and Stirling 1986). Polygyny is common among
mammals (Clutton-Brock and Harvey 1978) and sexual dimor-
phism is positively correlated with degree of polygyny (Ralls
1977; Alexander et al. 1979). Size dimorphism in brown bears
agrees with observations of larger male than female home
ranges in providing circumstantial evidence of polygyny.
Comparison between populations
In predicted weights of both sexes, and in the lengths of
females, the northwest Mackenzie bears were the largest at
most ages, the northwest Alaska bears were second largest,
and the northern Yukon bears were smallest (Table 3). Large
size in the Mackenzie bears may be due to the low elevation
and alluvial soil of the Mackenzie delta, which could lead to
greater productivity and to greater ease of digging out Arctic
ground squirrels (Spermophilusparryi); it might also be due to
reduced density of bears (and hence reduced competition) as a
result of hunting by local people, or to other, unknown factors.
Because of the slow growth of the northwest Alaska males,
predicted male lengths showed a different pattern: at ages of
less than 6 years, the Canadian populations were similar to
each other and longer than the Alaskan one, and at greater ages
 986 C A N . J . ZOOL. VOL. 66. 1988
they were shorter.
These three populations are all at the northern limit of the
range of the species. They range in size by only 4 - 5 % , except
that female weight ranges by 16% because o f the lightness o f
Yukon females. This variation is small compared with the con-
tinental range o f size in the species. Alaska Peninsula bears,
from skull size the largest in North America (Rausch 1963),
had spring weights 9 0 % and lengths 2 1 % (females) and 3 2 %
(males) greater than ours (Glenn 1980). Rausch (1963) sug-
gested that the northernmost populations o f brown bears
increased in body size from east to west, but o u r results d o not
confirm this.
Can. J. Zool. Downloaded from www.nrcresearchpress.com by Renmin University of China on 05/28/13
Brown bears in Jasper National Park in April-May were
+
9 0 kg and 159 c m (females 6 years; n = 3) and 2 0 9 kg and
195 c m (males 8 + years; n = 10) (data from appendices in
Russell et al. (1979)). F o r the interior Yukon, Pearson (1975)
gave mean weights of 9 5 kg (n = 2 1) and 139 kg (n = 4 0 ) for
females and males, respectively, but they were average
weights over the active season for bears 6 + years old, and
were not exactly comparable with asymptotes o f growth curves
fitted to spring weights.
Acknowledgements
W e thank the Canadian Wildlife Service for permission t o
use the data o n the Mackenzie and Yukon populations and the
Alaska Department o f Fish and G a m e for permission t o use the
data o n the Alaska population. Studies o n the Canadian popu-
For personal use only.
lations were supported by the Canadian Wildlife Service, and
the Polar Continental Shelf Project contributed logistic and
helicopter support o n the Tuktoyaktuk Peninsula. Alaskan
studies were supported by the Alaska Department o f Fish and
G a m e , and by Federal Aid in Wildlife Restoration Projects
W-17-11 and W-21-1, Jobs 4 . 1 4 R and 4.15R. W e thank all
those, pilots and others, w h o contributed t o the fieldwork, and
Dr. M . A . Ramsay and other reviewers for their contributions
to the manuscript.
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