American Society of Mammalogists

Allometric Relationships and Sexual Dimorphism in Captive Killer Whales (Orcinus orca)
Author(s): Steven T. Clark and Daniel K. Odell
Source: Journal of Mammalogy, Vol. 80, No. 3 (Aug., 1999), pp. 777-785
Published by: American Society of Mammalogists
Stable URL: https://www.jstor.org/stable/1383247
Accessed: 16-10-2019 04:35 UTC

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 ALLOMETRIC RELATIONSHIPS AND SEXUAL DIMORPHISM IN
CAPTIVE KILLER WHALES (ORCINUS ORCA)

STEVEN T. CLARK AND DANIEL K. ODELL

SeaWorld, Inc., Corporate Zoology, 7007 SeaWorld Drive, Orlando, FL 32821-8097

Morphometric data were collected on 30 captive killer whales (Orcinus orca, 10 males,
females) from 1984 to 1996 at SeaWorld parks to document allometric relationships an
sexual dimorphism. All nine characters examined exhibited negative allometric linear
lationships with total length. Proportional sexual dimorphism was apparent only in le
of flipper and height of dorsal fin among larger males. Absolute sexual dimorphism w
detected in all categories. Anterior and posterior length of flippers for males were be
approximated by a piecewise linear-regression model that suggested a greater rate of gro
in larger males. That may have been the result of onset of an adolescent growth spur
males and perhaps is necessary to maintain hydrodynamic stability. Girth measureme
(girth at anterior origin of dorsal fin, girth at axilla, girth at genital slit) regressed on
length were linear for males and smaller females (<500 cm total length). Increased var
ability of girth measurements and distance between mammary slits in females >500 c
apparently was due to pregnancies. A piecewise linear-regression described growth of do
fins of males. The growth coefficient was greater in larger males. Additionally, adult m
had greater dorsal fin ratios (height:base length) than females. The relationship of dista
from blowhole to tip of dorsal fin regressed on total length was significant. This morp
metric has applications for estimating length of killer whales during photogrammetric s
ies of wild animals.

Key words: Orcinus orca, killer whale, cetacean morphometrics, allometry,

taceans, dimorphism

Examining allometric relationshipsture

andand release programs (Tolley et al.,
sexual dimorphism throughout growth is Most of these studies have examined
1995).
important for understanding life history of cetaceans because it is difficult to
smaller
the killer whale (Orcinus orca). Addition- gather detailed morphometric information
ally, documentation of sexual dimorphism on larger species.
can provide insight into ontogenetic mile- SeaWorld has maintained a captive pop
stones, such as onset of sexual maturity in ulation of killer whales since 1965 (Duf-
male killer whales (Heimlich-Boran, 1986).
field et al., 1995). The dataset for this study
A number of species of cetaceans have been
consisted of 30 animals (10 males, 20 fe-
examined to document allometric relation-
males). Of those animals, 18 were wild
ships and sexual dimorphism. Many of
caught and 12 were captive-born. Avail-
those studies have gathered information
through photogrammetric techniques (Gui- ability of those animals provided a unique
net and Bouvier, 1995; Heimlich-Boran, opportunity to gather information on a large
1986; Ratnaswamy and Winn, 1993; Scott species of cetacean that may otherwise be
and Perryman, 1991), incidental or non-in- virtually impossible to gather in the wild.
cidental fisheries take (Amano and Miya- Our objective was to document allome-
zaki, 1993; Jefferson, 1989; Lockyer and tric relationships and sexual dimorphism in
Waters, 1986), examination of captive ani- killer whales. This information may provide
mals (Anli and Kaiya, 1992), or live-cap- valuable insight into the life history of kill-

Journal of Mammalogy, 80(3):777-785, 1999 777

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778 JOURNAL OF MAMMALOGY Vol. 80, No. 3

Blowhole to of determination (Read et al., 1993; Sokal and

tip of dorsal fin Rohlf, 1995).
Proportional sexual dimorphism (Perrin,
1975) was examined using adult and captive-
born animals and was analyzed with t-tests or
approximate t-tests (depending upon homoge-
Blowhole neity or heterogeneity of the variances, respec-
%. 1
tively) of parameter estimates. Differences were
considered significant at P < 0.05. Specifically,
these analyses investigated if slopes and Y-in-
FIG. 1.-Blowhole to tip of dorsal fin mor- tercepts of regression lines were coincident be-
phometric used in this study as modified from tween sexes. Two-sample t-tests were used to
examine mean differences in size between male-
Heimlich-Boran (1986).
female pairs of measurements -as indication of
absolute dimorphism (Perrin, 1975). Measure-
ments for analyses of absolute dimorphism wer
er whales both in marine zoological parks obtained by averaging measurements for each
and in the wild. adult animal. The average of those measure-
ments was then calculated for each sex within
MATERIALS AND METHODS each category.
Killer whales were measured at 2-4-week in-
RESULTS
tervals. At these times, a suite of measurements
were recorded based upon Norris (1961). In ad- Among adult animals only, absolute sex-
dition, three measurements unique to this
ual dimorphism existed in all measure-
study-height of dorsal fin 1, length from blow-
ments; males were significantly larger (Ta-
hole to tip of dorsal fin, and dorsal-fin ratio-
were collected and are defined below. Variablesble 1). Due to lack of data, proportional
sexual dimorphism of dorsal-fin ratio could
included a variety of straight-line and girth mea-
surements and were as follows: total body not be examined. However, absolute sexual
length, anterior length of flipper, posterior lengthdimorphism indicated that average dorsal-
of flipper, width of flukes, length from rostrum fin ratio of males was significantly greater
to anterior origin of flipper, girth at anterior or-than that of females.
igin of dorsal fin, girth at axilla, girth at genital All allometric growth relationships ex-
slit, distance between mammary slits, height ofamined exhibited negative allometry
dorsal fin 1 (from dorsal fin origin to tip), height
(slopes <1; Table 2). Proportional sexual
of dorsal fin 2 (Norris, 1961), dorsal fin ratio
dimorphism was apparent only in height of
(=height of dorsal fin 2/length of dorsal fin
dorsal fin in males >510 cm total length
base), and blowhole to dorsal fin tip (along line
directly from blowhole to tip of dorsal fin tip; and in anterior and posterior flipper length
Fig. 1-modified from Heimlich-Boran, 1986).among males >435 cm total length. The t-
For animals with bent dorsal fins, measurement tests for homogeneity of slopes and Y-in-
was taken along the fin and used as the height tercepts between sexes for all other cate-
of the fin had it been straight. gories showed no statistically significant
Allometric relationships were examined using difference; therefore, regression lines were
linear or piecewise linear-regression models and considered coincident. The only exceptions
took the form of model II regressions. However,
were length of flippers and height of dorsal
the high coefficients of determination (r2) of
fin among larger males.
most analyses allowed us to use model I regres-
All allometric plots for females were
sion techniques. All analyses presented in this
study were done using the SYSTAT statistical characterized by a distinct cluster of points
package (SPSS, Inc., 1996). Because data were at total lengths greater than ca. 550 cm.
longitudinal, jackknife techniques were used to Those clusters consisted of data from only
estimate regression parameters and coefficients two females that were considerably larger

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August 1999 CLARK AND ODELL-ALLOMETRY OF KILLER WHALES 779

than the remainder of females in the dataset

and were of North Pacific origin.
The relationship of anterior length of
4)r

6 0-4 ZN
N0N tN
9 z---' m .
flipper to total length was linear for fe-
c) r)z t- r) 000c
males (Fig. 2a). A piecewise linear-regres-
sion model was more appropriate in char-
acterizing allometric growth of flippers for
males (Fig. 2b). According to this model,
P-4

oo7ce7o&66
a break point occurred at ca. 435 cm total
length where the rate of growth increased.
w r- Nz00
A similar relationship existed between pos-
CI
0r Ci0 6~ t0
terior length of flippers and total length. In
both relationships, rate of growth in both
anterior and posterior lengths of flippers
became greater in larger males (>435 cm
CI
total length) than females. For both sexes,
the relationship of width of fluke to total
?6 t< ; ? ffi 06 4 length was linear. Allometry of distance
. 0 N o N , N - from rostrum to anterior origin of flippers
to total length was linear for both sexes.
t 4
All girth measurements (girth at anterior
origin of dorsal fin, girth at axilla, girth at
C?I 00 N ? ri C N 00 N C, genital slit) exhibited similar linear rela-
N ? In 0 tI I N 0NIn
tionships with total length. Among fe-
1" ,-- ,-- 0 ttq?'1 ', 6, males, greater variability occurred when
total lengths were >500 cm (Fig. 2c). That
variability was not evident in nulliparous
females (Fig. 2d). Concerning girth at gen-
CI
ital slit morphometric, growth coefficient
~. o 4
for males appeared greater than that of nul-
liparous females (males, 0.61; nulliparous
females, 0.53), although that difference
o
was not statistically significant. Distance
between mammary slits in females was lin-
ear, although the relationship was not as
strong (r2 = 0.32 and 0.60 for all and nul-
a . 0
liparous females, respectively) as others
examined; therefore, allometric equations
were suspect. Similar to the girth measure-
ments, there was a greater amount of vari-
CI
ability in females larger than ca., 500 cm
0
total length. A distinct cluster of points
4 -o 4-
that occurred at total lengths >550 cm
consisted almost exclusively of measure-
0 0
0 0
ments from the only two nulliparous killer
whales. Those animals were of North Pa-
- c 0 cific origin while the other females origi
nated from the North Atlantic. Greater to-
tal lengths of those females were due to

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 TABLE 2.-Allometric growth relationships and analyses of proportional sexual dimorphism in adult and c
parks. All measurements were regressed on total length (TL). Homogeneity of slopes or Y-intercepts refers to
tests for homogenousness (** = P < 0.01, *** = P < 0.001, ns = not significant).
Males Femal

Measurement Equation n r2 Equation n r2

Anterior length of flipper 10 0.97 Y = -8.62 t 0.17X
-435 cm TL Y = 10.66 + 0.1
>435 cm TL Y = -106.76 + 0.39
Posterior length of flipper 10 0.95 Y = -6.76 + 0.14X
-435 cm TL Y = 2.71 + 0.11
>435 cm TL Y - -84.07 + 0.3
Width of flukes Y = 1.45 + 0.28X 10 0.94 Y = 11.71 + 0.
Length from rostrum to origin of flipper Y = 42.00 + 0.12X 10 0.85 Y = 35.
Girth at anterior origin of dorsal fin Y = 26.20 + 0.58X 10 0.96
All females Y = 28.84 + 0.57X 10
Nulliparous females Y = 14.41 + 0.59X 1
Girth at axilla Y = 45.46 + 0.50X 10 0.97
All females Y = 31.75 + 0.52X 10
Nulliparous females Y = 1.26 + 0.58X 1
Girth at genital slit Y = -43.80 + 0.61X 10 0.92
All females Y = -24.57 + 0.50X 10
Nulliparous females Y = -40.30 + 0.53X 1
Distance between mammary slits
All females Y = -9.02 + 0.05X 20
Nulliparous females Y = -3.31 + 0.03X 1
Height of dorsal fin 1 10 0.85 Y = -8.75 + 0.17X
-510 cm TL Y = -1.95 + 0.12
>510 cm TL Y = -198.48 + 0.5
Blowhole to tip of dorsal fin Y = 2.53 + 0.41X 10 0.87 Y = 7.48 +
a Coefficient of growth for males was greater than that of females.
b Coefficient of growth for females was greater than that of males.

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August 1999 CLARK AND ODELL-ALLOMETRY OF KILLER WHALES 781

125 200

00 - females
4 10 males
C 150

75 -

100

50-

50

o 25- b

0 a
250 350 450 550 650 250 340 430 520 610 700

Total length (cm) Total length (cm)

-14 500 1 1 1 44 500 1 1 1

all nulliparous
W females females

400- - ?
400-

0 300
300-

200

200
r.l

Id
1200 . Sd
100 100

250
Vr 250350
350450
450 550
550 650
650 250 350 450 550 650

Total length (cm)

~3Total length (cm) Total length (cm)

300 j 200

-u males . males

- 250 150

S200 44 100

150 o 50

? e

100 l 0
200 300 400 500 600 700 200 300 400 500 600 700

Total length (cm) Total length (cm)

FIG. 2 (a-f).-Relationships for various morphometrics regressed on total leng

whales at SeaWorld parks (10 males, 20 females).

their genetic origins. Duffield andmales,Miller

a piecewise linear-regression model
(1988) reported that killer whales described
from the the data with a breakpoint at ca.
Pacific are larger than those from510 the
cmAt-
total length; growth rate of males
lantic. Height of dorsal fin regressed on to-beyond that point (Fig. 2e). For
increased
tal length was linear for females. For males and females, the relationship of the

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782 JOURNAL OF MAMMALOGY Vol. 80, No. 3

length from blowhole Among to tip

males, the of
substantial dorsal
increase in fi
and total length was growth linear
coefficients of (Fig. 2f).
pectoral flippers at
ca. 435 cm total length may have been the
DISCUSSION result of the beginnings of an adolescent
The dearth of published information
growth spurt. Christensen (1984) postulated on
aspects of allometric that male
growthkiller whales exhibit
in an increase
either ca
tive or wild killer whales in rate of growth precludes
of total lengthus when fro
the
making any direct reach comparisons
ca. 550-610 cm long, which to corre-
previo
investigations. However, lates to an estimated
related age of 15studies
years based ex
amining growth parameters upon tooth-growth, layer-group
of other counts odo
tocetes, as they pertain (tooth-growth,to layer-group
aging, counts
have are de- sug
gested that captive fined animals
as "repeating or do semi-repeating
not appear pat-
to differ appreciably from
tern of adjacent groupswildof incremental anima
Cheal and Gales (1992) growth found
layers within in bottlenose
the dentine, cemen-
dolphins (Tursiops truncatus) tum, or bone which is defined > 3 as ayears
count- o
age that growth in able totalunit"-Report
length of the International
was not a
fected by captivity, Whaling
which Commission, is 1980:48-49).
similar Per- to r
sults obtained by Kasuya et al. (1986) haps, increased rate of growth of flippers
among finless porpoises (Neophocaena also was attributable to this growth spurt.
phocaenoides) and bottlenose dolphins. Because size and shape of pectoral flippers
Furthermore, examination of weight:lengthof cetaceans is important for efficient aquat-
ratios of bottlenose dolphins suggested ic maneuverability (Fish and Battle, 1995),
there was a similarity between captive andit is reasonable to assume that if growth rate
wild animals (Ridgway and Fenner, 1982). in total lengths of males was to suddenly
Using these studies as precedence, it can be increase, the animal's components of hydro-
assumed that data from our study has viable dynamic steering (i.e., the pectoral flippers)
applicability for understanding dynamics ofwould likewise increase in size. In addition,
allometric growth in captive and wild killer Harmer (1927) suggested that accelerated
whales. growth in phalangeal cartilages of pectoral
To some degree, all allometric relation- flippers of males caused the increased
ships were characterized by increased vari- length in pectoral flippers and was related
ability in the dependent variable as total to growth of adolescents.
length increased. This probably was attrib- Our overall results of negative allometry
utable to inherent differences in size among for growth in length of flippers were similar
to the conclusions of Ratnaswamy and
adults (i.e., individual variability of adults).
Apart from length of flippers and height Winn (1993) working with fin whales (Ba-
of dorsal fin between females and largerlaenoptera physalus). In their study, ante-
males, sexual dimorphism was not appar- rior length of flipper was characterized by
ent. These results are similar to those of negative allometry, while posterior length
Hersh et al. (1990) for bottlenose dolphins exhibited isometric allometry. They attrib-
and Anli and Kaiya (1992) for the baiji uted (Li- their findings to differing food habits
potes vexillifer). Varying degrees of sexual or requirements of the animals as they
dimorphism were reported by Amano grow. and Likewise, Amano and Miyazaki
Miyazaki (1993) in Dall's porpoises (Pho- (1993) found negative allometry in poste-
coenoides dalli) and bottlenose dolphinsrior by length of flippers of Dall's porpoises.
Tolley et al. (1995). Overall, our study In in-
contrast to our study, width of flukes in
dicated that adult males end up with larger Dall's porpoises displayed positive allome-
appendages than females, which wastry ex-(Amano and Miyazaki, 1993). This spe-
pected. cies appears to be an exception; other ce-

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August 1999 CLARK AND ODELL-ALLOMETRY OF KILLER WHALES 783

50

E calf
born
45

calf

40 bornm

E
E 3calf
35 born

calf
30 L 3T0 born

25

150 650 1,150 1,650 2,150 2,650 3,150 3,650 4,150 4,650
Time (days)
FIG. 3.-Relationship between distance between mammary slits and time (days
during four pregnancies (animal 7806 from Duffield et al., 1995). Distance betw
increased during pregnancy and decreased rapidly after parturition.

taceans displayed a pattern of negative al-

ative allometry were in contrast to the result
lometry as was the case in our killer whales
obtained by Amano and Miyazaki (1993) in
(Kasuya, 1972; Perrin, 1975; Sergeant P. dalli et
for girth at anterior origin of dorsal
al., 1973). fin but concurred with their findings con-
Among girth and mammary measure- cerning girth at axilla.
ments, some of the increased variability Althoughin not statistically significant,
larger females probably was relatedmales to preg-
had a greater coefficient of growth in
nancy. As the fetus develops and grows, girth at the
the genital slit than did females. The
mother becomes increasingly larger lack of in
statistical significance could be a re-
girth. This increase peaks just prior sult
to of increased variability in larger fe-
par-
turition and is followed by a decrease malesafter
that may obscure any differences.
the young was born. Following parturition, Larger girths of males observed may be at-
girth and distance between mammary slits
tributed to increased muscle mass. Tolley et
declines to about pre-pregnancy dimensions al.(1995) suggested that larger size of these
(e.g., animal 7806 from Duffield et al., posterior muscles could provide increased
1995; Fig. 3). Indeed, nulliparous females strength to flukes either for incidences in-
did not show cyclic increase and decrease volving conflicts related to dominance by
in these measurements. males or need for increased swimming
Comparison of growth coefficients be- speeds.
tween sexes for girth at anterior origin of As expected, our male killer whales pos-
dorsal fin and girth at axilla indicated a lack
sessed significantly larger dorsal fins and
of sexual dimorphism. Our results of neg- dorsal-fin ratios than did females. Some in-

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784 JOURNAL OF MAMMALOGY Vol. 80, No. 3

vestigators have hypothesized that exami

LITERATURE CITED
nation of dorsal-fin ratio may provide som
AMANO, M., AND N. MIYAZAKI. 1993. External mor-
indication of sexual maturity
phology of dalli):
of Dall's porpoise (Phocoenoides males
Heimlich-Boran (1986)
growth andsuggested that
sexual dimorphism. Canadian Journal of ju
venile males may beZoology, 71:1124-1130.
separated from ado-
ANLI, G., AND Z. KAIYA. 1992. Sexual dimorphism in
lescent males when their dorsal-fin ratio is
the baiji, Lipotes vexillifer. Canadian Journal of Zo-
> 1.0. Likewise, Olesiuk et al. (1990) statedology, 70:1484-1493.
CHEAL, A. J., AND N. J. GALES. 1992. Growth, sexual
that a ratio ?1.4 may indicate onset of sex-
maturity and food intake of Australian Indian Ocean
ual maturity of males. Lack of data did notbottlenose dolphins (Tursiops truncatus). Australian
allow us to study change in dorsal-fin ratio
Journal of Zoology, 40:215-223.
as males and females grow. However, re- CHRISTENSEN, I. 1984. Growth and reproduction of kill-
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garding absolute sexual dimorphism, our
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DUFFIELD, D. A., AND K. W. MILLER. 1988. Demo-
was statistically significantly larger than
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Morphology, 225:51-60.
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305-315.
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ACKNOWLEDGMENTS KASUYA, T., T. TOBAYAMA, T. SAIGA, AND T. KATAOK
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